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Systematics, ecology and biogeography of the neotropical butterfly genus Adelpha (lepidoptera: nymphalidae: limenitidinae

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Systematics, ecology and biogeography of the neotropical butterfly genus Adelpha (lepidoptera: nymphalidae: limenitidinae
Creator:
Willmott, Keith R ( Keith Richard )
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English
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vii, 724 leaves : ill. ; 29 cm.

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Butterflies ( jstor )
Cell lines ( jstor )
Colors ( jstor )
Ecology ( jstor )
Female animals ( jstor )
Forest habitats ( jstor )
Instars ( jstor )
Species ( jstor )
Syntypes ( jstor )
Taxa ( jstor )
Adelpha ( lcsh )
Dissertations, Academic -- Entomology and Nematology -- UF ( lcsh )
Entomology and Nematology thesis, Ph.D ( lcsh )
City of Sarasota ( local )
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bibliography ( marcgt )
non-fiction ( marcgt )

Notes

Thesis:
Thesis (Ph.D.)--University of Florida, 1999.
Bibliography:
Includes bibliographical references (leaves 700-723).
General Note:
Typescript.
General Note:
Vita.
Statement of Responsibility:
by Keith Richard Willmott.

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SYSTEMATICS, ECOLOGY AND BIOGEOGRAPHY OF THE NEOTROPICAL
BUTTERFLY GENUS ADELPHA (LEPIDOPTERA: NYMPHALIDAE: LIMENITIDINAE)









BY

KEITH RICHARD WILLMOTT


A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE
UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY

UNIVERSITY OF FLORIDA


1999














ACKNOWLEDGMENTS


I am very grateful to P. Ackery and J. Reynolds (BMNH), Drs. L. and J. Miller (AME), Dr. J. Pierre (MNHN), Dr. W. Mey and M. NuB (ZMHU), Dr. J. Miller and Dr. E. Quinter (AMNH), Dr. P. Perkins (MCZ), Dr. J. Heppner (FSCA), Dr. G. Lamas (MUSM), Drs. R. Robbins and D. Harvey (USNM) and the curators at the BMB for giving me access to the Adelpha collections in their care and for the loan of material for dissection. Dr. Lrbl of the MHNG and A. Lvovsky of the ZISP were very helpful in identifying and supplying me with photographs of Adelpha types in their respective collections. I thank D. Trembath, M. Perceval, L. Constantino, J. Salazar, Dr. E. SchmidtMumm and J. LeCrom for allowing me to view and take notes from their private collections and for their generous hospitality. I especially thank A. Neild for providing accommodation in London on numerous occasions, for donating specimens of Adelpha, for his continuous help and correspondence and willingness to discuss ideas. R. Hesterberg generously loaned his specimen of A. hesterbergi to me for study. I thank the librarians at the BMNH entomological library for their help and for allowing me to photocopy or photograph important pages of several rare books.

I am greatly indebted to Dr. G. Lamas, who freely shared his peerless knowledge of neotropical butterfly taxonomy and nomenclature, providing invaluable help and advice on numerous questions and photographs of several type specimens, and whose unpublished checklist of Adelpha was the foundation for my research into the genus. I thank Dr. M. Parsons for help in navigating the maze of Adobe Photoshop and Aldus Freehand, Dr. A. Aiello for her continuous helpful correspondence, Dr. J. Mallet for donating slides of early stages, and Drs. W. Haber, D. Janzen, P. DeVries and A. Orellana









for unpublished information on Adelpha hostplants. This dissertation benefitted greatly from the valuable comments and suggestions of Dr. T. Emmel, J. Heppner, J. Lloyd, L. Miller and J. Reiskind, to all of whom I extend my appreciation. I am also very grateful to Dr. T. Emmel for providing superb facilities and for supporting me during my time at UF with research assistantships, funded by the U.S. Fish and Wildlife Service and private donations, and for all his help and encouragement throughout my time in Florida. I also thank the following for assisting with the costs of field work in Ecuador: Mr. I. Willmott, Mrs. M. Willmott, Christ's College Cambridge University, Albert Reckitt Charitable Trust (C. T.), Poulton Fund Oxford University, Round Table Trust, Lindeth C. T., Catherine Cookson Foundation, Morton C. T., Royal Entomological Society, Butler C. T., Mr. D. Exell, Peter Nathan C. T., Harry Crook Foundation, Douglas Heath Eves C. T., R. and M. Foreman C. T., Northern Bank, Banbridge Academy, C. Bruce, Hickley Valtone Ltd., Vera Trinder Ltd., Agfa, Phoenix Mountaineering, Balfour-Browne Fund, Worts Fund, Sigma Xi the Scientific Research Society (1996) and Equafor; field and museum research in 1997/8 was funded by National Geographic Society Research and Exploration Grant No. 5751-96. Florida Agricultural Experiment Station provided facilities and funded publication of my research work on Adelpha. I also thank INEFAN, the Pontificia Universidad Cat6lica and the Museo de Ciencias Naturales, Quito, and in particular Dra. Germania Est~vez, for arranging the necessary permits for field research in Ecuador.

Finally, I thank both my parents for their continuous support throughout my doctoral studies and before, my dear friend Julia Robinson-Dean for her fine company, help and tolerance during two field trips to Ecuador, and my lifelong friend Jason Hall for his continuous advice and healthy criticism during my work on Adelpha, for collecting many rare species, for his splendid humor in the field on many trips throughout the tropics, and for first introducing me to the delights of lepidopterology.














TABLE OF CONTENTS

Page
ACKNOW LEDGM ENTS .................................................................................................. ii

ABSTRACT ....................................................................................................................... vi

CHAPTERS

1 INTRODUCTION .............................................................................................. 1

2 REVISION OF THE GENUS ADELPHA ......................................................... 5

Introduction ......................................................................................................... 5
M aterials and M ethods ....................................................................................... 6
Systematics ....................................................................................................... 17
Biology .................................................................................................................. 58
Review of Adelpha ........................................................................................... 98

3 PHYLOGENY OF THE GENUS ADELPHA .................................................... 664

Introduction ......................................................................................................... 664
M ethods ............................................................................................................... 665
Results and Discussion ....................................................................................... 672

4 SUM M ARY AND CONCLUSIONS ................................................................. 681

APPENDICES

A LIST OF TAXA DISSECTED ............................................................................ 683

B NOMINA NUDA AND NAMES ERRONEOUSLY INCLUDED IN
ADELPHA ................................................................................................ 689

C NOMENCLATURAL CHANGES, ERRORS AND MISIDENTIFICATIONS OF
FIGURED ADELPHA IN POPULAR WORKS .................................... 690

D DATA FOR FIGURED SPECIMENS OF ADELPHA, FIGS. 10-94 ................ 693








E CHARACTER MATRIX FOR PHYLOGENY OF ADELPHA ......................... 698

REFERENCES ............................................................................................................... 700

BIOGRAPHICAL SKETCH .......................................................................................... 724














Abstract of Dissertation Presented to the Graduate School of the University of Florida in
Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy

SYSTEMATICS, ECOLOGY AND BIOGEOGRAPHY OF THE NEOTROPICAL
BUTTERFLY GENUS ADELPHA (LEPIDOPTERA: NYMPHALIDAE: LIMENITIDINAE)

By

Keith Richard Willmott

May 1999

Chairman: Dr. Thomas C. Emmel
Major Department: Entomology and Nematology


A revision of the diverse Neotropical nymphalid butterfly genus Adelpha is presented, which includes a review of the systematics, ecology and biogeography of all constituent taxa. I discuss the relationships of the genus to other New and Old World species and genera of the tribe Limenitidini and define the genus based on wing pattern characters. I present a synonymic checklist which includes all 362 described names, and recognise 85 valid species and 207 valid taxa. Eight new subspecies are described, and a new species and four subspecies are recognised but undescribed due to a lack of material. The status of all names is critically examined, the location of type material is discussed and notes on the identification of all taxa are presented in addition to a dichotomous key for identification of adults of all species. Adults of all taxa are figured and male and female genitalia of the great majority of species. Lectotypes are designated for 29 names, and neotypes for two names.
I summarise information from the literature on hostplants utilised by the genus and the morphology and behavior of the immature stages. Notes are provided on the









habitats, phenology, rarity and adult behavior of all taxa based on published data and approximately two years field work in six Neotropical countries. I also list the locality data and depository of 20,730 specimens of Adelpha examined and discuss the known and expected geographic and elevational ranges of all taxa.

A phylogeny for all species within the genus is presented based on wing pattern and adult morphological characters of both sexes. While there are several well defined clades, the phylogeny lacks high resolution and strong support for nodes throughout due to a pervasive lack of morphological characters. Nevertheless, there is much evidence that adult characters do support phylogenetic groups similar to those derived from analysis of characters from immature stages. I discuss in particular the problems of homology assessment of wing pattern characters which have led to past taxonomic confusion. Dorsal wing patterns show abundant evidence of convergence among sympatric taxa, and I discuss the evidence for mimicry within Adelpha.













CHAPTER 1
INTRODUCTION


The Neotropical region is justly famed for its rich biological diversity, and nowhere is this more apparent than in its insect fauna, particularly butterflies. Many of the most visually spectacular species occur within the largest single family, the Nymphalidae, whose species inhabit a very wide diversity of habitats and exhibit great variety in their life history strategies. This variety offers a potentially rich area for research into many aspects of evolutionary biology, and just a few of the fields in which nymphalids have made a particularly important contribution include plant-insect interactions (e.g. Brown, 1987), the study of speciation processes (Jiggins et al., 1996; Davies et al., 1997), the phenomenon of mimicry (Bates, 1862; Miiller, 1879) and studies into the historical biogeography of the neotropics (Brown, 1982). In the recent surge of interest in the conservation of biodiversity, one nymphalid subfamily, the Ithomiinae, has been suggested as an ideal indicator taxon of tropical lowland forest butterfly diversity (Beccaloni and Gaston, 1994).

The higher level systematics of the Nymphalidae are still keenly debated, and there is no consensus as to the relationships between subfamilies or even as to their total number. The most recent classification of Harvey (1991) recognises 11 neotropical subfamilies, including the Libytheinae, Danainae, Heliconiinae, Ithomiinae, Brassolinae, Morphinae, Satyrinae, Charaxinae, Apaturinae, Nymphalinae and Limenitidinae. Paradoxically, despite the wealth of information that nymphalid butterflies have provided, biological research to date has largely concentrated on only a few groups, such as the Danainae, Heliconiinae and Ithomiinae. These subfamilies contain species that are









typically common, slow flying and therefore easy to study in the field, and easily manipulated under artificial conditions. There exist several modem systematic treatments of these subfamilies (Fox, 1956, 1960, 1967; Fox and Real, 1971; Michener, 1942; Brown, 1981; Ackery and Vane-Wright, 1984), but it is only recently that explicitly derived phylogenies have been developed for the Heliconiinae (Brower and Egan, 1996; Penz, 1999) and Danainae (Ackery and Vane-Wright, 1984), while none is yet available for the Ithomiinae. The Brassolinae, Morphinae and Charaxinae, all popular groups with amateur collectors, have been the subject of various generic monographs (Bristow, 1981, 1982, 1991; Blandin, 1988, 1993; Comstock, 1961; Barselou, 1983). Higgins (1981) reviewed a large section of the Nymphalinae, while the species-level taxonomy of the Satyrinae has remained a virtual terra incognita since its treatment by Weymer (19101912).

Of the remaining subfamilies, two are monotypic, leaving the large and heterogeneous Limenitidinae. As yet there is no evidence to suggest that this subfamily comprises a natural (monophyletic) group of taxa, and it has been retained by modem taxonomists largely for convenience (Harvey, 1991; Neild, 1996). It is, however, in this subfamily that Neotropical nymphalids reach their greatest diversity in both morphology and species, and probably life history strategies. The monumental work of Seitz ([1907]1925) and the pictorial reviews of D'Abrera (1987) are, for many genera, the only available fairly comprehensive references. A number of other genera have been revised, including Callicore (Dillon, 1948), Ectima, Hamadryas, Myscelia, Catonephele, Eunica, Epiphile, Asterope, Nessaea (Jenkins, 1983, 1984, 1985ab, 1986, 1987, 1989, 1990) and Perisama (Attal and Crosson du Cormier, 1996), and these revisions have often illustrated the existence of both undescribed and unrecognised species even in small, well collected genera, a result that is certainly much more widely applicable than is currently acknowledged.









It is thus apparent that systematic study, the foundation for ecological research, is still urgently needed in many nymphalid groups. A further concern is the continual loss of biological diversity in tropical countries and the pressing need to catalogue and understand the distribution of such diversity. Among nymphalid butterflies, one of the most conspicuous members in all neotropical forest habitats is the genus Adelpha, which is certainly the most speciose genus of the neotropical Limenitidinae, and possibly of the entire subfamily Nymphalidae. Many Adelpha species are common, they occur in a very wide diversity of habitats and their immature stages exhibit a great diversity of behavior, morphology and hostplant utilisation (Moss, 1933; Aiello, 1984; Ackery, 1988). Previous studies of the immature stages demonstrated that adults with dorsal wing patterns which appeared superficially similar were often apparently unrelated (Moss, 1933; Aiello, 1984), leading to the suggestion that this adult similarity was due to mimicry (Aiello, 1984).
The genus thus offers great promise for biological study, but this has remained unrealised largely due to the problems in identification of many extremely similar species and a lack of understanding of their relationships, and the taxonomic chaos caused by this similarity and the availability of over 350 published names for Adelpha taxa. As a result, Adelpha has become renowned more for its taxonomic problems than its potential biological interest; Fruhstorfer (1915) states that "except the Phyciodes and certain Euthaliidae there is, therefore, hardly any genus of the Rhopalocera offering greater difficulties to the systematical and synoptical treatment", while DeVries (1987) echoed this sentiment, saying "the butterflies that compose the genus Adelpha are, in my opinion, the most difficult and trying taxonomically of all the nymphalids". Fruhstorfer (1915) was the last to treat the entire genus, but offered few reliable characters for separating species and his work is replete with misidentifications.

The objectives of this study are, therefore, to provide a stable nomenclature for the genus through a critical examination of the status of all available names and to









provide information to enable certain identification of all Adelpha taxa. I also attempt to create a systematic arrangement that reflects modem species concepts and our greatly improved knowledge of the distribution of taxa. As a baseline for future research, I summarise all of the available information on the immature and adult biology of Adelpha species. I also analyse the distribution of diversity and endemism within the genus and discuss its implications for the conservation of Adelpha, and probably other butterfly taxa. Finally I derive a preliminary phylogeny for the genus to serve as a heuristic tool for field workers and to test the hypothesis that similar dorsal wing patterns in many sympatric Adelpha are the result of convergent evolution through mimicry.














CHAPTER 2
REVISION OF THE GENUS ADELPHA


Introduction
The genus Adelpha is the only member of the tribe Limenitidini to occur in the Neotropical region, and, like its Old World relatives, it has always been a bane to systematists. Estimates of the number of species in the genus have ranged as high as 100 (Aiello, 1991), while there are no published reviews of the genus which enable accurate identification of its constituent species. Adelpha species occur from far northwestern U.S.A. throughout Central and South America to Uruguay, and are common and conspicuous members of the forest butterfly fauna in a wide variety of life zones, from sea level to 3000m. They constitute the most diverse genus of nymphalid butterflies in most lowland forests, and may be important models for Batesian mimics in other butterfly subfamilies. The larvae feed on a diverse assemblage of plants (Aiello, 1984; Ackery, 1988) and the morphology of the immature stages is highly variable (Mfiller, 1886; Moss, 1933; Aiello, 1984, 1991; Otero & Aiello, 1996).

The last published systematic revision of the entire genus Adelpha was that of Fruhstorfer (1915), followed by a pictorial review of the genus by D'Abrera (1987). While undertaking a study of Ecuadorian Rhopalocera in 1993, it became apparent that identification of the majority of Adelpha taxa was not possible using published sources, and that there was no general consensus as to the number of species in the genus or their relationships. With the capture of undescribed species of Adelpha by myself and Jason Hall in Ecuador in 1993, 1994 and 1995, it seemed that a systematic review of the genus was desperately needed as a baseline for further research into this diverse group of










butterflies. In addition, there has been a growing interest in recent years in the monitoring

and conservation of Neotropical butterfly faunas, and the first step to cataloguing the

distribution of such tropical biodiversity is a firm taxonomic basis. Since Adelpha is

perhaps the largest genus in the Nymphalidae, and is widespread and common in all

forested habitats throughout the Neotropics, it is also an ideal genus to examine general

trends in diversity and endemism in nymphalid butterflies.



Materials and Methods

Specimens Examined

A total of 20,730 specimens of Adelpha were examined and identified in public

and private collections, including 16,426 males and 4,304 females. The collection

acronyms used throughout the text are listed below, and those in which I have examined

material are marked with an asterisk. Other collections examined from which specimens

are not listed include the Museo de Ciencias Naturales and Pontificia Universidad

Cat6lica, both in Quito, Ecuador.


ADF Alberto Diiz Frances collection, Mexico, Mexico
* AFEN Andrew F. E. Neild collection, London, UK
AJ Artur Jasinski collection, Piast6w, Poland
* AME Allyn Museum of Entomology, Sarasota, FL, USA
* AMNH American Museum of Natural History, New York, NY, USA
AO Andr6s Orellana collection, Mrida, Venezuela
* BMB Booth Museum of Natural History, Brighton, UK
* BMNH Natural History Museum, London, UK
(M=Main coil.; R=Rothschild coil.; T=Type coil.; Moss=Moss coil.; A&B=Adams & Bernard
coil.)
* DAT David Trembath collection, Surrey, UK
DLF De la Fuente collection, Caracas, Venezuela
EF Eurides Furtado collection, Diamantino, Brazil
* ESM Ernesto W. Schmidt-Mumm collection, Bogoti, Colombia
FIML Fundaci6n e Instituto Miguel Lillo, Tucumnin, Argentina
* FSCA Florida State Collection of Arthropods, Division of Plant Industry, Gainesville, FL, USA
GTA George T. Austin collection, Las Vegas, NV, USA
HS Harold & Jos6 L. Skinner collection, Margarita, Venezuela
JB Joffri Blanco collection, San Crist6bal, Venezuela
JCS Juan Carlos de Sousa collection, Barquisimeto, Venezuela
* JFL Jean F. Le Crom collection, Bogoti, Colombia
JYG Jean-Yves Gallard collection, French Guiana









* KWJH Keith R. Willmott & Jason P. W. Hall collection, Gainesville, FL, USA
LCB L. & C. Br~vignon collection, French Guiana
* LMC Luis M. Constantino collection, Cali, Colombia
MALUZ Museo de Artr6podos Terrestres, La Universidad de Zulia, Venezuela
* MCZ Museum of Comparative Zoology, Harvard University, Cambridge, MA, USA
MHNG Muse d'Histoire Naturelle de Gen~ve, Geneva, Switzerland
* MHNUC Museo de Historia Natural, Universidad de Caldas, Manizales, Colombia
MIZA Museo del Instituto de Zoologia Agricola, Universidad Central de Venezuela, Maracay,
Venezuela
* MJP Mike J. Perceval collection, Surrey, UK
* MJWC Matthew J. W. Cock collection, Reading, UK
* MNHN Musrun National d'Histoire Naturelle, Paris, France
* MUSM Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru
MZUJ Muzeum Zoologiczne Uniwersytetu Jagiellofiskiego, Krak6w, Poland
* PJD Philip J. DeVries collection, Eugene, OR, USA
R Romero Family collection, Maracay, Venezuela RDM Roberto de la Maza collection, Mexico, Mexico
RM Raymond Murphy collection, Torquay, UK
RNHL Rijksmuseum van Natuurlijke Histoire, Leiden, Netherlands
SMTD Staatliches Museum ffir Tierkunde, Dresden, Germany
TP Tomasz Pyrcz collection, Warsaw, Poland; incorporated into the MZUJ
UFP Universidade Federal do Parani, Curitiba, Brazil
UNAM Instituto de Biologia, Universidad Nacional Aut6noma de Mexico, Mexico, Mexico
* USNM National Museum of Natural History, Smithsonian Institution, Washington, DC, USA
ZISP Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia
ZMUC Zoologisk Museum, Universitets Copenhagen, Copenhagen, Denmark
* ZMHU Zoologisches Museum, Humboldt Universitit, Berlin, Germany
ZSBS Zoologisches Sammlung des Bayerischen Staates, Munich, Germany



I list the specimens examined under each taxon by country in an order which

corresponds to their geographical position, starting with the USA, then Mexico, east to

Panama, from Venezuela south along the Andes to Argentina, Brazil, Paraguay, Uruguay,

Guyana, French Guiana, Surinam, Trinidad and the Caribbean. I list provinces, states or

departments within each country, and localities within each of these subdivisions, in

alphabetical order. I regard words such as "Rio", "Quebrada", "Mount", "Volcin",

"Puerto", "via" and "El/La" as part of the name, but not relational phrases such as "n",

"vic..", "env.", "region", "above", "head", all compass points (N., E., S., W. etc.) and

"km." or "mi." from a locality. Collection sites within each country were located using

atlases in addition to Brown (1941), Lamas (1976), Brown (1979) and Jenkins (1983,

1984, 1985ab, 1986, 1987, 1989, 1990). Sites that I was unable to precisely locate are

listed at the end of each country subdivision, country or under "country unknown". I have









used the names of localities as given on the original label, and not altered them when the locality has undergone a subsequent change of name, as in the case of ParAi to Belrm. Countries, states, provinces or departments represented only by specimens with label data that I consider erroneous are enclosed in quotes. For each taxon I list the number, sex (m

- male, f- female), collection locality and altitude when given, date, any notes of interest on accompanying labels, and the depository of all specimens examined. In a number of cases specimens were examined which are transitional between subspecies, or exhibit other significant external phenotypic variation, and these are marked with various symbols keyed immediately at the top of the list of specimens for each taxon. For each species I list the total number of specimens examined, and the number of each sex in parentheses for each taxon and country. In a very few cases I have listed specimens examined only by photograph, where these are unique or of very rare taxa, and I state where this is the case in the taxonomic discussion.


Identification of Type Specimens

One reason for confusion regarding the application of Adelpha names to taxa is the difficulty in identification of type specimens, and I have attempted to locate all extant type material of available Adelpha names. The great majority of names were based on type series rather than a single type specimen, and there is therefore the possibility of mixed type series for many taxa. Most early authors gave no indication of the number of specimens on which they were basing their descriptions, and their material must therefore be regarded as syntypic, even if only a single specimen is known. I designate lectotypes or neotypes for all names that are not sufficiently well described to apply unambiguously to a particular taxon, and lectotypes for those that clearly apply to several taxa on the basis of the syntypic series.
It appears that no authors labeled Adelpha material on which they based names as type specimens prior to the Felder brothers in the 1860s, and the same is true of many









subsequent authors. In the absence of an original figure sufficiently detailed to identify the specimen on which it was based, or a specimen labeled specifically as a type by the original author, I believe there is an element of doubt surrounding the validity of all other type material. I have therefore tried to base my application of a name first and foremost on the original description, and to examine critically all labeled and unlabeled potential type material in the light of this description. I have been able to locate type material in the collections examined of almost all described Adelpha taxa. Notable exceptions are certain types of very early authors such as Linnaeus, Cramer, Godart, Drury and Walch, and these specimens are, in most instances, probably lost or destroyed. The Linnaeus Lepidoptera types are currently under study by Honey (pers. comm.) and located in London, Cramer types are either in the RNHL or the BMNH (Vane-Wright, 1975), Godart types should be in the MNHN (Horn & Kahle, 1935-37), Drury types should be in the BMNH, and the whereabouts of Walch's collection is unknown (Horn & Kahle, 1935-37). Of these, only the descriptions of Linnaeus and Walch are too vague to permit unambiguous identification, and I discuss these problems under the appropriate taxa.

The collections of most of those authors who subsequently published descriptions of Adelpha are now at the BMNH, including those of Hewitson, Butler, Bates, Boisduval, the Felder brothers, Godman and Salvin, and Fruhstorfer (Horn & Kahle, 1935-37). In the case of these authors, I identified type specimens using the original description and figure, specimen label data indicating the collection of origin, the locality, the collector and any other important information, particularly hand-written labels of the original author identifying the taxon. Horn & Kahle (1935-37) figure examples of hand-written labels of Hewitson, Butler, the Felders, Boisduval and Fruhstorfer, in addition to other lepidopterists who published on Adelpha. The Felders also applied a particular type of label to their type specimens, which bears a gold and a black stripe and the name, and recognising their type material presents no difficulty. Identification of Fruhstorfer type material is, however, often problematic, for several reasons. It appears that the original









locality labels on many Fruhstorfer specimens were subsequently removed and new, typed labels indicating only the country of origin substituted, probably prior to the sale of the collection. Thus the originally cited type locality often does not match the label data. Putative Fruhstorfer type specimens may bear either or both of a red, typed "TYPE" label and a label with the identification of the taxon written in Fruhstorfer's hand. Often only one of these labels is present, but several specimens that are not types also bear such labels. Occasional specimens lack either of these labels and have been identified as syntypes through other information in the original description. Fruhstorfer described a number of taxa based on specimens in Rothschild's collection at Tring Museum, which is now also housed at the BMNH, separate from the main collection. A small number of Fruhstorfer's types are to be found at various other European museums, and I am grateful to Gerardo Lamas for providing photographs of such specimens.

Given the uncertainty surrounding the validity of type material of many older names, I have tried to list full specimen label data for such types. Full label data are enclosed by quotes, with data on separate labels separated by a "//". Occasionally I have added supplementary information on the type of label in square parentheses. I use the abbreviations HT, AT, ST, PT, LT and PLT to refer to holotype, allotype, syntype, paratype, lectotype and paralectotype respectively. I regard the type locality of a taxon as being that cited in the original description, even if it is clearly erroneous and regardless of any alternative indication on the label of a type specimen, which may have been replaced at a later date. The type locality is cited for each taxon as "TL", while for names where I have designated a lectotype or neotype, I also give the original type locality as "OTL". I have enclosed in square parentheses type localities that were not cited in the original description itself, but in the title or indirectly given elsewhere in the work (in the case of Fruhstorfer (1913, 1915), localities were given in the text, published in 1915, for names that appeared first in 1913), in addition to the country, if this was not originally given.









Field Work

In addition to museum research, I conducted field work in six Neotropical countries, including Mexico (3 weeks), Guatemala (1 week), Costa Rica (4 weeks), Venezuela (3 weeks), Bolivia (3 weeks), and Ecuador (17 months). Many Adelpha specimens in museums are vaguely and/or unreliably labeled, and establishing certain sympatry or otherwise of phenotypes is difficult when based exclusively on such data. The purpose of this field work was to gather reliable information on the sympatry of phenotypes, to collect series of specimens to study the stability or variability of wing pattern characters and to make and record ecological observations of possible taxonomic importance. I decided to concentrate my efforts in a single country, Ecuador, which contains 69% of the species in the genus, since it seemed more useful to gain a deep understanding of the fauna of a single country rather than a superficial acquaintance with the faunas of several. Indeed this proved to be an effective strategy, and some of the most intractable taxonomic problems in the genus, such as the true diversity and identification of members of the Adelpha serpa group, and the taxonomic status of A. iphiclus and A. iphicleola, were solved through collections made in eastern Ecuador. In addition to making collections of specimens, I attempted to record as accurately as possible the elevational distributions of species throughout the country, to examine patterns of endemism with altitude. I also made observations on adult food sources and the microhabitat preferences, perching and flight behavior of species, to ascertain whether phenotypically similar species occur microsympatrically, as predicted if Adelpha are involved in mimicry.

I conducted field work in Ecuador on both slopes of the Andes, from sea level on the western slope and 250m on the eastern slope to above the upper elevational limit of Adelpha species, approximately 3000m. Sites included as wide a range of topography, vegetation and microhabitat types as possible, including both flat and undulating terrain, paths along ridge tops, hill sides and river sides, in primary to heavily disturbed









secondary growth habitats, ranging from dry deciduous forest to pluvial forest, premontane rain and montane cloud forest. Sites were visited from 6.30 a.m. to 5 p.m., when Adelpha activity ceased. I made supplementary observations of individuals using a pair of Canon 8x32mm waterproof binoculars, and used two techniques to capture specimens, an entomological net with extension handles of up to 5.5m, and baited traps. Traps were of the Van Someron-Rydon design as figured by Sourakov & Emmel (1995) and Austin & Riley (1995), and were deployed in a variety of microhabitats, including forest edges, river sides, forest light gaps and the forest subcanopy. Traps were set at a variety of heights above the ground up to 15m, by use of polyethylene string looped over branches by a lead fishing weight. I used two types of bait; three to five day old fermenting banana, and one to three week old rotting fish, of a variety of fresh and saltwater species. Different species of fish produced no noticeable alteration in the attractiveness of the bait. Traps were patrolled approximately once per hour, and carefully lowered to check for individuals resting on the outside of the netting, a common occurrence in Adelpha. Once I was familiar with the fauna, I was able to identify and release the great majority of specimens in the field, and these sight records are listed under Additional locality data below each taxon. Specimens of taxonomic interest were killed by a pinch to the thorax and stored in glassine stamp envelopes in a wooden entomological box, and subsequently prepared in the U.S.A.


Morphological Study

Adults. The wing patterns of all species were studied through examination of material in collections. I examined the male genitalia of all species with the exception of A. herbita, of which there are no known specimens, and Adelpha sp. nov., of which the single specimen I have seen has already been dissected and the dissected genitalia subsequently lost. I have tried to examine the nominate subspecies of each species, and any others that exhibit significant phenotypic differences and/or have been regarded in the









past as distinct species. In the majority of cases, I examined several specimens of each species to assess intraspecific variation. Due to the rarity of females of many species, my choice of dissections was limited, both of the species I studied and the number of specimens. Specimens dissected are listed in Appendix A. I also examined the wing venation for all species except A. herbita and A. sp. nov., and the morphology of the legs, antennae and labial palpi for representative species from all species groups. Appendages and genitalia were prepared for dissection and study by soaking in hot 10% KOH for 1530 minutes, being subsequently stored in glycerol. Drawings were made using Wild M4A stereomicroscope at 30x magnification and a Wild camera lucida. Wing venation was studied and drawn using the same equipment as the genitalia. Terminology for the wing venation follows Comstock & Needham (1918), and I refer to wing cells by the vein that lies immediately above (anterior) to that cell. Genitalic terminology follows Klots (1956), and I use the term "clunicula" (of Fruhstorfer (1915)) to refer to the dorsally directed projection on the inner edge of the male genitalic valvae, which is typically adorned with numerous spines on the inner surface.

Immature stages. I was able to obtain preserved material of various immature stages of a few species on loan from individuals and public institutions. Such material typically consisted of cast head capsules, larval skins and pupal cases, and in a very few cases, dried larvae. The morphology was studied using the same microscope as for adult material; due to the rarity of preserved material, it was not possible to study it using a Scanning Electron Microscope. Color slides of the immature stages for several species were given to me by Jim Mallet, and published illustrations and sketches were also examined.


Taxonomic concepts

In many cases the taxonomy that I have adopted, at both the specific and subspecific level, differs significantly from that of previous workers. At present, there is









much debate as to what constitutes a species, both in theoretical and practical terms, and even as to whether the term species has any true meaning at all. The criteria used to define a species largely depend on which species concept one adopts and the primary goal of the classification that one hopes to achieve. Many also believe that the term subspecies also has no clear definition and is meaningless (Collins, 1992). It is my belief that the primary purpose of a classification, at least at the level of species and subspecies, is to be of use to biologists who are involved in research on the organisms. A classification can be of use in two principal ways; it allows effective communication among researchers and ready identification of taxa, and it is heuristic, that is it can be used to predict unknown biological traits in taxa when the traits are known in a related taxon.
There are two principal competing species concepts, the biological species concept, as championed by Mayr (1942), and the phylogenetic species concept, as discussed by Wiley (1981); other species concepts are discussed by Collins (1992). Of these two concepts, only the biological provides a means to test whether or not a taxon constitutes a species, through its sympatry and the existence of a consistent genetic and phenotypic gap with relation to its sister taxon. The phylogenetic concept argues that any taxonomically diagnosable monophyletic clade may be regarded as a species, but provides no guidelines as to where to draw the line between the subspecies, species, genus or higher level taxon. I regard the criterion of monophyly of species as unhelpful in classifying Adelpha, since it would either lead to the recognition of many more "species", thus obscuring their true relationships, or cause probably genetically isolated and potentially sympatric taxa to be regarded as the same species (see discussion under A. ethelda and A. epione). However, it is also true that while it is usually clear what is meant by the term species within a single locality, there is no way to determine whether isolated populations, which may or may not differ phenotypically, represent distinct biological species, and this is perhaps the most frequently cited criticism of the biological species concept.









There has been relatively little research directed towards the genetic distinctness of isolated sister taxa, but there is evidence that even parapatric populations that are known to hybridise may maintain their genetic integrity in the hybrid zone (Jiggins et al., 1996; Davies et al., 1997), and thus be regarded as distinct evolutionary lineages. Such research is lacking for all Adelpha, and I have therefore used characters that are reliable in distinguishing sympatric species, and that show little or no variation throughout the range of monotypic species, as evidence as to whether or not two allopatric populations should be treated as distinct species. In the great majority of cases, I have treated isolated populations that are closely allopatric and occur in similar habitats at similar elevations, and that do not differ from each other by any "reliable" character (as defined above), as constituting the same species, while remaining aware that all such taxonomic hypotheses require testing in the field. The uniting of such populations as a single species provides two important pieces of information: that they are not known to be sympatric, and that they are considered sister taxa, even in the absence of a synapomorphy (such species were termed "paraspecies" by Ackery & Vane-Wright (1984)). They therefore almost certainly also share similar biological traits and the classification is more heuristic. This approach is essentially the same as the biogeographical species concept utilised by Tyler et al. (1994), and I regard it as the most useful first approach to classifying Adelpha. In one case I have departed from this approach, namely in my treatment of Adelpha epione and Adelpha ethelda, and I discuss my reasons for this under each respective species.

My approach to classifying Adelpha at the species level has led to the grouping in some cases of taxa which have very distinct dorsal wing patterns and have in the past been placed in completely different species groups. Examples include A. ximena ximena and A. ximena mossi, A. messana messana and A. messana delphicola, and A. malea malea, A. malea aethalia and A. malea fundania. Typically these taxa show little variation over their range and specimens transitional to other subspecies, if known, originate from a relatively very narrow zone of apparent hybridisation. Another classic









example is the common A. lycorias; individuals of A. lycorias lara are phenotypically indistinguishable from Trinidad to northern Argentina, yet in eastern Colombia specimens clearly transitional to A. lycorias melanippe occur, which is also phenotypically stable throughout much of central Colombia. I retain subspecies names for these populations since it aids their identification and recognises that they may be demonstrated to be distinct species, and such cases are similar to many species in mimetic nymphalid genera in the subfamilies Heliconiinae and Ithomiinae.

In the examples mentioned above, if we assume that the dorsal wing patterns serve a mimetic function (see below), then narrow hybrid zones are explicable in terms of strong purifying selection by predators on either side of the zone of contact (Mallet, 1993; Joron & Mallet, 1998). At the other end of the spectrum occur taxa which exhibit continuous, clinal variation in several characters over their geographic range. While opposite ends of the cline may be phenotypically quite distinct, since there is no clear character to identify specimens from intermediate localities I regard the entire population as a single taxon. In other cases, even when there is some clinal variation, there is a consistent phenotypic gap which separates two populations. I believe it is worthwhile recognising such phenotypic gaps by application of subspecies names to the two populations in question, since it highlights geographic areas where the fragmentation of populations may frequently occur. Assessing whether or not geographic variation is discontinuous is complicated in many cases due to the lack of series of reliably and precisely labeled specimens in collections. I have tended to adopt a conservative approach to situations where I have insufficient material to ascertain the nature of geographic variation, since retaining subspecies names provides more information than synonymising them, and I leave assessment of their validity to field workers in each region.









Systematics

History of Classification

The first Adelpha species, probably all originating from the Guianas, were described by Linnaeus in 1758, in the genus Papilio. Linnaeus described three names, two in his phalanx Barbarus, and one in the Phalerati section of the phalanx Nymphalis, both groups based on wing shape and variable wing pattern characters. There are taxonomic problems surrounding all three of these names, discussed under Adelpha cytherea and Adelpha iphiclus. Probably more species were represented in collections in the latter half of the eighteenth century than is reflected by the description of names, since early authors regarded even relatively distinct taxa, such as Adelpha naxia, A. iphiclus and A. serpa, as constituting the same species. Following single descriptions in the genus Papilio, accompanied by figures, by Walch (1775) and Cramer (1777, 1780), Latreille (1809) described leucophthalma in the genus Nymphalis Latreille, 1804. Hiibner ([1819]: 42) introduced the name Adelpha for eight listed species, including A. mesentina, A. iphiclus (as A. basilis), A. serpa diadochus (as A. iphicla), A. plesaure (described shortly after by Hibner (1823)), A. cocala and A. cytherea. Htibner also listed Adelpha elea, described by Linnaeus as A. eleus, which is here considered a synonym of A. cytherea, and A. phliase, which is in fact a member of the family Riodinidae, Synargis phliassus (Cramer). He omitted A. capucinus, which has only been mentioned twice since its original description prior to this revision, and A. leucophthalma, possibly because it lacked the vertical dorsal bands typical of the other Adelpha. As can been seen by the inclusion of a riodinid, Huibner based the genus purely on wing pattern and shape, and failed to name a type species, placing it in his Familia D, which contained genera with a typically brown ground color to the wings. Scudder (1875) subsequently selected the first named species, Papilio mesentina, as the type species of Adelpha. Hiibner's genus Adelpha was largely ignored by subsequent authors, with the notable exception of several German lepidopterists, and Godart ([1824]: 335-342) soon after treated the species in the genus









Nymphalis Latreille, with a number of other unrelated Nymphalidae. Godart however provided an important review of the previous taxonomy of all species (except of course A. capucinus) with supplementary descriptions, and made the first major contribution to the description of Adelpha taxa, with five new species. Two years later, Boisduval (1836: pl. 8, fig. 4) introduced the new generic name Heterochroa, along with a new species, Heterochroa serpa. There was no description accompanying the color plate or indication of other included species, and serpa therefore becomes the type species of Heterochroa by monotypy (Hemming, 1967). Geyer (1837) was the first author, with the exception of Htibner, and the last for several decades, to use the name Adelpha, when he described the species bredowii. Although the generic name of this species has oscillated almost continuously since its original description between Adelpha, Heterochroa and Limenitis Fabricius, the species is most closely related to members of the A. serpa group, as discussed below, and belongs in the genus Adelpha as conceived here.

The year 1847 saw the publication of Hewitson's first paper on Neotropical butterflies and the start of a period of intensive activity in Neotropical taxonomy, including the genus Adelpha (see Fig. 1). Hewitson (1847) described eight valid species in Heterochroa, from throughout the Neotropics, including the formerly unexplored Andean region, and three years later in 1850 a further four. The number of described species in the genus almost doubled during this time. His descriptions were notable for their great attention to detail and their accurate figures, and he was the first to appreciate the great importance of ventral wing pattern characters, particularly in the forewing discal cell, in distinguishing species. Westwood (1850: 276) provided the first detailed description of the external morphology of Adelpha, though he also mistakenly used the name Heterochroa, listing Adelpha as a synonym. He did not list any character to define the genus, but stated that it was closely related to Limenitis, and possibly the apaturine genus Apatura. He listed the majority of described species and their synonyms, with the exception of capucinus and bredowii, the latter being placed in the genus Limenitis. The





19


name marse was also listed under Heterochroa, but Westwood (1850: 304) subsequently noted that it was in fact a member of the genus Doxocopa.


E3 taxa
0 species


o 0
t t- tt t


0 N 4I


Time


Fig. 1. Total number of valid described species and subspecies of Adelpha since the time of Linnaeus (1758).


The name Heterochroa gained widespread acceptance in the lepidopterological community in the two decades following the publication of Hewitson's and Westwood's works, during which time the number of described species of Adelpha almost doubled again. This was due largely to the papers of Bates (1864, 1865, 1866), Butler (1866), Hewitson (1867), and the Felder brothers (1861, 1862, 1867). In particular, Felder &








Felder's (1867) report on the collections made during the voyage of the Frigate "Novara" around the Earth is notable for containing the descriptions of nine valid Adelpha species, a number second only to Fruhstorfer (1913, 1915). The majority of these species originated along the northern coast of South America, with a number from mountainous areas in Venezuela and Colombia. However, the lack of figures in this work resulted in much taxonomic confusion in years to come.
Kirby (1871) finally correctly restored the name Adelpha and placed Heterochroa in synonymy, and provided a thorough synonymic catalogue of all published names in the genus. Although many of his taxonomic conclusions were highly questionable, this catalogue provided the basis for future research. Throughout the 1870s Butler was the most noticeable author of new Adelpha taxa, and the next important work to treat a large portion of the genus was that of Godman & Salvin's (1884) review of Central American species. They described in further detail the external morphology and appear to have been the first authors to have examined the male genitalia, making two principal observations; that all species were remarkably uniform, and that species of Limenitis were scarcely distinguishable. They also figured many species, the majority of which were correctly identified, and arranged the species in the form of a key. One might regard this as the first attempt to define species groups within the genus, but this was not a stated goal and many of the resulting species groups are artificial. Nevertheless, their arrangement, beginning with species with entirely black hindwings and oblique or straight orange forewing bands, such as A. lycorias melanthe and A. boreas, was followed by almost all subsequent authors (Fruhstorfer, 1915; Hall, 1938; DeVries, 1987; Neild, 1996) and is reflected in the curation of most major museum collections. They also united for the first time the members of the A. serpa group.
A relatively static period followed, until the arrival of Fruhstorfer (1907, 1908), culminating in the best known work on Adelpha, the revision of the genus in the Macrolepidoptera of the World (Fruhstorfer, 1915). Fruhstorfer's revision is at times









inexplicably careless, at times surprisingly insightful. He also failed to find any single character to define the genus irrespective of Limenitis, and his arrangement of species was based largely on a variable character of the forewing venation, dorsal wing pattern, and the presence or absence of the clunicula in the male genitalia. He recognised 91 species, described eleven valid species and 28 valid subspecies, and a very large number of additional names based on insignificant wing pattern variation. He appears to have examined some type specimens of earlier authors, yet still made a great number of errors and misidentifications, many of which have been propagated through the literature until very recently. The plates are still widely used to identify Adelpha taxa, yet they too are replete with mistakes and incorrect names. Fruhstorfer's most important contribution was the recognition and description of a number of superficially similar southeast Brazilian species, and he was the first to recognise that "orange and white banded" Adelpha contained many more species than had been previously appreciated, several of which he described.

It is, however, arguable whether the plethora of new names authored by Fruhstorfer, many with breathtaking brevity, resulted in more subsequent taxonomic confusion than clarification. Hall (1938) made an attempt to assess the validity of many of the names after Fruhstorfer's collection, including most of the type material, was transferred to the British Museum. Hall (1938) synonymised many of the subspecific names and also authored several himself, including two species. There followed a period of remarkable calm, a period of thirty-five years in which not a single valid Adelpha taxon was described (see Fig. 1), as attention perhaps turned more to the biology of species and the higher level classification of butterflies. Two papers by Beutelspacher (1975, 1976), in which three species of Mexican Adelpha were described, marked a brief period of interest in the group, during which a new Colombian species was also described (Steinhauser & Miller, 1977). The description of a remarkably distinct new species from









Costa Rica by DeVries & Chac6n (1982) gave indication that there were still undescribed Adelpha taxa awaiting discovery.

With the description of two highly distinctive species from Ecuador, Willmott & Hall (1995) began a recent surge of interest in the group, at a time which is proving to be a veritable renaissance period for butterfly taxonomy in all groups (see Fig. 1). Neild (1996) followed with the description of nine new subspecies from Venezuela, in one of the most taxonomically important works to date on the genus. Neild (1996) was the first to recognise that the "A. ixia" of previous authors actually included three separate species, A. messana, A. malea ixia and A. barnesia leucas, and was one of the very few modem authors to show an appreciation of the value of ventral wing pattern characters in species identification, in particular among certain phenotypically very similar Amazonian species. Willmott & Hall's (1999) description of a further two new species and seven new subspecies from Ecuador, and the number of new taxa described in this work, demonstrate that our understanding of the diversity of Adelpha is still far from complete.


Generic Relationships
Adelpha is currently placed in the subtribe Limenitiditi, tribe Limenitidini, and subfamily Limenitidinae of the family Nymphalidae (Harvey, 1991). As mentioned earlier, there is no evidence that the Limenitidinae constitutes a monophyletic group (Harvey, 1991), and the relationships of the Limenitidini to the Pantropical Cyrestidini, the Neotropical Coeini and Biblidini, the latter including most of the remaining Neotropical species, are not known.

The tribe Limenitidini includes a large number of species and genera distributed in both tropical and temperate regions throughout the globe, and is in all probability an ancient lineage. Chermock (1950) defined the group on the basis of a single character, the preservation of the first anal vein (IA) as a short spur at the base of the forewing cubital vein (see Fig. 2a). This character, however, is not present in one of the genera, Neptis, and


























JuIa NI-


d





k9


-11
0


I -I


Fig. 2. Morphology of adult Adelpha. a: A. serpa serpa, wing venation; b,c,d: base of forewing veins M2, Ml, R3-5 and R2, b, A. lycorias lara, c, A. lycorias spruceana, d, A. lycorias melanthe; e,f: forewing discocellular vein, e, A. capucinus capucinus, f, A. boreas boreas; g,h: A. mesentina, female genitalia, g, dorsal view, h, lateral view; ij: A. mesentina, male genitalia, i, lateral view, j, inside of valva; kl: juxta, posterior, lateral and ventral views (left to right), k, A. alala completa, 1, A. serpa celeio; m: A. diocles creton, posterior view of gnathos; n: A. serpa celerio, lateral view of aedeagus; o: A. serpa celerio, sclerotised aedeagal pad, ventral (left) and lateral (right) view; p,q: lateral view of eye and labial palpi, p, A. alala negra, q, A. mesentina.


6:.. ilk









also occurs in certain members of the Heliconiinae, as noted by Michener (1942) and Brown & Heineman (1972), including the genera Philaethria, Dryas and Dryadula (Penz, pers. comm.). Harvey (1991) also defined the tribe on the basis of this character, while noting that a further possible character which may prove to be a synapomorphy for the tribe is the structure of the eggs (Aiello, 1984). All eggs known to date are composed of concave, hexagonal facets with setae at the interstices. Harvey (1991) stated that the eggs of members of the Partheniti were unknown, but I can confirm, from dissection of the female of Parthenos sylvia, that they also have this characteristic structure. Harvey (1991) recognised four subtribal divisions corresponding to groups of various taxonomic rank recognised by earlier authors (Chermock, 1950, and references therein; Eliot, 1978), including the Neptiti, Limenitiditi, Partheniti and Euthaliiti, all of which were defined by Eliot (1978) on the basis of wing venation. The character given by Eliot (1978) to define the Limenitiditi is the origin of the hindwing humeral vein at or before the origin of vein Rs-M1-M2. However, wing venation is likely to be highly variable (it is even in a single species of Adelpha, A. lycorias, see Fig. 2), and there is no evidence whether the characters of Eliot represent synapomorphies or symplesiomorphies. While the Euthaliiti and Neptiti probably constitute monophyletic groups, on the basis of the highly distinctive early stages, particularly the larvae, in the former, and the adult morphology and early stages of the latter (Harvey, 1991), there is no evidence that the Partheniti or Limenitiditi are monophyletic; they merely include the members of the Limenitidini that do not belong in the tribes Euthaliiti and Neptiti.

The Limenitiditi consists of a number of tropical and temperate species whose generic relationships are probably the most poorly understood of the entire subfamily. Most are placed in the catch-all genus Limenitis, while numerous generic names have been proposed (see Chermock, 1950), a number of which are no doubt valid. Within the New World the only representatives of the Limenitiditi, and indeed the Limenitidini, are









Adelpha and four North American species placed either in the genus Limenitis or Basilarchia.


The Genus Adelpha

Characteristics

The genus Adelpha includes medium to large butterflies which have essentially sexually monomorphic wing patterns, consisting of contrasting ventral and dorsal surfaces (see Figs. 10-94). The dorsal surface always has a dark brown ground color and almost always a postdiscal band on one or both wings, which may be vertical or oblique, and orange, white, or red in color. The ventral surface typically has a black ground color which is usually obscured by much reddish brown coloring, and in addition to the postdiscal bands which appear on the dorsal surface it has a number of lighter postdiscal and submarginal lines and other more basal pattern elements (see Fig. 3). Despite the superficially large differences in dorsal pattern between species, all are composed of relatively few pattern elements, particularly the postdiscal band, the postdiscal series and, on the forewing of some species, the submarginal series.

Identification of the origin of various areas of the dorsal wing pattern in terms of the ventral pattern elements is often important in distinguishing Adelpha species. In almost all species the ventral pattern elements are visible, and more primitive species, such as members of the A. alala group (see Table 1), tend to have a greater complement of unmodified pattern elements. The pattern elements that I have recognised in Fig. 3 differ from those recognised by Schwanwitsch (1924) and Nijhout (1991), who both state that the nymphalid ground color is pale and pattern elements are dark. Thus, the pale postdiscal and submarginal series and postdiscal band, which play the most important role in species identification, actually constitute ground color, while the darker areas between are the "pattern". Although Nijhout (1991) rightly points out that few authors have attempted to recognise the system of Schwanwitsch (1924) in description of









butterfly wing patterns, I have chosen a terminology for the wing pattern of Adelpha that allows discussion and comparison of wing patterns with maximum clarity. It is much simpler to refer to the pale areas of the ventral wing surface in Adelpha as pattern elements, while acknowledging that they are not by the definition Schwanwitsch (1924). It is, of course, also possible to recognise the pattern elements described by Schwanwitsch (1924) and Nijhout (1991) in Adelpha; more basal elements are obvious, while the distal band of the central symmetry system borders the basal edge of the postdiscal band, and the dark areas between the postdiscal series, the outer postdiscal and inner submarginal series, and the submarginal series correspond respectively to elements g, h (border ocelli) and I (parafocal element) (Nijhout, 1991: Fig. 2.17).


postdiscal band
(upper, above M3) I
postcellular markings
postcellular bar fourth cell bar third cell bara


Fig. 3. Wing pattern elements of Adelpha, indicating terminology adopted in the text.









Morphologically Adelpha is remarkably homogeneous, and detailed descriptions of the wing venation, antenna, eyes, labial palpi and legs are given by Westwood (1850) and Godman & Salvin (1884). The wing venation (Fig. 2a) varies little in the genus, except that the discocellular vein may or may not be present on the forewing, while its juncture with vein M3 may be above or below the origin of vein Cul (Fig. 2e,f). The position of the base of vein R2 is also variable within A. lycorias (Fig. 2b-d). The eyes are variably setose, with montane species of the A. alala group having particularly long and dense setae, while certain species, such as A. bredowii and A. nea, have naked eyes. The labial palpi may or may not have a pronounced lateral black stripe (Figs. 2p,q), while the thorax is either entirely pale in color, or has dark bands where the legs rest against the body. Otherwise all of these structures show little other morphological variation throughout the genus.

The male (Figs. 2ij, 95-177) and female genitalia (Figs. 2g,h, 178-249) are also both relatively homogeneous. The male genitalia show interspecific differences particularly in the shape of the valva and distribution of terminal spines, and may or may not have a pronounced projection from the inner edge of the valva (Fig. 2j), termed the "clunicula" by Fruhstorfer (1915). The uncus, tegumen, gnathos and aedeagus vary little, except that members of the A. serpa group have a sclerotised pad in the aedeagus on the dorsal surface of the vesica. The juxta is also relatively uniform, with slight differences in the A. serpa group (Fig. 2k,l). The female genitalia vary somewhat in size, even considering size differences of the adults, but also vary little morphologically. The lamella postvaginalis and antevaginalis are wrinkled plates which are variably sclerotised, the ostium bursae is always a simple ring which tends to be wider in members of the A. serpa group, the ductus seminalis always originates dorsally near the ostium bursae and the relative lengths of the ductus bursae and corpus bursae vary little. The corpus bursae typically has a pair of sclerotised bands (Fig. 2g,h) which may be dorsal, on the right hand side or ventral, with both inter and intraspecific variation in the position and spacing of









the bands. The degree of sclerotisation also varies, as does the size of the spines on the inner surface of the bands, with members of the A. serpa group having the most heavily sclerotised and coarsely spined bands. The terminal tergite of the female genitalia is variably sclerotised at the anterio-ventral edge, in one species (A. thoasa) forming a small pouch. Similar structures occur in the Asian genus Neptis.

While Adelpha has been regarded as a distinct genus by the great majority of authors, Chermock (1950) placed it as a subgenus of Limenitis, in which he was followed by authors in the following few decades. This was due to the entire absence of morphological characters by which to define Adelpha, and the retention of the genus by all previous authors has been on the basis of its isolated geographical distribution and the overall similarity of most species. Despite an intensive examination of the external and internal morphology of both males and females, I have also been unable to find any morphological characters to define the genus. I have, however, located several wing pattern characters which suggest that the genus is monophyletic. All species of Adelpha have a dark brown longitudinal line extending from the base of cell 2A to the middle of the anal margin in this cell, with the exception of several species clearly closely related to species that do have the character. In many species this line may be fused to a band of color which fills the anterior half of the cell, and in several it is broken in the basal half of the wing. Species that lack this line also have all the markings in the basal half of the wing reduced, and presumably the line has been secondarily lost. This line appears to be either an intervenal stripe (Nijhout, 1991) or the posterior half of a venal stripe along vein 2A that has become detached from the vein and displaced posteriorly. Almost all species also have a venal stripe at vein 3A on the ventral hindwing, which may also be split along the vein into two halves and variously modified or reduced, being almost invisible in members of the A. alala group. Almost all species of Adelpha have the postdiscal series fused on the dorsal surface of the forewing to form a subapical marking; in other limenitidine genera the postdiscal series are clearly distinct. Again, the species of Adelpha










in which the postdiscal series are not fused are derived and closely related to species that do have this character. A further possible synapomorphy is the streak at the base of the ventral hindwing discal cell, apparently corresponding to the basal symmetry system of Nijhout (1991).



Taxonomy and identification

I recognise 85 species and 207 taxa in the genus Adelpha, and I present a synonymic checklist below in an order from primitive to advanced, largely following the phylogeny presented in Chapter 3. I include names proposed as quadrinomials even though such names are excluded by the Code, since in at least one instance (syrna Fruhstorfer, a synonym of A. seriphia godmani) subsequent authors have made the name available. Since I do not claim my bibliographic review to be exhaustive, this may also be true of other names, and all such names are listed in the Review of the genus below with their associated type material. I also discuss the spelling and status of Fruhstorfer names in the introduction to the generic Review below. One species and four subspecies are undescribed, three of these due to a lack of material, and two due to their impending description by other authors.

Adeipha Hibner ([1819]: 42).
Type species by selection by Scudder (1875: 102): Papilio mesentina Cramer (1777: 102, pl. 162, figs.
B, C).
=Heterochroa Boisduval (1836: pl. 8, fig. 4)
Type species by monotypy: Heterochroa serpa Boisduval (1836: pl. 8, fig. 4)
("-" denotes a subspecies, "--" a synonym and "--" an unavailable name) bredowii Geyer, 1837 nea (Hewitson, 1847)
-eulalia (Doubleday, [1848]) --campeda Fruhstorfer, 1915
--guatemalensis (Carpenter & Hobby, 1945) syn. -sentia Godman & Salvin, 1884 stat. nov. nov. paraena (Bates, 1865)
-californica (Butler, 1865) -massilia (C. Felder & R. Felder, 1867) diocles Godman & Salvin, 1878 -reyi Neild, 1996
-creton Godman, 1901 -lecromi Willmott, 1999 ssp. nov. herbita Weymer, 1907 radiata Fruhstorfer, 1915 zea (Hewitson, 1850) -myrlea Fruhstorfer, 1915
--serpentina Fruhstorfer, 1915 -gilletella Brivignon, 1995
--tarpeia Fruhstorfer, 1915 -aiellae Willmott & Hall, 1999 paroeca (Bates, 1864) -explicator Wilmott & Hall, 1999
--emathia (R. Felder, 1869) stat. rest. serpa (Boisduval, 1836)









--damon Fruhstorfer, 1913
-ornamenta Fruhstorfer, 1915
-celerio (Bates, 1864)
--diademeta Fruhstorfer, 1913
--phintias Fruhstorfer, 1913
-duiliae Fruhstorfer, 1913
-diadochus Fruhstorfer, 1915
--timehri Hall, 1938
-florea Br6vignon, 1995 seriphia (C. Felder & R. Felder, 1867)
-pione Godman & Salvin, 1884
-aquillia Fruhstorfer, 1915
--naryce Fruhstorfer, 1915
-therasia Fruhstorfer, 1915
-egregia R6ber, 1927
-godmani Fruhstorfer, 1913
--syrna Fruhstorfer, 1915
-barcanti Willmott, 1999 ssp. nov. hyas (Doy~re, [ 1840])
-viracocha Hall, 1938
-hewitsoni Willmott & Hall, 1999 a/ala (Hewitson, 1847)
-completa Fruhstorfer, 1907
--titia Fruhstorfer, 1915
-negra (C. Felder & R. Felder, 1862)
--ehrhardi Neuburger, 1907 syn. nov.
--albifida Fruhstorfer, 1907 syn. nov.
--cora Fruhstorfer, 1907 syn. nov.
--fillo Fruhstorfer, 1907 syn. nov.
--negrina Fruhstorfer, 1913 syn. nov.
---praecaria Fruhstorfer, 1915
--privigna Fruhstorfer, 1915 syn. nov. aricia (Hewitson, 1847)
-serenita Fruhstorfer, 1915 stat. rest.
-portunus Hall, 1938 corcyra (Hewitson, 1847)
-aretina Fruhstorfer, 1907
-collina (Hewitson, 1847) stat. rest.
-epidamna (C. Felder & R. Felder, 1867)
-dognini Willmott, 1999 ssp. nov.
-salazari Willmott, 1999 ssp. nov. tracta (Butler, 1872) pithys (Bates, 1864)
--vodena Fruhstorfer, 1915 stat. nov. donysa (Hewitson, 1847)
--roela (Boisduval, 1870)
-albifilum Steinhauser, 1974 stat. nov. fessonia (Hewitson, 1847)
-lapitha Hall, 1929 stat. nov.
-cestus (Hewitson, 1847)
-ernestoi Willmott, 1999 ssp. nov. gelania (Godart, [1824])
-arecosa (Hewitson, 1847) calliphane Fruhstorfer, 1915 poitius Hall, 1938 mythra (Godart, [1824])
--zeba (Hewitson, 1850) basiloides (Bates, 1865)


--lydia (Butler, 1865) stat. rest.
--lemnia (C. Felder & R. Felder, 1867)
--lativittata Staudinger, 1886 syn. nov.
--caelia Fruhstorfer, 1915 syn. nov. plesaure Hibner, 1823
--heredia Fruhstorfer, 1915
--antoniae Fruhstorfer, 1915
-phliassa (Godart, [1824])--euboea (C. Felder & R Felder, 1867)
--implicata Fruhstorfer, 1915
--bartolme Fruhstorfer, 1915 syn. nov.
--cerachates Fruhstorfer, 1915
--sirona Fruhstorfer, 1915
-pseudomalea Hall, 1938
-symona Kaye, 1925 gavina Fruhstorfer, 1915 falcipennis Fruhstorfer, 1915
-perga Fruhstorfer, 1915 stat. rest. thoasa (Hewitson, 1850)
--silia Frubstorfer, 1913
-cuyaba Fruhstorfer, 1915
-gerona (Hewitson, 1867) stat. rest.
--brevifascia Talbot, 1928
-calliphiclea (Butler, [1870]) stat. nov.
-manilia Fruhstorfer, 1915
--zalma Fruhstorfer, 1915 thessalia (C. Felder & R. Felder, 1867)
-indefecta Fruhstorfer, 1913 stat. nov.
--mincia Hall, 1938 syn. nov.
-cesilas Fruhstorfer, 1915
--ampla Hayward, 1935 syn. nov. iphiclus (Linnaeus, 1758)
-basilea (Cramer, 1777) stat. rest.
--basilis Hiibner, [ 1819]
-funalis Fruhstorfer, 1915
--exanima Fruhstorfer, 1915
--pharae Fruhstorfer, 1915
-ephesa (Mnktri~s, 1857)
--gellia Fruhstorfer, 1915
--abylina Fruhstorfer, 1915
-estrecha Willmott & Hall, 1999 iphicleola (Bates, 1864)
--massilides Fruhstorfer, 1915
-iphimedia Fruhstorfer, 1915
--phera Fruhstorfer, 1915
-thessalita Willmott & Hall, 1999
-gortyna Fruhstorfer, 1915
-leucates Fruhstorfer, 1915
--daceleia Fruhstorfer, 1915 abyla (Hewitson, 1850) melona (Hewitson, 1847)
--arete (Mnftri~s, 1857) syn. nov.
--cibyra Fruhstorfer, 1915 stat. nov.
-leucocoma Fruhstorfer, 1915 stat. rest.
--meridionalis Fruhstorfer, 1915 syn. nov.
--biedermanni Fruhstorfer, 1915 syn. nov.
--nonsecta Kaye, 1925 syn. nov.
-pseudarete Fruhstorfer, 1915 stat. nov.









-deborah Weeks, 1901 stat. rest.
-neildi Willmott, 1999 ssp. nov. ethelda (Hewitson, 1867)
-zalmona (Hewitson, 1871) stat. nov.
-sophax Godman & Salvin, 1878 stat. nov.
-eponina Staudinger, 1886 stat. nov.
-volupis Fruhstorfer, 1915
-galbao Br6vignon, 1995 stat. nov.
-ssp. nov.
epione (Godart, [1824])
-agilla Frubstorfer, 1907 syma (Godart, [1824])
--pravitas Fruhstorfer, 1913 syn. nov. cytherea (Linnaeus, 1758)
-eleus (Linnaeus, 1758) stat. rest.
--lanilla Fruhstorfer, 1913
-aea (C. Felder & R. Felder, 1867)
--herennia Fruhstorfer, 1915
-daguana Fruhstorfer, 1913
--tarratia Fruhstorfer, 1913 syn. nov.
--despoliata Fruhstorfer, 1915 syn. nov.
-insularis Fruhstorfer, 1913
-marcia Fruhstorfer, 1913
-olbia (C. Felder & R. Felder, 1867)
-nahua Grose-Smith, 1898
-wernickei R6ber, 1923 viola Fruhstorfer, 1913 stat. nov.
-pseudococala Hall, 1933 stat. nov. salmoneus (Butler, 1866)
-emilia Fruhstorfer, 1908
-colada (C. Felder & R. Felder, 1867)
-salmonides Hall, 1938 demialba (Butler, 1872) epizygis Fruhstorfer, 1915
-epona Fruhstorfer, 1915 fabricia Fruhstorfer, 1913
--hilareia Fruhstorfer, 1915 stat. nov.
--mamaea Fruhstorfer, 1915 syn. nov. capucinus (Walch, 1775)
--juruana Butler, 1877 stat. nov.
-phylacides Frustorfer, 1913 stat. nov.
-georgias Fruhstorfer, 1915 stat. nov.
-velia (C. Felder & R. Felder, 1867) stat. nov.
--veliada Fruhstorfer, 1915
-gutierrezi Neild, 1996 stat. nov. barnesia Schaus, 1902
--pseudomessana Fruhstorfer, 1913 syn. nov.
-leucas Fruhstorfer, 1915 stat. nov.
--paula Bargmann, 1928 syn. nov.
-trinita Kaye, 1914 stat. nov. diazi Beutelspacher, 1975 hesterbergi Willmott & Hall, 1999 abia (Hewitson, 1850)
--rufilia Fruhstorfer, 1915 naxia (C. Felder & R. Felder, 1867)
--dominula M6schler, 1877 syn. nov.
--epiphicla Godman & Salvin, 1884 stat. nov.
--mucia Fruhstorfer, 1915 syn. nov.


--diatreta Fruhstorfer, 1915
--zynia Fruhstorfer, 1915 syn. nov.
--canuleia Fruhstorfer, 1915 syn. nov.
-hieronica Fruhstorfer, 1913
-oteroi Neild, 1996 heraclea (C. Felder & R. Felder, 1867)
--fugela Fruhstorfer, 1915
-makkeda (Hewitson, 1871)
-antonii Neild, 1996 sp. nov.
malea (C. Felder & R. Felder, 1861)
-ixia (C. Felder & R. Felder, 1867) stat. nov.
--himera (C. Felder & R. Felder, 1867) syn. nov.
-fundania Fruhstorfer, 1915 stat. nov.
-aethalia (C. Felder & R. Felder, 1867)
--davisii (Butler, 1877) syn. nov.
--metaxa Fruhstorfer, 1913 stat. nov.
--frusina Fruhstorfer, 1913 syn. nov.
--sarana Fruhstorfer, 1913 stat. nov.
--trinina Kaye, 1914 syn. nov.
--incomposita Austin & Mielke, 1993 syn. nov.
-goyama Schaus, 1902 stat. nov.
--leopardus Fruhstorfer, 1913 boeotia (C. Felder & R. Felder, 1867)
--fulica Fruhstorfer, 1915 syn. nov.
-oberthurii (Boisduval, 1870)
--jacquelinae Steinhauser & Miller, 1977 syn. nov.
-fidicula Fruhstorfer, 1915 altamazona Austin & Jasinski, 1999 ximena (C. Felder & R. Felder, 1862)
-mossi Hall, 1933
--willmotti Neild, 1996 syn. nov. delinita Fruhstorfer, 1913
-albina Fruhstorfer, 1915 syn. nov.
-utina Hall, 1938
--escalantei Steinhauser & Miller, 1977 syn. nov. pollina Fruhstorfer, 1915
--uta Fruhstorfer, 1915 syn. nov. erotia (Hewitson, 1847)
--lerna (Hewitson, 1847) stat. rest.
--aeolia (C. Felder & R Felder, 1867) stat. rest.
--permagna Fruhstorfer, 1913 stat. rest.
--leonina Fruhstorfer, 1913
--archidona Fruhstorfer, 1913
-caphira (Hewitson, 1869) stat. rest.
--deleta Fruhstorfer, 1913 syn. nov. phylaca (Bates, 1866)
-pseudaethalia Hall, 1938
-joffrei Neild, 1996 messana (C. Felder & R. Felder, 1867)
--aufidia Fruhstorfer, 1915 stat. nov.
-delphicola Fruhstorfer, 1909 stat. nov. thesprotia (C. Felder & R. Felder, 1867)
--praetura Fruhstorfer, 1913
--nava Fruhstorfer, 1913 syn. nov.
--ophellas Fruhstorfer, 1915 syn. nov.
--zunilaces Fruhstorfer, 1915 syn. nov. mesentina (Cramer, 1777)









--chancha Staudinger, 1886 syn. nov. lycorias (Godart, [1824])
--isis (Drury, 1782) preocc. (Drury, 1773)
--divina Fruhstorfer, 1907 stat. nov.
--pseudagrias Fruhstorfer, 1908 syn. nov.
-wallisii (Dewitz, 1877) stat. nov.
--hypsenor Godman & Salvin, 1879
-fassli Fruhstorfer, 1915 syn. nov.
-lara (Hewitson, 1850)
--mainas Frubstorfer, 1915
-melanthe (Bates, 1864)
-melanippe Godman & Salvin, 1884 stat. nov.
-spruceana (Bates, 1864) stat. nov. attica (C. Felder & R. Felder, 1867)
--lesbia Staudinger, 1886
--oronoco Weeks, 1906
-carmela Fruhstorfer, 1915
-serita Fruhstorfer, 1915
-hemileuca Willnott & Hall, 1999 leuceria (Druce, 1874)
-juanna Grose-Smith, 1898 stat. nov. leucerioides Beutelspacher, 1975 erymanthis Godman & Salvin, 1884
-adstricta Fruhstorfer, 1915
-ssp. nov.
-ssp. nov.
sichaeus (Butler, 1866)
-privata Fruhstorfer, 1913
--leucopetra Fruhstorfer, 1913 stat. nov.
-irisa Fruhstorfer, 1913 syn. nov. rothschildi Fruhstorfer, 1913
--pallida R6ber, 1927 stilesiana DeVries & Chac6n, 1982 boreas (Butler, 1866)
--tizona (C. Felder & R. Felder, 1867) syn. nov.
-tizonides Fruhstorfer, 1908 stat. nov.
--verenda Fruhstorfer, 1913
--opheltes Fruhstorfer, 1915 syn. nov.
-kayei Hall, 1939 cocala (Cramer, 1780)
--urraca (C. Felder & R. Felder, 1862)
--saparua Fruhstorfer, 1913
--urracina Fruhstorfer, 1913 stat. nov.
--lorzina Fruhstorfer, 1915 syn. nov.
-caninia Fruhstorfer, 1915
-lorzae (Boisduval, 1870)
--cocalina Fruhstorfer, 1913 stat. nov.
--fufia Fruhstorfer, 1913 syn. nov.
--fufina Fruhstorfer, 1915 syn. nov.
-didia Fruhstorfer, 1915 stat. rest.
---chlide d'Almeida, 1931
-orellanai Neild, 1996 felderi (Boisduval, 1870)


--falcata Godman & Salvin, 1878 stat. rest.
--jarias Fruhstorfer, 1915 syn. nov. leucophthalma (Latreille, [ 1809])
--mephistopheles (Butler, 1869)
--tegeata Fruhstorfer, 1915
-irminella Strand, 1918 stat. nov.
--iaere Hall, 1929 syn. nov.
-smalli Willmott, 1999 ssp. nov. irmina (Doubleday, [ 1848])
-tumida (Butler, 1873)
--wilhelmina Fruhstorfer, 1907 saundersii (Hewitson, 1867)
--leutha Fruhstorfer, 1915
-helepecki Weeks, 1901 stat. rest.
-frontina Hall, 1935 lamasi Willmott & Hall, 1999 salus Hall, 1935
-emmeli Wilhnott & Hall, 1999
-ssp. nov.
shuara Willmott & Hall, 1995 argentea Willmott & Hall, 1995 coryneta (Hewitson, 1874) jordani Fruhstorfer, 1913
--ernestina Fruhstorfer, 1913 syn. nov. zina (Hewitson, 1867)
--restricta Fruhstorfer, 1913
--leucacantha Fruhstorfer, 1915
--manetho Hall, 1938 syn. nov.
-inachia Staudinger, 1886
---inachia Fruhstorfer, 1915
-lacina (Butler, 1872)
-irma Fruhstorfer, 1907 stat. nov.
--nadja Fruhstorfer, 1907
--nicetas Hall, 1938 syn. nov.
-enope Hall, 1938 stat. nov.
-desousai Neild & Orellana, 1996 stat. nov.
-pyrczi Neild & Orellana, 1996 stat. nov. milleri Beutelspacher, 1976 justina (C. Felder & R. Felder, 1861)
--praevalida Fruhstorfer, 1915
-justinella Fruhstorfer, 1907
-valentina Fruhstorfer, 1915 stat. nov.
-maira Willmott, 1999 ssp. nov.
---maira Fruhstorfer, 1915
-inesae Orellana, 1996 olynthia (C. Felder & R. Felder, 1867)
--olynthina Fruhstorfer, 1907 syn. nov.
--levicula Fruhstorfer, 1913 syn. nov.
--olynthina Fruhstorfer, 1915 syn. nov.
--theaena Fruhstorfer, 1915 syn. nov.
--zopyra Fruhstorfer, 1915 stat. nov. levona Steinhauser & Miller, 1977









Key to species of Adelpha based on adult wing pattern

I believe that the simplest way to identify Adelpha is through comparison of specimens with the plates and text accounts, but I recognise that many find a key to be a useful device. The purpose of this key is to allow the non-specialist to identify specimens without needing to resort to dissection or examination of microscopic structures, or analysis of wing pattern element homologies, which requires some experience with the genus. I have therefore expressly not necessarily used the terminology of Fig. 3 for wing patterns, but I have used more widespread wing pattern terminology, such as the term "postdiscal band" to refer to the entire dorsal band rather than just the portion homologous to the element shown on Fig. 3. The key is not arranged strictly phylogenetically and in most cases the characters employed are not considered to be specific synapomorphies; indeed, a number of species have been keyed out more than once due to differences in the most obvious wing pattern characters. I discuss the characters that I consider to define species under the species accounts in the generic Review below, and the Key is to be used in conjunction with the plates, the ranges and further identifying characters given in the accounts for each taxon. I have in some cases listed additional characters to confirm identification, but those given first are the most reliable. It should be noted that, in terms of variation, "usually" refers to variation between subspecies, and "typically" refers to variation within a single taxon.


1. DHW with white or orange markings in postdiscal area ....................................... 2

DHW entirely brown in postdiscal area (orange tornal spot may be present) .......... 87 2(1). VFW with pale distal half of discal cell, between dark cell band and discocellular

band, divided into three (usually) by basally pointing, "V"-shaped black line (line may not quite touch basal red-orange band, in which case it divides area into two, a basal, hourglass shape and distal, much smaller triangle); very base of VHW discal








cell always entirely white; dark streak absent in base of VFW discal cell ............................................................................................................................... 3

VFW with pale distal half of discal cell divided into two by a straight, vertical central dark line, a horizontal dark line, or undivided; very base of VHW discal cell entirely white or with a dark line or darker shading; dark streak almost always present in base of VFW discal cell ...................................................................... 14

3(2).VHW with orange/yellow postdiscal line distal of white postdiscal band extending

from tomal orange/yellow spot in cell Cu2 to costa; basal edge of white postdiscal band usually bordered by a red-orange band in most taxa; continuous black line from costa to vein 3A through discal cell usually present ...................................... 4

VHW with postdiscal area distal of white postdiscal band lacking orange/yellow band, with isolated orange tornal spot in cell Cu2; basal edge of white postdiscal band not bordered by a red-orange band; no continuous black line from costa to vein 3A through discal cell ....................................................................................... herbita

4(3).VHW with basal portion of vein 3A white, distal portion lined with black which

continues as a continuous dark line to costa .......................................................... 5

VHW with vein 3A entirely black to wing base, not part of a line crossing discal cell ........................................................................................................................... zea

5(4). VFW with undivided whitish submarginal dashes in each cell except Cu2,

particularly clear in cells Cul and M I ................................................................... 6

VFW with whitish submarginal dashes divided into two in every cell, particularly clear in cells Cul and M 1 ....................................................................................... 8

6(5). VFW with distal edges of silvery-white postdiscal blocks in cell Cul and Cu2

aligned; VFW costa whitish immediately distal of postcellular bar ...................... 7

VFW with distal edge of silvery-white postdiscal block in cell Cul much more distal than that in cell Cu2; VFW costa brown immediately distal of postcellular bar .... .... .........�.... ............. �.... ..... ...... '. ... ...................................... ...... n ea









7(6). DFW with approximately equally sized, orange and white postdiscal spots in cell

M 3 ................................................................................................................... p aroeca

DFW with either an orange or a white postdiscal spot in cell M3, but not both ..................................................................................................................p araena

8(5). DFW orange postdiscal marking absent in cells M1 and R5 ........................... seriphia

DFW orange postdiscal marking present in cells M1 and R5 ............................... 9

9(8).VFW pale orange postdiscal marking in cells M2-R5, and/or pale silver grey to

brownish shading in cell M3 just distal of white postdiscal spot, with clear, dark lines parallel to veins intruding distally at least half way into pale marking and/or to edge of white postdiscal spot (in cell M3) .......................................................... 10

VFW with both pale orange postdiscal marking and pale coloring in cell M3 distal of white postdiscal spot with no dark lines parallel to veins intruding distally, or these only enter a short way, or entire marking is dark orange and darker lines cannot be seen clearly ............................................................................................... 11

10(11).VFW costa pale yellowish or whitish at base; pale markings distal of red-orange

postdiscal band on VHW parallel to band ........................................................ hyas

VFW costa red-orange at base, at least along distal edge of costal vein; pale markings distal of red-orange postdiscal band on VHW noticeably closer to band in cells Rs and Sc+R1 than in cells M2 and Ml seriphia 11(9). DFW with white postdiscal marking in cell M3 well developed, always at least

half width of marking in cell Cul ..................................................................... 12

DFW with white postdiscal marking in cell M3 absent or a tiny dot, much reduced compared to size of marking in cell Cul ....................................................... radiata

12( 1).VHW with two rows of whitish submarginal markings separated by a sharply

defined dark brown line; whitish markings may or may not be divided in two in each cell by a dark brown line parallel to the veins .......................................... 13









VHW with two rows of whitish submarginal markings almost merged, separated only by a faint, poorly defined darker line; whitish markings clearly divided in two in each cell by a dark brown line parallel to the veins .................................. radiata

13(12).DFW with orange subapical marking vertically orientated, more tall than wide;

white postdiscal spots in cells Cul and M3 strongly diagonally displaced; VHW with white submarginal markings much nearer to orange postdiscal band in cells Rs and Sc+RI than in cells M3-M1; Central America and Andes ............. seriphia

DFW with orange subapical marking horizontally orientated, more wide than tall; white postdiscal spots in cells Cul and M3 usually not strongly diagonally displaced; VHW with white submarginal markings approximately parallel to orange postdiscal band throughout wing; Central and South America to southeastern Brazil, Paraguay ........................................................................... serpa

14(2). DFW with continuous, even white postdiscal band from costa to anal margin,

unbroken by dark veins ................................................................................ fessonia

DFW postdiscal band variable in extent and color, but never white and unbroken from costa to anal margin, and may be entirely absent ...................................... 15

15(14).VFW discal cell with pale area between red-orange cell bands divided in two by a

vertical, clearly defined red-brown to black line .............................................. 16

VFW discal cell with pale area between red-orange cell bands undivided or (very rarely) with an indistinct, poorly defined darker central vertical line which obliterates most of the pale area ....................................................................... 24

16(15). DFW discal cell with red scaling at base and between cell bars 1 and 2 ........... 17

DFW discal cell entirely dark brown or with some scattered orange brown scaling "' ".. ........ ����...... ""... ..... "�......... � ....oo..o..... o... . ooo ............................................... 18
17(16). DFW with a large subapical orange marking .............................................. bredowii

DFW lacking orange markings ..................................................................... gelania

18(16).DFW with postdiscal band in cell Cu2 entirely, or at least in basal half, white ..... 19









DFW with postdiscal band entirely orange in cell Cu2 .................................... viola

19(18).DFW with area between postdiscal band in cell M3 and costa dark brown, or with

indistinct, whitish shading, or entirely orange ................................................. 20

DFW with area between postdiscal band in cell M3 and costa dark brown with two well defined white ovals in cells Ml and R5 ............................................. alala

20(19).DFW with usually relatively narrow orange postdiscal band which remains

isolated from more posterior white postdiscal band ........................................ 21

DFW with very broad orange postdiscal band of which basal edge merges smoothly with basal edge of more posterior white postdiscal band ............ cytherea

21(20).VHW with silvery submarginal line and distal edge in cells M3-Rs approximately

straight, with a uniform red-brown border lacking white marginal scaling in the m iddle of each cell; Andes ............................................................................... 22

VHW with silvery submarginal line and distal edge in cells M3-Rs distinctly scalloped, with sparse white marginal scaling in the middle of each cell; Central A m erica ............................................................................................................ 23

22(21). VFW with pale subapical marking in cells M2-R5 composed of basally pointing,

"V"-shaped dashes; basal edge of marking in cells M2-R5 slightly convex; Andes from extreme northern Peru-Venezuela ........................................................ corcyra

VFW with pale subapical marking in cells M2-R5 composed of straight dashes; basal edge of marking in cells M2-R5 straight; Andes from extreme northern Peru-B olivia ..................................................................................................... aricia

23(21).DFW with orange subapical marking approximately equal in width to white

postdiscal band; VHW discal cell with ground color entirely whitish gray; VFW with red-brown line entirely surrounded by whitish gray scaling between distal edge of white postdiscal band and costa; base of cell Cul with whitish gray scaling; hindwing always slightly produced at tomus ..................................... pithys









DFW with orange subapical marking broader than white postdiscal band; VHW discal cell with ground color reddish brown between cell bars 1 and 2, and 3 and postcellular; VFW with red-brown line between distal edge of white postdiscal band and costa usually lacking whitish gray scaling between it pale yellow subapical marking; base of cell Cu 1 reddish brown; hindwing usually rounded at tom us .............................................................................................................. donysa

24(15). DFW lacking red scaling in discal cell and at discocellulars ............................ 25

DFW with red band crossing middle of discal cell and over discocellulars...diocles 25(24). DFW with postdiscal band present and completely white to vein M3, spot in cell

Cul w ell developed .......................................................................................... 26

DFW with some orange scaling in postdiscal band between anal margin and vein M3, or band is absent in cell Cul (a tiny, isolated whitish brown dot may be present at base of cell Cul), or DFW entirely dark brown ............................... 46

26(25).DHW with orange tomal spot present ............................................................... 27

DHW with orange tornal spot absent ............................................................... 44

27(26).DFW with a large orange subapical marking, lacking additional well developed

orange subapical spots distally (tiny dots may be present near distal edge in cells M 1 an d R 5) ....................................................................................................... 28

DFW with two well developed orange subapical spots in cells R5 and R4 separated from a more basal, much larger orange marking by an area of dark brown at least as wide as the more basal marking ............................................ naxia

28(27).VFW with uneven, indistinct, pale orange, silver-gray and/or reddish brown

markings in postdiscal area, not forming well defined straight stripes ............. 29

VFW with a pair of well defined, straight, vertical postdiscal stripes between cells Cul-R5, a basal white stripe and a distal orange stripe, bordered distally and basally by black ................................................................................................. sym a









29(28).DFW with orange subapical marking not touching white postdiscal marking in

cell M 3 ................................................................................................................... 30

DFW with orange subapical marking touching distal edge of white postdiscal m arking in cell M 3 ............................................................................................ 45

30(29). DFW with white postdiscal band terminating at vein M3 ................................ 31

DFW with white postdiscal band extending fuly to vein M2, sometimes as dashes in cells M 1 and R5 ..................................................................................... basiloides

31(30). VFW with area immediately distal of discocellular veins orange to reddish brown,

then dark brown line (postcellular bar), then silver gray ................................. 32

VFW with area immediately distal of discocellular veins orange, then very faint slightly darker line (postcellular bar), then orange ......................................... mythra

32(31).VFW with white postdiscal band continuing to costa as grayish white band

anterior of vein M3, distal edge of which is aligned with distal edge of white postdiscal band in cells Cul and M 3 ................................................................. 33

VFW with white postdiscal band broken in cell M2; if there is whitish shading in cell M2, distal edge is not aligned with distal edge of white postdiscal band in cells Cul and M 3 .............................................................................................. 34

33(32). VHW white postdiscal band immediately bordered distally by very thin dark

brown line, then red-brown band as broad as white postdiscal band, then one to two indistinct silver gray lines ....................................................................... poltius

VHW white postdiscal band immediately bordered distally by thin red brown band, then thin silver gray line, then red brown, then two silver gray lines .................................................................................................................. callip h an e
34(32). VFW with orange-brown to red-brown discal cell bands parallel; basal area of cell

M2 mostly filled with silver gray scaling or with a well defined silver gray triangle; VHW discal cell bands separate and at least as far apart as width of cell bands ...................................................................................................................... 35









VFW with orange-brown discal cell bands converging posteriorly; basal area of cell M2 dark brown or reddish brown, or with faint, poorly defined paler gray scaling in anterior half only; VHW discal cell bands merged into a single band or nearly so .......................................................................................................... thoasa

35(34).Ventral surface marginal border of both wings almost entirely orange-brown or

red-brown, with very faint white dashes only in cell Cu2 and trace in cell Cul... 36 Ventral surface marginal border of both wings orange-brown or red-brown with clear white dashes in most cells ....................................................................... 37

36(35).VHW with elongate tomus, so that basal pair of black tornal spots are nearer

distal margin than white postdiscal band in cell Cu2; white postdiscal band in cells Rs and Sc+R1 bordered distally by thin dark brown line then paler yellowish brow n line ................................................................................................ falcipennis

VHW without elongate tornus, basal pair of black tornal spots are equidistant between distal margin and white postdiscal band in cell Cu2; white postdiscal band in cells Rs and Sc+R1 bordered distally by thin red brown line then thicker silver gray line ................................................................................................. gavina
37(35).VFW with pale orange subapical marking in cells M2 and MI not split by

continuous line of darker red-brown scaling near distal edge, marking split only in cell M2; DFW orange subapical marking may only just extend into cell M2, extend to terminate broadly at vein M3, or extend (rarely) as orange dashes into cell C u l .................................................................................................................. 38

VFW with pale orange subapical marking in cells M2 and M1 split by continuous line of darker red-brown scaling near distal edge; DFW orange subapical marking always extends broadly posteriorly at least as far as vein M3, often broadly into cell C u l ........................................................................................................ thessalia
38(37).Hindwing margin not produced into a short "tail" at vein Cul; DHW distal half

dark brown with slightly paler brown submarginal lines; DFW orange subapical









marking does not extend distally to reach inner paler brown submarginal line in cell M 1; not Jam aica .......................................................................................... 39

Hindwing margin produced into a short "tail" at vein Cu 1; DHW with thin whitish submarginal line; DFW orange subapical marking extends distally to reach inner paler brown submarginal line in cell M 1; Jamaica ........................................... abyla

39(38).DFW orange subapical marking extending to broadly border vein M3, or

extending into cell MI as a triangle of varying size with basal edge of marking straight or smoothly concave ............................................................................ 40

DFW orange subapical marking much broader in cell Ml than M2, extending into cell M2 only at distal edge as a triangular, posteriorly pointing dash, basal edge of marking not straight but kinked at vein M2 ................................................. iphiclus

40(39). DFW white postdiscal band tapering slightly from anal margin to cell M3; base of

eastern Andes S. Colombia-Peru ............................................................... iphicleola

DFW white postdiscal band not tapering from anal margin to cell M3; not base of eastern Andes S. Colombia-Peru ..................................................................... 41

41(40). DFW orange subapical marking broadly bordering vein M3 ............................ 42

DFW orange subapical marking extending into cell M2 only as a small triangle, often touching but not broadly bordering vein M3 .......................................... 43

42(41).DFW with distal edge of orange subapical marking in each of cells M2-R5

slightly concave in some or all cells; white postdiscal band of approximately equal width to orange subapical marking; VFW with edges of pale orange subapical marking sharply defined, particularly basal edge, with red brown scaling closely confined to veins Cu2 and Ml; VHW with first pale grayish line distal of white postdiscal band brighter whitish at vein M3 and closer to second distal pale grayish line ................................................................................................. iphicleola

DFW with distal edge of orange subapical marking in each of cells M2-R5 straight or slightly convex; white postdiscal band approximately half width of









orange subapical marking; VFW with edges of pale orange subapical marking poorly defined, typically with diffuse red brown scaling, with red brown scaling broadly along veins Cu2 and M1; VHW with first pale grayish line distal of white postdiscal band of uniform color and roughly parallel to second distal pale grayish line in cells Cu l-M 2 ..................................................................................... iphiclus

43(41).DFW with distal edge of orange subapical marking from vein MI to costa slightly

concave; VFW with edges of pale orange subapical marking sharply defined, particularly basal edge, with red brown scaling closely confined to veins Cu2 and Ml; VHW with first pale grayish line distal of white postdiscal band brighter whitish at vein M3 and closer to second distal pale grayish line ............... iphicleola

DFW with distal edge of orange subapical marking from vein M1 to costa straight or slightly convex; VFW with edges of pale orange subapical marking poorly defined, typically with diffuse red brown scaling, with red brown scaling broadly along veins Cu2 and Ml; VHW with first pale grayish line distal of white postdiscal band of uniform color and roughly parallel to second distal pale grayish line in cells Cul-M 2 ..................................................................................... iphiclus

44(26).DFW with orange subapical marking broadest in cells MI and R5, tapering to a

point in cell Cul ........................................................................................... coryneta

DFW with orange subapical marking broadest in cell M2, absent or present as indistinct orange scaling in cell M 3 ................................................................ federi

45(29).DFW orange subapical marking extending right across cell Ml to broadly end at

costa; dark red-brown line cutting pale VFW orange subapical marking in half is even in w idth ....................................................................................................... abia

DFW with only basal half of orange subapical marking ending at costa, distal half ending mid cell Ml; dark red-brown line cutting pale VFW orange subapical marking in half is much broader in cell M I than M2 ................................... epizygis

46(25). DFW with some orange or white markings ...................................................... 47









DFW entirely dark brown ..............................................................................justina

47(46).DHW with either white or partially white hindwing postdiscal markings, or if

entirely orange, isolated subapical spots are present on DFW .......................... 48

DHW with continuous, uniformly orange postdiscal band which extends from tornus to costa, then across DFW to costa, DFW subapical spots absent ...... ethelda 48(47).DFW with postdiscal band in cells Cu2 to M3 vertical, or, if inclined, orange

subapical spots are present at least in cells R5 and R4 or DHW is entirely brown ........................................................49

DFW with oblique orange band from tomus to costa, orange subapical spots entirely absent, DHW with central white spot .................................. leucophthalma

49(48). VFW discal cell silvery gray or whitish with clear brown to black basal streak,

cell bars one and two distinct and separated by reddish to orange brown bar, a pale band between second and third cell bars .......................................................... 50

VFW discal cell silvery gray with basal streak absent or very faint black scaling, cell bars one and two coalesced to form a black band, a round silver gray spot in anterior half of cell between second and third cell bars .............................. argentea

50(49). DHW postdiscal band or markings either partially or totally white .................. 51

DHW postdiscal band uniformly orange .......................................................... 69

51(50).DFW with orange subapical spot in cell R4, and/or two sets of orange postdiscal

to subapical markings in either or both of cells Ml and R5 ............................ 52

DFW with no orange subapical spot in cell R4, and only one orange postdiscal to subapical marking in cells M 1 and R5 ............................................................ 71

52(51).VHW with white postdiscal band bordered basally by reddish or orange brown

line, then whitish or gray in discal cell, then similarly colored reddish or orange brow n line .............................................................................................................. 53









VHW with white postdiscal band bordered basally by continuous black/dark brown line, then continuous orange band, both extending from near tomus to at least vein Sc+R 1 .................................................................................................... 73

53(52).DFW with postdiscal band variably orange and white to vein Cul, joined to

postdiscal and subapical orange markings ........................................................ 54

DFW with postdiscal band white to vein Cul, isolated from postdiscal and subapical orange markings .............................................................................. 74

54(53). VFW with inclined or vertical postdiscal band, if the latter, band is completely or

in its basal half very pale orange, not dissimilar in color from white VHW postdiscal band ................................................................................................... 55

VFW with vertical, matt orange postdiscal band, distinctly different in color from V HW postdiscal band ...................................................................................... cocala

55(54). VFW postdiscal band inclined or vertical, if the latter, very pale orange, similar in

color to white VHW postdiscal band, with darker vertical red brown lines variably present; silver gray submarginal markings not particularly pronounced in cells MI and R5, silver gray apical markings typically present in cells R4 and R3 ...... 56 VFW postdiscal band vertical, distinctly two-tone, with basal half very pale orange, similar in color to white VHW postdiscal band, distal half entirely orange brown; two prominent, bright whitish subapical spots in cells M1 and R5, no pale apical spots in cells R4 and R3 ........................................................................ cocala

56(55).DFW with postdiscal band always reaching anal margin, basal edge of postdiscal

band in cells Cu2 and Cul perpendicular to anal margin ................................. 57

DFW with postdiscal band usually not reaching anal margin, basal edge of postdiscal band in cells Cu2 and Cul inclined to anal margin .......................... 78

57(56). VFW with anterior dash of inner submarginal pair of silver gray dashes in cell

Cu2 of approximately equal width or slightly wider than posterior dash ...... 58









VFW with anterior dash of inner submarginal pair of silver gray dashes in cell Cu2 of approximately twice width of posterior dash; silver gray marginal dashes absent except in cell Cu2; ventral colors predominantly red-brown and graysilver ....................................................................................................................... 79

58(57).VFW with very pale orange postdiscal markings in cells Cul and M3 of equal or

differing widths, split by dark lines, either distinct or diffuse, each marking followed distally by one to two silver gray dashes; basal edge of pale postdiscal marking in cell Cu2 perpendicular to anal margin; pale silver gray marginal dashes/spots present in cell Cu2 ....................................................................... 59

VFW with very pale orange postdiscal markings in cells Cul and M3 of equal widths, not split by dark lines, each marking followed distally by only a single silver gray dash; basal edge of pale postdiscal marking in cell Cu2 inclined to anal margin; pale silver gray marginal dashes/spots absent in cell Cu2 ................... 80

59(58).VHW distal of white postdiscal band not entirely red-brown except for row of

close, paired silver dashes from costa to tomus, basally indented in cell M2; DFW with orange subapical spot in cell M1 absent to pronounced; postdiscal band in cell Cu2 white or orange, same thickness or narrower than band in cell Cul ....... 60 VHW distal of white postdiscal band entirely red-brown except for row of close, paired silver dashes from costa to tomus, basally indented in cell M2, and two white tomal dashes in cell Cu2; DFW with orange subapical spot in cell Ml absent or very faint orange scales; postdiscal band in cell Cu2 white and half thickness of band in cell Cul .......................................................................... m illeri

60(59).DFW with distal edge of postdiscal band in cell Cul straight or slightly convex;

VFW with pale postdiscal band usually distally displaced from postcellular bar, very rarely almost touching postcellular bar, in cell Ml; pale postdiscal markings in cells Cul and M3 ma or may not be split by vertical, straight, dark red-brown scaling .................................................................................................................... 6 1









DFW with distal edge of postdiscal band in cell Cul concave; VFW with pale postdiscal band almost touching postcellular bar in cell M1; pale postdiscal markings in cells Cul and M3 split by vertical, straight, dark red-brown scaling ....................................................................................................................... x im en a

61(60).DFW with distal edge of orange subapical spot in cell M1 distally displaced with

respect to distal edge of orange postdiscal band in cell M2 (or spot absent), spot clearly isolated from band; VFW with two long pale dashes, extending across cell, or single round spot, in cell Cul distal of pale postdiscal block ....................... 62
DFW with distal edge of orange subapical spot in cell Ml aligned with distal edge of orange postdiscal band in cell M2, spot fused or nearly so (vein M3 dark brown) with band; VFW with two short pale dashes, not extending across cell, in cell CuI distal of pale postdiscal block ..................................................... capucinus
62(61).VFW with pale postdiscal band either split by an even line, which cuts right

through band and ends at distal edge of band at vein Cu2, variably split, or not split ......................................................................................................................... 63

VFW with pale postdiscal band split by posteriorly narrowing line, starting in cell M2 and terminating within marking at vein Cu2 .................................... altamazona

63(62). VFW pale postdiscal band not completely divided in two by a red-brown line, or

line is slightly concave; DFW orange subapical spot in cell MI absent, smaller than, or of equivalent size to that in cell R5; VFW pale subapical spot in cell Ml not divided with darker red brown scaling; VHW with pale, thin silver gray line distal of white postdiscal band even throughout or much thicker whitish scaling in cells Rs and Sc+R l ............................................................................................ 64

VFW pale postdiscal band divided in two by distinct, vertical, straight red-brown line; DFW orange subapical spot in cell Ml pronounced and as large as that in cell R5 or larger; VFW pale, subapical spot in cell MI typically divided with









darker red brown scaling; VHW with pale, thin silver gray line distal of white postdiscal band even throughout .................................................................... boeotia

64(63).VFW with straight silver gray submarginal dash in cell M3 or marking absent;

VHW with paler lines and darker bands of variable color and width distal to white postdiscal band, all approximately parallel and evenly spaced; DHW with tornal orange spot of varying size in cell Cu2 typically present .................................. 65

VFW with basally pointing, "V"-shaped silver gray submarginal marking in cell M3; VHW with thin silver gray line distal to white postdiscal band, then dark red band constricted in cell M2, then line of close, paired silver-gray dashes basally displaced in cell M2, then marginal silver gray dashes; DHW with tornal orange spot in cell Cu2 absent ...................................................................................jordani

65(64).VFW with pale postdiscal marking divided into a basal block and two distal

dashes ..................................................................................................................... 66

VFW with pale postdiscal marking divided into a basal block and one round distal spot ......................................................................................................................... 8 1

66(65).VHW with first two pale gray lines distal of white postdiscal band separate

throughout w ing ................................................................................................. 67

VHW with first two pale gray lines distal of white postdiscal band fused to form a single large whitish spot in cell Sc+R1 .......................................................... pollina

67(66). VHW with first pale line distal of white postdiscal band of even width and color

throughout w ing ................................................................................................ 68

VHW with first pale line distal of white postdiscal band much thicker in cells Rs and Sc+R1 than in remainder of wing ............................................................... 86

68(67). VFW with submarginal silver gray dashes of approximately even size in each cell,

forming a line parallel to distal margin ............................................................ malea









VFW with submarginal silver gray dashes of varying size, that in M3 absent or much reduced compared with that in cell Cul, and basally displaced with respect to rem aining dashes .......................................................................................fabricia

69(50) VFW with line of silver gray submarginal dashes notably basally displaced in

cells Cul, M3 and M2, most displaced in M2; west of Andes ......................... 70

VFW with submarginal silver gray dashes forming a line parallel to distal margin; eastern M erida range in Venezuela .................................................................. malea

70(69).Hindwing elongate and triangular; VHW with second pale line distal of white

postdiscal band almost uniform in color, sometimes slightly paler in cells Rs and Sc+R1; DFW with basal edge of orange postdiscal band continuously curving through vein M 3 ........................................................................................... leuceria

Hindwing squarish; VHW with second pale line distal of white postdiscal band consisting of large, silvery, scalloped spots in cells Rs and Sc+R1 and much thinner and paler throughout remainder of wing; DFW with basal edge of orange postdiscal band displaced at vein M3 ..................................................... leucerioides

71(51). DFW orange postdiscal band with distal edge more distal in cell M2 than in cell

M 1 .......................................................................................................................... 72

DFW orange postdiscal band with distal edge more distal in cell Ml than in cell M 2 ................................................................................................................p lesaure

72(71) VFW postdiscal band matt orange brown; VFW distal ground color reddish

brown; pronounced silver white subapical spots in cells M l and R5, former larger, apical spots absent in cells R4 and R3 ............................................................ cocala
VFW postdiscal band white or very pale orange; VFW distal ground color dark brown; silver white subapical spots in cells M1 and R5, former sometimes smaller and often absent, apical spots present in cells R4 and R3 ............................. melona

73(52) DHW with postdiscal band entirely white; VFW with pale postdiscal markings in

cells M3 and M2 split in two by vertical orange-brown line .......................... shuara









DHW with postdiscal band orange, or tinged with orange, particularly at distal edge, in cells Rs and Sc+R1; VFW with undivided pale postdiscal marking in cell M3, that in M2 occasionally with very faint darker brown scaling through middle .......................................................................................................................... attica

74(53) VFW with single pale yellowish white spot between white postdiscal block in cell

Cul and inner, thick silver gray submarginal dash .......................................... 75

VFW with two pale dashes between white postdiscal block in cell Cul and inner, thick silver gray submarginal dash ................................................................... 76

75(74).VHW with spots of first row of pale whitish gray markings distal of white

postdiscal band largest in cells MI and Rs ....................................................... erotia

VHW with spots of first row of pale whitish gray markings distal of white postdiscal band even dashes in all cells, or slightly larger in cells Rs and Sc+RI ..................................................................................................................... m essan a

76(74) DFW with orange subapical spot in cell M2 larger than or equivalent in size to

that in cell Ml; distal edge of orange subapical spot in cell Ml aligned with distal edge of orange postdiscal band in cell M2, spot fused or nearly so (vein M3 may be dark brown) with band; VFW with silver submarginal marking in cell M3 absent or basally displaced with respect to remaining line of submarginal m arkings ................................................................................................................. 77

DFW with orange subapical spot in cell M2 absent or much smaller than that in cell MI; if present, orange subapical spot in cell MI isolated from orange postdiscal band in cell M2; VFW with silver submarginal markings forming a row parallel to distal m argin ............................................................................... m alea

77(76).DFW with orange subapical spot in cell R4 present; VFW with silver gray

submarginal marking in cell M3 present; forewing with cell M3 of approximately even width throughout, may be very slightly wider at base than at distal margin ..................................................................................................................... barn esia









DFW with orange subapical spot in cell R4 absent; VFW with silver gray submarginal marking in cell M3 absent; forewing with cell M3 noticeably wider near base than at distal m argin .......................................................................... diazi

78(56). VFW with distal edge of pale orange postdiscal band nearer distal margin in cell

Cu2 than in cell Cul; VHW with each dash of most distal submarginal row of silver gray dashes bordered basally by dark gray then another silver gray dash.zina VFW with distal edge of pale orange postdiscal band nearer distal margin in cell Cul than in cell Cu2; VHW with each dash of most distal submarginal row of silver gray dashes bordered basally by dark gray, then orange-brown .......... justina

79(57).VFW with pale orange postdiscal band cut by red-brown line in cells Cul and

M3; DFW with postdiscal band narrow at costa and cut by dark veins; DHW distal margin fringe with white scaling in each cell; VFW with silvery gray spot distal to postcellular bar in cell Ml; E. Andes ............................................. sichaeus

VFW with pale orange postdiscal band not cut by red-brown line in cells Cul and M3; DFW with postdiscal band broad at costa and not cut by dark veins; DHW distal margin fringe entirely black; VFW with no silvery gray spot distal to postcellular bar in cell Ml; Costa Rica-W. Andes .................................. hesterbergi

80(58).DHW white postdiscal band narrow, of almost even width throughout wing,

tapering towards tornus; VFW with silver gray submarginal marking in cell M3 "U "-shaped ................................................................................................... olynthia

DHW white postdiscal band of moderate width, tapering towards costa and tornus, broadest in cells M2 and Ml; VFW with silver gray submarginal marking in cell M 3 a straight, inclined dash .................................................................justina

81(65).VHW with spots forming first row of pale markings distal of white postdiscal

band of even width throughout wing, or largest in cells Ml and Rs, or in cells Rs and Sc+Rl, this row bordered distally by dark red-brown/brown band of even width or slightly broader in cell Ml than M2; VFW with black area between









white/pale orange postdiscal block in cell Cu2 and silver gray submarginal spots undivided ................................................................................................................ 82

VHW with spots forming first row of pale markings distal of white postdiscal band largest in cell M2, then M3, this row bordered distally by dark brown band much broader in cell Ml than M2; VFW with black area between white/pale orange postdiscal block in cell Cu2 and silver gray submarginal spots divided by vertical pale orange/orange dash ................................................................... delinita

82(81).VFW with silver gray apical spots in cells R4 and R3; VHW with four silver gray

dashes distal of white postdiscal band in cell Rs; pale orange postdiscal marking in cell M3 not divided by orange-brown line ................................................... 83

VFW within no silver gray apical spots in cells R4 and R3; VHW with three silver gray dashes and a pale red-brown dash distal of white postdiscal band in cell Rs; pale orange postdiscal marking in cell M3 typically divided in half by vertical orange-brown line .................................................................................... erym anthis

83(82).VFW with silver gray submarginal dash in cell M3 of similar size and shape to

that in cell Cul; VHW ground color almost uniform orange-brown to red-brown ...... -o o..... oo. . oo* .......... oo �........... o.o.. o... ............................................ 84
VFW with silver gray submarginal dash in cell M3 absent or much fainter and thinner compared to that in cell Cul; VHW ground color typically dark brown with red-brown to orange-brown band between first and second pale lines distal of white postdiscal band and orange-brown marginal ovals at each vein ...... thesprotia 84(83).VHW with spots of first row of pale whitish gray markings distal of white

postdiscal band even dashes in all cells, or slightly larger in cells Rs and Sc+Rl 85 VHW with spots of first row of pale whitish gray markings distal of white postdiscal band largest in cells M I and Rs ....................................................... erotia

85(84). VFW with first discal cell bar slightly inclined to costa, and second and fourth cell

bars concave; ventral pale lines distal of postdiscal band distinctly silvery gray;









VHW with first and second discal cell bars broadly spaced, noticeably paler in area between, equidistant between third discal cell bar and basal red-brown line ...................................................................................................................... p hylaca

VFW with discal cell bars approximately straight, particularly second cell bar, fourth cell bar slightly convex; ventral pale lines distal of postdiscal band silvery white; VHW with first and second discal cell bars close together, almost forming a single dark band, band closer to third discal cell bar than to basal red-brown line ..................................................................................................................... m essan a

86(67).DFW with orange in cell M3 filling base of cell; VFW with white postdiscal

block in cell Cu2 narrower than orange postdiscal block in this cell on DFW; pale orange postdiscal marking in cell M3 split by orange brown line near base and near distal margin; pale orange dashes in cell Cu 1 extend across entire cell; VHW with first two pale lines distal of white postdiscal band pale silver gray, broader in cells Rs and Sc+R 1 ...................................................................................... heraclea

DFW with orange in cell M3 not filling base of cell; VFW with width of white postdiscal block in cell Cu2 equal to width of orange postdiscal block in this cell on DFW; pale orange postdiscal marking in cell M3 entire except for some indistinct orange scaling near base; pale orange dashes in cell Cul only in anterior half of cell; VHW with first two pale lines distal of white postdiscal band pale gray, except white in cells Rs and Sc+Rl ..................................................... sp. nov.

87(1). DFW with some orange or white markings ...................................................... 88

DFW entirely dark brown ................................................................................... zina

88(87) DFW with postdiscal band vertical or inclined, of uniform color, not broken in

cell M 3 ................................................................................................................... 89

DFW postdiscal band vertical, broken in cell M3, a thin white band from anal margin to cell M3, a very broad orange band from cell Cul to costa ............ donysa









89(88). VHW with distal half varying shades of red/orange to brown, with some pale rows

of markings, no dark brown lines parallel to veins bisecting each cell; VFW discal cell with a single silvery white/silvery gray band opposite middle of cell Cul ..... 90 VHW with distal half almost entirely brown or yellowish brown, with dark brown lines parallel to veins bisecting each cell; VFW discal cell with a pair of silvery white spots opposite middle of cell Cul, an anterior and a posterior spot, separated by a black line ................................................................................... 99

90(89). DFW with postdiscal orange or white markings in cells M3 and M2 not filling cell

to distal m argin ................................................................................................. 91

DFW with postdiscal orange markings in cells M3 and M2 filling cell to distal m argin ......................................................................................................... stilesiana

91(90).DFW with only a single white or orange postdiscal to submarginal marking in

cells Cul and M 3 .............................................................................................. 92

DFW with three white postdiscal to submarginal markings in each of cells Cul and M 3 ........................................................................................................ dem ialba

92(91). DFW with vertical or inclined orange postdiscal band .................................... 93

DFW with oblique white postdiscal band ............................................................ 100

93(92).VFW pale silver gray area in distal half of discal cell undivided by a well-defined

darker line, rarely with indistinct, incomplete line of black scaling; VFW with or without pale postdiscal marking in cell Cu2 .................................................. 94

VFW pale silver gray area in distal half of discal cell completely divided by well defined, dark brown line; VFW with pale postdiscal marking in cell Cu2 .......... 101 94(93). VHW with silver gray postdiscal band absent, broken, or continuous and

extending throughout wing, bordered basally anterior of vein M3 by dark redbrown line then silver gray scaling, at least anterior of vein M2 ...................... 95









VHW with continuous silver gray postdiscal band extending throughout wing, bordered basally anterior of vein M3 by dark brown line then broad orange band .............................................................................................................................. 102

95(94). DFW with orange postdiscal markings just distal of discocellulars in cells Ml and

R5 at least twice as broad as orange subapical spots in those cells, or orange subapical spots absent ........................................................................................ 96

DFW with orange postdiscal markings just distal of discocellulars in cells Ml and R5 of approximately equal width or narrower than orange subapical spots in those cells, or postdiscal m arkings absent ..................................................................... 103

96(95).VFW with some submarginal silver gray dashes distal of pale orange postdiscal to

submarginal band in cells Cu2 to M2 ............................................................... 97

VFW with no submarginal silver gray dashes distal of pale orange postdiscal to submarginal band in cells Cu2 to M 2 .................................................................. 106

97(96). VHW with area between first and second discal cell bars whitish gray, with

variable amounts of reddish scaling at inner edges of cell bars ....................... 98

VHW with area between first and second discal cell bars orange brown ............ 109

98(97).DFW with distal edge of orange postdiscal band vertical and parallel to distal

m argin in cells Cul and M 3 .......................................................................... .ximena

DFW with distal edge of orange postdiscal band inclined, much closer to distal m argin in cell Cul than M 3 ............................................................................ boreas

99(89).DFW with orange subapical spots in cells M1 and R5 ............................. mesentina

DFW with subapical spots in cells MI and R5 white or absent ................... lycorias

100(92).VHW with postdiscal band uniformly silver white from costa to tomus; area

posterior to vein 3A silver white lined with orange-brown; W. Colombia ...................................................................................................................... eth elda
VHW with postdiscal band steely gray with white spot at costa in cell Sc+R1; area posterior to vein 3A orange; E. Andes to S. E. Brazil ........................... epione









101(93).DFW with vertical orange postdiscal band parallel to distal margin .............. tracta

DFW with oblique orange postdiscal band extending from costa to tomus ................................................................................................................ salm oneus

102(94).DFW postdiscal orange band oblique, extending from costa to tomus ........ ethelda

DFW postdiscal orange band vertical, extending from costa to anal margin ...................................................................................................................... m elon a

103(95).VHW with silver white postdiscal band from costa to tornus present ............... 104

VHW with silver white postdiscal band from costa to tomus absent, postdiscal area entirely red-brow n ....................................................................................... 110

104(103). VFW with pale postdiscal spots in cells M3 and M2 not split in half by darker

red-brow n line ................................................................................................. 105

VFW with pale postdiscal spots in cells M3 and M2 split in half by darker redbrow n line ..................................................................................................... salus

105(104).DFW with basal edge of postdiscal band from anal margin to cell M3 slightly

inclined to anal margin; VFW discal cell with thin, well defined dark brown basal streak; silver submarginal dash in cell M3 reduced or absent; two silvery gray marginal spots present in cell Cu2 ............................................... rothschildi

DFW with basal edge of postdiscal band from anal margin to cell M3 perpendicular to anal margin; VFW discal cell with broad, poorly defined orange basal streak; silver submarginal dash in cell M3 of similar size to remaining submarginal dashes; silvery gray marginal spots absent in cell Cu2 .................................................................................................................... levon a
106(96). DFW with distal edge of orange postdiscal band projecting distally in cell M2

....... '... . ...."....... �............ --"......... ... ,..... ............. .............................107
DFW with distal edge of orange postdiscal band indented basally in cell M2 ...................................................................................................... leucop hthalm a









107(106). VFW pale yellow postdiscal markings in cells M2-R5 close to postcellular bar

(distance from basal edge of markings to bar much less than width of m arkings) ......................................................................................................... 108

VFW pale yellow postdiscal markings in cells M2-R5 not very close to postcellular bar (distance from basal edge of markings to bar equal to width of m arkings) .................................................................................................... lam asi

108(107). VHW predominantly yellow; VFW with no white postdiscal spot in cell Cu2

.............................................................................................................. saundersii

VHW predominantly dark orange-brown; VFW with white postdiscal spot in cell C u2 ...................................................................................................... irm ina

109(97). VHW with continuous silver gray postdiscal band with even basal edge from

costa to cell Cu2 (band may be darkened anterior of vein M1) .................... salus

VHW with silver gray to whitish postdiscal spots, occasionally forming a band from costa to cell Cul, but not entering Cu2, always with uneven basal edge .�.. ������� o�� ���o�, ���.... � ................. o�..... ...... . .......................................zina

110(103). VFW with distal edge of pale orange marking in cell Cul nearer distal margin

than that of marking in cell Cu2 .................................................................justina
VFW with distal edge of pale orange marking in cell Cu2 nearer to distal margin than that of marking in cell Cul ......................................................... zina


Species groups

Given the general lack of resolution and weak support for nodes in the phylogeny derived in Chapter 3, due to the lack of morphological characters, I feel it would be premature to attempt to divide the entire genus into species groups. Nevertheless, I have indicated in Tables 2-5, with horizontal lines, groups which may be monophyletic or at least contain species likely to share morphological and biological traits, although the latter groups may still be paraphyletic. In addition, there are several strongly or relatively












strongly supported, probably monophyletic species groups which I refer to in the text, and these are listed and defined in Table 1, with the constituent species given in the cladogram (Fig. 250).






Table 1. Synapomorphies and other typical characters of species groups within Adelpha. Species included in each group are given in Chapter3, Fig. 250.


Group: Synapomorphies: A. serpa Male genitalia: valve lacks well developed clunicula; aedeagus has sclerotised, spiny pad on the dorsal surface of the vesica. Wing pattern: DFW discal cell has basal streak placed near costa, enclosing anteriorly a dense patch of reddish scales; basal streak in VHW and VFW discal cell absent; welldefined black line extending from base of VHW to tip of humeral vein.
A. alala Wing pattern: third discal cell bar present on VFW; dark lines bordering vein 3A on VHW greatly faded and reduced.

A. iphiclus Postdiscal series on VFW broaden anteriorly, being further from inner submarginal series in cell MI than cell M2.

A. phylaca Wing pattern: densely packed, pale whitish scales along base of vein Rs on DHW. Male genitalia: clunicula broad; valva with terminal teeth extending along ventral edge of valva; saccus large, almost same length as valva. Female genitalia: corpus bursae and ductus bursae large; sclerotised bands placed far apart. A. capucinus Subapical orange spot in cell MI on DFW aligned with distal edge of orange marking in cell M2; postdiscal series fused on VFW in cell M3 but separate in cell Cul. A. cocala Female genitalia: corpus bursae lacking sclerotised bands, rounded, typically with ductus bursae of approximately even width.


Other typical characters: Male genitalia: typically with poorly developed ventral and dorsal medial lobes. Female genitalia: bands on corpus bursae heavily sclerotised with coarse teeth. Immature stages: pupa often with shimmering silver or gold areas.





Wing pattern: ventral colors all faded; outer submarginal series on VHW absent. Male genitalia: valva with few, large terminal spines. Immature stages: larval scoli greatly reduced. Wing pattern: postdiscal and submarginal series on VFW typically visible throughout; upper postdiscal band on DFW absent.

Wing pattern: postdiscal series fused in cell Cul on VFW and fused to postdiscal band in cell M3; Male genitalia: valva with pronounced dorsal and ventral medial lobes. Immature stages: pupa with prominent, hooked dorsal projection on thorax; final instar larva pale with dark area laterally on thorax-A2.

Male genitalia: valva broad throughout, not tapering, squared at tip with terminal spines; clunicula long; saccus relatively short compared to valva.


Ecological characters: Typically geographically widespread; often found perching in large clearings near rivers; relatively little geographic differentiation of wing pattern; hostplants diverse, not Rubiaceae.






Occur in high elevation cloud forests; larval hostplants Caprifoliaceae; known larvae construct shelters from hostplant leaves.

Especially diverse in southeastern Brazil; often in secondary growth areas; relatively little geographic differentiation of wing pattern; hostplants all Rubiaceae. Usally occur in lowland primary forest; perch along forest edges or in canopy; hostplants diverse, not in Rubiaceae.






Typically confined to lowland forest.


Generally small species, orange tomal Uusually occurring in premontane spot on DHW often absent, thorax cloud forest habitats, many species and abdomen relatively narrow. with restricted ranges; strongly mimetic, with sharp regional
differentiation of dorsal wine nattem.









Biology

Habitats, Phenology and Rarity

Adelpha species occur in a very wide variety of forested life zones, and microhabitats within life zones. They have been recorded from sea level to 3000m. While higher species numbers are found in wet forests, certain Adelpha are restricted to drier life zones, ranging from semi deciduous to deciduous forest (A. fessonia), xeric Acacia scrub (A. fessonia lapitha), and mixed pine-hardwood forest (A. gelania). A large number of species are also confined to cloud forest habitats in the Andes and other mountainous areas.

There is relatively little information available on the adult phenology of Adelpha species. My research in Ecuador indicates that species and individual abundance is highest during the wet season and early dry season. These observations are in contrast to those of DeVries (1987), who states that most individuals are more abundant in the dry season. However, through the use of traps I have found several species to be abundant in Costa Rica during the wet season (Hall & Willmott, 1993), and believe that species appear to be more abundant in the dry season since there are less collecting days lost due to rain and males more frequently leave the forest to puddle along roads or rivers. DeVries et al. (1997), in a trapping study, report in eastern Ecuador that nymphalid species and individual abundance in eastern Ecuador is highest in the wet season.

In many instances, individuals were encountered in numbers over a period of a couple of days, but never seen again despite repeated visits to a site. This was the case with A. hesterbergi, A. lamasi and A. levona in northwestern Ecuador. This suggests the possibility that pupal eclosion may perhaps be triggered by some environmental cue, such as heavy rains. Aiello (pers. comm.) reports to me a recent outbreak of A. iphiclus in Panama, which she believes to be linked to an unusually dry period causing high mortality in predators and parasitoids of the immature stages, which usually control population numbers.









The majority of Adelpha species seem to be present throughout the year, even if abundance is seasonal, but A. bredowii eulalia has been reported to have two distinct broods in the U.S.A., from May to June, then from August to October (Ferris & Brown, 1982; Cary, 1994). The larvae are in diapause during the winter (Ferris & Brown, 1982). It is very probable that species in the more strongly seasonal, more temperate regions of eastern and southern Brazil to Argentina, Paraguay and Uruguay, may have similar life cycles. There are, however, no published observations and museum specimen data are too few to draw any conclusions.

While several Adelpha species are common to abundant, many are infrequently encountered and 14 species are represented by fewer than 25 specimens in all of the collections examined (see Fig. 4). One species, A. herbita, is known only from the single female specimen on which the original illustration is based, and the whereabouts of this specimen, if it still exists, is currently unknown. For almost all species there are more known specimens of males than females, which overall outnumber females in collections by approximately 4:1. This probably reflects the propensity for males to occur in groups (see Adult ecology below) and to be attracted to a wider variety of nutrient sources. The rarity of many species is due to their small geographic range and/or confinement to cloud forest habitats, where they may be locally not uncommon, and it is this life zone that most recently described species inhabit (Beutelspacher, 1976; Steinhauser & Miller, 1977; DeVries & Chac6n, 1982; Willmott & Hall, 1995, 1999). Such habitats have historically been poorly collected due to problems of access from the steepness of the terrain, and the difficulties of collecting in almost perpetual rain and cloud. They continue to provide new species in many other groups of butterflies (Hall & Willmott, 1995; Willmott et al., in prep.) and their Adelpha fauna still remains relatively poorly known. Certain species, such as A. nea, are extremely widespread but very rare throughout their range. This species is known from eastern Ecuador from two specimens, one captured at Chichicorrumi in 1993, but despite numerous subsequent visits to this locality it has not









been collected there again. At the other end of the spectrum are species that are everywhere common and typically encountered in numbers. Such is the case with Adelpha cytherea, which alone contributed almost 8% of the specimens examined.

The perceived rarity of species, and the number of specimens in collections, presumably almost never reflects their true abundance in the field. The use of traps in Ecuador has demonstrated certain species, such as A. pollina, A. heraclea and A. boreas, to be widespread and common in many localities, despite the low numbers of specimens in collections. These species are never encountered outside of traps and presumably spend their time in the forest canopy.


Iiiil~iil ,11,,,,1


Number of specimens/species

Fig. 4. Abundance of Adelpha species in each class of number of specimens recorded.









Hostplants

Their exist hostplant records for at least 39 species of Adelpha, comprising 19 plant families, 61 genera and 94 species (see Table 2), and the genus has one of the widest host breadths of any nymphalid genus (Ackery, 1988). The most important single contributors to our knowledge of Adelpha hostplants are MUller (1886), Moss (1933) and Aiello (1984), with a number of records by Janzen, Haber, Marquis and DeVries (L_: DeVries, 1986). Many of the identifications of Adelpha species in the hostplant literature are unreliable or erroneous, and I have tried to correct as many of these misidentifications as possible, based on descriptions of the adult, the locality and voucher material. The latter is sadly lacking for the majority of species, rendering a number of records questionable to almost worthless.

Many Adelpha appear to be highly polyphagous; A. serpa, for example, has been recorded from 13 genera of seven plant families. Certain families, such as Moraceae, Urticaceae and Tiliaceae, are characteristic of species from phylogenetically distant groups, and it is unclear what factor of hostplant biology mediates the host choice. The family Rubiaceae is responsible for the great majority of records and appears throughout the genus, with rubiaceous feeders generally being unrecorded on other families.

Despite there being hostplant records for almost 50% of the species in the genus, a number are unreliable and a great many are concentrated into two species groups, the A. serpa and A. phylaca group. No doubt this is due to the larvae feeding on plants which grow in disturbed areas in lowland regions, habitats most frequented by biologists. It is thus not possible at the present to draw conclusions about broad evolution of hostplant choice within the genus. It is, however, of possible evolutionary interest that the most primitive group in the genus, the A. alala group, has been recorded predominantly on the family Caprifoliaceae; this family is otherwise unknown in the genus, but is a common hostplant of other Asian and European limenitidines (Ackery, 1988) which may be close relatives of Adelpha.










Table 2. Recorded hostplants for Adelpha taxa.


Species: Subspecies: Records Host genus: Host species: Host family: Locality: Source: Reported as: bredowii californica Quercus spp. USA: Cal Dyar (1891) Heterochroa


chrysolepis Fagaceae


chrysolepis
spp.

agrifolia douglasii kelloggi virginiana chrysolepis
sp.
sp.
arizonica emoryi gambeli vaccinifolia


gambelii agrifolia wizlizenii turbinella chrysophylla


USA: Cal Comstock (1927); Tietz (1972), Howe (1975);
Orsak (1977), Domfeld (1980), Christensen
(1981), Scott (1986) Mexico Brown et al. (1992) USA: Cal Comstock (1927); Ehrlich & Ehrlich (1961);
Ackery (1988)
Mexico Brown et al. (1992) USA Tietz (1972); Scott (1986)


Quercus


Quercus Quercus Quercus Quercus Quercus Quercus Quercus Quercus Quercus Quercus Quercus Quercus
Quercus


Tietz (1972); Scott (1986) Tietz (1972) Emmel & Emmel (1973) Ferris & Brown (1980) Beutelspacher (1980) Bailowitz & Brock (1991) N. McFarland (in: Bailowitz & Brock, 1991) Harry (1994) Scott (1986) Scott (1986) Scott (1986) Scott (1986) Scott (1986) Scott (1986)


USA USA USA: Cal USA
Mexico USA: Ariz USA: Ariz USA: Ut USA USA
USA USA USA USA


Quercus Quercus Quercus Quercus Castanopsis


eulalia


californica/ eulalia


californica Heterochroa californica


Heterochroa b. californica Heterochroa b. californica Heterochroa b. californica









Limenitis bredowii Limenitis bredowii Limenitis bredowii Limenitis bredowii Limenitis bredowii Limenitis bredowii


*paraena paraena Miconia minutiflora Melastornataceae Brazil: Pa Moss (1933); Aiello (1984), Ackery (1988) serpa Qualea multiflora Vochysiaceae Brazil: DF Diniz & Morais (1997)










Table 2. Continued.
Species: Subspecies: Records Host genus: Host species: Host family: Locality: Source: Reported as:
* lecromil I Combretum decandrum Combretaceae Panama Aiello (1984); Neild (1996) nr. paraena
massilia
*radiata aiellae Cespedezia spathulata Ochnaceae Colombia Constantino (1998) serpa serpa serpa Melastoma sp. Melastomataceae Brazil Hoffmann (1936) serpa
(?) Miconicae - Brazil: SC MOller (1886); Aiello (1984), Ackery (1988) serpa
(?) Miconia sp. Brazil Brown (1992) serpa
* diadochus Isertia longiflora Rubiaceae Brazil Moss (1933); Aiello (1984), Ackery (1988), paraena Neild (1996)
* Remiqia amazonica Brazil: Pa Moss (1933); Aiello (1984), Ackery (1988), paraena Neild (1996)
* celerio 2 Ochroma pyramidale Bombacaceae Panama Coley & Aiello (in: Aiello, 1984); DeVries celerio (1987), Ackery (1988), Neild (1996)
* Urera sp. Urticaceae Costa Rica DeVries (in: Aiello, 1984); DeVries (1986, celerio 1987), Ackery (1988), Neild (1996)
* Myriocarpa longipes Costa Rica DeVries, specimen data; DeVries (1987), celerio Neild (1996)
Heliocarpus americanus Tiliaceae Venezuela A. M. Orellana (pers. comm.); Neild (1996) celerio celerio
* Melastomataceae El Salvador Specimen data (AME) leg. S. Steinhauser
(?) Conostegia xalapensis Melastomataceae Mexico: Jal. Comstock & Vizquez (1961); Aiello (1984), celerio Ackery (1988)
(?) Miconia sp. Mexico: Jal. Comstock & Vizquez (1961); Aiello (1984) celerio
(?) Miconia argentea Costa Rica Mallet (in: Aiello, 1984); DeVries (1986), celerio Ackery (1988)
(?) 4 Miconia argentea Costa Rica Janzen & Hallwachs (pers. comm.) celerio
(?) I Cecropia peltala Moraceae Panama Coley (in: Aiello, 1984); Ackery (1988) celerio seriphia pione Cavendishia sp. Ericaceae Venezuela A. H. Orellana (pers. comm.) hyas hyas Casearia sylvestris Flacourtiaceae Uruguay Biezanko et a. (1966) lex paraguariensis Aquifoliaceae Brazil: SP Silva et a. (1968); Aiello (1984), Ackery serpa hyas (1988: as serpa), Brown (1992)
*alala completa Viburnum tinoides Caprifoliaceae Venezuela Otero & Aiello (1996) alala










Table 2. Continued.
Species: Subspecies: Records Host genus: Host species: Host family: Locality: Source: Reported as:
*tracta I Rhondeletia sp. Rubiaceae Costa Rica Haber, specimen in the PJD (reported as "Rubiaceae" in: Aiello, 1984); DeVries (1986,
1987), Ackery (1988)
I Rubiaceae Costa Rica Chac6n (in: Aiello, 1984); DeVries (1986, 1987), Ackery (1988)
many Viburnum costaricanum Caprifoliaceae Costa Rica Haber (pers. comm.) fessonia fessonia I Randia echinocarpa Rubiaceae Costa Rica Janzen (in: Aiello, 1984); DeVries (1986), Ackery (1988), Neild (1996)
1 Randia karstenii Costa Rica Janzen (in: Aiello, 1984); DeVries (1986, 1987); Ackery (1988), Neild (1996)
Randia sp. El Salvador Scott (1986) Limenitisfessonia Randia armata Costa Rica Dyer (1997)
9 Randia aculeata Costa Rica Janzen & Hallwachs (pers. comm.) 9 Randia monantha Costa Rica Janzen & Hallwachs (pets. comm.) 15 Randia rhagocarpa USA: Tex Specimen data, FSCA, leg. R. W. Bosese ?? probably Doxocopa sp. Celtis lindheimeri USA: Tex Scott (1986); Neck (1996) Limenitisfessonia ?? probably Doxocopa sp. Celtis laevigata Ulmaceae USA: Tex Neck (1996)
(?) calliphane flex paraguariensis Aquifoliaceae Argentina Hayward (1940a, 1969b)
(?) Vitex montevidensis Verbenaceae Brazil Silva et al. (1968); Aiello (1984), Ackery (1987)
(?) Vitex sp. Brazil: RJ Zikin & Zikin (1968); Brown (1992) mythra Bathysa nr. barbinervis Rubiaceae Brazil Miller (1886); Aiello (1984), Ackery (1988), Brown (1992)
Rubus sp. Rosaceae ? Fountaine, unpubl. watercolors (in: Ackery, 1988)
basiloides 7 Alibertia edulis Rubiaceae Panama Aiello (1984); Neild (1996) 30 Alibertia edulis Costa Rica Janzen (in: DeVries, 1986); Ackery (1988), Janzen & Hallwachs (pers. comm.)
5 Amaioua corymbosa Panama Aiello (1984); Ackery (1988), Neild (1996) 3 Bertiera guianensis Panama Aiello (1984); Ackery (1988), Neild (1996) Ixora nicaraguensis Panama Mallet (in: Aiello, 1984)









Table 2. Continued.
Species: Subspecies: Records Host genus: Host species: Host family: Locality: Source: Reported as:
* Ixora nicaraguensis Costa Rica Marquis (in: DeVries, 1986, 1987); Ackery (1988), Neild (1996)
8 Ixora floribunda Costa Rica Janzen & Hallwachs (pers. comm.) Faramea sp. Costa Rica DeVries (1987) 45 Lindenia rivalis Costa Rica Janzen & Hallwachs (pers. comm.) 5 Randia aculeata Costa Rica Janzen & Hallwachs (pers. comm.) plesaure plesaure Bathysa? sp. Rubiaceae Brazil: SC Mlller (1886); Aiello (1984), Ackery (1988), Brown (1992), Neild (1996)
- Rubiaceae Brazil: SC Mlller (1886)
* phliassa Alibertia edulis Brazil: Pa Moss (1933); Aiello (1984), Ackery (1988), phliassa Neild (1996)
* Berteira guianensis Brazil: Pa Moss (1933); Aiello (1984), Ackery (1988), phliassa Neild (1996)
thoasa gerona Wlex paraguariensis Aquifoliaceae Argentina Hayward (1969)
*iphiclus iphiclus Uncaria guianensis Rubiaceae Brazil: Pa Moss (1933: as Ourouparia); Aiello (1984), iphicla Ackery (1988), Neild (1996)
ephesa Bathysa sp. Brazil Silva et at. (1968); Aiello (1984), Ackery ephicla ephesa (1988)
iphicleola iphimedia Calycophyllum sp. Rubiaceae Cuba Dewitz (1879); Aiello (1984) Heterochroa basilea
Calycophyllum candidissimum Cuba Riley (1975); Alayo & Hemrndez (1981), iphicla Aiello (1984), Ackery (1988), Neild (1996)
Calycophyllum candidissimum Cuba Bruner et al. (1975) iphicla Calcyophyllum candidissimum Cuba Smith el al. (1994) iphicla iphiclus Schoenleinia sp. Rubiaceae Neotropics Moller (1878) basilea /iphicleola
Calycophyllum sp. Brazil Moller (1886); Ackery (1988) basilea 2 Calycophyllum candidissimum Panama Aiello (1984); DeVries (1986), Neild (1996) iphicla 82 Calycophyllum candidissimum Costa Rica Janzen (in: Aiello, 1984); DeVries (1986, iphicla iphicleola 1987), Neild (1996), Janzen & Hallwachs
(pers. comm.)
Bathysa nr. barbinervis Brazil: SC Moller (1886); Aiello (1984), Brown (1992), iphicla Neild (1996)










Table 2. Continued.
Species: Subspecies: Records Host genus: Host species: Host family: Locality: Source: Reported as:
2 Uncaria tomentosa Panama Aiello (1984); DeVries (1987), Ackery iphicla (1988), Neild (1996)
Gonzalaguina spicata Trinidad Barcant (1970); Aiello (1984), Ackery (1988), iphicla Riley (1975), Smith et al. (1994)
4 Alseis blackiana Panama Aiello (1984); Ackery (1988), Neild (1996) iphicla 3 Antirrhoea trichantha Panama Aiello (1984); Ackery (1988), Neild (1996) iphiclus Antirrhoea sp. Panama Orellana (pers. comm.) iphicla I Isertia haenkeana Panama Aiello (1984); DeVries (1987), Ackery iphicla (1988), Neild (1996)
2 Rondeletia panamensis Panama Aiello (1984); Ackery (1988) iphicla 6 Guettarda macrosperma Costa Rica Janzen & Hallwachs (pers. comm.) iphiclus Sipania sp. ? Fountaine, unpub. watercolours (in: Ackery, iphicla 1988)
abyla Guettarda argentea Rubiaceae Jamaica Milner & Turner (in: Smith et al., 1994) Mussaendra treutleri (exot.) Jamaica Blake (in: Ellwood & Harvey, 1990) ethelda sophax Sabicea aspera Rubiaceae Costa Rica DeVries (in: Aiello, 1984); DeVries (1986, zalmona zalmona 1987), Ackery (1988), Neild (1996)
*melona leucocoma Malanea sp. Rubiaceae Brazil: Pa Moss (1933); Aiello (1984), Ackery (1988), melona Neild (1996)
syma Rubus fructicosus Rosaceae Brazil Maller (1886); Hayward (1 969b), Aiello (1984), Ackery (1988)
Rubus fructicosus Uruguay Biezanko el al. (1966) Rubus fructicosus Brazil: Pr Brown (1992) (probably distinct record) ? Rubus sp. ("zarza") Argentina Hayward (1931) Rubus sp. Brazil Hoffmann (1936, 1937) Blackberry Brazil Hoffmann (1936) Sarandy Rubiaceae Brazil: RS D'Almeida (1928) Cephalanthus glabratus Brasil: RS Biezanko (1949); Hayward (1969, 1973) Cephalanthus sarandi (= syn. Brazil Silva et al. (1968); Aiello (1984), Ackery of above?) (1988)
*cytherea cytherea Sabicea aspera Rubiaceae Brazil Moss (1933); Aiello (1984), Ackery (1988) cytherea










Table 2. Continued.
Species: Subspecies: Records Host genus: Host species: Host family: Locality: Source: Reported as:
* marcia Sabicea villosa Costa Rica DeVries (in: Aiello, 1984); DeVries (1986, 1987), Ackery (1988), Neild (1996)
Sabicea villosa Costa Rica Mallet (in: DeVries, 1986)
2 Sabicea villosa Costa Rica Janzen & Hallwachs (pers. comm.) daguana 1 Sabicea panamensis Panama Aiello (1984); Ackery (1988), Neild (1996) cytherea marcia 2 Sabicea villosa Panama Aiello (1984) cytherea marcia
*viola pseudococala Sabicea aspera Rubiaceae Brazil: Pa Moss (1933); Aiello (1984), Ackery (1988), pseudococala Neild (1996)
? viola Sabicea sp. Brazil: SC Mlller (1886); Aiello (1984) sp. .r, cocala salmoneus salmonides 2 Sabicea panamensis Rubiaceae Panama Aiello (1984); DeVries (1987), Neild (1996) salmoneus demialba Rondeletia sp. Rubiaceae Costa Rica Haber (in: Aiello, 1984); DeVries (1987), Ackery (1988)
Rhondeletia buddleioides Costa Rica Haber (pers. comm.) Gonzalaguina rosea Costa Rica Haber (pers. comn.) Rondeletia brenesii Costa Rica Haber (pers. comm.) epizygis Vitex sp. Verbenaceae Brazil: RI Zikin & Zikfin (1968), Brown (1992) Vitex montevidensis Brazil Silva eta. (1968); Aiello (1984), Ackery (1988)
*capucinus capucinus ? False cacao not Malpighiaceae Brazil: Pa Moss (1933); Aiello (1984), Neild (1996) thesprotia ? velia Tetrapterys sp. Malpighiaceae Bracil: SC Moller (1886); Aiello (1984), Ackery (1988), erotia var.
Neild (1996)
? abia Vitex sp. Verbenaceae Brazil: SC Moller (1886); Aiello (1984), Ackery (1988), abia Brown (1992)
*heraclea heraclea Vitex triflora Verbenaceae Brazil Moss (1933); Aiello (1984), Ackery (1988) jordani
* 2 Vitex cooperi Costa Rica DeVries (1986, 1987); Neild (1996) Piper arieianum Piperaceae Costa Rica Marquis (in: DeVries, 1986)
(?) erotia erotia (f. lerna) Sligmaphyllon sp. Malpighiaceae Costa Rica DeVries (pers. comm.) lerna phylaca pseudaethalia 4 Cecropia obtusifolia Moraceae Panama Aieilo (1984); Ackery (1988), Aiello (1991), phylaca aethalia Neild (1996)










Table 2. Continued. Species: Subspecies:


9

9

*messana


*thesprotia


messana


*mesenhna


Host family:


Records Host genus:

2 Cecropia I Cecropia
Luehea

Cecropia
Luehea
Cecropia Bombax

Cecropia

Pourouma

Cecropia Cecropia
Chodanthus
Malanea

Pourouma


Host species: peltata

sp.
seemannii

sp.
seemanii peltata munguba

sp.

sp.

sp.
adenopus
splendens
sp.

sp.


Locality:

Panama Panama Costa Rica Costa Rica Panama Costa Rica Brazil: Pa Brazil: Pa Brazil: Pa Paraguay Paraguay Paraguay
captivity, Brazil: Pa Brazil: Pa


lycorias lycorias Cecropia pachystachia Moraceae Brazil: SC Moller (1886); Silva et a. (1968), Hayward isis (1969b, 1973), Aiello (1984), Ackery (1988), Brown (1992)
Coussapoa schottii Brazil: SC Mailer (1886); Silva et a (1968), Hayward isis (1969b), Aiello (1984), Ackery (1988), Brown (1992)
Pourouma acutifora Brazil: SC Moller (1886); Silva et al. (1968), Hayward isis (1969b), Aiello (1984), Ackery (1988), Brown (1992)
lara - "Cecropiaceae" Trinidad Cock (in: Aiello, 1984); Ackery (1988), Neild lara (1996)
Cecropia peltata Venezuela Orellana (pers. comm.); Neild (1996) lara


Source: Repo
Coley & Aiello (in: Aiello, 1984); DeVries phyla (1987), Ackery (1988), Neild (1996) Aiello (1984) phyla Mallet (in: Aiello, 1984); DeVries (1986a, boeot 1987), Ackery (1988), Neild (1996) DeVries (1987); Neild (1996) boeol Aiello (1991); Neild (1996) ixia 1 DeVries (1987); Neild (1996) phyla Moss (1933); Aiello (1984), Ackery (1988), delph Neild (1996)
Moss (1933); Aiello (1984), Ackery (1988), delph Neild (1996)
Moss (1933); Aiello (1984), Ackery (1988), delph Neild (1996)
Jorgensen (1934) jurua Jorgensen (1924) jurua Jorgensen (1924) jurua Moss (1933) delpi

Moss (1933); Aiello (1984), Ackery (1988), Neild (1 996)


rted as:

ca aethalia 'ca aethalia ia oberthuri ia boeotia
eucas rca
icola ricola iicola ha
na na
hicola


Tiliaceae Moraceae Tiliaceae Moraceae Bombacaceae Moraceae


Rubiaceae Moraceae










Table 2. Continued.
Species: Subspecies: Records Host genus: Host species: Host family: Locality: Source: Reported as:
wallisii Cecropia peltata Colombia Constantino (1 997b, 1998) lara wallisii
melanthe Cecropia sp. Costa Rica DeVries (in: Aiello, 1984); DeVries (1986, melanthe 1987), Ackery (1988)
1 Cecropia peltata Costa Rica Janzen & Hallwachs (pers. comm.) Trema micrantha Ulmaceae Costa Rica DeVries (in: Aiello, 1984), DeVries (1986, melanthe 1987), Ackery (1988)
1 Trema micrantha Panama Aiello (1984); Aiello (1991) melanthe 23 Trema micrantha Costa Rica Janzen & Hallwachs (pers. comm.) melanthe Urera sp. (x2) Urticaceae Costa Rica DeVries (in: Aiello, 1984); DeVries (1986, melanthe 1987), Ackery (1988)
* Myriocarpa longipes Costa Rica DeVries (1986, 1987); Ackery (1988) melanthe leuceria leuceria - Rubiaceae Costa Rica DeVries (pers. comm.) cocala lorzae I Calycophyllum candidissimum Rubiaceae Panama Aiello (1984); Neild (1996) cocala 2 Calycophyllum candidissimum Costa Rica Janzen (in: Aiello, 1984); DeVries (1986, cocala 1987), Ackery (1988), Janzen & Hallwachs
(pers. comm.)
I Chomelia psilocarpa Panama Aiello (1984); DeVries (1987), Ackery cocala (1988), Neild (1996)
1 Uncaria tomentosa Panama Aiello (1984); DeVries (1987), Ackery cocala (1988), Neild (1996)
1 Warscewiczia coccinea Panama Aiello (1984), Ackery (1988), Neild (1996) cocala Pentagonia macrophylla Costa Rica DeVries (in: Aiello, 1984); DeVries (1986, cocala 1987), Ackery (1988), Neild (1996)
* 2 Pentagonia maxoni Costa Rica Specimen data in the PJD
* Psychotria sp. Costa Rica Marquis (in: Aiello, 1984); DeVries (1986, cocala 1987), Ackery (1988)
Genipa sp. Costa Rica DeVries (1987); Neild (1996) cocala
* cocala Malanea sp. Brazil: Pa Moss (1933); Aiello (1984), Ackery (1987), cocala Neild (1996)
* Emmotum nitens ? Icacinaceae Brazil: Pa Moss (1933); Aiello (1984), Ackery (1987), cocala Neild (1996)
(?) didia/caninia Rubiaceae Brazil: SP Brown (1992) cocala









Table 2. Continued.
Species: Subspecies: Records Host genus: Host species: Host family: Locality: Source: Reported as: ?? almost certainly A. viola Sabicea Rubiaceae ? Fountaine, unpubl. watercolors (in: Ackery, cocala 1988)
leucophthalma leucophthalma Pentagonia wendlandia Rubiaceae Costa Rica Young (1975); Ackery (1988)
- Rubiaceae Costa Rica DeVries (in: Aiello, 1984); DeVries (1986)
- Rubiaceae Panama Aiello (1984)
Pentagonia spp. Colombia Andrade (1994)
(?) boreas Satyria sp. Ericaceae Costa Rica Marquis (in: Aiello, 1984); DeVries (1986, 1987), Ackery (1988)
Satyria sp. Colombia Constantino (1 997b)
(?) Chomelia bispinosa Rubiaceae Costa Rica DeVries (in: Aieilo, 1984); DeVries (1986), Ackery (1988)
zina zina Alt. lvd. plant unknown Panama Aiello (1984) justinajustina

?? - identification very dubious; ? - identification uncertain, either original taxon name reported is incorrect, being either a combination of subspecific and specific epithets of different species, or a name of a taxon which does not occur in the area of the record, or vague information only is given as to the taxon; (?) - identification is slightly uncertain since the taxon is very similar to other sympatric taxa and usually misidentified, but the taxon to which the record is assigned here is either much more common than similar sympatric taxa, or there is relatively little reason to regard it as misidentified; * voucher specimen or photograph examined. References followed by a ";" are deemed to be the original source. In the absence of a specific statement to the contrary, I conservatively regard later records as citations of previous records; e.g. many records in DeVries (1987) lack information on the source of the record, and while many may be valid new records in Costa Rica, since it is not possible to ascertain this I have listed them as citations of earlier records. Single lines separate species, double lines separate species groups.









Immature Stages

The pioneering work of Muller (1886) in southeastern Brazil set the stage for studies of the immature life histories of Adelpha butterflies (see Table 3). Miller reared, described and figured certain immature stages for 7 species of Adelpha, and his descriptions still remain some of the most detailed published to date. Moss (1927) reared 11 species within 6 months of his arrival in Par, Brazil, and provided the next major contribution to our knowledge (Moss, 1933), followed by Aiello (1984), and both of these authors figured larvae and pupae. Aiello (1984) summarised almost all of the published information and her paper remains the most important work to date on Adelpha early stages. There are a number of other reports of Adelpha early stages in the literature, but all suffer from the same problems of identification as for hostplant records, and the vast majority offer no illustrations, only brief descriptions from which taxonomic inferences can seldom be made.

Oviposition. Females typically seek oviposition sites throughout the late morning and middle of the day, and the majority of Adelpha hostplants selected by females occur in recent openings in forest, and along the edges of forest and rivers. However, such hostplants are probably only those most easily located by the biologist, and it is highly probable that many species oviposit in the canopy. The female of A. iphiclus alights upon a leaf and walks backwards, searching for a suitable oviposition site with the tip of the abdomen. After oviposition the female takes flight, but may land on the same leaf and deposit further eggs. Eggs are deposited beneath the surface of leaves by the abdomen being curled under from the leaf upperside (Aiello, 1984). Reports of egg placement are rare in the literature and summarised by Aiello (1984). The majority of species deposit a single egg at the tip of a leaf on the upperside, while certain species typically oviposit next to a damaged portion of leaf (Aiello, 1984; DeVries, 1986). Up to four eggs may be deposited on a single leaf (Aiello, 1984), but often the plant chosen is a seedling and only a single egg is laid per plant.









Eggs. The eggs of all Adelpha species known to date are typical of the tribe, having hexagonal, concave facets with setae at the interstices (see Aiello, 1984).

First instar larvae. There are almost no detailed descriptions or figures of first instar larvae, which according to Aiello (1984) are all identical in form and some shade of brown or gray. The head bears setae but none of the chalazae of later instars, and only pale bumps on the body where future scoli are situated; I use the terminology of Petersen (1962) and Aiello (1984), and refer to thr first three thoracic segments as T1-T3, and remaining abdominal segments as Al-A10. After their first meal the larvae take on the color of their hostplant, though the head remains brown.

Second to fourth instar larvae. From the second instar on the head is ornamented with chalazae and numerous round, tiny pits, which may be of contrasting color to the head capsule. The body begins to develop scoli, arranged in three rows, subdorsal, supraspiracular and subspiracular. Typically the subdorsal scoli on T2 & 3 and A2, 7 & 8 are slightly larger than others. Some species undergo color and pattern changes, while in certain species, such as A. basiloides, the color depends on the hostplant (Aiello, 1984).

Fifth instar larvae. Fifth instar larvae are diverse in form, color and pattern. All species have dense chalazae around the head capsule, the size of which may vary interspecifically (Aiello, 1984). The head capsule may also have darker vertical stripes (Mfiller, 1886; Aiello, 1984). The body scoli reach their peak of development, and exist in two main forms, termed terete and flattened by Aiello (1984). In almost all species they are reduced on the prothorax and first abdominal segment, while in known members of the A. alala group they are almost obsolete on all segments with the exception of the third thoracic segment (Otero & Aiello, 1996; pers. obs.). Terete scoli bear 3-5 radiating spines either at the apex, or at intervals along the length of the scolus, and may be short and thick or club-shaped and sparsely spined (Aiello, 1984). Flattened scoli are characteristic of more derived members of the A. serpa group, and have a leaf-like appearance. The









spines are flattened, arranged in two opposite rows and may be densely packed, or even appear fused (MUller, 1886). The majority of larvae have lateral, inclined darker stripes on the abdomen, while in members of the A. phylaca group these stripes are absent and there is instead a large, lateral block of dark coloration from TI to A2 (Moss, 1933; Aiello, 1984). The larvae of A. alala is almost uniformly green (Otero & Aiello, 1996).
Pupa. Pupae are also variable in morphology, but all have the posterior wing margins protruding as keels, and the second thoracic and abdominal segments are expanded to form a dorsal projection (Miller, 1886; Moss, 1933; Aiello, 1984). The thoracic projection is smaller and directed posteriorly, while the abdominal projection may be elongated to form a very large, laterally flattened hook (MUller, 1886; Moss, 1933; Aiello, 1984). The head typically bears a pair of laterally pointing projections which are variable in size, shape and orientation (Aiello, 1984). The pupal color varies from pearly white, yellowish, brown, green, or copper to a shimmering gold or silver, and the sutures may be black. All pupae apparently give the impression of being empty or diseased (Aiello, 1984).
Development time. Aiello (1984) summarises the sparse available information on development time, which ranges from five to over thirteen weeks. Development time is rather variable within species and may depend on the hostplant (Aiello, 1984). A. basiloides is known to occasionally pass through six instars, which nevertheless take no longer to develop than the usual five.

Larval behavior. Adelpha larvae, upon hatching, eat some or all of the egg shell and begin to feed on the leaf tip, leaving the midrib intact, a behavior observed in many nymphalids (MUller, 1886; Moss, 1933; Aiello, 1984). Larvae rest when not feeding facing away from the leaf stem on the exposed midribs, and extend them by means of fecal material bound with silk, often constructing similar supports on lateral veins. Several supports may be constructed during each instar. After moulting to the final instar, larvae rest on the upper surface of the leaves (Maller, 1886; Moss, 1933; Aiello, 1984).









First to fourth instar larvae of Adelpha also engage in a distinctive activity first reported by Miller (1886), accumulating frass, and sometimes bits of dead leaf material, bound with silk, to form a mass surrounding the base of the support or suspended beneath. Adelpha basiloides also consistently constructs a small, usually curved larva-shaped mass on the leaf surface, away from the leaf edge or usual mass, which closely resembles the larva. This behavior has also been observed once in A. iphiclus (Aiello, 1984). The behavior of A. alala is unique within the genus; from the late first instar onwards the larva constructs a shelter by making two opposing cuts in the leaf, perpendicular to the midrib, then pulling the two leaf flaps down and fastening them with silk. The larva rests in this shelter when not feeding, and the shelter entrance typically has a curtain of small pieces of leaf material left by the feeding first instar. Larvae utilise these shelters until the late fourth instar, after which they rest on the leaf surface. This shelter, which presumably confers a protective benefit, may explain the greatly reduced scoli of all instars. Similar shelter construction is known in the Old World species Limenitis camilla and Ladoga reducta (Aiello, 1984), which utilise the same hostplant family (Ackery, 1988), supporting the primitive position of the A. alala group within the genus. Larvae of all species rest in a variety of positions, summarised in Aiello (1984).










Table 3. Summary of published and unpublished early stage information for Adelpha taxa.


Taxon: Egg: Larva: Pupa: Oviposition: Larval behaviour: Source: Locality: Reported as: i 2 3 4 5
bredowfi
californica D Edwards (1873) USA: Cal Limenitis californica
D;T D D D D D D N,P D Dyar(1891) USA:Cal Heterochroa californica
D F b/w, lateral, dorsal & Comstock (1927) USA: Cal Heterochroa ventral; D californica F F D; T D; T Ph D; T D; T N, P D Comstock & USA: Cal Heterochroa
b/w; b/w; b/w; Dammers (1932) bredowii
D; T D;T D; T californica F b/w, lateral F b/w, lateral Emmel & Emmel USA: Cal (1973)
eulalia D D Emmel & Emmel USA: Cal (1973)
* Phc Phc Phc Phc Phc Phc Phc, lateral Harry(1994) USA:Ut
eulalial D D D Scott (1986) USA Limenitis californica bredowii paraena
* paraena F b/w, dorsal, head F b/w, lateral and dorsal D Moss (1933) Brazil: Pa serpa capsule; D anterior ; D
lecromil T F b/w, lateral (Fig. 4: F b/w, lateral and dorsal Aiello (1984) Panama nr. paraena
massilia nr. PAR); D; T anterior 6 (fig. 6, nr. PAR); D; T
serpa
* diadochus F b/w, lateral; D D D Moss (1933) Brazil: Pa paraena
serpa D; D T P D Hoffmann (1937) Brazil: SC
(?) D; F b/w, head D P Moller (1886) Brazil: SC serpa capsule, taf. 2, fig.
21, scoli, taf. 2, fig. 7
* celerio T T F b/w head capsule F b/w, dorsal anterior V.; D; T Aiello (1984) Panama celerio min. (Fig. 2); D; T
* D; PJD: head capsule F b/w, dorsal (Fig. 24, F); N, N/plant D DeVries (1986, 1987) Costa Rica celerio PJD: pupal case, T
(?) F b/w; D F b/w; D P Comstock & El Salvador celerio Vizquez (1961)
hyas
hyas D Hoffmann (1937) Brazil: SC radiata
* aiellae D D Constantino (1998) Colombia serpa











Table 3. Continued.
Taxon: Egg: Larva: Pupa: Oviposition: Larval behaviour: Source: Locality: Reported as:
1 2 3 4 5
alala
completa D D D D Ph b/w; F b/w, head F b/w, lateral and dorsal view; P; N, N/plant Ph b/w leaf shelter; Otero & Aiello Venezuela alala capsule; D D D (1996)
*tracta PJD: head capsule PJD: pupal case, T D Haber, Chac6n (in: Costa Rica DeVries, 1986)
fessonia
fessonia D Janzen (in: DeVries, Costa Rica fessonia 1986)
* FSCA: head capsules FSCA: pupal cases R. W. Bosese, USA: Tex
-total T -specimens mythra D Mflller (1886) Brazil: SC
*basiloides D;T D;T D;T T T F b/w, lateral (Fig. 4) F b/w, lateral and dorsal D D Aiello (1984) Panama min. and head capsule anterior %; D; T (Fig. 2); D; T, + T for
6d' instar
PJD: head capsule PJD: pupal case D Marquis, Mallet & Costa Rica Janzen (in: DeVries,
1986)
USNM: head capsules USNM: pupal cases Small, specimens Panama plesaure
pleasure D D; Fb/w, ventral (Taf. 4, fig. P Miller (1886) Brazil: SC 13)
phliassa F b/w, lateral; D F b/w, lateral and dorsal D Moss (1933) Brazil: Pa phliassa anterior %; D
iphiclus
(?) ephesa D MOller (1886) Brazil: SC ephesa
(?) D; T D; T D;T D; T D; T D; T D; T N, P D D'Almeida (1933) Brazil: RJ iphicla ephesa
min.
* iphiclus F b/w, lateral F b/w, lateral and dorsal Moss (1933) Brazil: Pa iphicla anterior Ya
iphiclusi D Janzen (in: DeVries, Costa Rica iphicla iphicleola 1986) iphicleola D P Mailer (1886) Brazil: SC iphicla D;T D;T T T T F b/w, lateral (Fig. 4) F b/w, lateral and dorsal P Aiello (1984) Panama iphicla and head capsule anterior 1A; D; T
(Fig. 2); D; T
D DeVries (1987) Costa Rica iphiclus
* USNM: head capsules USNM: pupal cases Small, specimens Panama abyla D D Swainson (1901) Jamaica Heterochroa iphicla
D D D D N, P Milner & Turner (In: Jamaica Smith et al., 1994)










Table 3. Continued.
Taxon: Egg: Larva: Pupa: Oviposition: Larval behaviour: Source: Locality: Reported as:
1 2 3 4 5
abyla F c F c Blake, unpublished Jamaica
watercolour in
BMNH
ethelda
sophax D D D "as in A. cytherea". N, N/plant D DeVries (1986, 1987) Costa Rica zalmona

sophax
melona
leucocoma F b/w, lateral, head F b/w, lateral and dorsal Moss (1933) Brazil: Pa capsule; D anterior %; D
syma D MOller (1886) Brazil: SC D; T D; T ----D; T... D; T D; T P; N/plant D Hoffmann (1937) Brazil: SC cytherea
* cytherea D F b/w, lateral; D F b/w, lateral and dorsal D Moss (1933) Brazil: Pa
anterior 2
* marcia D D D "similar to boreas"; PJD: N, N/plant D DeVries (1986, 1987) Costa Rica
pupal case
* daguana D;T D;T T T T F b/w, lateral (Fig. 4) F b/w, lateral and dorsal Aiello (1984) Panama cytherea min. and head capsule anterior V2; D; T marcia (Fig. 2); D; T
USNM: head capsules USNM: pupal cases Small, specimens Panama ?) aea D P Moller (1886) Brazil: SC cytherea ?
viola
* pseudococala F b/w, lateral; D F b/w, lateral and dorsal Moss (1933) Brazil: Pa pseudococala anterior 1/2
(?) viola D; F b/w, lateral, taf. p Moller (1886) Brazil: SC sp. nr. cocala 3, fig. 2
? viola D D D D N, P D'Almeida (1931) Brazil: RJ cocala viola salmoneus
* salmonides D;T D;T D;T D;T D;T F b/w, lateral (Fig. 4) F b/w, lateral and dorsal P Aiello (1984) Panama salmoneus min. and head capsule anterior 'A; D; T (Fig. 2); D; T
D "similar to A. zalmona" DeVries (1987) Costa Rica demialba D Haber (in: DeVries, Costa Rica

1986)
D DeVries (1987) Costa Rica capucinus
* capucinus D D D F biw, dorsal; D F b/w, lateral and dorsal D Moss (1933) Brazil: Pa thesprotia
anterior 'A
? velia F b/w scolus, taf. 2, Moller (1886) Brazil: SC erotia var.
?ig. 8
? abia D D Moller (1886) Brazil: SC










Table 3. Continued.
Taxon: Egg: Larva: Pupa: Oviposition: Larval behaviour: Source: Locality: Reported as:
1 2 3 4 5 heraclea
* heraclea D F b/w, lateral; D F b/w, lateral and dorsal D Moss (1933) Brazil: Pa jordani anterior '4; D
* D F b/w, lateral (Fig. D; PJD: pupal case, T N, N/plant D DeVries (1986, 1987) Costa Rica heraclea 24, E); D; PJD: head capsule
erotia
* erotia (f. Pupa with large dorsal hook Aiello (pers. comm.) Panama
lerna)
phylaca
* pseudaethalia D D; T T T T F b/w, lateral (Fig. 4) F b/w, lateral and dorsal P Aiello (1984) Panama phylaca and head capsule anterior 'A; D; T pseudaelhalia (Fig. 2); D; T
Ph b/w lateral Aiello (1991) Panama ? D D DeVries (1987) Costa Rica boeotia boeotia messana
messana D Ph b/w lateral; F b/w F b/w, lateral and dorsal D Aiello (1991) Panama ixia leucas lateral; D; T anterior 1A; D
*thesprotia D F b/w, lateral F b/w, lateral and dorsal D Moss (1933) Brazil: Pa delphicola
anterior 'A
*mesenina F b/w, lateral F b/w, c, lateral and dorsal Moss (1933) Brazil: Pa anterior VA
lycorias
lycorias D D;T D;T D;T D;T F b/w lateral, taf. 3, F b/w, lateral view, taf 4, fig. P Maller (1886) Brazil: SC isis fig. 5, scoli, taf. 2, 8; D; T fig. 3; D; T
BMNH: Pupal case Specimen Brazil melanthe D D; T T min. T T F b/w, lateral (Fig. 4) F b/w, lateral and dorsal P Aiello (1984) Panama melanthe min. and head capsule anterior A; D; T (Fig. 2); D; T
D F b/w, lateral (Fig. F b/w, lateral (Fig. 24, C); D; N, N/plant D DeVries (1986, 1987) Costa Rica melanthe 24, D); D; PJD: head PJD: pupal case capsule
Ph b/w lateral Aiello (1991) Panama melanthe
* USNM: all instars, head capsules USNM: pupal cases Small, specimens Panama
wallisii D D D D Constantino (1998) Colombia lara wallisii cocala
* cocala D F b/w, lateral; D F b/w, lateral and dorsal D Moss (1933) Brazil: Pa anterior IA
* Iorzae D D; T T T T F b/w, lateral (Fig. 4) F b/w, lateral and dorsal P Aiello (1984) Panama cocala and head capsule anterior %; D; T (Fig. 2); D; T











Table 3. Continued.
Taxon: Egg: Larva: Pupa: Oviposition: Larval behaviour: Source: Locality: Reported as:
1 2 3 4 5
* lorzae PJD: PJD: D; PJD: head D; PJD: pupal case, T P; N, N/plant D DeVries (1986) Costa Rica T T capsule, T
USNM: head capsules USNM: pupal cases Small, specimens Panama

leucophthalma
leucophthalma Ph Ph Ph b/w, Ph Ph Ph b/w, lateral, Ph b/w, lateral, ventral; D; T P; N, N/plant D; Ph resting Young (1975) Costa Rica leucophthalma
b/w; b/w, d-lat. b/w, b/w, dorsal, head capsule; posture tegeata
D; T d-lat. d-lat. lateral D; T for all larval stages
T T T D "indistinguishable D; T Aiello & Small (in: Panama from cocala"; T Aiello, 1984) N, N/plant D DeVries (1986) Costa Rica leucophthalma mephistopheles
(?bereas
boreas D D N, N/plant D DeVries (1986, 1987) Costa Rica boreas tizona & b. opheltes
zina
* zina D; T T T T F b/w, lateral (Fig. F b/w, lateral and dorsal Aiello & Small (in: Panama justinajustina 5);D; T anterior /; D; T Aiello, 1984)
Unidentifiable
? radiata D; F b/w, lateral (Fig. D Aiello (1984) Panama nr. celerio
aiellae 4: CEL)
? sp. D Pupa spiny and gold, with two Several larvae Leaf midrib left Jorgensen (1922) Paraguay calliphiclea head horns. Thorax with a on same plant. intact, frass pile high keel, prolonged toward accumulated, the thorax, forming a large pupation on same "nose". Pupation lasts 20 days. leaf. ? sp. D D D Blanchard (1928) Argentina calliphiclea [possibly report of
Jorgensen (1922)]


Ph - photograph; F - figure; D - description; T - development time; c - color; b/w - black and white; d-lat. - dorso-lateral; P
position on leaf; N - number of eggs laid per oviposition event; N/plant - number of eggs per plant. ? - indicates identification uncertain, several similar species present at the record locality; (?) indicates identification uncertain, but probably this species; either original identification definitely incorrect, or several similar species present which would not have been recognised by author, with that listed here overwhelmingly more common than other(s). * adult voucher specimen or photograph examined. Single lines separate species, double lines separate species groups. Confirmed records are chronological within each species, records which are have an element of doubt are listed last. Specimens in collections on which publications have been based are listed in the same row as the publication information.









Adult Ecology

Nutrition

Adult Adelpha have been recorded at a variety of nutrient sources, of which the most numerically important is rotting carrion. This nutrient source has been mentioned occasionally in the literature (Adler, 1982; Sourakov & Enmel, 1995; Austin & Riley, 1995), but its pervasive utilisation by the great majority of the tropical forest butterfly fauna still remains unappreciated. Authors continue to place unwarranted stress on the importance of rotting fruit in the diet of tropical butterflies (DeVries et al., 1997), while most ignore rotting carrion completely. Of the 54 species recorded by myself during field work in Ecuador, 47 species (87%) were strongly attracted to rotting fish, and a great number of these were not observed outside of traps. Of the species not recorded on this nutrient source, only A. shuara is sufficiently common to believe that it might genuinely not utilise this source. I recorded only nine species on rotting fruit, and this low number is consistent with other studies (DeVries et al., 1997). Many Adelpha have also been recorded feeding at damp sand and mud, particularly at urine, a behavior known as puddling. While most Adelpha have not been observed at nectar sources, the species that have been reported tend to be in more primitive groups within the genus, though an explanation for this is lacking.

Hall & Willmott (1999) discuss the possible nutritive benefits from feeding on rotting fish, which appears to offer similar nutrients to puddling (Downes, 1973). Sodium ions have been shown to stimulate puddling (Arms et al., 1974), and this is a probable nutrient obtained from rotting fish, which is fed on exclusively by males. Sodium has been shown to be donated to the female in the spermatophore during copulation, and is probably important for the higher neuromuscular activity of perching males (Downes, 1973; Arms et al., 1974; Hall & Willmott, 1999). In addition, though there is no evidence to date, males may also be seeking amino acids for maintenance of their thoracic muscle









mass. Rotting fruits and nectar are probably both sources of carbohydrates, the energy source for powered flight.

In addition to providing information on the nutrient requirements of adult Adelpha, I cannot stress enough the importance of trapping with rotting carrion for anyone conducting faunal inventories in the Neotropics. In most localities, particularly primary forest, Adelpha species are seldom if ever observed, and certainly not the rarer species. Males of the recently described species A. hesterbergi and A. lamasi are known from Ecuador only through the use of such baits. Our understanding of the geographic and elevational distribution of Adelpha would be inestimably greater if researchers in all regions used rotting carrion baits. I greatly hope that researchers will take note of the information on trapping techniques given in Sourakov & Emmel (1995) and Austin & Riley (1995), and use it to contribute to our knowledge of Adelpha distribution.


Perching and courtship

Males of many Adelpha species have been observed "perching"; perching is defined as males occupying a specific area and flying out to investigate passing butterflies in search of females (Scott, 1976). Perching males typically maintain one to several perches, where they wait from early morning to early afternoon, usually in bright sun. Perches may be up to 30m apart, but while the male often flies from one perch to the next and back, I have never seen them move through a habitat as suggested by DeVries (1987). On the contrary, I have found perching species to be typically highly localised within a habitat, using the same bush for up to four consecutive years (A. argentea), and often not being encountered elsewhere within a site. Perching height varies from less than a meter above the ground (A. cytherea) to 15m (A. leuceria) and almost certainly higher, and while varying to some extent within species, there nevertheless appears to be a favored height range for each species. Males have been recorded perching in several areas, including open clearings or primary forest along rivers, forest edges and ridge top light









gaps. While some species occur only in one of these microhabitats (A. shuara, A. hyas, A. argentea, A. serpa, along rivers; A. cocala, A. phylaca, on ridge tops and along forest edges), a number may be found in both (A. salmoneus, A. iphicleola). In addition, there are many common species for which there are no perching observations, and these probably perch high in the forest canopy. Perching males rest on the tip of the leaf upperside, with the wings typically closed or almost so, and are very wary. Flight between perches is typically gliding, with the wings held angled slightly downwards.

Despite having observed perching males on numerous occasions, I have never observed courtship or copulation, and know of no published observations on either of these activities. Almost nothing is known about mate recognition, but due to the great variation in wing pattern between some subspecies, and the great similarity between many species, it is very probable that wing pattern cues play a small, or no part. This hypothesis is borne out by the possible hybrid specimens between the closely related but phenotypically very distinct species A. mesentina and A. thesprotia, and A. cocala and A. irmina. Instead, mate recognition and courtship in Adelpha may be pheromonally mediated; male Adelpha have a dense area of darker scales at the base of the ventral forewing, in cell Cu2, which is lacking in females, and although these scales are of a similar structure to remaining wing scales, they may have some role in pheromone dissemination.


Mimicry
The close resemblance on the dorsal surface of several Adelpha species, which proved to be unrelated when their early stages were examined, led Aiello (1984) to suggest that Adelpha were involved in mimicry. She tentatively suggested that species which fed on rubiaceous plants might form the models for other species. Unfortunately there is very little information available on the palatability of Adelpha species, as is the case with all Neotropical butterflies. The experiments of Srygley & Chai (1990)









demonstrated that 100% of specimens of A. cytherea and A. serpa celerio, the former a proposed model of Aiello (1984), were entirely consumed by caged jacamars, while more recently Pinheiro (1996) also found the few species that he tested to be palatable to kingbirds in Brazil. Nevertheless, there is now a sufficient body of evidence available to say without any doubt that Adelpha species are involved in mimicry, in the sense that dorsal wing patterns have converged onto a common pattern, both with other Adelpha and with species from other butterfly families. This is apparent both at the level of subspecies, and also at the species level, as illustrated by an examination of Figs. 10-94, the order of which is based on the phylogeny derived in Chapter 3. All that remains is to isolate the selective agent driving the wing pattern convergence. It is also worth noting that while the study of Pinheiro (1996) failed to find evidence for unpalatability in Adelpha, several species in classically unpalatable genera, such as Heliconius, were consumed with equal impunity.
The Adelpha fauna of the Apure region in Venezuela, whence a number of new subspecies were recently described by Neild (1996), constitutes possibly the most compelling evidence of mimicry. No less than seven subspecies (A. cytherea nahua, A. cocala orellanai, A. malea malea, A. heraclea antonii, A. phylaca joffrei, A. capucinus gutierrezi and A. plesaure pseudomalea) occurring only in this small region have independently evolved an orange band bordering and touching the distal edge of the white band on the dorsal hindwing, a pattern that occurs outside of this region only in one subspecies from the neighboring Catatumbo region. Moreover, the female of Doxocopa lavinia, which closely resembles Adelpha throughout its range, also has the same hindwing pattern in this area (Neild, pers. comm.). Another strong example of independent convergence of a dorsal wing pattern is the narrowing of the white dorsal bands in a number of unrelated species that are superficially similar on the dorsal surface and which occur sympatrically in western Ecuador. Such species include A. iphiclus, A. iphicleola, A. erotia f. lerna, A. basiloides and A. barnesia. All of these species develop








particularly broad white bands in the Rancho Grande region of northern Venezuela. There are numerous other examples of the close correspondence of ranges of subspecies in which the dorsal wing patterns change synchronously from one region to the next, discussed under each taxon. In a number of cases similar wing patterns are derived from the fusion and modification of different wing pattern elements, indicating that they are clearly not homologous, such as in the species A. naxia and A. iphiclus.

As with other mimetic groups, several mimicry "rings" may occur sympatrically (Beccaloni, 1997a; Joron & Mallet, 1998). Maintenance of diverse mimetic assemblages has been suggested to be due to microhabitat partitioning (DeVries & Lande, 1999) through associated ecological factors, including hostplant height (Beccaloni, 1997b) and roosting height (Mallet & Gilbert, 1995), and either of these and/or some other ecological factor may be important in Adelpha. It is my experience that males of Adelpha species with similar dorsal wing patterns typically also perch in similar microhabitats; for example, in eastern Ecuador I have found A. thoasa manilia, A. serpa diadochus, A. hyas hewitsoni and A. iphicleola thessalita perching in a single river side clearing on the same bushes. "Orange and white banded" Adelpha, by contrast, have very few perching records, but their numerical abundance (pers. obs.) in traps in primary forest suggests that they perch in the canopy or subcanopy.

Mimicry of Adelpha also appears to extend beyond the genus, and other species with very similar dorsal wing patterns to Adelpha include the females of most Doxocopa species, and the riodinid Synargis phliassus (Cramer), which Hfibner ([1819]) even included in the genus in his original description of Adelpha. Finally, it is worth noting that the simple, contrasting color patterns of the dorsal wing surfaces of Adelpha are certainly not cryptic, particularly given the habit of many species of resting (when not perching) with the wings open. More data are urgently needed on the palatability of Adelpha species to a wider variety of potential predators to more fully understand the exact nature of the mimicry system.









Biogeography
Diversity. Adelpha species range from far northwestern U.S.A. south to Uruguay, and while the majority of species are strictly continental, four species occur in the Greater Antilles, of which two are endemic. Tabulating species and subspecies diversity by political boundaries (Table 4) is a simple first step to revealing broad biogeographic patterns, and it is important in terms of the allocation of resources for biodiversity conservation (McNeely et al., 1990). However, it is also apparent that sampling effort has a great effect on perceived taxa diversity in different countries, and as a step towards understanding the extent of this effect in Adelpha and other butterfly groups, and to more fully elucidate true taxa diversity, I also tabulate the number of expected taxa from each country in addition to those already recorded.
Expected taxa are predicted either through extrapolation of a known range spanning either side of a country (in areas such as the Andes and Central America), or through examination of the ranges of more common species known to be sympatric in similar habitats and at similar elevations. Clearly, expected taxa will be underestimated for all countries at the periphery of biogeographic regions, since the first criterion is not possible to apply, and these countries include Mexico, Venezuela, Ecuador (west), Bolivia and Uruguay. The countries with the most complete faunal knowledge, excluding island areas and countries with very low species diversity, are Mexico, Costa Rica, Panama, Ecuador, Peru, Bolivia and Brazil. Countries whose fauna is particularly poorly known include most Central American countries between Mexico and Costa Rica, the Guianas and Uruguay.
There is an obvious latitudinal gradient in taxa diversity, which peaks in the countries of Colombia and Ecuador. Although Ecuador has the highest recorded species diversity of any country (56 species), this is due to extensive sampling there during the course of my research by myself and Jason Hall. Both Colombia and Ecuador are predicted to contain 59 species, or 69% of the entire genus, and it is possible that an









additional species (A. fessonia) may be found in the isolated dry forests of southwestern Ecuador, which support other dry forest species with disjunct ranges (Willmott & Hall, unpubl.). By contrast, Peru and Brazil, often cited as the most species-rich countries in the world for butterflies (Brown, 1991), are significantly less diverse, with 46 and 42 species respectively, and reasons for this are discussed further below.

Since Adelpha species occur in a wide variety of habitats at a broad range of elevations, and show much variation in ecological traits such as hostplant specificity and microhabitat preference, they are probably reasonably accurate indicators of diversity in other groups of butterflies (Brown, 1996). However, certain more sedentary groups, such as the Satyrinae and Ithomiinae, are likely to exhibit higher levels of endemism, probably more similar to the patterns evident from an analysis of the distribution of Adelpha taxa diversity. Again, Colombia proves to be clearly the most diverse country in terms of taxa, with a predicted total of 110 (53% of the genus), followed by Ecuador with a predicted total of 81 (39% of the genus). Of interest is the high number predicted for Venezuela, seldom mentioned in considerations of the distribution of biodiversity (e.g. Brown, 1991), which ranks third with 75 (36% of the genus).

An explanation for these patterns of diversity may be gleaned from an examination of the distribution of species by biogeographic region (Table 5). I use the biogeographic regions as defined by Brown (1982ab), with several of the subdivisions of Brown (1979) and others that appear useful to understand the distribution of Adelpha diversity. The reason for the high species and taxa diversity of Venezuela, Colombia and Ecuador is clear; all three countries contain faunas derived from three of the four principal biogeographic regions (Transandean, Andes and Amazon), while countries such as Peru and Brazil span only two regions (Andes and Amazon, and Amazon and Atlantic, respectively). Despite the high endemism of the Brazilian Atlantic region (11 species), this does not compensate for the great diversity of Andean and Transandean faunas.










Table 4. Distribution of Adelpha taxa by country.


> 0i

0 0

a ~ 0.~ 0 0 > ) 2 , 1.


XX
X
XX
X
XX XX



XX X ? X ?


X? ?XX X?
?X X


XXX
? XX
?


Taxon
bredowil californica eulalia bredowii diodes creton diodes herbita zea
paroeca nea
sentia nea
paraena massilia lecromi reyi paraena radiata
aiellae explicator radiata myrlea gilletella serpa
celerio duiliae diadochus serpa seriphia godmani egregia
seriphia pione aquillia therasia barcanti hyas hewitsoni viracocha


XX

X
?XX
? XX
? ?


XXXXXXXX
XXXXXXXX



XX?X? ?XX XX?X? ?XX


X
XXXX


XXXX ? XXXX ? XXXXX


XXX X ? ?
X
X ? ?


X X ?X


X X


X ? ? X


? ? X
X


XXXXXX


X
XXXX


XXXXX
X X
X


X
XXX


X ? X
XXX



XXX
? ?X




? ?X X ? X X ? X


XX
XX


_ yas X X X X alala X X X X X X alala X X completa X X negra X X X X X aricia ? X X portunus ? X serenita X aricia X X corcyra X X X X corcyra X X


XXX?X? XXX? X?


XXXXX XX









Table 4. Continued.

Taxon J 5 m Lu C au > w CL n < a o U
salazari X
collina X X aretina X X
dognini X X tracta X X X pithys X X ? X donysa XX X X
donysa X X
albIilum X X
fessonia X XXX X XXXXXX? fessonia XX XX X X X X X ernestoi X
cestus ?X
lapitha ? X
gdania X X X
gelania X X
arecosa X cafllphane XX X ? poius X m.thra X X X ? basiloides XXXXXXXXXXXX plesaure XX X X X X X X X X
pseudomalea X X
phliassa x x x x x x x x x
plesaure X
symona X ? ? X X gavina X falcipennis X X X ? thoasa X X X XXX X X ? X ?
manilia x x x x
thoasa x x x x gerona x x x
calliphiclea ? X ? thessalia X ? X XXX X X X
thessalia x ? x x x
cesilas X X x
indefecta x x x x
iphidus XX X ? X X XXX X X X X ? XX X X X X
iphiclus x x x ? x x x x x x x x x x x x x X
estrecha ? x
ephesa ? x x
iphicteola XXXXXXXX X X X X X X X ? ? ? X X
iphicleola x x x x x x x x
gortyna ? x x X thessalita ? x x leucates x x x x ? ? ? x iphimedia X abyla X melona ? X X X X X X X X X x x x x
neildi ? x deborah ? x x x leucocoma x x x x x x x x x x melona x pseudarete x x x ethelda X ? ? ? ? XX X XX ? ? X ssp. nov. x ? ? ?
sophax ? x X eponina x









Table 4. Continued.

D~~~~t r._0< uw0< Taxon :02m j zuw c 3 nU
zalmona X X
ethelda X X
galbao X ? ? X epione X ? X X X ? ? X
agilla X ? X XX
epione ? ? X
syma X X X X cotherea X X X ? X X X X X X X X X X ? X X X X X
marcia X X X ? X X X
daguana X X X X X
nahua X
olbia X X X
cytherea X X X X X X X X X
aea ? ? X
insularis X viola X X X X ? X X X X
pseudococala X X X X ? X X X X
riola X salmoneus X X X ? X X X X X X X X X
salmonides X XX X X XX
emilia X X
salmoneus X X
colada X X X X demialba ? X X ? epiois X X XX
epizygis X X X X
epona X
fabricia X XX X X X X ? ? capucinus X X X X X ? ? X X X X
gutierrezi ? X
capucinus x x x x x x x x x
velia X ? ? X
barnesia X X X X X ? X X X x x x x X X
leucas X X X X X ? XX X X X
barnesia ? ? ? X X X
trinila x diazi x
hesterbergi X ? ? X abia XXX ? naxia X X X ? X ? X X X x x x x X ? X ? X
naxia X X X ? X ? X X X X X X X X ? X ?
oteroi ? X
hieronica X heraclea ? X ? ? ? X X X X X X X ? X
heraclea ? X ? ? ? X X X x ? X x ? x
makkeda ? X antonii ? X
sp. nov. X malea X X ? ? X X X X X X X X x x x x X fundania x x? ?x x x x X ? ixia x malea ? X aethalia x ? x x x x x goyama x x x x boeotia X X ?X ? X ? X X X X X X X X X ? X oberthurii x x ? ? x ? x x boeotia x x x x x x x x ? X










Table 4. Continued.

Taxono
fidicula X
altamazona ? ? ? X ? X X ? ? ximena X X X X X X ? ? X
ximena XX X X X
mossi ? X ? X X X ? ? X delinita X X ? ? X ? X X X X X X X
utina X X? ? X ? X X
delinita XX X X X
poilina ? X X X X X ? ? X erod X ? ? ? ? X X X X X X X X X X ? X
erotia X ? ? ? ? X X X X X X X X X X ? X
caphira X X
phylaca X X X X X ? X X X X X
phylaca X X X XX
pseudaethalia X X ? X XX X X X X
joffrei ? X
messana ? X ? ? ? ? X X X X X X X X ? ? X ?
messana ? X ? ? ? ? X X X X X
delphicola X XX X X X ? ? X ? thesproda X X X X X X X X X X X mesentina X X X X X X X X X X lycorias X X X X XX X XX X X X X X X X ? X
melanthe XX X X X X XX
wallisii X
spruceana X X
melanippe X
lara X X X X X X X
lycorias X X X ? attica ? XXX X X X X
hemileuca X X
attica X X X X X X X leuceria X X ? X X X X X X X
leuceria X X ? X X X X X
juanna X X
leucerioides X X
erymanthis X ? ? X XX X X
ssp. nov. x
erymanthis X ? ? X X X
adstricta X
ssp. nov. X
sichaeus X X X X X rothschUdi X X stilesiana X X boreas ? X X X X X X X ? X ? X
boreas ? X X X X X X X
kayei X ? X ? X cocala X X ? ? X X XX XX X X X X X X X
lorzae XX 9. ? X X X X X X X
orellanai ? X
cocala X X X X X X X X X
didia X caninia X felderi X X X X leucophthalma X X X X X
leucophthalma X X X X
smalli X
irminella ? X










Table 4. Continued.

Taxon = mw zu up > w m m U, F on irmina X X X X X X
irmina X X X
tumida X X X X saundersii X X X X
frontina X X
saundersii X X X
helepecki X X lamasi X X salus X ? ? ? X X X X
ssp. nov. X ? ? ? X X
salus X
emmeli ? X
shuara X X ? argentea X X X ? coryneta ? X X jordani ? X X X zina X X X X X X X X
lacina X X X
zina X X X X
inachia X
desousai X pyrczi X
enope X
irma X X X X mileri X ?
justina X X X X X
justina X X
maira ?
justinella X inesae X
valentina X X X X olynthia X ? X X X levona X X
Z~

Taxa
Total recorded 4 36 28 14 13 21 18 35 40 86 66 74 52 44 20 19 58 5 18 12 21 11 1 1 2 1 1 Total expected 4 38 34 24 28 31 32 38 43 1to 75 81 57 50 24 24 61 10 27 26 26 11 1 2 2 1 1 Recordedas% too 95 82 58 46 68 56 92 93 78 88 91 91 88 83 79 95 50 67 46 81 too too 50 too too too of expected
Expected as% 1.9 18 16 12 14 15 15 18 21 53 36 39 28 24 12 12 29 4.8 13 13 13 5.3 0.5 1 1 0.5 0.5 of genus
Species
Total recorded 3 32 27 14 12 20 18 34 36 53 40 56 46 39 18 18 42 5 18 12 20 11 1 1 2 1 1 Total expected 3 34 33 23 27 30 32 37 37 59 45 59 48 45 22 23 43 10 26 25 25 11*1 2 2 1 1
Recorded as% too 94 82 61 44 67 56 92 97 90 89 95 96 87 82 78 97.7 50 69 48 80 too too 50 too too too of expected
Expected as% 3.5 40 39 27 32 35 38 44 44 69 53 69 56 53 26 27 51 12 31 29 29 13 1.2 2.4 2.4 1.2 1.2 of genus








naxia x x ? x heraclea x x x x sp. nov. E malea x x ? x x boeotia x x x x x altamazona E ximena E delinita x x x x polling E
erotia x x x x x phylaca x x x
messana X X X X thesprotia E mesenfina E IWcOr-s x x x attica x x x leucerta x x x leucernoides E
erymanthis x x x sichaeus E rothschila& E stilesiana E boreas x x x cocala x x x x x felden E leucophthalma x x innina E saundersih x x lamasi E salus x x x shuara E argenrea E coryneta E jordani E zina x x x milleri E justina X x obwnhia x x levona E
Transandean Andes Armazon Brazilian Caribbean Atlantic



Go
Species I

Endemic/subregion 7 3 0 3 7 5 11 1 0 I ENDEMIC is 10 5 11 2
Total/subregion 33 36 17 19 22 27 1 1 2
(22)
TOTAL 44 31 27 31 4 x = recorded, ? = expected; E - endemic


Table 5. Distribution of Adelpha species by biogeographic region.
Transandean Andes Amazon Brazilian Caribbean Atlantic




Species . / /

bredowi E
herbitaE ZeaE
paroeca x x
nea x x ? x paraena x x ? x x radiata X X X X serpa x x x X x seriphia x x x x ha xcs XX alula x x arl cta E corcyraz x x tracia E
pihs E
do ~a E
fessonta x x ? x gelaia E
calliphane E polhius E mythra E basiloides x x x plesaure x x gavi a E falcipennis E thoava X X thessalia X X lphiwlus x x x x x tphwcleola X x x X x x
abylaE melona x x x x ethelda x x X x epiore X X
syma E
,threa x x x x x viola x x salmoneus x x x x demialba E
epI--Ygls E fabricia x x capucinus x x barnesta x x x x x draft x
hesterbergi x x abia E









An analysis of the distribution of diversity in Adelpha with elevation (Fig. 5) shows the expected pattern of a decrease in diversity with altitude, and a peak at around 700-1000m. However, this peak in diversity does not necessarily reflect the local diversity within particular regions, but may be due to the greater endemism of species in low elevation premontane forests, and the extension of many lowland species into the lower elevation zone.


1000 1500 2000
Elevation above sea level/m


Fig. 5. Variation in species diversity with elevation throughout the range of Adelpha.




To examine the effect of elevation on local species diversity I tabulated data for Ecuadorian species on either slope of the Andes, thus eliminating the effects of changing latitude and mean diurnal temperature on elevational ranges. I have included only data collected by myself and Jason Hall, and by other reliable workers. While it is clear that several species, particularly on the western slope, have wider ranges than the known




Full Text

PAGE 1

SYSTEMATICS, ECOLOGY AND BIOGEOGRAPHY OF THE NEOTROPICAL BUTTERFLY GENUS ADELPHA (LEPIDOPTERA: NYMPHALIDAE: LMENITIDINAE) BY KEITH RICHARD WILLMOTT A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 1999

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ACKNOWLEDGMENTS I am very grateful to P. Ackery and J. Reynolds (BMNH), Drs. L. and J. Miller (AME), Dr. J. Pierre (MNHN), Dr. W. Mey and M. Nuj3 (ZMHU), Dr. J. Miller and Dr. E. Quinter (AMNH), Dr. P. Perkins (MCZ), Dr. J. Heppner (FSCA), Dr. G. Lamas (MUSM), Drs. R. Robbins and D. Harvey (USNM) and the curators at the BMB for giving me access to the Adelpha collections in their care and for the loan of material for dissection. Dr. Lobl of the MHNG and A. Lvovsky of the ZISP were very helpful in identifying and supplying me with photographs of Adelpha types in their respective collections. I thank D. Trembath, M. Perceval, L. Constantino, J. Salazar, Dr. E. SchmidtMumm and J. LeCrom for allowing me to view and take notes from their private collections and for their generous hospitality. I especially thank A. Neild for providing accommodation in London on numerous occasions, for donating specimens of Adelpha, for his continuous help and correspondence and willingness to discuss ideas. R. Hesterberg generously loaned his specimen of .4. hesterbergi to me for study. I thank the librarians at the BMNH entomological library for their help and for allowing me to photocopy or photograph important pages of several rare books. I am greatly indebted to Dr. G. Lamas, who freely shared his peerless knowledge of neotropical butterfly taxonomy and nomenclature, providing invaluable help and advice on numerous questions and photographs of several type specimens, and whose unpublished checklist of Adelpha was the foundation for my research into the genus. I thank Dr. M. Parsons for help in navigating the maze of Adobe Photoshop and Aldus Freehand, Dr. A. Aiello for her continuous helpful correspondence. Dr. J. Mallet for donating slides of early stages, and Drs. W. Haber, D. Janzen, P. DeVries and A. Orellana ii

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for unpublished information on Adelpha hostplants. This dissertation benefitted greatly from the valuable comments and suggestions of Dr. T. Emmel, J. Heppner, J. Lloyd, L. Miller and J. Reiskind, to all of whom I extend my appreciation. I am also very grateftil to Dr. T. Emmel for providing superb facilities and for supporting me during my time at UF with research assistantships, funded by the U.S. Fish and Wildlife Service and private donations, and for all his help and encouragement throughout my time in Florida. I also thank the following for assisting with the costs of field work in Ecuador: Mr. I. Wilhnott, Mrs. M. Willmott, Christ's College Cambridge University, Albert Reckitt Charitable Trust (C. T.), Poulton Fund Oxford University, Round Table Trust, Lindeth C. T., Catherine Cookson Foundation, Morton C. T., Royal Entomological Society, Butler C. T., Mr. D. Exell, Peter Nathan C. T., Harry Crook Foundation, Douglas Heath Eves C. T., R. and M. Foreman C. T., Northern Bank, Banbridge Academy, C. Bruce, Hickley Valtone Ltd., Vera Trinder Ltd., Agfa, Phoenix Mountaineering, Balfour-Browne Fund, Worts Fund, Sigma Xi the Scientific Research Society (1996) and Equafor; field and museum research in 1997/8 was fianded by National Geographic Society Research and Exploration Grant No. 5751-96. Florida Agricultural Experiment Station provided facilities and fimded publication of my research work on Adelpha. I also thank INEFAN, the Pontificia Universidad Catolica and the Museo de Ciencias Naturales, Quito, and in particular Dra. Germania Estevez, for arranging the necessary permits for field research in Ecuador. Finally, I thank both my parents for their continuous support throughout my doctoral studies and before, my dear fiiend Julia Robinson-Dean for her fine company, help and tolerance during two field trips to Ecuador, and my lifelong fiiend Jason Hall for his continuous advice and healthy criticism during my work on Adelpha, for collecting many rare species, for his splendid humor in the field on many trips throughout the tropics, and for first introducing me to the delights of lepidopterology. iii

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TABLE OF CONTENTS Page ACKNOWLEDGMENTS ii ABSTRACT vi CHAPTERS 1 INTRODUCTION 1 2 REVISION OF THE GENUS ADELPHA 5 Introduction 5 Materials and Methods 6 Systematics 17 Biology 58 Review of Adelpha 98 3 PHYLOGENY OF THE GENUS ADELPHA 664 Introduction 554 Methods 555 Results and Discussion 672 4 SUMMARY AND CONCLUSIONS 681 APPENDICES A LIST OF TAXA DISSECTED 683 B NOMINA NUDA AND NAMES ERRONEOUSLY INCLUDED IN ADELPHA 689 C NOMENCLATURAL CHANGES, ERRORS AND MISIDENTmCATIONS OF FIGURED ADELPHA IN POPULAR WORKS 690 D DATA FOR FIGURED SPECIMENS OF ADELPHA, FIGS. 10-94 693 iv

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E CHARACTER MATRIX FOR PHYLOGENY OF ADELPHA 698 REFERENCES 700 BIOGRAPHICAL SKETCH 724 V

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Abstract of Dissertation Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy SYSTEMATICS, ECOLOGY AND BIOGEOGRAPHY OF THE NEOTROPICAL BUTTERFLY GENUS ADELPHA (LEPIDOPTERA: NYMPHALIDAE: LIMENITIDINAE) By Keith Richard Wilhnott May 1999 Chairman: Dr. Thomas C. Emmel Major Department: Entomology and Nematology A revision of the diverse Neotropical nymphalid butterfly genus Adelpha is presented, which includes a review of the systematics, ecology and biogeography of all constituent taxa. I discuss the relationships of the genus to other New and Old World species and genera of the tribe Limenitidini and define the genus based on wing pattern characters. I present a synonymic checklist which includes all 362 described names, and recognise 85 valid species and 207 valid taxa. Eight new subspecies are described, and a new species and four subspecies are recognised but undescribed due to a lack of material. The status of all names is critically examined, the location of type material is discussed and notes on the identification of all taxa are presented in addition to a dichotomous key for identification of adults of all species. Adults of all taxa are figured and male and female genitalia of the great majority of species. Lectotypes are designated for 29 names, and neotypes for two names. I summarise information fi-om the literature on hostplants utilised by the genus and the morphology and behavior of the immature stages. Notes are provided on the vi

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habitats, phenology, rarity and adult behavior of all taxa based on published data and approximately two years field work in six Neotropical countries. I also list the locality data and depository of 20,730 specimens of Adelpha examined and discuss the known and expected geographic and elevational ranges of all taxa. A phylogeny for all species within the genus is presented based on wing pattern and adult morphological characters of both sexes. While there are several well defined clades, the phylogeny lacks high resolution and strong support for nodes throughout due to a pervasive lack of morphological characters. Nevertheless, there is much evidence that adult characters do support phylogenetic groups similar to those derived fi-om analysis of characters fi-om immature stages. I discuss in particular the problems of homology assessment of wing pattern characters which have led to past taxonomic confiision. Dorsal wing patterns show abundant evidence of convergence among sympatric taxa, and I discuss the evidence for mimicry within Adelpha. vii

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CHAPTER 1 INTRODUCTION The Neotropical region is justly famed for its rich biological diversity, and nowhere is this more apparent than in its insect faima, particularly butterflies. Many of the most visually spectacular species occur within the largest single family, the Nymphalidae, whose species inhabit a very wide diversity of habitats and exhibit great variety in their life history strategies. This variety offers a potentially rich area for research into many aspects of evolutionary biology, and just a few of the fields in which nymphalids have made a particularly important contribution include plant-insect interactions (e.g. Brown, 1987), the study of speciation processes (Jiggins et al., 1996; Davies et al., 1997), the phenomenon of mimicry (Bates, 1862; Muller, 1879) and studies into the historical biogeography of the neotropics (Brown, 1982). In the recent surge of interest in the conservation of biodiversity, one nymphalid subfamily, the Ithomiinae, has been suggested as an ideal indicator taxon of tropical lowland forest butterfly diversity (Beccaloni and Gaston, 1994). The higher level systematics of the Nymphalidae are still keenly debated, and there is no consensus as to the relationships between subfamilies or even as to their total number. The most recent classification of Harvey (1991) recognises 11 neotropical subfamilies, including the Libytheinae, Danainae, Heliconiinae, Ithomiinae, Brassolinae, Morphinae, Satyrinae, Charaxinae, Apaturinae, Nymphalinae and Limenitidinae. Paradoxically, despite the wealth of information that nymphalid butterflies have provided, biological research to date has largely concentrated on only a few groups, such as the Danainae, Heliconiinae and Ithomiinae. These subfamilies contain species that are 1

PAGE 9

2 typically common, slow flying and therefore easy to study in the field, and easily manipulated under artificial conditions. There exist several modem systematic treatments of these subfamihes (Fox, 1956, 1960, 1967; Fox and Real, 1971; Michener, 1942; Brown, 1981; Ackery and VaneWright, 1984), but it is only recently that explicitly derived phylogenies have been developed for the Heliconiinae (Brower and Egan, 1996; Penz, 1999) and Danainae (Ackery and Vane-Wright, 1984), while none is yet available for the Ithomiinae. The Brassolinae, Morphinae and Charaxinae, all popular groups with amateur collectors, have been the subject of various generic monographs (Bristow, 1981, 1982, 1991; Blandin, 1988, 1993; Comstock, 1961; Barselou, 1983). Higgins (1981) reviewed a large section of the Nymphalinae, while the species-level taxonomy of the Satyrinae has remained a virtual terra incognita since its treatment by Weymer (19101912). Of the remaining subfamilies, two are monotypic, leaving the large and heterogeneous Limenitidinae. As yet there is no evidence to suggest that this subfamily comprises a natural (monophyletic) group of taxa, and it has been retained by modem taxonomists largely for convenience (Harvey, 1991; Neild, 1996). It is, however, in this subfamily that Neotropical nymphalids reach their greatest diversity in both morphology and species, and probably life history strategies. The monumental work of Seitz ([1907]1925) and the pictorial reviews of D'Abrera (1987) are, for many genera, the only available fairly comprehensive references. A number of other genera have been revised, including Callicore (Dillon, 1948), Ectima, Hamadryas, Myscelia, Catonephele, Eunica, Epiphile, Asterope, Nessaea (Jenkins, 1983, 1984, 1985a,b, 1986, 1987, 1989, 1990) and Perisama (Attal and Crosson du Cormier, 1996), and these revisions have often illustrated the existence of both undescribed and unrecognised species even in small, well collected genera, a result that is certainly much more widely applicable than is currently acknowledged.

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3 It is thus apparent that systematic study, the foundation for ecological research, is still urgently needed in many nymphalid groups. A further concern is the continual loss of biological diversity in tropical countries and the pressing need to catalogue and understand the distribution of such diversity. Among nymphalid butterflies, one of the most conspicuous members in all neotropical forest habitats is the genus Adelpha, which is certainly the most speciose genus of the neotropical Limenitidinae, and possibly of the entire subfamily Nymphalidae. Many Adelpha species are common, they occur in a very wide diversity of habitats and their immature stages exhibit a great diversity of behavior, morphology and hostplant utilisation (Moss, 1933; Aiello, 1984; Ackery, 1988). Previous studies of the immature stages demonstrated that adults with dorsal wing patterns which appeared superficially similar were often apparently unrelated (Moss, 1933; Aiello, 1984), leading to the suggestion that this adult similarity was due to mimicry (Aiello, 1984). The genus thus offers great promise for biological study, but this has remained unrealised largely due to the problems in identification of many extremely similar species and a lack of understanding of their relationships, and the taxonomic chaos caused by this similarity and the availability of over 350 published names for Adelpha taxa. As a result, Adelpha has become renowned more for its taxonomic problems than its potential biological interest; Fruhstorfer (1915) states that "except the Phyciodes and certain Euthaliidae there is, therefore, hardly any genus of the Rhopalocera offering greater difficulties to the systematical and synoptical treatment", while DeVries (1987) echoed this sentiment, saying "the butterflies that compose the genus Adelpha are, in my opinion, the most difficult and trying taxonomically of all the nymphalids". Fruhstorfer (1915) was the last to treat the entire genus, but offered few reliable characters for separating species and his work is replete with misidentificafions. The objectives of this study are, therefore, to provide a stable nomenclature for the genus through a critical examination of the status of all available names and to

PAGE 11

4 provide information to enable certain identification of all Adelpha taxa. I also attempt to create a systematic arrangement that reflects modem species concepts and our greatly improved knowledge of the distribution of taxa. As a baseline for future research, I summarise all of the available information on the immature and adult biology of Adelpha species. I also analyse the distribution of diversity and endemism within the genus and discuss its implications for the conservation of Adelpha, and probably other butterfly taxa. Finally I derive a preliminary phylogeny for the genus to serve as a heiuistic tool for field workers and to test the hypothesis that similar dorsal wing patterns in many sympatric Adelpha are the result of convergent evolution through mimicry.

PAGE 12

CHAPTER 2 REVISION OF THE GENUS ADELPHA Introduction The genus Adelpha is the only member of the tribe Limenitidini to occur in the Neotropical region, and, like its Old World relatives, it has always been a bane to systematists. Estimates of the number of species in the genus have ranged as high as 100 (Aiello, 1991), while there are no published reviews of the genus which enable accurate identification of its constituent species. Adelpha species occur fi-om far northwestern U.S.A. throughout Central and South America to Uruguay, and are common and conspicuous members of the forest butterfly fauna in a wide variety of life zones, firom sea level to 3000m. They constitute the most diverse genus of nymphalid butterflies in most lowland forests, and may be important models for Batesian mimics in other butterfly subfamilies. The larvae feed on a diverse assemblage of plants (Aiello, 1984; Ackery, 1988) and the morphology of the immature stages is highly variable (Miiller, 1886; Moss, 1933; Aiello, 1984, 1991; Otero & Aiello, 1996). The last published systematic revision of the entire genus Adelpha was that of Fruhstorfer (1915), followed by a pictorial review of the genus by D'Abrera (1987). While undertaking a study of Ecuadorian Rhopalocera in 1993, it became apparent that identification of the majority of Adelpha taxa was not possible using published sources, and that there was no general consensus as to the number of species in the genus or their relationships. With the capture of undescribed species of Adelpha by myself and Jason Hall in Ecuador in 1993, 1994 and 1995, it seemed that a systematic review of the genus was desperately needed as a baseline for further research into this diverse group of 5

PAGE 13

6 butterflies. In addition, there has been a growing interest in recent years in the monitoring and conservation of Neotropical butterfly faunas, and the first step to cataloguing the distribution of such tropical biodiversity is a firm taxonomic basis. Since Adelpha is perhaps the largest genus in the Nymphalidae, and is widespread and common in all forested habitats throughout the Neotropics, it is also an ideal genus to examine general trends in diversity and endemism in nymphalid butterflies. Materials and Methods Specimens Examined A total of 20,730 specimens of Adelpha were examined and identified in pubhc and private collections, including 16,426 males and 4,304 females. The collection acronyms used throughout the text are listed below, and those in which I have examined material are marked with an asterisk. Other collections examined fi-om which specimens are not listed include the Museo de Ciencias Naturales and Pontificia Universidad Catolica, both in Quito, Ecuador. Alberto Diaz Frances collection, Mexico, Mexico Andrew F. E. Neild collection, London, UK Artur Jasinski collection, Piastow, Poland Allyn Museum of Entomology, Sarasota, FL, USA American Museum of Natural History, New York, NY, USA Andres Orellana collection, Merida, Venezuela Booth Museimi of Natural History, Brighton, UK Natural History Museum, London, UK (M=Main coll.; R=Rothschild coll.; T=Type coll.; Moss=Moss coll.; A&B=Adams & Bernard coll.) David Trembath collection, Surrey, UK De la Fuente collection, Caracas, Venezuela Eurides Furtado collection, Diamantino, Brazil Ernesto W. Schmidt-Mumm collection, Bogota, Colombia Fundacion e Instituto Miguel Lillo, Tucuman, Argentina Florida State Collection of Arthropods, Division of Plant Industry, Gainesville, FL, USA George T. Austin collection. Las Vegas, NV, USA Harold & Jose L. Skinner collection, Margarita, Venezuela Jofl&e Blanco collection, San Cristobal, Venezuela Juan Carlos de Sousa collection, Barquisimeto, Venezuela Jean F. Le Crom collection, Bogota, Colombia JeanYves Gallard collection, French Guiana ADF * AFEN AJ * AME * AMNH AO * BMB * BMNH * DAT DLF EF * ESM FIML * FSCA GTA HS JB JCS * JFL JYG

PAGE 14

7 KWJH Keith R. Willmott & Jason P. W. Hall collection, Gainesville, FL, USA LCB L. & C. Brevignon collection, French Guiana LMC Luis M. Constantino collection, Cali, Colombia MALUZ Museo de Artropodos Terrestres, La Universidad de Zulia, Venezuela MCZ Museum of Conparative Zoology, Harvard University, Cambridge, MA, USA MHNG Musee d'Histoire Naturelle de Geneve, Geneva, Switzerland MHNUC Museo de Historia Natural, Universidad de Caldas, Manizales, Colombia MIZA Museo del Institute de Zoologia Agricola, Universidad Central de Venezuela, Maracay, Venezuela MJP Mike J. Perceval collection, Surrey, UK MJWC Matthew J. W. Cock collection, Reading, UK MNHN Museum National d'Histoire Naturelle, Paris, France MUSM Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru MZUJ Muzeum Zoologiczne Uniwersytetu Jagielloiiskiego, Krakow, Poland PJD Philip J. DeVries collection, Eugene, OR, USA R Romero Family collection, Maracay, Venezuela RDM Roberto de la Maza collection, Mexico, Mexico RM Raymond Murphy collection, Torquay, UK RNHL Rijksmuseimi van Natuurlijke Histoire, Leiden, Netherlands SMTD Staatliches Musexmi fur Tierkunde, Dresden, Germany TP Tomasz Pyrcz collection, Warsaw, Poland; incorporated into the MZUJ UFP Universidade Federal do Parana, Curitiba, Brazil UNAM Instituto de Biologia, Universidad Nacional Autonoma de Mexico, Mexico, Mexico USNM National Museum of Natural History, Smithsonian Institution, Washington, DC, USA ZISP Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia ZMUC Zoologisk Museum, Universitets Copenhagen, Copenhagen, Denmark ZMHU Zoologisches Museum, Humboldt Universitat, Berlin, Germany ZSBS Zoologisches Sammlung des Bayerischen Staates, Munich, Germany I list the specimens examined under each taxon by country in an order which corresponds to their geographical position, starting with the USA, then Mexico, east to Panama, from Venezuela south along the Andes to Argentina, Brazil, Paraguay, Uruguay, Guyana, French Guiana, Surinam, Trinidad and the Caribbean. I list provinces, states or departments within each country, and localities within each of these subdivisions, in alphabetical order. I regard words such as "Rio", "Quebrada", "Mount", "Volcan", "Puerto", "via" and "El/La" as part of the name, but not relational phrases such as "nr.", 'Vic", "env.", "region", "above", "head", all compass points (N., E., S., W. etc.) and "km." or "mi." from a locality. Collection sites within each country were located using atlases in addition to Brown (1941), Lamas (1976), Brown (1979) and Jenkins (1983, 1984, 1985a,b, 1986, 1987, 1989, 1990). Sites that I was unable to precisely locate are listed at the end of each country subdivision, country or under "country unknown". I have

PAGE 15

8 used the names of localities as given on the original label, and not altered them when the locality has undergone a subsequent change of name, as in the case of Para to Belem. Countries, states, provinces or departments represented only by specimens with label data that I consider erroneous are enclosed in quotes. For each taxon I list the number, sex (m male, ffemale), collection locality and altitude when given, date, any notes of interest on accompanying labels, and the depository of all specimens examined, hi a number of cases specimens were examined which are transitional between subspecies, or exhibit other significant external phenotypic variation, and these are marked with various symbols keyed immediately at the top of the list of specimens for each taxon. For each species I list the total number of specimens examined, and the number of each sex in parentheses for each taxon and country, hi a very few cases I have listed specimens examined only by photograph, where these are unique or of very rare taxa, and I state where this is the case in the taxonomic discussion. Identiflcation of Type Specimens One reason for confusion regarding the application of Adelpha names to taxa is the difficulty in identification of type specimens, and I have attempted to locate all extant type material of available Adelpha names. The great majority of names were based on type series rather than a single type specimen, and there is therefore the possibility of mixed type series for many taxa. Most early authors gave no indication of the number of specimens on which they were basing their descriptions, and their material must therefore be regarded as syntypic, even if only a single specimen is known. I designate lectotypes or neotypes for all names that are not sufficiently well described to apply unambiguously to a particular taxon, and lectotypes for those that clearly apply to several taxa on the basis of the syntypic series. It appears that no authors labeled Adelpha material on which they based names as type specimens prior to the Felder brothers in the 1860s, and the same is true of many

PAGE 16

9 subsequent authors. In the absence of an original figure sufficiently detailed to identify the specimen on which it was based, or a specimen labeled specifically as a type by the original author, I believe there is an element of doubt surrounding the validity of all other type material. I have therefore tried to base my application of a name first and foremost on the original description, and to examine critically all labeled and unlabeled potential type material in the light of this description. I have been able to locate type material in the collections examined of almost all described Adelpha taxa. Notable exceptions are certain types of very early authors such as Lirmaeus, Cramer, Godart, Drury and Walch, and these specimens are, in most instances, probably lost or destroyed. The Linnaeus Lepidoptera types are currently under study by Honey (pers. comm.) and located in London, Cramer types are either in the RNHL or the BMNH (VaneWright, 1975), Godart types should be in the MNHN (Horn & Kahle, 1935-37), Drury types should be in the BMNH, and the whereabouts of Walch's collection is imknown (Horn & Kahle, 1935-37). Of these, only the descriptions of Linnaeus and Walch are too vague to permit unambiguous identification, and I discuss these problems under the appropriate taxa. The collections of most of those authors who subsequently published descriptions oi Adelpha are now at the BMNH, including those of Hewitson, Butler, Bates, Boisduval, the Felder brothers, Godman and Salvin, and Fruhstorfer (Horn & Kahle, 1935-37). In the case of these authors, I identified type specimens using the original description and figure, specimen label data indicating the collection of origin, the locality, the collector and any other important information, particularly hand-written labels of the original author identifying the taxon. Horn & Kahle (1935-37) figure examples of hand-written labels of Hewitson, Butler, the Felders, Boisduval and Fruhstorfer, in addition to other lepidopterists who published on Adelpha. The Felders also applied a particular type of label to their type specimens, which bears a gold and a black stripe and the name, and recognising their type material presents no difficulty. Identification of Fruhstorfer type material is, however, of^en problematic, for several reasons. It appears that the original

PAGE 17

10 locality labels on many Fruhstorfer specimens were subsequently removed and new, typed labels indicating only the country of origin substituted, probably prior to the sale of the collection. Thus the originally cited type locality often does not match the label data. Putative Fruhstorfer type specimens may bear either or both of a red, typed "TYPE" label and a label with the identification of the taxon written in Fruhstorfer' s hand. Often only one of these labels is present, but several specimens that are not types also bear such labels. Occasional specimens lack either of these labels and have been identified as syntypes through other information in the original description. Fruhstorfer described a number of taxa based on specimens in Rothschild's collection at Tring Museum, which is now also housed at the BMNH, separate fi-om the main collection. A small number of Fruhstorfer' s types are to be found at various other European museums, and I am gratefiil to Gerardo Lamas for providing photographs of such specimens. Given the uncertainty surrounding the validity of type material of many older names, I have tried to list fiill specimen label data for such types. Full label data are enclosed by quotes, with data on separate labels separated by a "//". Occasionally I have added supplementary information on the type of label in square parentheses. I use the abbreviations HT, AT, ST, PT, LT and PLT to refer to holotype, allotype, syntype, paratype, lectotype and paralectotype respectively. I regard the type locality of a taxon as being that cited in the original description, even if it is clearly erroneous and regardless of any ahemative indication on the label of a type specimen, which may have been replaced at a later date. The type locality is cited for each taxon as "TL", while for names where I have designated a lectotype or neotype, I also give the original type locality as "OTL". I have enclosed in square parentheses type localities that were not cited in the original description itself, but in the title or indirectly given elsewhere in the work (in the case of Fruhstorfer (1913, 1915), localities were given in the text, published in 1915, for names that appeared first in 1913), in addition to the country, if this was not originally given.

PAGE 18

11 Field Work In addition to museum research, I conducted field work in six Neotropical countries, including Mexico (3 weeks), Guatemala (1 week), Costa Rica (4 weeks), Venezuela (3 weeks), Bolivia (3 weeks), and Ecuador (17 months). Many Adelpha specimens in museimis are vaguely and/or unreliably labeled, and establishing certain sympatry or otherwise of phenotypes is difficult when based exclusively on such data. The purpose of this field work was to gather reliable information on the sympatry of phenotypes, to collect series of specimens to study the stability or variability of wing pattern characters and to make and record ecological observations of possible taxonomic importance. I decided to concentrate my efforts in a single country, Ecuador, which contains 69% of the species in the genus, since it seemed more usefial to gain a deep understanding of the fauna of a single country rather than a superficial acquaintance with the faunas of several. Indeed this proved to be an effective strategy, and some of the most intractable taxonomic problems in the genus, such as the true diversity and identification of members of the Adelpha serpa group, and the taxonomic status of A. iphiclus and A. iphicleola, were solved through collections made in eastern Ecuador. In addition to making collections of specimens, I attempted to record as accurately as possible the elevational distributions of species throughout the country, to examine patterns of endemism with altitude. I also made observations on adult food sources and the microhabitat preferences, perching and flight behavior of species, to ascertain whether phenotypically similar species occur microsympatrically, as predicted if Adelpha are involved in mimicry. I conducted field work in Ecuador on both slopes of the Andes, fi-om sea level on the western slope and 250m on the eastern slope to above the upper elevational limit of Adelpha species, approximately 3000m. Sites included as wide a range of topography, vegetation and microhabitat types as possible, including both flat and undulating terrain, paths along ridge tops, hill sides and river sides, in primary to heavily disturbed

PAGE 19

12 secondary growth habitats, ranging from dry deciduous forest to pluvial forest, premontane rain and montane cloud forest. Sites were visited from 6.30 a.m. to 5 p.m., when Adelpha activity ceased. I made supplementary observations of individuals using a pair of Canon 8x32mm waterproof binoculars, and used two techniques to capture specimens, an entomological net with extension handles of up to 5.5m, and baited fraps. Traps were of the Van Someron-Rydon design as figured by Sourakov & Emmel (1995) and Austin & Riley (1995), and were deployed in a variety of microhabitats, including forest edges, river sides, forest light gaps and the forest subcanopy. Traps were set at a variety of heights above the ground up to 15m, by use of polyethylene string looped over branches by a lead fishing weight. I used two types of bait; three to five day old fermenting banana, and one to three week old rotting fish, of a variety of fresh and saltwater species. Different species of fish produced no noticeable alteration in the atfractiveness of the bait. Traps were patrolled approximately once per hour, and carefiilly lowered to check for individuals resting on the outside of the netting, a common occurrence in Adelpha. Once I was familiar with the fauna, I was able to identify and release the great majority of specimens in the field, and these sight records are listed under Additional locality data below each taxon. Specimens of taxonomic interest were killed by a pinch to the thorax and stored in glassine stamp envelopes in a wooden entomological box, and subsequently prepared in the U.S.A. Morphological Study Adults. The wing patterns of all species were studied through examination of . material in collections. I examined the male genitalia of all species with the exception of A. herbita, of which there are no known specimens, and Adelpha sp. nov., of which the single specimen I have seen has already been dissected and the dissected genitalia subsequently lost. I have tried to examine the nominate subspecies of each species, and any others that exhibit significant phenotypic differences and/or have been regarded in the

PAGE 20

13 past as distinct species. In the majority of cases, I examined several specimens of each species to assess intraspecific variation. Due to the rarity of females of many species, my choice of dissections was limited, both of the species I studied and the number of specimens. Specimens dissected are listed in Appendix A. I also examined the wing venation for all species except A. herbita and A. sp. nov., and the morphology of the legs, antennae and labial palpi for representative species from all species groups. Appendages and genitalia were prepared for dissection and study by soaking in hot 10% KOH for 1530 minutes, being subsequently stored in glycerol. Drawings were made using Wild M4A stereomicroscope at 30x magnification and a Wild camera lucida. Wing venation was studied and drawn using the same equipment as the genitalia. Terminology for the wing venation follows Comstock 8c Needham (1918), and I refer to wing cells by the vein that Hes immediately above (anterior) to that cell. Genitalic terminology follows Klots (1956), and I use the term "clunicula" (of Fruhstorfer (1915)) to refer to the dorsally directed projection on the inner edge of the male genitalic valvae, which is typically adorned with numerous spines on the inner surface. Immature stages. I was able to obtain preserved material of various immature stages of a few species on loan from individuals and public institutions. Such material typically consisted of cast head capsules, larval skins and pupal cases, and in a very few cases, dried larvae. The morphology was studied using the same microscope as for adult material; due to the rarity of preserved material, it was not possible to study it using a Scanning Electron Microscope. Color slides of the immature stages for several species were given to me by Jim Mallet, and published illustrations and sketches were also examined. Taxonomic concepts In many cases the taxonomy that I have adopted, at both the specific and subspecific level, differs significantly from that of previous workers. At present, there is

PAGE 21

14 much debate as to what constitutes a species, both in theoretical and practical terms, and even as to whether the term species has any true meaning at all. The criteria used to define a species largely depend on which species concept one adopts and the primary goal of the classification that one hopes to achieve. Many also believe that the term subspecies also has no clear definition and is meaningless (Collins, 1992). It is my belief that the primary purpose of a classification, at least at the level of species and subspecies, is to be of use to biologists who are involved in research on the organisms. A classification can be of use in two principal ways; it allows effective communication among researchers and ready identification of taxa, and it is heuristic, that is it can be used to predict unknown biological traits in taxa when the traits are known in a related taxon. There are two principal competing species concepts, the biological species concept, as championed by Mayr (1942), and the phylogenetic species concept, as discussed by Wiley (1981); other species concepts are discussed by Collins (1992). Of these two concepts, only the biological provides a means to test whether or not a taxon constitutes a species, through its sympatry and the existence of a consistent genetic and phenotypic gap with relation to its sister taxon. The phylogenetic concept argues that any taxonomically diagnosable monophyletic clade may be regarded as a species, but provides no guidelines as to where to draw the line between the subspecies, species, genus or higher level taxon. I regard the criterion of monophyly of species as unhelpfiil in classifying Adelpha, since it would either lead to the recognition of many more "species", thus obscuring their true relationships, or cause probably genetically isolated and potentially sympatric taxa to be regarded as the same species (see discussion under A. ethelda and A. epione). However, it is also true that while it is usually clear what is meant by the term species within a single locality, there is no way to determine whether isolated populations, which may or may not differ phenotypically, represent distinct biological species, and this is perhaps the most fi-equently cited criticism of the biological species concept.

PAGE 22

15 There has been relatively little research directed towards the genetic distinctness of isolated sister taxa, but there is evidence that even parapatric populations that are known to hybridise may maintain their genetic integrity in the hybrid zone (Jiggins et al., 1996; Davies et al, 1997), and thus be regarded as distinct evolutionary lineages. Such research is lacking for all Adelpha, and I have therefore used characters that are reliable in distinguishing sympatric species, and that show little or no variation throughout the range of monotypic species, as evidence as to whether or not two allopatric populations should be treated as distinct species. In the great majority of cases, I have treated isolated populations that are closely allopatric and occur in similar habitats at similar elevations, and that do not differ from each other by any "reliable" character (as defined above), as constituting the same species, while remaining aware that all such taxonomic hypotheses require testing in the field. The imiting of such populations as a single species provides two important pieces of information: that they are not known to be sympatric, and that they are considered sister taxa, even in the absence of a synapomorphy (such species were termed "paraspecies" by Ackery & VaneWright (1984)). They therefore almost certainly also share similar biological traits and the classification is more heuristic. This approach is essentially the same as the biogeographical species concept utilised by Tyler et al. (1994), and I regard it as the most usefial first approach to classifying Adelpha. In one case I have departed from this approach, namely in my freatment of Adelpha epione and Adelpha ethelda, and I discuss my reasons for this under each respective species. My approach to classifying Adelpha at the species level has led to the grouping in some cases of taxa which have very distinct dorsal wing patterns and have in the past been placed in completely different species groups. Examples include A. ximena ximena and A. ximena mossi, A. messana messana and A. messana delphicola, and A. malea malea, A. malea aethalia and A. malea fundania. Typically these taxa show little variation over their range and specimens transitional to other subspecies, if known, originate from a relatively very narrow zone of apparent hybridisation. Another classic

PAGE 23

16 example is the common A. lycorias; individuals of A. lycorias lara are phenotypically indistinguishable from Trinidad to northern Argentina, yet in eastern Colombia specimens clearly transitional to A. lycorias melanippe occur, which is also phenotypically stable throughout much of central Colombia. I retain subspecies names for these populations since it aids their identification and recognises that they may be demonstrated to be distinct species, and such cases are similar to many species in mimetic nymphalid genera in the subfamilies Heliconiinae and Ithomiinae. In the examples mentioned above, if we assume that the dorsal wing patterns serve a mimetic function (see below), then narrow hybrid zones are explicable in terms of strong purifying selection by predators on either side of the zone of contact (Mallet, 1993; Joron & Mallet, 1998). At the other end of the spectrum occur taxa which exhibit continuous, clinal variation in several characters over their geographic range. While opposite ends of the cline may be phenotypically quite distinct, since there is no clear character to identify specimens from intermediate localities I regard the entire population as a single taxon. In other cases, even when there is some clinal variation, there is a consistent phenotypic gap which separates two populations. I believe it is worthwhile recognising such phenotypic gaps by application of subspecies names to the two populations in question, since it highlights geographic areas where the fragmentation of populations may frequently occur. Assessing whether or not geographic variation is discontinuous is complicated in many cases due to the lack of series of reliably and precisely labeled specimens in collections. I have tended to adopt a conservative approach to situations where I have insufficient material to ascertain the nature of geographic variation, since retaining subspecies names provides more information than synonymising them, and I leave assessment of their validity to field workers in each region.

PAGE 24

17 Systematics History of Classification The first Adelpha species, probably all originating from the Guianas, were described by Linnaeus in 1758, in the genus Papilio. Linnaeus described three names, two in his phalanx Barbarus, and one in the Phalerati section of the phalanx Nymphalis, both groups based on wing shape and variable wing pattern characters. There are taxonomic problems surrounding all three of these names, discussed under Adelpha cytherea and Adelpha iphiclus. Probably more species were represented in collections in the latter half of the eighteenth century than is reflected by the description of names, since early authors regarded even relatively distinct taxa, such as Adelpha naxia, A. iphiclus and A. serpa, as constituting the same species. Following single descriptions in the genus Papilio, accompanied by figures, by Walch (1775) and Cramer (1777, 1780), Latreille (1809) described leucophthalma in the genus Nymphalis Latreille, 1804. Hiibner ([1819]: 42) introduced the name Adelpha for eight listed species, including A. mesentina, A. iphiclus (as A. basilis), A. serpa diadochus (as A. iphicla), A. plesaure (described shortly after by Hubner (1823)), A. cocala and A. cytherea. Hubner also listed Adelpha elea, described by Linnaeus as A. eleus, which is here considered a synonym oiA. cytherea, and A. phliase, which is in fact a member of the family Riodinidae, Synargis phliassus (Cramer). He omitted A. capucinus, which has only been mentioned twice since its original description prior to this revision, and A. leucophthalma, possibly because it lacked the vertical dorsal bands typical of the other Adelpha. As can been seen by the inclusion of a riodinid, Hubner based the genus purely on wing pattern and shape, and failed to name a type species, placing it in his Familia D, which contained genera with a typically brown ground color to the wings. Scudder (1875) subsequently selected die first named species, Papilio mesentina, as the type species of Adelpha. Hubner's genus Adelpha was largely ignored by subsequent authors, with the notable exception of several German lepidopterists, and Godart ([1824]: 335-342) soon after treated the species in the genus

PAGE 25

18 Nymphalis Latreille, with a number of other unrelated Nymphalidae. Godart however provided an important review of the previous taxonomy of all species (except of course A. capucinus) with supplementary descriptions, and made the first major contribution to the description of Adelpha taxa, with five new species. Two years later, Boisduval (1836: pi. 8, fig. 4) introduced the new generic name Heterochroa, along with a new species, Heterochroa serpa. There was no description accompanying the color plate or indication of other included species, and serpa therefore becomes the type species of Heterochroa by monotypy (Hemming, 1967). Geyer (1837) was the first author, with the exception of Hiibner, and the last for several decades, to use the name Adelpha, when he described the species bredowii. Although the generic name of this species has oscillated ahnost continuously since its original description between Adelpha, Heterochroa and Limenitis Fabricius, the species is most closely related to members of the A. serpa group, as discussed below, and belongs in the genus Adelpha as conceived here. The year 1847 saw the publication of Hewitson's first paper on Neotropical butterflies and the start of a period of intensive activity in Neotropical taxonomy, including the genus Adelpha (see Fig. 1). Hewitson (1847) described eight valid species in Heterochroa, fi-om throughout the Nootropics, including the formerly unexplored Andean region, and three years later in 1850 a further four. The number of described species in the genus almost doubled during this time. His descriptions were notable for their great attention to detail and their accurate figures, and he was the first to appreciate the great importance of ventral wing pattern characters, particularly in the forewing discal cell, in distinguishing species. Westwood (1850: 276) provided the first detailed description of the external morphology of Adelpha, though he also mistakenly used the name Heterochroa, listing Adelpha as a synonym. He did not list any character to define the genus, but stated that it was closely related to Limenitis, and possibly the apaturine genus Apatura. He listed the majority of described species and their synonyms, with the exception capucinus and bredowii, the latter being placed in the genus Limenitis. The

PAGE 26

19 name marse was also listed under Heterochroa, but Westwood (1850: 304) subsequently noted that it was in fact a member of the genus Doxocopa. Fig. 1 . Total number of valid described species and subspecies of Adelpha since the time of Linnaeus (1758). The name Heterochroa gained widespread acceptance in the lepidopterological community in the two decades following the publication of Hewitson's and Westwood's works, during which time the number of described species of Adelpha almost doubled again. This was due largely to the papers of Bates (1864, 1865, 1866), Butler (1866), Hewitson (1867), and the Felder brothers (1861, 1862, 1867). hi particular, Felder &

PAGE 27

20 Felder's (1867) report on the collections made during the voyage of the Frigate "Novara" around the Earth is notable for containing the descriptions of nine valid Adelpha species, a number second only to Fruhstorfer (1913, 1915). The majority of these species originated along the northern coast of South America, with a nimiber from mountainous areas in Venezuela and Colombia. However, the lack of figures in this work resulted in much taxonomic confusion in years to come. Kirby (1871) finally correctly restored the name Adelpha and placed Heterochroa in synonymy, and provided a thorough synonymic catalogue of all published names in the genus. Although many of his taxonomic conclusions were highly questionable, this catalogue provided the basis for fixture research. Throughout the 1870s Butler was the most noticeable author of new Adelpha taxa, and the next important work to treat a large portion of the genus was that of Godman & Salvin's (1884) review of Central American species. They described in fiulher detail the external morphology and appear to have been the first authors to have examined the male genitalia, making two principal observations; that all species were remarkably uniform, and that species of Limenitis were scarcely distinguishable. They also figured many species, the majority of which were correctly identified, and arranged the species in the form of a key. One might regard this as the first attempt to define species groups within the genus, but this was not a stated goal and many of the resulting species groups are artificial. Nevertheless, their arrangement, beginning with species with entirely black hindwings and oblique or straight orange forewing bands, such as A. lycorias melanthe and A. boreas, was followed by almost all subsequent authors (Fruhstorfer, 1915; Hall, 1938; DeVries, 1987; Neild, 1996) and is reflected in the curation of most major museum collections. They also united for the first time the members of the A. serpa group. A relatively stafic period followed, unfil the arrival of Fruhstorfer (1907, 1908), culminating in the best known work on Adelpha, the revision of the genus in the Macrolepidoptera of the World (Fruhstorfer, 1915). Fruhstorfer's revision is at times

PAGE 28

21 inexplicably careless, at times surprisingly insightful. He also failed to find any single character to define the genus irrespective of Limenitis, and his arrangement of species was based largely on a variable character of the forewing venation, dorsal wing pattern, and the presence or absence of the clunicula in the male genitalia. He recognised 91 species, described eleven valid species and 28 valid subspecies, and a very large number of additional names based on insignificant wing pattern variation. He appears to have examined some type specimens of earlier authors, yet still made a great number of errors and misidentifications, many of which have been propagated through the literature until very recently. The plates are still widely used to identify Adelpha taxa, yet they too are replete with mistakes and incorrect names. Fruhstorfer's most important contribution was the recognition and description of a number of superficially similar southeast Brazilian species, and he was the first to recognise that "orange and white banded" Adelpha contained many more species than had been previously appreciated, several of which he described. It is, however, arguable whether the plethora of new names authored by Fruhstorfer, many with breathtaking brevity, resulted in more subsequent taxonomic confusion than clarification. Hall (1938) made an attempt to assess the validity of many of the names after Fruhstorfer's collection, including most of the type material, was transferred to the British Museum. Hall (1938) synonymised many of the subspecific names and also authored several himself, including two species. There followed a period of remarkable calm, a period of thirty-five years in which not a single valid Adelpha taxon was described (see Fig. 1), as attention perhaps turned more to the biology of species and the higher level classification of butterflies. Two papers by Beutelspacher (1975, 1976), in which three species of Mexican Adelpha were described, marked a brief period of interest in the group, during which a new Colombian species was also described (Steinhauser & Miller, 1977). The description of a remarkably distinct new species fi-om

PAGE 29

22 Costa Rica by DeVries & Chacon (1982) gave indication that there were still undescribed Adelpha taxa awaiting discovery. With the description of two highly distinctive species from Ecuador, Willmott «& Hall (1995) began a recent surge of interest in the group, at a time which is proving to be a veritable renaissance period for butterfly taxonomy in all groups (see Fig. 1). Neild (1996) followed with the description of nine new subspecies from Venezuela, in one of the most taxonomically important works to date on the genus. Neild (1996) was the first to recognise that the ixia" of previous authors actually included three separate species, A. messana, A. malea ixia and A. barnesia leucas, and was one of the very few modem authors to show an appreciation of the value of ventral wing pattern characters in species identification, in particular among certain phenotypically very similar Amazonian species. Willmott & Hall's (1999) description of a fiuther two new species and seven new subspecies from Ecuador, and the number of new taxa described in this work, demonstrate that our imderstanding of the diversity of Adelpha is still far from complete. Generic Relationships Adelpha is currently placed in the subtribe Limenitiditi, tribe Limenitidini, and subfamily Limenitidinae of the family Nymphalidae (Harvey, 1991). As mentioned earlier, there is no evidence that the Limenitidinae constitutes a monophyletic group (Harvey, 1991), and the relationships of the Limenitidini to the Pantropical Cyrestidini, the Neotropical Coeini and Biblidini, the latter including most of the remaining Neotropical species, are not known. The tribe Limenitidini includes a large number of species and genera distributed in both tropical and temperate regions throughout the globe, and is in all probability an ancient lineage. Chermock (1950) defined the group on the basis of a single character, the preservation of the first anal vein (lA) as a short spur at the base of the forewing cubital vein (see Fig. 2a). This character, however, is not present in one of the genera, Neptis, and

PAGE 30

23 Fig. 2. Morphology of adult Adelpha. a: A. serpa serpa, wing venation; b,c,d: base of forewing veins M2, Ml, R3-5 and R2, b, A. lycorias lara, c, A. lycorias spruceana, d, A. lycorias melanthe; e,f: forewing discocellular vein, e, A. capucinus capucinus, f, A. boreas boreas; g,h: A. mesentina, female genitalia, g, dorsal view, h, lateral view; i,j: A. mesentina, male genitalia, i, lateral view, j, inside of valva; k,I: juxta, posterior, lateral and ventral views (left to right), k, A. alala completa, 1, A. serpa celerio; m: A. diodes creton, posterior view of gnathos; n: A. serpa celerio, lateral view of aedeagus; o: A. serpa celerio, sclerotised aedeagal pad, ventral (left) and lateral (right) view; p,q: lateral view of eye and labial palpi, p, A. alala negra, q, A. mesentina.

PAGE 31

24 also occurs in certain members of the Heliconiinae, as noted by Michener (1942) and Brown & Heineman (1972), including the genera Philaethria, Dryas and Dryadula (Penz, pers. comm.). Harvey (1991) also defined the tribe on the basis of this character, while noting that a further possible character which may prove to be a synapomorphy for the tribe is the structure of the eggs (Aiello, 1984). All eggs known to date are composed of concave, hexagonal facets with setae at the interstices. Harvey (1991) stated that the eggs of members of the Partheniti were unknown, but I can confirm, from dissection of the female of Parthenos sylvia, that they also have this characteristic structure. Harvey (1991) recognised four subtribal divisions corresponding to groups of various taxonomic rank recognised by earlier authors (Chermock, 1950, and references therein; Eliot, 1978), including the Neptiti, Limenitiditi, Partheniti and Euthaliiti, all of which were defined by Eliot (1978) on the basis of wing venation. The character given by Eliot (1978) to define the Limenitiditi is the origin of the hindwing humeral vein at or before the origin of vein RS-M1-M2. However, wing venation is likely to be highly variable (it is even in a single species of Adelpha, A. lycorias, see Fig. 2), and there is no evidence whether the characters of Eliot represent synapomorphies or symplesiomorphies. While the Euthaliiti and Neptiti probably constitute monophyletic groups, on the basis of the highly distinctive early stages, particularly the larvae, in the former, and the adult morphology and early stages of the latter (Harvey, 1991), there is no evidence that the Partheniti or Limenitiditi are monophyletic; they merely include the members of the Limenitidini that do not belong in the tribes Euthaliiti and Neptiti. The Limenitiditi consists of a number of tropical and temperate species whose generic relationships are probably the most poorly understood of the entire subfamily. Most are placed in the catch-all genus Limenitis, while numerous generic names have been proposed (see Chermock, 1950), a number of which are no doubt valid. Within the New World the only representatives of the Limenitiditi, and indeed the Limenitidini, are

PAGE 32

25 Adelpha and four North American species placed either in the genus Limenitis or Basilar chia. The Genus Adelpha Characteristics The genus Adelpha includes medium to large butterflies which have essentially sexually monomorphic wing patterns, consisting of contrzisting ventral and dorsal surfaces (see Figs. 10-94). The dorsal surface always has a dark brown ground color and almost always a postdiscal band on one or both wings, which may be vertical or oblique, and orange, white, or red in color. The ventral surface typically has a black ground color which is usually obscured by much reddish brown coloring, and in addition to the postdiscal bands which appear on the dorsal surface it has a number of lighter postdiscal and submarginal Hnes and other more basal pattern elements (see Fig. 3). Despite the superficially large differences in dorsal pattern between species, all are composed of relatively few pattern elements, particularly the postdiscal band, the postdiscal series and, on the forewing of some species, the submarginal series. Identification of the origin of various areas of the dorsal wing pattern in terms of the ventral pattem elements is often important in distinguishing Adelpha species. Li almost all species the ventral pattem elements are visible, and more primitive species, such as members of the A. alala group (see Table 1), tend to have a greater complement of unmodified pattem elements. The pattem elements that I have recognised in Fig. 3 differ from those recognised by Schwanwitsch (1924) and Nijhout (1991), who both state that the nymphalid ground color is pale and pattem elements are dark. Thus, the pale postdiscal and submarginal series and postdiscal band, which play the most important role in species identification, actually constitute ground color, while the darker areas between are the "pattem". Although Nijhout (1991) rightly points out that few authors have attempted to recognise the system of Schwanwitsch (1924) in description of

PAGE 33

26 butterfly wing patterns, I have chosen a terminology for the wing pattern of Adelpha that allows discussion and comparison of wing patterns with maximum clarity. It is much simpler to refer to the pale areas of the ventral wing surface in Adelpha as pattern elements, while acknowledging that they are not by the definition Schwanwitsch (1924). It is, of course, also possible to recognise the pattern elements described by Schwanwitsch (1924) and Nijhout (1991) in Adelpha; more basal elements are obvious, while the distal band of the central symmetry system borders the basal edge of the postdiscal band, and the dark areas between the postdiscal series, the outer postdiscal and inner submarginal series, and the submarginal series correspond respectively to elements g, h (border ocelli) and I (parafocal element) (Nijhout, 1991: Fig. 2.17). postcellular bar venal stripe intervenal stripe postdiscal band postdiscal band (upper, above M3) postcellular markings postcellular bar fourth cell bar third cell bar second cell bar first cell bar basal streak basal streak first cell bar second cell bar third cell bar inner postdiscal series outer postdiscal series inner submarginal series outer submarginal series postdiscal band (lower, below M3) iimer postdiscal series outer postdiscal series inner submarginal series outer submarginal series Fig. 3. Wing pattern elements oi Adelpha, indicating terminology adopted in the text. J

PAGE 34

27 Morphologically Adelpha is remarkably homogeneous, and detailed descriptions of the wing venation, antenna, eyes, labial palpi and legs are given by Westwood (1850) and Godman & Salvin (1884). The wing venation (Fig. 2a) varies little in the genus, except that the discocellular vein may or may not be present on the forewing, while its juncture with vein M3 may be above or below the origin of vein Cul (Fig. 2e,f). The position of the base of vein R2 is also variable within A. lycorias (Fig. 2b-d). The eyes are variably setose, with montane species of the A. alala group having particularly long and dense setae, while certain species, such as A. bredowii and A. nea, have naked eyes. The labial palpi may or may not have a pronounced lateral black stripe (Figs. 2p,q), while the thorax is either entirely pale in color, or has dark bands where the legs rest against the body. Otherwise all of these structures show little other morphological variation throughout the genus. The male (Figs. 2ij, 95-177) and female genitaUa (Figs. 2g,h, 178-249) are also both relatively homogeneous. The male genitalia show interspecific differences particularly in the shape of the valva and distribution of terminal spines, and may or may not have a pronounced projection fi-om the inner edge of the valva (Fig. 2j), termed the "clunicula" by Fruhstorfer (1915). The imcus, tegumen, gnathos and aedeagus vary little, except that members of the A. serpa group have a sclerotised pad in the aedeagus on the dorsal surface of the vesica. The juxta is also relatively imiform, with slight differences in the A. serpa group (Fig. 2k,l). The female genitalia vary somewhat in size, even considering size differences of the adults, but also vary little morphologically. The lamella postvaginalis and antevaginalis are wrinkled plates which are variably sclerotised, the ostium bursae is always a simple ring which tends to be wider in members of the A. serpa group, the ductus seminalis always originates dorsally near the ostium bursae and the relative lengths of the ductus bursae and corpus bursae vary little. The corpus bursae typically has a pair of sclerotised bands (Fig. 2g,h) which may be dorsal, on the right hand side or ventral, with both inter and intraspecific variation in the position and spacing of

PAGE 35

28 the bands. The degree of sclerotisation also varies, as does the size of the spines on the inner surface of the bands, with members of the A. serpa group having the most heavily sclerotised and coarsely spined bands. The terminal tergite of the female genitalia is variably sclerotised at the anterio-ventral edge, in one species (A. thoasa) forming a small pouch. Similar structures occur in the Asian genus Neptis. While Adelpha has been regarded as a distinct genus by the great majority of authors, Chermock (1950) placed it as a subgenus of Limenitis, in which he was followed by authors in the following few decades. This was due to the entire absence of morphological characters by which to define Adelpha, and the retention of the genus by all previous authors has been on the basis of its isolated geographical distribution and the overall similarity of most species. Despite an intensive examination of the external and internal morphology of both males and females, I have also been unable to find any morphological characters to define the genus. I have, however, located several wing pattern characters which suggest that the genus is monophyletic. All species of Adelpha have a dark brown longitudinal line extending fi-om the base of cell 2A to the middle of the anal margin in this cell, with the exception of several species clearly closely related to species that do have the character. In many species this line may be fiised to a band of color which fills the anterior half of the cell, and in several it is broken in the basal half of the wing. Species that lack this line also have all the markings in the basal half of the wing reduced, and presumably the line has been secondarily lost. This line appears to be either an intervenal stripe (Nijhout, 1991) or the posterior half of a venal stripe along vein 2A that has become detached fi-om the vein and displaced posteriorly. Almost all species also have a venal stripe at vein 3 A on the ventral hindwing, which may also be split along the vein into two halves and variously modified or reduced, being almost invisible in members of the A. alala group. Almost all species of Adelpha have the postdiscal series fiised on the dorsal surface of the forewing to form a subapical marking; in other limenitidine genera the postdiscal series are clearly distinct. Again, the species oi Adelpha

PAGE 36

29 in which the postdiscal series are not fused are derived and closely related to species that do have this character. A further possible synapomorphy is the streak at the base of the ventral hindwing discal cell, apparently corresponding to the basal symmetry system of Nijhout(1991). Taxonomy and identification I recognise 85 species and 207 taxa in the genus Adelpha, and I present a synonymic checklist below in an order fi^om primitive to advanced, largely following the phylogeny presented in Chapter 3. I include names proposed as quadrinomials even though such names are excluded by the Code, since in at least one instance {syma Fruhstorfer, a synonym of A. seriphia godmani) subsequent authors have made the name available. Since I do not claim my bibliographic review to be exhaustive, this may also be true of other names, and all such names are listed in the Review of the genus below with their associated type material. I also discuss the spelling and status of Fruhstorfer names in the introduction to the generic Review below. One species and four subspecies are undescribed, three of these due to a lack of material, and two due to their impending description by other authors. Adelpha Hubner ([1819]: 42). Type species by selection by Scudder (1875: 102): Papilio mesentina Cramer (1777: 102, pi. 162, figs. -guatemalensis (Carpenter & Hobby, 1945) syn. -sentia Godman & Salvin, 1884 stat. nov. B, C). =Heterochroa Boisduval (1836: pi. 8, fig. 4) Type species by monotypy: Heterochroa serpa Boisduval (1836: pi. 8, fig. 4) ("-" denotes a subspecies, "-" a synonym and " — " an unavailable name) bredowii Geyer, 1837 -eulalia (Doubleday, [1848]) nea (Hewitson, 1 847) —campeda FruhstoTfei, 1915 nov. -califomica (Butler, 1865) diodes Godman & Salvin, 1878 -creton Godman, 1901 herbita Weymer, 1907 zea (Hewitson, 1850) —serpentina Fruhstorfer, 1915 —/a/pe/a Fruhstorfer, 1915 paroeca (Bates, 1864) -emathia (R. Felder, 1869) stat. rest. paraena (Bates, 1865) -massilia (C. Felder & R. Felder, 1867) -rey/Neild, 1996 -lecromi Willmott, 1999 ssp. nov. radiata Fruhstorfer, 1915 -myrlea Fruhstorfer, 1915 -gilletella Brevignon, 1995 -aiellae Willmott & Hall, 1999 -explicator Wilmott & Hall, 1999 serpa (Boisduval, 1 836)

PAGE 37

—damon Fruhstorfer, 1913 -omamenta Fruhstorfer, 1915 -celerio (Bates, 1864) —diademeta Fruhstorfer, 1913 -phintias FmhstorfeT, 1913 -Juj/ioe Fruhstorfer, 1913 -rfjWoc/iM5 Fruhstorfer, 1915 -timehriHM, 1938 —florea Brevignon, 1995 seriphia (C. Felder & R. Felder, 1867) -pione Godman & Salvin, 1884 -a^M/Z/ja Fruhstorfer, 1915 —naryce Fruhstorfer, 1915 -therasia Fruhstorfer, 1915 -egregia Rober, 1927 -gotfmani Fruhstorfer, 1913 —syma Fruhstorfer, 1915 -barcanti Wilhnott, 1999 ssp. nov. Aj'as (Doyere, [1840]) -viracocha Hall, 1938 -hewitsoni Wilhnott & Hall, 1999 fl/fl/a (Hewitson, 1847) -completa Fruhstorfer, 1907 —rifta Fruhstorfer, 1915 -negra (C. Felder & R. Felder, 1862) — eAr/iarrfi Neuburger, 1907 syn. nov. —albifida Fruhstorfer, 1907 syn. nov. —cora Fruhstorfer, 1907 syn. nov. —fillo Fruhstorfer, 1907 syn. nov. —negrina Fruhstorfer, 1913 syn. nov. —praecaria Fruhstorfer, 1915 —privigna Fruhstorfer, 1915 syn. nov. aricia (Hewitson, 1 847) -serenita Fruhstorfer, 1915 stat. rest. -portunusli2i\\, 1938 corcyra (Hewitson, 1847) -aretina Fruhstorfer, 1907 -collina (Hewitson, 1 847) stat. rest. —epidamna (C. Felder & R. Felder, 1867) -dognini Wilhnott, 1999 ssp. nov. -salazari Willmott, 1999 ssp. nov. fracto (Butler, 1872) p/rAj's (Bates, 1864) —vodena Fruhstorfer, 1915 stat. nov. donysa (Hewitson, 1847) -roe/a (Boisduval, 1870) -albifilum Steinhauser, 1974 stat. nov. fessonia (Hewitson, 1 847) -lapitha Hall, 1929 stat. nov. -cestus (Hewitson, 1 847) -emestoi Willmott, 1999 ssp. nov. gelania (Godart, [1824]) -arecosa (Hewitson, 1 847) ca///p/iaii^ Fruhstorfer, 1915 poltius Hall, 1938 mythra (GodzTt, [1824]) -zeba (Hewitson, 1850) basiloides (Bates, 1865) -lydia (Butler, 1865) stat. rest. -lemnia (C. Felder & R. Felder, 1867) -lativittata Staudinger, 1886 syn. nov. -caelia Fruhstorfer, 1915 syn. nov. plesaureHuhnex, 1823 -Aererf/'a Fruhstorfer, 1915 —anto/Jiae Fruhstorfer, 1915 -phliassa (Godart, [1824])-euboea (C. Felder & R. Felder, 1867) —imp/icafa Fruhstorfer, 1915 -bartolme Fruhstorfer, 1915 syn. nov. —cerac/ia/e5 Fruhstorfer, 1915 —5/rona Fruhstorfer, 1915 -pseudomalea Hall, 1938 -symona Kaye, 1925 ^avino Fruhstorfer, 1915 /a/c/peii«/s Fruhstorfer, 1915 —perga Fruhstorfer, 1915 stat. rest. thoasa (Hewitson, 1850) -jiV/a Fruhstorfer, 1913 —cuyaba Fruhstorfer, 1915 -gerona (Hewitson, 1867) stat. rest. — brevifascia Talhot, 1928 -calliphiclea (Butler, [1870]) stat. nov. -?nan//w Fruhstorfer, 1915 —za/ffia Fruhstorfer, 1915 thessalia (C. Felder & R. Felder, 1867) -indefecta Fruhstorfer, 1913 stat. nov. —mincia Hall, 1938 syn. nov. -cesilas Fruhstorfer, 1915 —ampla Hayward, 1935 syn. nov. i/;/fic/«s (Linnaeus, 1758) —basilea (Cramer, 1777) stat. rest. -^)aii7« Hubner, [1819] --^Ma/is Fruhstorfer, 1915 —exaw/wa Fruhstorfer, 1915 —pAarae Fruhstorfer, 1915 -ephesa (Menetries, 1857) —gellia Fruhstorfer, 1915 —abylina Fruhstorfer, 1915 -estrecha Wilhnott & Hall, 1999 iphicleola (Bates, 1 864) —massilides FnihstoifeT, 1915 -j/7/i/me/!era Fruhstorfer, 1915 -thessalita Wilhnott & Hall, 1999 -gortyna Fruhstorfer, 1915 -/ewcafej Fruhstorfer, 1915 --
PAGE 38

31 -deborah Weeks, 1901 stat. rest. -neildi Willmott, 1999 ssp. nov. er/ieWa (Hewitson, 1867) -zalmona (Hewitson, 1871) stat. nov. -sophax Godman & Salvin, 1878 stat. nov. -eponina Staudinger, 1 886 stat. nov. — vo/m/jw Fruhstorfer, 1915 -galbao Brevignon, 1995 stat. nov. -ssp. nov. epione (Godart, [1824]) -agz7/a Fruhstorfer, 1907 syma (Godart, [1824]) —pravitas Fruhstorfer, 1913 syn. nov, cytherea (Linnaeus, 1758) —eleus (Linnaeus, 1758) stat. rest. — /an j7/a Fruhstorfer, 1913 -aea (C. Felder & R. Felder, 1867) —herennia Fruhstorfer, 1915 -rfagMflwa Fruhstorfer, 1913 —tarratia Fruhstorfer, 1913 syn. nov. —despoliata Fruhstorfer, 1915 syn. nov. -insularis Fruhstorfer, 1913 -ffiarcia Fruhstorfer, 1913 -olbia (C. Felder & R. Felder, 1867) -nahua Grose-Smith, 1898 —wemickei Koher, 1923 viola Fruhstorfer, 1913 stat. nov. -pseudococala Hall, 1933 stat. nov. salmoneus (Butler, 1 866) -emilia Fruhstorfer, 1908 -colada (C. Felder & R. Felder, 1867) -salmonides HdXl, 1938 demialba (Butler, 1872) cpi^gis Fruhstorfer, 1915 -epona Fruhstorfer, 1915 /fl6r/c/fl Fruhstorfer, 1913 —hilareia Fruhstorfer, 1915 stat. nov. —mamaea Fruhstorfer, 1915 syn. nov. capucinus (Walch, 1775) —juruana Butler, 1877 stat. nov. —phylacides Fruhstorfer, 1913 stat. nov. —georgias Fruhstorfer, 1915 stat. nov. -velia (C. Felder & R. Felder, 1867) stat. nov. —ve/iWa Fruhstorfer, 1915 -gutierrezi 'Neild, 1996 stat. nov. barnesia Schaus, 1902 —pseudomessana Fruhstorfer, 1913 syn. nov. -leucas Fruhstorfer, 1915 stat. nov. —paula Bargmann, 1928 syn. nov. -trinita Kaye, 1914 stat. nov. rf/ozi Beutelspacher, 1975 hesterbergi Wilhnott & Hall, 1999
PAGE 39

-chancha Staudinger, 1886 syn. nov. lycorias (Godart, [1824]) —isis (Drury, 1782) preocc. (Drury, 1773) —divina Fruhstorfer, 1907 stat. nov. —pseudagrias Fruhstorfer, 1908 syn. nov. -wallisii (Dewitz, 1877) stat. nov. —hypsenor Godman & Salvin, 1 879 —fassli Fruhstorfer, 1915 syn. nov, -/ara (Hewitson, 1850) —/nainaj Fruhstorfer, 1915 -melanthe {B&tes, 1864) -melanippe Godman & Salvin, 1884 stat. nov. -spruceana (Bates, 1864) stat. nov. attica (C. Felder & R. Felder, 1867) —lesbia Staudinger, 1886 —oronoco Weeks, 1906 —ca/7we/a Fruhstorfer , 1915 —ienYa Fruhstorfer, 1915 -hemileuca Wilhnott & Hall, 1999 leuceria (Druce, 1 874) -juanna Grose-Smith, 1898 stat. nov. leucerioides Beutelspacher, 1975 erymanthis Godman & Salvin, 1884 -oi/i/rjcto Fruhstorfer, 1915 -ssp. nov. -ssp. nov. sicAae«s (Butler, 1866) —/jnvafa Fruhstorfer, 1913 —leucopetra Fruhstorfer, 1913 stat. nov. —irisa Fruhstorfer, 1913 syn. nov. ra/AscA/M Fruhstorfer, 1913 -pallida Rober, 1927 stilesiana DeVries & Chacon, 1982 Aortas (Butler, 1866) —tizona (C. Felder & R. Felder, 1 867) syn. nov. —tizonides Fruhstorfer, 1908 stat. nov. —verewifa Fruhstorfer, 1913 —opheltes Fruhstorfer, 1915 syn. nov. -^aye/Hall, 1939 cocala (Cramer, 1780) -urraca (C. Felder & R. Felder, 1862) —iaparwa Fruhstorfer, 1913 —urracina Fruhstorfer, 1913 stat. nov. —lorzina Fruhstorfer, 1915 syn. nov. -cani'nia Fruhstorfer, 1915 -lorzae (Boisduval, 1 870) —cocalina Fruhstorfer, 1913 stat. nov. —fufia Fruhstorfer, 1913 syn. nov. —fiifina Fruhstorfer, 1915 syn. nov. -didia Fruhstorfer, 1915 stat. rest. — cA/jWe d'Ahneida, 1931 -orellanai Neild, 1996 /eWm (Boisduval, 1870) —falcata Godman & Salvin, 1 878 stat. rest. —j arias Fruhstorfer, 1915 syn. nov. teucophthalma (Latreille, [1809]) —mephistopheles (Buder, 1869) —tegeata Fruhstorfer, 1915 -irminella Strand, 1918 stat. nov. —iaere Hall, 1929 syn. nov. -5ma/// Wilhnott, 1999 ssp. nov. irmina (Doubleday, [1848]) -mw/i/a (Butler, 1873) —wilhelmina Fruhstorfer, 1907 saundersii (Revntson, 1867) —leutha Fruhstorfer, 1915 -helepecki Weeks, 1901 stat. rest. -frontina Hall, 1935 lamasi Wilhnott & Hall, 1999 sa/Ms Hall, 1935 -emmeli Wilhnott & Hall, 1 999 -ssp. nov. shuara Wilhnott & Hall, 1 995 argentea Wilhnott & Hall, 1995 coryneta (Hewitson, 1874) yorrfa/ii Fruhstorfer, 1913 —ernestina Fruhstorfer, 1913 syn. nov. zina (Hewitson, 1867) —restricta Fruhstorfer, 1913 ~/eMcaca/i//ia Fruhstorfer, 1915 —manetho Hall, 1938 syn. nov. -inachia Staudinger, 1 886 — znac/ija Fruhstorfer, 1915 -/acma (Butler, 1872) -irma Fruhstorfer, 1907 stat. nov. —nadja Fruhstorfer, 1907 —nicetas Hall, 1938 syn. nov. -enope Hall, 1938 stat. nov. -desousai Neild & Orellana, 1996 stat. nov. -pyrczi Neild & Orellana, 1996 stat. nov. /wi/teri Beutelspacher, 1976 Justina (C. Felder & R. Felder, 1861) —praevalida Fruhstorfer, 1915 -7M5rine//a Fruhstorfer, 1907 -Valentino Fruhstorfer, 1915 stat. nov. -maira Wilhnott, 1999 ssp. nov. — ffiaz'ra Fruhstorfer, 1915 -inesae Orellana, 1996 olynthia (C. Felder & R. Felder, 1867) —olynthina Fruhstorfer, 1907 syn. nov. —levicula Fruhstorfer, 1913 syn. nov. —olynthina Fruhstorfer, 1915 syn. nov. —theaena Fruhstorfer, 1915 syn. nov. —zopyra Fruhstorfer, 1915 stat. nov. levona Steinhauser & Miller, 1977

PAGE 40

33 Key to species otAdelpha based on adult wing pattern I believe that the simplest way to identify Adelpha is through comparison of specimens with the plates and text accounts, but I recognise that many find a key to be a useful device. The purpose of this key is to allow the non-specialist to identify specimens without needing to resort to dissection or examination of microscopic structiu^es, or analysis of wing pattern element homologies, which requires some experience with the genus. I have therefore expressly not necessarily used the terminology of Fig. 3 for wing patterns, but I have used more widespread wing pattern terminology, such as the term "postdiscal band" to refer to the entire dorsal band rather than just the portion homologous to the element shown on Fig. 3. The key is not arranged strictly phylogenetically and in most cases the characters employed are not considered to be specific synapomorphies; indeed, a number of species have been keyed out more than once due to differences in the most obvious wing pattern characters. I discuss the characters that I consider to define species under the species accounts in the generic Review below, and the Key is to be used in conjunction with the plates, the ranges and further identifying characters given in the accounts for each taxon. I have in some cases listed additional characters to confirm identification, but those given first are the most reliable. It should be noted that, in terms of variation, "usually" refers to variation between subspecies, and "typically" refers to variation within a single taxon. 1 . DHW with white or orange markings in postdiscal area 2 DHW entirely brown in postdiscal area (orange tomal spot may be present) 87 2(1). VFW with pale distal half of discal cell, between dark cell band and discocellular band, divided into three (usually) by basally pointing, "V"-shaped black line (line may not quite touch basal red-orange band, in which case it divides area into two, a basal, hourglass shape and distal, much smaller triangle); very base of VHW discal

PAGE 41

34 cell always entirely white; dark streak absent in base of VFW discal cell 3 VFW with pale distal half of discal cell divided into two by a straight, vertical central dark line, a horizontal dark line, or undivided; very base of VHW discal cell entirely white or with a dark line or darker shading; dark streak almost always present in base of VFW discal cell 14 3(2).VHW with orange/yellow postdiscal line distal of white postdiscal band extending from tomal orange/yellow spot in cell Cu2 to costa; basal edge of white postdiscal band usually bordered by a red-orange band in most taxa; continuous black line from costa to vein 3A through discal cell usually present 4 VHW with postdiscal area distal of white postdiscal band lacking orange/yellow band, with isolated orange tomal spot in cell Cu2; basal edge of white postdiscal band not bordered by a red-orange band; no continuous black line from costa to vein 3 A through discal cell herbita 4(3). VHW with basal portion of vein 3A white, distal portion lined with black which continues as a continuous dark line to costa 5 VHW with vein 3A entirely black to wing base, not part of a line crossing discal cell zea 5(4). VFW with undivided whitish submarginal dashes in each cell except Cu2, particularly clear in cells Cul and Ml 6 VFW with whitish submarginal dashes divided into two in every cell, particularly clear in cells Cul and Ml 8 6(5). VFW with distal edges of silvery-white postdiscal blocks in cell Cul and Cu2 aligned; VFW costa whitish immediately distal of postcellular bar 7 VFW with distal edge of silvery-white postdiscal block in cell Cul much more distal than that in cell Cu2; VFW costa brown immediately distal of postcellular bar

PAGE 42

35 7(6). DFW with approximately equally sized, orange and white postdiscal spots in cell M3 paroeca DFW with either an orange or a white postdiscal spot in cell M3, but not both .paraena 8(5). DFW orange postdiscal marking absent in cells Ml and R5 seriphia DFW orange postdiscal marking present in cells Ml and R5 9 9(8). VFW pale orange postdiscal marking in cells M2-R5, and/or pale silver grey to brownish shading in cell M3 just distal of white postdiscal spot, with clear, dark lines parallel to veins intruding distally at least half way into pale marking and/or to edge of white postdiscal spot (in cell M3) 10 VFW with both pale orange postdiscal marking and pale coloring in cell M3 distal of white postdiscal spot with no dark lines parallel to veins intruding distally, or these only enter a short way, or entire marking is dark orange and darker lines cannot be seen clearly 1 1 10(1 1). VFW costa pale yellowish or whitish at base; pale markings distal of red-orange postdiscal band on VHW parallel to band hyas VFW costa red-orange at base, at least along distal edge of costal vein; pale markings distal of red-orange postdiscal band on VHW noticeably closer to band in cells Rs and Sc+Rl than in cells M2 and Ml seriphia 11(9). DFW with white postdiscal marking in cell M3 well developed, always at least half width of marking in cell Cul 12 DFW with white postdiscal marking in cell M3 absent or a tiny dot, much reduced compared to size of marking in cell Cul radiata 12(11). VHW with two rows of whitish submarginal markings separated by a sharply defined dark brown line; whitish markings may or may not be divided in two in each cell by a dark brown line parallel to the veins 13

PAGE 43

36 VHW with two rows of whitish submarginal markings ahnost merged, separated only by a faint, poorly defined darker line; whitish markings clearly divided in two in each cell by a dark brown line parallel to the veins radiata 13(1 2). DFW with orange subapical marking vertically orientated, more tall than wide; white postdiscal spots in cells Cul and M3 strongly diagonally displaced; VHW with white submarginal markings much nearer to orange postdiscal band in cells Rs and Sc+Rl than in cells M3-M1; Central America and Andes seriphia DFW with orange subapical marking horizontally orientated, more wide than tall; white postdiscal spots in cells Cul and M3 usually not strongly diagonally displaced; VHW with white submarginal markings approximately parallel to orange postdiscal band throughout wing; Central and South America to southeastern Brazil, Paraguay serpa 14(2). DFW with continuous, even white postdiscal band firom costa to anal margin, unbroken by dark veins fessonia DFW postdiscal band variable in extent and color, but never white and unbroken fi"om costa to anal margin, and may be entirely absent 1 5 15(14). VFW discal cell with pale area between red-orange cell bands divided in two by a vertical, clearly defined red-brown to black line 16 VFW discal cell with pale area between red-orange cell bands imdivided or (very rarely) with an indistinct, poorly defined darker central vertical line which obliterates most of the pale area 24 16(15).DFW discal cell with red scaling at base and between cell bars 1 and 2 17 DFW discal cell entirely dark brown or with some scattered orange brown scaling 18 17(1 6). DFW with a large subapical orange marking bredowii DFW lacking orange markings gelania 18(16).DFW with postdiscal band in cell Cu2 entirely, or at least in basal half, white 19

PAGE 44

37 DFW with postdiscal band entirely orange in cell Cu2 viola 19(18).DFW with area between postdiscal band in cell M3 and costa dark brown, or with indistinct, whitish shading, or entirely orange 20 DFW with area between postdiscal band in cell M3 and costa dark brown with two well defined white ovals in cells Ml and R5 alala 20(1 9). DFW with usually relatively narrow orange postdiscal band which remains isolated fi"om more posterior white postdiscal band 21 DFW with very broad orange postdiscal band of which basal edge merges smoothly with basal edge of more posterior white postdiscal band cytherea 21(20). VHW with silvery submarginal line and distal edge in cells M3-Rs approximately straight, with a uniform red-brown border lacking white marginal scaling in the middle of each cell; Andes 22 VHW with silvery submarginal line and distal edge in cells M3-Rs distinctly scalloped, with sparse white marginal scaling in the middle of each cell; Central America. 23 22(21).VFW with pale subapical marking in cells M2-R5 composed of basally pointing, "V"-shaped dashes; basal edge of marking in cells M2-R5 slightly convex; Andes fi-om extreme northern PeruVenezuela corcyra VFW with pale subapical marking in cells M2-R5 composed of straight dashes; basal edge of marking in cells M2-R5 straight; Andes fi-om extreme northern Peru-Bolivia aricia 23(21). DFW with orange subapical marking approximately equal in width to white postdiscal band; VHW discal cell with ground color entirely whitish gray; VFW with red-brown line entirely surrounded by whitish gray scaling between distal edge of white postdiscal band and costa; base of cell Cul with whitish gray scaling; hindwing always slightly produced at tomus pithys

PAGE 45

38 DFW with orange subapical marking broader than white postdiscal band; VHW discal cell with ground color reddish brown between cell bars 1 and 2, and 3 and postcellular; VFW with red-brown line between distal edge of white postdiscal band and costa usually lacking whitish gray scaling between it pale yellow subapical marking; base of cell Cul reddish brown; hindwing usually rounded at tomus donysa 24(1 5). DFW lacking red scaling in discal cell and at discocellulars 25 DFW with red band crossing middle of discal cell and over discocellulars.. .rfwc/gs 25(24). DFW with postdiscal band present and completely white to vein M3, spot in cell Cul well developed 26 DFW with some orange scaling in postdiscal band between anal margin and vein M3, or band is absent in cell Cul (a tiny, isolated whitish brown dot may be present at base of cell Cul), or DFW entirely dark brown 46 26(25).DHW with orange tomal spot present 27 DHW with orange tomal spot absent 44 27(26). DFW with a large orange subapical marking, lacking additional well developed orange subapical spots distally (tiny dots may be present near distal edge in cells Ml andR5) 28 DFW with two well developed orange subapical spots in cells R5 and R4 separated from a more basal, much larger orange marking by an area of dark brown at least as wide as the more basal marking naxia 28(27). VFW with uneven, indistinct, pale orange, silver-gray and/or reddish brown markings in postdiscal area, not forming well defined straight stripes 29 VFW with a pair of well defined, straight, vertical postdiscal stripes between cells Cul-R5, a basal white stripe and a distal orange stripe, bordered distally and basally by black syma

PAGE 46

39 29(28). DFW with orange subapical marking not touching white postdiscal marking in cell M3 30 DFW with orange subapical marking touching distal edge of white postdiscal marking in cell M3 45 30(29). DFW with white postdiscal band terminating at vein M3 31 DFW with white postdiscal band extending fully to vein M2, sometimes as dashes in cells Ml and R5 basiloides 31(30).VFW with area immediately distal of discocellular veins orange to reddish brown, then dark brown line (postcellular bar), then silver gray 32 VFW with area immediately distal of discocellular veins orange, then very faint shghtly darker line (postcellular bar), then orange mythra 32(31). VFW with white postdiscal band continuing to costa as grayish white band anterior of vein M3, distal edge of which is aligned with distal edge of white postdiscal band in cells Cul and M3 33 VFW with white postdiscal band broken in cell M2; if there is whitish shading in cell M2, distal edge is not aligned with distal edge of white postdiscal band in cells Cul andM3 34 33(32). VHW white postdiscal band immediately bordered distally by very thin dark brown line, then red-brown band as broad as white postdiscal band, then one to two indistinct silver gray lines poltius VHW white postdiscal band immediately bordered distally by thin red brown band, then thin silver gray line, then red brown, then two silver gray lines calliphane 34(32). VFW with orange-brown to red-brown discal cell bands parallel; basal area of cell M2 mostly filled with silver gray scaling or with a well defined silver gray triangle; VHW discal cell bands separate and at least as far apart as width of cell

PAGE 47

40 VFW with orange-brown discal cell bands converging posteriorly; basal area of cell M2 dark brown or reddish brown, or with faint, poorly defined paler gray scaling in anterior half only; VHW discal cell bands merged into a single band or nearly so thoasa 35(34). Ventral surface marginal border of both wings ahnost entirely orange-brown or red-brown, with very faint white dashes only in cell Cu2 and trace in cell Cul... 36 Ventral surface marginal border of both wings orange-brown or red-brown with clear white dashes in most cells 37 36(35). VHW with elongate tomus, so that basal pair of black tomal spots are nearer distal margin than white postdiscal band in cell Cu2; white postdiscal band in cells Rs and Sc+Rl bordered distally by thin dark brown line then paler yellowish brown line falcipennis VHW without elongate tomus, basal pair of black tomal spots are equidistant between distal margin and white postdiscal band in cell Cu2; white postdiscal band in cells Rs and Sc+Rl bordered distally by thin red brown line then thicker silver gray line .gavina 37(35). VFW with pale orange subapical marking in cells M2 and Ml not split by continuous line of darker red-brown scaling near distal edge, marking split only in cell M2; DFW orange subapical marking may only just extend into cell M2, extend to terminate broadly at vein M3, or extend (rarely) as orange dashes into cell Cul 3g VFW with pale orange subapical marking in cells M2 and Ml split by continuous line of darker red-brown scaling near distal edge; DFW orange subapical marking always extends broadly posteriorly at least as far as vein M3, often broadly into Cul thessalia 38(37). Hindwing margin not produced into a short "tail" at vein Cul; DHW distal half dark brown with slightly paler brown submarginal lines; DFW orange subapical

PAGE 48

41 marking does not extend distally to reach inner paler brown submarginal line in cell Ml; not Jamaica 39 Hindwing margin produced into a short "tail" at vein Cul; DHW with thin whitish submarginal line; DFW orange subapical marking extends distally to reach inner paler brown submarginal line in cell Ml; Jamaica abyla 39(38).DFW orange subapical marking extending to broadly border vein M3, or extending into cell Ml as a triangle of varying size with basal edge of marking straight or smoothly concave 40 DFW orange subapical marking much broader in cell Ml than M2, extending into cell M2 only at distal edge as a triangular, posteriorly pointing dash, basal edge of marking not straight but kinked at vein M2 iphiclus 40(39). DFW white postdiscal band tapering slightly from anal margin to cell M3; base of eastern Andes S. Colombia-Peru iphicleola DFW white postdiscal band not tapering from anal margin to cell M3; not base of eastern Andes S. Colombia-Peru 41 4 1(40). DFW orange subapical marking broadly bordering vein M3 42 DFW orange subapical marking extending into cell M2 only as a small triangle, often touching but not broadly bordering vein M3 43 42(41). DFW with distal edge of orange subapical marking in each of cells M2-R5 slightly concave in some or all cells; white postdiscal band of approximately equal width to orange subapical marking; VFW with edges of pale orange subapical marking sharply defined, particularly basal edge, with red brown scaling closely confined to veins Cu2 and Ml; VHW with first pale grayish line distal of white postdiscal band brighter whitish at vein M3 and closer to second distal pale grayish line iphicleola DFW with distal edge of orange subapical marking in each of cells M2-R5 sfraight or slightly convex; white postdiscal band approximately half width of

PAGE 49

42 orange subapical marking; WW with edges of pale orange subapical marking poorly defined, typically with diffuse red brown scaling, with red brown scaling broadly along veins Cu2 and Ml; VHW with first pale grayish line distal of white postdiscal band of uniform color and roughly parallel to second distal pale grayish 43(41).DFW with distal edge of orange subapical marking fi-om vein Ml to costa slightly concave; VFW with edges of pale orange subapical marking sharply defined, particularly basal edge, with red brown scaling closely confined to veins Cu2 and Ml; VHW with first pale grayish line distal of white postdiscal band brighter DFW with distal edge of orange subapical marking fi-om vein Ml to costa straight or slightly convex; VFW with edges of pale orange subapical marking poorly defined, typically with diffixse red brown scaling, with red brown scaling broadly along veins Cu2 and Ml; VHW with first pale grayish line distal of white postdiscal band of uniform color and roughly parallel to second distal pale grayish line in cells Cul-M2 iphiclus 44(26). DFW with orange subapical marking broadest in cells Ml and R5, tapering to a point in cell Cul coryneta DFW with orange subapical marking broadest in cell M2, absent or present as indistinct orange scaling in cell M3 felderi 45(29). DFW orange subapical marking extending right across cell Ml to broadly end at costa; dark red-brown line cutting pale VFW orange subapical marking in half is even in width abia DFW with only basal half of orange subapical marking ending at costa, distal half ending mid cell Ml; dark red-brown line cutting pale VFW orange subapical marking in half is much broader in cell Ml than M2 epizygis 46(25). DFW with some orange or white markings 47 line in cells Cul-M2 iphiclus whitish at vein M3 and closer to second distal pale grayish line, iphicleola

PAGE 50

43 DFW entirely dark brown .justina 47(46). DHW with either white or partially white hindwing postdiscal markings, or if entirely orange, isolated subapical spots are present on DFW 48 DHW with continuous, uniformly orange postdiscal band which extends from tomus to costa, then across DFW to costa, DFW subapical spots absent ethelda 48(47). DFW with postdiscal band in cells Cu2 to M3 vertical, or, if inclined, orange subapical spots are present at least in cells R5 and R4 or DHW is entirely brown 49 DFW with oblique orange band from tomus to costa, orange subapical spots entirely absent, DHW with central white spot leucophthalma 49(48). VFW discal cell silvery gray or whitish with clear brown to black basal streak, cell bars one and two distinct and separated by reddish to orange brown bar, a pale band between second and third cell bars 50 VFW discal cell silvery gray with basal streak absent or very faint black scaling, cell bars one and two coalesced to form a black band, a round silver gray spot in anterior half of cell between second and third cell bars argentea 50(49). DHW postdiscal band or markings either partially or totally white 51 DHW postdiscal band uniformly orange 69 5 1(50). DFW with orange subapical spot in cell R4, and/or two sets of orange postdiscal to subapical markings in either or both of cells Ml and R5 52 DFW with no orange subapical spot in cell R4, and only one orange postdiscal to subapical marking in cells Ml and R5 71 52(51).VHW with white postdiscal band bordered basally by reddish or orange brown line, then whitish or gray in discal cell, then similarly colored reddish or orange brown line 53

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44 VHW with white postdiscal band bordered basally by continuous black/dark brown line, then continuous orange band, both extending from near tomus to at least vein Sc+Rl 73 53(52). DFW with postdiscal band variably orange and white to vein Cul, joined to postdiscal and subapical orange markings 54 DFW with postdiscal band white to vein Cul, isolated from postdiscal and subapical orange markings 74 54(53). VFW with inclined or vertical postdiscal band, if the latter, band is completely or in its basal half very pale orange, not dissimilar in color from white VHW postdiscal band 55 VFW with vertical, matt orange postdiscal band, distinctly different in color from VHW postdiscal band cocala 55(54). VFW postdiscal band inclined or vertical, if the latter, very pale orange, similar in color to white VHW postdiscal band, with darker vertical red brown lines variably present; silver gray submarginal markings not particularly pronounced in cells Ml and R5, silver gray apical markings typically present in cells R4 and R3 56 VFW postdiscal band vertical, distinctly two-tone, with basal half very pale orange, similar in color to white VHW postdiscal band, distal half entirely orange brown; two prominent, bright whitish subapical spots in cells Ml and R5, no pale apical spots in cells R4 and R3 cocala 56(55). DFW with postdiscal band always reaching anal margin, basal edge of postdiscal band in cells Cu2 and Cul perpendicular to anal margin 57 DFW with postdiscal band usually not reaching anal margin, basal edge of postdiscal band in cells Cu2 and Cul inclined to anal margin 78 57(56). VFW with anterior dash of inner submarginal pair of silver gray dashes in cell Cu2 of approximately equal width or slightly wider than posterior dash 58

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45 VFW with anterior dash of inner submarginal pair of silver gray dashes in cell Cu2 of approximately twice width of posterior dash; silver gray marginal dashes absent except in cell Cu2; ventral colors predominantly red-brovm and graysilver 79 58(57). VFW with very pale orange postdiscal markings in cells Cul and M3 of equal or differing widths, split by dark lines, either distinct or diffuse, each marking followed distally by one to two silver gray dashes; basal edge of pale postdiscal marking in cell Cu2 perpendicular to anal margin; pale silver gray marginal dashes/spots present in cell Cu2 59 VFW with very pale orange postdiscal markings in cells Cul and M3 of equal widths, not split by dark lines, each marking followed distally by only a single silver gray dash; basal edge of pale postdiscal marking in cell Cu2 inclined to anal margin; pale silver gray marginal dashes/spots absent in cell Cu2 80 59(58). VHW distal of white postdiscal band not entirely red-brown except for row of close, paired silver dashes from costa to tomus, basally indented in cell M2; DFW with orange subapical spot in cell Ml absent to pronounced; postdiscal band in cell Cu2 white or orange, same thickness or narrower than band in cell Cul 60 VHW distal of white postdiscal band entirely red-brown except for row of close, paired silver dashes from costa to tomus, basally indented in cell M2, and two white tomal dashes in cell Cu2; DFW with orange subapical spot in cell Ml absent or very faint orange scales; postdiscal band in cell Cii2 white and half thickness of band in cell Cul milleri 60(59). DFW with distal edge of postdiscal band in cell Cul sfraight or slightly convex; VFW with pale postdiscal band usually distally displaced from postcellular bar, very rarely ahnost touching postcellular bar, in cell Ml; pale postdiscal markings in cells Cul and M3 ma or may not be split by vertical, straight, dark red-brown scaling 61

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46 DFW with distal edge of postdiscal band in cell Cul concave; VFW with pale postdiscal band almost touching postcellular bar in cell Ml; pale postdiscal markings in cells Cul and M3 split by vertical, straight, dark red-brown scaling ximena 6 1(60). DFW with distal edge of orange subapical spot in cell Ml distally displaced with respect to distal edge of orange postdiscal band in cell M2 (or spot absent), spot clearly isolated from band; VFW with two long pale dashes, extending across cell, or single round spot, in cell Cul distal of pale postdiscal block 62 DFW with distal edge of orange subapical spot in cell Ml aligned with distal edge of orange postdiscal band in cell M2, spot fused or nearly so (vein M3 dark brown) with band; VFW with two short pale dashes, not extending across cell, in cell Cul distal of pale postdiscal block capucinus 62(61). VFW with pale postdiscal band either split by an even line, which cuts right through band and ends at distal edge of band at vein Cu2, variably split, or not split 63 VFW with pale postdiscal band split by posteriorly narrowing line, starting in cell M2 and terminating within marking at vein Cu2 altamazona 63(62). VFW pale postdiscal band not completely divided in two by a red-brown line, or line is slightly concave; DFW orange subapical spot in cell Ml absent, smaller than, or of equivalent size to that in cell R5; VFW pale subapical spot in cell Ml not divided with darker red brown scaling; VHW with pale, thin silver gray line distal of white postdiscal band even throughout or much thicker whitish scaling in cells RsandSc+Rl 64 VFW pale postdiscal band divided in two by distinct, vertical, straight red-brown line; DFW orange subapical spot in cell Ml pronounced and as large as that in cell R5 or larger; VFW pale, subapical spot in cell Ml typically divided with

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47 darker red brown scaling; VHW with pale, thin silver gray line distal of white postdiscal band even throughout boeotia 64(63). VFW with straight silver gray submarginal dash in cell M3 or marking absent; VHW with paler lines and darker bands of variable color and width distal to white postdiscal band, all approximately parallel and evenly spaced; DHW with tomal orange spot of varying size in cell Cu2 typically present 65 VFW with basally pointing, "V"-shaped silver gray submarginal marking in cell M3; VHW with thin silver gray line distal to white postdiscal band, then dark red band constricted in cell M2, then line of close, paired silver-gray dashes basally displaced in cell M2, then marginal silver gray dashes; DHW with tomal orange spot in cell Cu2 absent Jordani 65(64). VFW with pale postdiscal marking divided into a basal block and two distal dashes 66 VFW with pale postdiscal marking divided into a basal block and one roimd distal spot 81 66(65). VHW with first two pale gray lines distal of white postdiscal band separate throughout wing 67 VHW with first two pale gray lines distal of white postdiscal band fiased to form a single large whitish spot in cell Sc+Rl pollina 67(66). VHW with first pale line distal of white postdiscal band of even width and color throughout wing 68 VHW with first pale line distal of white postdiscal band much thicker in cells Rs and Sc+Rl than in remainder of wing 86 68(67). VFW with submarginal silver gray dashes of approximately even size in each cell, forming a line parallel to distal margin malea

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48 WW with submarginal silver gray dashes of varying size, that in M3 absent or much reduced compared with that in cell Cul, and basally displaced with respect to remaining dashes .fabricia 69(50) VFW with line of silver gray submarginal dashes notably basally displaced in cells Cul, M3 and M2, most displaced in M2; west of Andes 70 VFW with submarginal silver gray dashes forming a line parallel to distal margin; eastern Merida range in Venezuela malea 70(69). Hindwing elongate and triangular; VHW with second pale line distal of white postdiscal band almost uniform in color, sometimes slightly paler in cells Rs and Sc+Rl; DFW with basal edge of orange postdiscal band continuously curving through vein M3 leuceria Hindwing squarish; VHW with second pale line distal of white postdiscal band consisting of large, silvery, scalloped spots in cells Rs and Sc+Rl and much thinner and paler throughout remainder of wing; DFW with basal edge of orange postdiscal band displaced at vein M3 leucerioides 71(51). DFW orange postdiscal band with distal edge more distal in cell M2 than in cell Ml 72 DFW orange postdiscal band with distal edge more distal in cell Ml than in cell M2 .plesaure 72(71) VFW postdiscal band matt orange brown; VFW distal ground color reddish brown; pronounced silver white subapical spots in cells Ml and R5, former larger, apical spots absent in cells R4 and R3 cocala VFW postdiscal band white or very pale orange; VFW distal ground color dark brown; silver white subapical spots in cells Ml and R5, former sometimes smaller and often absent, apical spots present in cells R4 and R3 melona 73(52) DHW with postdiscal band entirely white; VFW with pale postdiscal markings in cells M3 and M2 split in two by vertical orange-brown line shuara

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49 DHW with postdiscal band orange, or tinged with orange, particularly at distal edge, in cells Rs and Sc+Rl; VFW with undivided pale postdiscal marking in cell M3, that in M2 occasionally with very faint darker brown scaling through middle attica 74(53) VFW with single pale yellowish white spot between white postdiscal block in cell Cul and inner, thick silver gray submarginal dash 75 VFW with two pale dashes between white postdiscal block in cell Cul and inner, thick silver gray submarginal dash 76 75(74). VHW with spots of first row of pale whitish gray markings distal of white postdiscal band largest in cells Ml and Rs erotia VHW with spots of first row of pale whitish gray markings distal of white postdiscal band even dashes in all cells, or slightly larger in cells Rs and Sc+Rl messana 76(74) DFW with orange subapical spot in cell M2 larger than or equivalent in size to that in cell Ml; distal edge of orange subapical spot in cell Ml aligned with distal edge of orange postdiscal band in cell M2, spot fused or nearly so (vein M3 may be dark brown) with band; VFW with silver submarginal marking in cell M3 absent or basally displaced with respect to remaining line of submarginal markings 77 DFW with orange subapical spot in cell M2 absent or much smaller than that in cell Ml; if present, orange subapical spot in cell Ml isolated fi-om orange postdiscal band in cell M2; VFW with silver submarginal markings forming a row parallel to distal margin malea 77(76). DFW with orange subapical spot in cell R4 present; VFW with silver gray submarginal marking in cell M3 present; forewing with cell M3 of approximately even width throughout, may be very slightly wider at base than at distal margin barnesia

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50 DFW with orange subapical spot in cell R4 absent; VFW with silver gray submarginal marking in cell M3 absent; forewing with cell M3 noticeably wider near base than at distal margin diazi 78(56). VFW with distal edge of pale orange postdiscal band nearer distal margin in cell Cu2 than in cell Cul; VHW with each dash of most distal submarginal row of silver gray dashes bordered basally by dark gray then another silver gray dash.zina VFW with distal edge of pale orange postdiscal band nearer distal margin in cell Cul than in cell Cu2; VHW with each dash of most distal submarginal row of silver gray dashes bordered basally by dark gray, then orange-brown justina 79(57). VFW with pale orange postdiscal band cut by red-brown line in cells Cul and M3; DFW with postdiscal band narrow at costa and cut by dark veins; DHW distal margin fringe with white scaling in each cell; VFW with silvery gray spot distal to postcellular bar in cell Ml; E. Andes sichaeus VFW with pale orange postdiscal band not cut by red-brown line in cells Cul and M3; DFW with postdiscal band broad at costa and not cut by dark veins; DHW distal margin fringe entirely black; VFW with no silvery gray spot distal to postcellular bar in cell Ml; Costa Rica-W. Andes hesterbergi 80(58). DHW white postdiscal band narrow, of almost even width throughout wing, tapering towards tomus; VFW with silver gray submarginal marking in cell M3 "IT'-shaped olynthia DHW white postdiscal band of moderate width, tapering towards costa and tomus, broadest in cells M2 and Ml; VFW with silver gray submarginal marking in cell M3 a straight, inclined dash justina 8 1(65). VHW with spots forming first row of pale markings distal of white postdiscal band of even width throughout wing, or largest in cells Ml and Rs, or in cells Rs and Sc+Rl, this row bordered distally by dark red-brown/brown band of even width or slightly broader in cell Ml than M2; VFW with black area between

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51 white/pale orange postdiscal block in cell Cu2 and silver gray submarginal spots imdivided 82 VHW with spots forming first row of pale markings distal of white postdiscal band largest in cell M2, then M3, this row bordered distally by dark brown band much broader in cell Ml than M2; VFW with black area between white/pale orange postdiscal block in cell Cu2 and silver gray submarginal spots divided by vertical pale orange/orange dash delinita 82(81). VFW with silver gray apical spots in cells R4 and R3; VHW with four silver gray dashes distal of white postdiscal band in cell Rs; pale orange postdiscal marking in cell M3 not divided by orange-brown line 83 VFW within no silver gray apical spots in cells R4 and R3; VHW with three silver gray dashes and a pale red-brown dash distal of white postdiscal band in cell Rs; pale orange postdiscal marking in cell M3 typically divided in half by vertical orange-brown line erymanthis 83(82). VFW with silver gray submarginal dash in cell M3 of similar size and shape to that in cell Cul; VHW ground color almost uniform orange-brown to red-brown 84 VFW with silver gray submarginal dash in cell M3 absent or much fainter and thinner compared to that in cell Cul; VHW ground color typically dark brown with red-brown to orange-brown band between first and second pale lines distal of white postdiscal band and orange-brown marginal ovals at each vein thesprotia 84(83). VHW with spots of first row of pale whitish gray markings distal of white postdiscal band even dashes in all cells, or slightly larger in cells Rs and Sc+Rl 85 VHW with spots of first row of pale whitish gray markings distal of white postdiscal band largest in cells Ml and Rs erotia 85(84). VFW with first discal cell bar slightly inclined to costa, and second and fourth cell bars concave; ventral pale lines distal of postdiscal band distinctly silvery gray;

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52 VHW with first and second discal cell bars broadly spaced, noticeably paler in area between, equidistant between third discal cell bar and basal red-brown line phylaca WW with discal cell bars approximately straight, particularly second cell bar, fourth cell bar slightly convex; ventral pale lines distal of postdiscal band silvery white; VHW with first and second discal cell bars close together, ahnost forming a single dark band, band closer to third discal cell bar than to basal red-brown line messana 86(67). DFW with orange in cell M3 filling base of cell; WW with white postdiscal block in cell Cu2 narrower than orange postdiscal block in this cell on DFW; pale orange postdiscal marking in cell M3 split by orange brown line near base and near distal margin; pale orange dashes in cell Cul extend across entire cell; WIW with first two pale lines distal of white postdiscal band pale silver gray, broader in cells Rs and Sc+Rl heraclea DFW with orange in cell M3 not filling base of cell; WW with width of white postdiscal block in cell Cu2 equal to width of orange postdiscal block in this cell on DFW; pale orange postdiscal marking in cell M3 entire except for some indistinct orange scaling near base; pale orange dashes in cell Cul only in anterior half of cell; VHW with first two pale lines distal of white postdiscal band pale gray, except white in cells Rs and Sc+Rl sp. nov. 87(1). DFW with some orange or white markings 88 DFW entirely dark brown zina 88(87) DFW with postdiscal band vertical or inclined, of uniform color, not broken in cell M3 89 DFW postdiscal band vertical, broken in cell M3, a thin white band fi-om anal margin to cell M3, a very broad orange band fi-om cell Cul to costa donysa

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53 89(88). VHW with distal half varying shades of red/orange to brown, with some pale rows of markings, no dark brown lines parallel to veins bisecting each cell; VFW discal cell with a single silvery white/silvery gray band opposite middle of cell Cul 90 VHW with distal half almost entirely brown or yellowish brown, with dark brown lines parallel to veins bisecting each cell; VFW discal cell with a pair of silvery white spots opposite middle of cell Cul, an anterior and a posterior spot, separated by a black line 99 90(89). DFW with postdiscal orange or white markings in cells M3 and M2 not filling cell to distal margin 91 DFW with postdiscal orange markings in cells M3 and M2 filling cell to distal margin stilesiana 9 1(90). DFW with only a single white or orange postdiscal to submarginal marking in cells Cul andM3 92 DFW with three white postdiscal to submarginal markings in each of cells Cul and M3 demialba 92(91). DFW with vertical or inclined orange postdiscal band 93 DFW with oblique white postdiscal band 100 93(92). VFW pale silver gray area in distal half of discal cell undivided by a well-defined darker line, rarely with indistinct, incomplete line of black scaling; VFW with or without pale postdiscal marking in cell Cu2 94 VFW pale silver gray area in distal half of discal cell completely divided by well defined, dark brown line; VFW with pale postdiscal marking in cell Cu2 101 94(93). VHW with silver gray postdiscal band absent, broken, or continuous and extending throughout wing, bordered basally anterior of vein M3 by dark redbrown line then silver gray scaling, at least anterior of vein M2 95

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54 VHW with continuous silver gray postdiscal band extending throughout wing, bordered basally anterior of vein M3 by dark brown line then broad orange band 102 95(94). DFW with orange postdiscal markings just distal of discocellulars in cells Ml and R5 at least twice as broad as orange subapical spots in those cells, or orange subapical spots absent 96 DFW with orange postdiscal markings just distal of discocellulars in cells Ml and R5 of approximately equal width or narrower than orange subapical spots in those cells, or postdiscal markings absent 103 96(95). VFW with some submarginal silver gray dashes distal of pale orange postdiscal to submarginal band in cells Cu2 to M2 97 WW with no submarginal silver gray dashes distal of pale orange postdiscal to submarginal band in cells Cu2 to M2 106 97(96). VHW with area between first and second discal cell bars whitish gray, with variable amounts of reddish scaling at inner edges of cell bars 98 VHW with area between first and second discal cell bars orange brown 109 98(97). DFW with distal edge of orange postdiscal band vertical and parallel to distal margin in cells Cul and M3 jcimena DFW with distal edge of orange postdiscal band inclined, much closer to distal margin in cell Cul than M3 boreas 99(89). DFW with orange subapical spots in cells Ml and R5 mesentina DFW with subapical spots in cells Ml and R5 white or absent lycorias 100(92). VHW with postdiscal band uniformly silver white from costa to tomus; area posterior to vein 3A silver white lined with orange-brown; W. Colombia ethelda VHW with postdiscal band steely gray with white spot at costa in cell Sc+Rl; area posterior to vein 3 A orange; E. Andes to S. E. Brazil epione

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55 101(93).DFW with vertical orange postdiscal band parallel to distal margin tracta DFW with oblique orange postdiscal band extending from costa to tomus salmoneus 102(94).DFW postdiscal orange band obhque, extending from costa to tomus ethelda DFW postdiscal orange band vertical, extending from costa to anal margin melona 1 03(95). VHW with silver white postdiscal band from costa to tomus present 104 VHW with silver white postdiscal band from costa to tomus absent, postdiscal area entirely red-brown 1 10 104(103). VFW with pale postdiscal spots in cells M3 and M2 not spht in half by darker red-brown line 105 WW with pale postdiscal spots in cells M3 and M2 split in half by darker redbrown line salus 105(104).DFW with basal edge of postdiscal band from anal margin to cell M3 slightly inclined to anal margin; VFW discal cell with thin, well defined dark brown basal streak; silver submarginal dash in cell M3 reduced or absent; two silvery gray marginal spots present in cell Cu2 rothschildi DFW with basal edge of postdiscal band from anal margin to cell M3 perpendicular to anal margin; VFW discal cell with broad, poorly defined orange basal sfreak; silver submarginal dash in cell M3 of similar size to remaining submarginal dashes; silvery gray marginal spots absent in cell Cu2 levona 106(96). DFW with distal edge of orange postdiscal band projecting distally in cell M2 107 DFW with distal edge of orange postdiscal band indented basally in cell M2 leucophthalma

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56 107(106). VFW pale yellow postdiscal markings in cells M2-R5 close to postcellular bar (distance from basal edge of markings to bar much less than width of markings) 108 VFW pale yellow postdiscal markings in cells M2-R5 not very close to postcellular bar (distance from basal edge of markings to bar equal to width of markings) lamasi 108(107). VHW predominantly yellow; VFW with no white postdiscal spot in cell Cu2 saundersU VHW predominantly dark orange-brown; VFW with white postdiscal spot in cell Cu2 irmina 109(97). VHW with continuous silver gray postdiscal band with even basal edge from costa to cell Cu2 (band may be darkened anterior of vein Ml) salus VHW with silver gray to whitish postdiscal spots, occasionally forming a band from costa to cell Cul, but not entering Cu2, always with uneven basal edge una 1 10(103). VFW with distal edge of pale orange marking in cell Cul nearer distal margin than that of marking in cell Cu2 justina VFW with distal edge of pale orange marking in cell Cu2 nearer to distal margin than that of marking in cell Cul zina Species groups Given the general lack of resolution and weak support for nodes in the phylogeny derived in Chapter 3, due to the lack of morphological characters, I feel it would be premature to attempt to divide the entire genus into species groups. Nevertheless, I have indicated in Tables 2-5, with horizontal lines, groups which may be monophyletic or at least contain species likely to share morphological and biological traits, although the latter groups may still be paraphyletic. In addition, there are several sfrongly or relatively

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57 strongly supported, probably monophyletic species groups which I refer to in the text, and these are listed and defined in Table 1, with the constituent species given in the cladogram (Fig. 250). Table 1 . Synapomorphies and other typical characters of species groups within Adelpha. Species included in each group are given in Chapters, Fig. 250. Group: Synapomorphies: Other typical characters: Ecological characters: A. serpa A. alala A. iphiclus A. phylaca A. capucinus A. cocala Male genitalia: valve lacks well developed clunicula; aedeagus has sclerotised, spiny pad on the dorsal surface of the vesica. Wing pattern: DFW discal cell has basal streak placed near costa, enclosing anteriorly a dense patch of reddish scales; basal streak in VHW and VfW discal cell absent; welldefined black line extending from base of VHW to tip of humeral vein. Wing pattern: third discal cell bar present on VFW; dark lines bordering vein 3A on VHW greatly faded and reduced. Postdiscal series on VFW broaden anteriorly, being further from inner submarginal series in cell Ml than cell M2. Wing pattern: densely packed, pale whitish scales along base of vein Rs on DHW. Male genitalia: clunicula broad; valva with terminal teeth extending along ventral edge of valva; saccus large, almost same length as valva. Female genitalia: corpus bursae and ductus bursae large; sclerotised bands placed far apart. Subapical orange spot in cell Ml on DFW aligned with distal edge of orange marking in cell M2; postdiscal series fused on VFW in cell M3 but separate in cell Cul . Female genitalia: corpus bursae lacking sclerotised bands, rounded, typically with ductus bursae of approximately even width. Male genitalia: typically with poorly developed ventral and dorsal medial lobes. Female genitalia: bands on corpus bursae heavily sclerotised with coarse teeth. Immature stages: pupa often with shimmering silver or gold areas. Wing pattern: ventral colors all faded; outer submarginal series on VHW absent. Male genitalia: valva with few, large terminal spines. Immature stages: larval scoli greatly reduced. Wing pattern: postdiscal and submarginal series on VFW typically visible throughout; upper postdiscal band on DFW absent. Wing pattern: postdiscal series fused in cell Cul on VFW and fused to postdiscal band in cell M3; Male genitalia: valva with pronounced dorsal and ventral medial lobes. Immature stages: pupa with prominent, hooked dorsal projection on thorax; final instar larva pale with dark area laterally on thorax-A2. Male genitalia: valva broad throughout, not tapering, squared at tip with terminal spines; clunicula long; saccus relatively short compared to valva. Generally small species, orange tomal spot on DHW often absent, thorax and abdomen relatively narrow. Typically geographically widespread; often found perching in large clearings near rivers; relatively little geographic differentiation of wing pattern; hostplants diverse, not Rubiaceae. Occur in high elevation cloud forests; larval hostplants Caprifoliaceae; known larvae construct shelters from hostplant leaves. Especially diverse in southeastern Brazil; often in secondary growth areas; relatively little geographic differentiation of wing pattern; hostplants all Rubiaceae. Usally occur in lowland primary forest; perch along forest edges or in canopy; hostplants diverse, not in Rubiaceae. Typically confined to lowland forest. Uusually occurring in premontane cloud forest habitats, many species with restricted ranges; strongly mimetic, with sharp regional differentiation of dorsal wing pattern.

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58 Biology Habitats, Phenology and Rarity Adelpha species occur in a very wide variety of forested life zones, and microhabitats within life zones. They have been recorded from sea level to 3000m. While higher species numbers are found in wet forests, certain Adelpha are restricted to drier life zones, ranging from semi deciduous to deciduous forest {A. fessonia), xeric Acacia scrub {A. fessonia lapitha), and mixed pine-hardwood forest {A. gelania). A large number of species are also confined to cloud forest habitats in the Andes and other mountainous areas. There is relatively little information available on the adult phenology of Adelpha species. My research in Ecuador indicates that species and individual abundance is highest during the wet season and early dry season. These observations are in contrast to those of DeVries (1987), who states that most individuals are more abundant in the dry season. However, through the use of fraps I have found several species to be abundant in Costa Rica during the wet season (Hall & Willmott, 1993), and believe that species appear to be more abundant in the dry season since there are less collecting days lost due to rain and males more frequently leave the forest to puddle along roads or rivers. DeVries et al. (1997), in a frapping study, report in eastern Ecuador that nymphalid species and individual abundance in eastern Ecuador is highest in the wet season. hi many instances, individuals were encoimtered in numbers over a period of a couple of days, but never seen again despite repeated visits to a site. This was the case with A. hesterbergi, A. lamasi and A. levona in northwestern Ecuador. This suggests the possibility that pupal eclosion may perhaps be triggered by some environmental cue, such as heavy rains. Aiello (pers. comm.) reports to me a recent outbreak of A. iphiclus in Panama, which she believes to be linked to an unusually dry period causing high mortality in predators and parasitoids of the immature stages, which usually control population numbers.

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59 The majority of Adelpha species seem to be present throughout the year, even if abundance is seasonal, but A. bredowii eulalia has been reported to have two distinct broods in the U.S.A., from May to June, then from August to October (Ferris & Brown, 1982; Gary, 1994). The larvae are in diapause during the winter (Ferris &. Brown, 1982). It is very probable that species in the more sfrongly seasonal, more temperate regions of eastern and southern Brazil to Argentina, Paraguay and Uruguay, may have similar life cycles. There are, however, no published observations and museum specimen data are too few to draw any conclusions. While several Adelpha species are common to abundant, many are infrequently encountered and 14 species are represented by fewer than 25 specimens in all of the collections examined (see Fig. 4). One species, A. herbita, is known only from the single female specimen on which the original illustration is based, and the whereabouts of this specimen, if it still exists, is currently unknown. For almost all species there are more known specimens of males than females, which overall outnumber females in collections by approximately 4:1. This probably reflects the propensity for males to occur in groups (see Adult ecology below) and to be attracted to a wider variety of nutrient sources. The rarity of many species is due to their small geographic range and/or confinement to cloud forest habitats, where they may be locally not uncommon, and it is this life zone that most recently described species inhabit (Beutelspacher, 1976; Steinhauser «&. Miller, 1977; DeVries & Chacon, 1982; Willmott & Hall, 1995, 1999). Such habitats have historically been poorly collected due to problems of access from the steepness of the terrain, and the difficulties of collecting in almost perpetual rain and cloud. They continue to provide new species in many other groups of butterflies (Hall & Willmott, 1995; Willmott et al., in prep.) and their Adelpha fauna still remains relatively poorly known. Certain species, such as A. nea, are extremely widespread but very rare throughout their range. This species is known from eastern Ecuador from two specimens, one captured at Chichicorrumi in 1993, but despite numerous subsequent visits to this locality it has not

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60 been collected there again. At the other end of the spectrum are species that are everywhere common and typically encountered in numbers. Such is the case with Adelpha cytherea, which alone contributed almost 8% of the specimens examined. The perceived rarity of species, and the number of specimens in collections, presumably almost never reflects their true abundance in the field. The use of traps in Ecuador has demonstrated certain species, such as A. pollina, A. heraclea and A. boreas, to be widespread and common in many localities, despite the low numbers of specimens in collections. These species are never encountered outside of traps and presumably spend their time in the forest canopy. 14-, Number of specimens/species Fig. 4. Abundance of Adelpha species in each class of number of specimens recorded.

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61 Hostplants Their exist hostplant records for at least 39 species of Adelpha, comprising 19 plant families, 61 genera and 94 species (see Table 2), and the genus has one of the widest host breadths of any nymphalid genus (Ackery, 1988). The most important single contributors to our knowledge of Adelpha hostplants are Miiller (1886), Moss (1933) and Aiello (1984), with a number of records by Janzen, Haber, Marquis and DeVries (in: DeVries, 1986). Many of the identifications of Adelpha species in the hostplant literature are unreliable or erroneous, and I have tried to correct as many of these misidentifications as possible, based on descriptions of the adult, the locality and voucher material. The latter is sadly lacking for the majority of species, rendering a number of records questionable to almost worthless. Many Adelpha appear to be highly polyphagous; A. serpa, for example, has been recorded fi-om 13 genera of seven plant families. Certain families, such as Moraceae, Urticaceae and Tiliaceae, are characteristic of species fi-om phylogenetically distant groups, and it is unclear what factor of hostplant biology mediates the host choice. The family Rubiaceae is responsible for the great majority of records and appears throughout the genus, with rubiaceous feeders generally being unrecorded on other families. Despite there being hostplant records for almost 50% of the species in the genus, a number are unreliable and a great many are concentrated into two species groups, the A. serpa and A. phylaca group. No doubt this is due to the larvae feeding on plants which grow in disturbed areas in lowland regions, habitats most frequented by biologists. It is thus not possible at the present to draw conclusions about broad evolution of hostplant choice within the genus. It is, however, of possible evolutionary interest that the most primitive group in the genus, the A. alala group, has been recorded predominantly on the family Caprifoliaceae; this family is otherwise unknown in the genus, but is a common hostplant of other Asian and European limenitidines (Ackery, 1988) which may be close relatives of Adelpha.

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71 Immature Stages The pioneering work of Miiller (1886) in southeastern Brazil set the stage for studies of the immature life histories of Adelpha butterflies (see Table 3). Miiller reared, described and figured certain immature stages for 7 species of Adelpha, and his descriptions still remain some of the most detailed published to date. Moss (1927) reared 1 1 species within 6 months of his arrival in Para, Brazil, and provided the next major contribution to our knowledge (Moss, 1933), followed by Aiello (1984), and both of these authors figured larvae and pupae. Aiello (1984) summarised almost all of the published information and her paper remains the most important work to date on Adelpha early stages. There are a number of other reports of Adelpha early stages in the literature, but all suffer fi-om the same problems of identification as for hostplant records, and the vast majority offer no illustrations, only brief descriptions fi"om which taxonomic inferences can seldom be made. Oviposition. Females typically seek oviposition sites throughout the late morning and middle of the day, and the majority of Adelpha hostplants selected by females occur in recent openings in forest, and along the edges of forest and rivers. However, such hostplants are probably only those most easily located by the biologist, and it is highly probable that many species oviposit in the canopy. The female of A. iphiclus alights upon a leaf and walks backwards, searching for a suitable oviposition site with the tip of the abdomen. After oviposition the female takes flight, but may land on the same leaf and deposit fiirther eggs. Eggs are deposited beneath the surface of leaves by the abdomen being curled under fi-om the leaf upperside (Aiello, 1984). Reports of egg placement are rare in the literature and summarised by Aiello (1984). The majority of species deposit a single egg at the tip of a leaf on the upperside, while certain species typically oviposit next to a damaged portion of leaf (Aiello, 1984; DeVries, 1986). Up to four eggs may be deposited on a single leaf (Aiello, 1984), but often the plant chosen is a seedling and only a single egg is laid per plant.

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72 Eggs. The eggs of all Adelpha species known to date are typical of the tribe, having hexagonal, concave facets with setae at the interstices (see Aiello, 1984). First instar larvae. There are almost no detailed descriptions or figures of first instar larvae, which according to Aiello (1984) are all identical in form and some shade of brown or gray. The head bears setae but none of the chalazae of later instars, and only pale bumps on the body where future scoli are situated; I use the terminology of Petersen (1962) and Aiello (1984), and refer to thr first three thoracic segments as T1-T3, and remaining abdominal segments as Al-AlO. After their first meal the larvae take on the color of their hostplant, though the head remains brown. Second to fourth instar larvae. From the second instar on the head is ornamented with chalazae and numerous round, tiny pits, which may be of contrasting color to the head capsule. The body begins to develop scoli, arranged in three rows, subdorsal, supraspiracular and subspiracular. Typically the subdorsal scoli on T2 & 3 and A2, 7 & 8 are slightly larger than others. Some species undergo color and pattern changes, while in certain species, such as A. basiloides, the color depends on the hostplant (Aiello, 1984). Fifth instar larvae. Fifth instar larvae are diverse in form, color and pattern. All species have dense chalazae around the head capsule, the size of which may vary interspecifically (Aiello, 1984). The head capsule may also have darker vertical stripes (Mtiller, 1886; Aiello, 1984). The body scoli reach their peak of development, and exist in two main forms, termed terete and flattened by Aiello (1984). In almost all species they are reduced on the prothorax and first abdominal segment, while in known members of the A. alala group they are almost obsolete on all segments with the exception of the third thoracic segment (Otero & Aiello, 1996; pers. obs.). Terete scoli bear 3-5 radiating spines either at the apex, or at intervals along the length of the scolus, and may be short and thick or club-shaped and sparsely spined (Aiello, 1984). Flattened scoli are characteristic of more derived members of the A. serpa group, and have a leaf-like appearance. The

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73 spines are flattened, arranged in two opposite rows and may be densely packed, or even appear fused (MuUer, 1886). The majority of larvae have lateral, inclined darker stripes on the abdomen, while in members of the A. phylaca group these stripes are absent and there is instead a large, lateral block of dark coloration from Tl to A2 (Moss, 1933; Aiello, 1984). The larvae of A. alala is almost uniformly green (Otero & Aiello, 1996). Pupa. Pupae are also variable in morphology, but all have the posterior wing margins protruding as keels, and the second thoracic and abdominal segments are expanded to form a dorsal projection (Muller, 1886; Moss, 1933; Aiello, 1984). The thoracic projection is smaller and directed posteriorly, while the abdominal projection may be elongated to form a very large, laterally flattened hook (Muller, 1886; Moss, 1933; Aiello, 1984). The head typically bears a pair of laterally pointing projections which are variable in size, shape and orientation (Aiello, 1984). The pupal color varies from pearly white, yellowish, brown, green, or copper to a shimmering gold or silver, and the sutures may be black. All pupae apparently give the impression of being empty or diseased (Aiello, 1984). Development time. Aiello (1984) summarises the sparse available information on development time, which ranges from five to over thirteen weeks. Development time is rather variable within species and may depend on the hostplant (Aiello, 1984). A. basiloides is known to occasionally pass through six instars, which nevertheless take no longer to develop than the usual five. Larval behavior. Adelpha larvae, upon hatching, eat some or all of the egg shell and begin to feed on the leaf tip, leaving the midrib intact, a behavior observed in many nymphalids (Muller, 1886; Moss, 1933; Aiello, 1984). Larvae rest when not feeding facing away from the leaf stem on the exposed midribs, and extend them by means of fecal material bound with silk, often constructing similar supports on lateral veins. Several supports may be constructed during each instar. After moulting to the final instar, larvae rest on the upper surface of the leaves (Muller, 1886; Moss, 1933; Aiello, 1984).

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74 First to fourth instar larvae of Adelpha also engage in a distinctive activity first reported by Muller (1886), accumulating frass, and sometimes bits of dead leaf material, bound with silk, to form a mass surrounding the base of the support or suspended beneath. Adelpha basiloides also consistently constructs a small, usually curved larva-shaped mass on the leaf surface, away fi-om the leaf edge or usual mass, which closely resembles the larva. This behavior has also been observed once in A. iphiclus (Aiello, 1984). The behavior of A. alala is unique within the genus; from the late first instar onwards the larva constructs a shelter by making two opposing cuts in the leaf, perpendicular to the midrib, then pulling the two leaf flaps down and fastening them with silk. The larva rests in this shelter when not feeding, and the shelter entrance typically has a curtain of small pieces of leaf material left by the feeding first instar. Larvae utilise these shelters until the late fourth instar, after which they rest on the leaf surface. This shelter, which presumably confers a protective benefit, may explain the greatly reduced scoli of all instars. Similar shelter construction is known in the Old World species Limenitis Camilla and Ladoga reducta (Aiello, 1984), which utilise the same hostplant family (Ackery, 1988), supporting the primitive position of the A. alala group within the genus. Larvae of all species rest in a variety of positions, summarised in Aiello (1984).

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80 Adult Ecology Nutrition Adult Adelpha have been recorded at a variety of nutrient sources, of which the most numerically important is rotting carrion. This nutrient source has been mentioned occasionally in the Uterature (Adler, 1982; Sourakov & Emmel, 1995; Austin & Riley, 1995), but its pervasive utilisation by the great majority of the tropical forest butterfly fauna still remains unappreciated. Authors continue to place unwarranted stress on the importance of rotting frm\ in the diet of tropical butterflies (DeVries et ai, 1997), while most ignore rotting carrion completely. Of the 54 species recorded by myself during field work in Ecuador, 47 species (87%) were strongly attracted to rotting fish, and a great number of these were not observed outside of traps. Of the species not recorded on this nutrient source, only A. shuara is sufficiently common to believe that it might genuinely not utilise this source. I recorded only nine species on rotting fruit, and this low number is consistent with other studies (DeVries et ai, 1997). Many Adelpha have also been recorded feeding at damp sand and mud, particularly at urine, a behavior known as puddling. While most Adelpha have not been observed at nectar sources, the species that have been reported tend to be in more primitive groups within the genus, though an explanation for this is lacking. Hall & Wilhnott (1999) discuss the possible nutritive benefits fi-om feeding on rotting fish, which appears to offer similar nutrients to puddling (Downes, 1973). Sodium ions have been shown to stimulate puddling (Arms et ai, 1974), and this is a probable nutrient obtained fi-om rotting fish, which is fed on exclusively by males. Sodium has been shown to be donated to the female in the spermatophore during copulation, and is probably important for the higher neuromuscular activity of perching males (Downes, 1973; Arms et ai, 1974; Hall & Wilhnott, 1999). hi addition, though there is no evidence to date, males may also be seeking amino acids for maintenance of their thoracic muscle

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81 mass. Rotting fruits and nectar are probably both sources of carbohydrates, the energy source for powered flight. In addition to providing information on the nutrient requirements of adult Adelpha, I cannot stress enough the importance of trapping with rotting carrion for anyone conducting faunal inventories in the Neotropics. hi most localities, particularly primary forest, Adelpha species are seldom if ever observed, and certainly not the rarer species. Males of the recently described species A. hesterbergi and A. lamasi are known from Ecuador only through the use of such baits. Our understanding of the geographic and elevational distribution oi Adelpha would be inestimably greater if researchers in all regions used rotting carrion baits. I greatly hope that researchers will take note of the information on frapping techniques given in Sourakov & Emmel (1995) and Austin «& Riley (1995), and use it to contribute to our knowledge of Adelpha distribution. Perching and courtship Males of many Adelpha species have been observed "perching"; perching is defined as males occupying a specific area and flying out to investigate passing butterflies in search of females (Scott, 1976). Perching males typically maintain one to several perches, where they wait from early morning to early afternoon, usually in bright sun. Perches may be up to 30m apart, but while the male often flies from one perch to the next and back, I have never seen them move through a habitat as suggested by DeVries (1987). On the contrary, I have foimd perching species to be typically highly localised within a habitat, using the same bush for up to four consecutive years {A. argentea), and often not being encountered elsewhere within a site. Perching height varies from less than a meter above the ground (A. cytherea) to 15m {A. leuceria) and almost certainly higher, and while varying to some extent within species, there nevertheless appears to be a favored height range for each species. Males have been recorded perching in several areas, including open clearings or primary forest along rivers, forest edges and ridge top light

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82 gaps. While some species occur only in one of these microhabitats (A. shuara, A. hyas, A. argentea, A. serpa, along rivers; A. cocala, A. phylaca, on ridge tops and along forest edges), a number may be found in both {A. salmoneus, A. iphicleola). In addition, there are many common species for which there are no perching observations, and these probably perch high in the forest canopy. Perching males rest on the tip of the leaf upperside, with the wings typically closed or almost so, and are very wary. Flight between perches is typically gliding, with the wings held angled slightly downwards. Despite having observed perching males on numerous occasions, I have never observed courtship or copulation, and know of no published observations on either of these activities. Almost nothing is knovm about mate recognition, but due to the great variation in wing pattern between some subspecies, and the great similarity between many species, it is very probable that wing pattern cues play a small, or no part. This hypothesis is borne out by the possible hybrid specimens between the closely related but phenotypically very distinct species A. mesentina and A. thesprotia, and A. cocala and A. irmina. Instead, mate recognition and courtship in Adelpha may be pheromonally mediated; male Adelpha have a dense area of darker scales at the base of the ventral forewing, in cell Cu2, which is lacking in females, and although these scales are of a similar structure to remaining wing scales, they may have some role in pheromone dissemination. Mimicry The close resemblance on the dorsal surface of several Adelpha species, which proved to be unrelated when their early stages were examined, led Aiello (1984) to suggest that Adelpha were involved in mimicry. She tentatively suggested that species which fed on rubiaceous plants might form the models for other species. Unfortunately there is very little information available on the palatability Adelpha species, as is the case with all Neotropical butterflies. The experiments of Srygley & Chai (1990)

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83 demonstrated that 100% of specimens of A. cytherea and A. serpa celerio, the former a proposed model of Aiello (1984), were entirely consumed by caged jacamars, while more recently Pinheiro (1996) also found the few species that he tested to be palatable to kingbirds in Brazil. Nevertheless, there is now a sufficient body of evidence available to say without any doubt that Adelpha species are involved in mimicry, in the sense that dorsal wing patterns have converged onto a common pattern, both with other Adelpha and with species from other butterfly families. This is apparent both at the level of subspecies, and also at the species level, as illustrated by an examination of Figs. 10-94, the order of which is based on the phylogeny derived in Chapter 3. All that remains is to isolate the selective agent driving the wing pattern convergence. It is also worth noting that while the study of Pinheiro (1996) failed to find evidence for unpalatability in Adelpha, several species in classically unpalatable genera, such as Heliconius, were consimied with equal impunity. The Adelpha fauna of the Apure region in Venezuela, whence a number of new subspecies were recently described by Neild (1996), constitutes possibly the most compelling evidence of mimicry. No less than seven subspecies (A. cytherea nahua, A. cocala orellanai, A. malea malea, A. heraclea antonii, A. phylaca joffrei, A. capucinus gutierrezi and A. plesaure pseudomalea) occurring only in this small region have independently evolved an orange band bordering and touching the distal edge of the white band on the dorsal hindwing, a pattem that occurs outside of this region only in one subspecies from the neighboring Catatumbo region. Moreover, the female of Doxocopa lavinia, which closely resembles Adelpha throughout its range, also has the same hindwing pattem in this area (Neild, pers. comm.). Another sfrong example of independent convergence of a dorsal wing pattem is the narrowing of the white dorsal bands in a number of unrelated species that are superficially similar on the dorsal surface and which occur sympatrically in westem Ecuador. Such species include A. iphiclus, A. iphicleola, A. erotia L lerna, A. basiloides and A. barnesia. All of these species develop

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84 particularly broad white bands in the Rancho Grande region of northern Venezuela. There are numerous other examples of the close correspondence of ranges of subspecies in which the dorsal wing patterns change synchronously from one region to the next, discussed under each taxon. In a number of cases similar wing patterns are derived from the fusion and modification of different wing pattern elements, indicating that they are clearly not homologous, such as in the species A. naxia and A. iphiclm. As with other mimetic groups, several mimicry "rings" may occur sympatrically (Beccaloni, 1997a; Joron & Mallet, 1998). Maintenance of diverse mimetic assemblages has been suggested to be due to microhabitat partitioning (DeVries & Lande, 1999) through associated ecological factors, including hostplant height (Beccaloni, 1997b) and roosting height (Mallet & Gilbert, 1995), and either of these and/or some other ecological factor may be important in Adelpha. It is my experience that males of Adelpha species with similar dorsal wing patterns typically also perch in similar microhabitats; for example, in eastern Ecuador I have found A. thoasa manilia, A. serpa diadochus, A. hyas hewitsoni and A. iphicleola thessalita perching in a single river side clearing on the same bushes. "Orange and white banded" Adelpha, by contrast, have very few perching records, but their numerical abundance (pers. obs.) in traps in primary forest suggests that they perch in the canopy or subcanopy. Mimicry of Adelpha also appears to extend beyond the genus, and other species with very similar dorsal wing patterns to Adelpha include the females of most Doxocopa species, and the riodinid Synargis phliassus (Cramer), which Hiibner ([1819]) even included in the genus in his original description of Adelpha. Finally, it is worth noting that the simple, contrasting color patterns of the dorsal wing surfaces of Adelpha are certainly not cryptic, particularly given the habit of many species of resting (when not perching) with the wings open. More data are urgently needed on the palatability of Adelpha species to a wider variety of potential predators to more frilly understand the exact nature of the mimicry system.

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85 Biogeography Diversity. Adelpha species range from far northwestern U.S.A. south to Uruguay, and while the majority of species are strictly continental, four species occur in the Greater Antilles, of which two are endemic. Tabulating species and subspecies diversity by political boundaries (Table 4) is a simple first step to revealing broad biogeographic patterns, and it is important in terms of the allocation of resources for biodiversity conservation (McNeely et al., 1990). However, it is also apparent that sampling effort has a great effect on perceived taxa diversity in different countries, and as a step towards understanding the extent of this effect in Adelpha and other butterfly groups, and to more fully elucidate true taxa diversity, I also tabulate the number of expected taxa from each country in addition to those already recorded. Expected taxa are predicted either through extrapolation of a known range spanning either side of a country (in areas such as the Andes and Central America), or through examination of the ranges of more common species known to be sympatric in similar habitats and at similar elevations. Clearly, expected taxa will be underestimated for all countries at the periphery of biogeographic regions, since the first criterion is not possible to apply, and these countries include Mexico, Venezuela, Ecuador (west), Bolivia and Uruguay. The countries with the most complete faunal knowledge, excluding island areas and coimtries with very low species diversity, are Mexico, Costa Rica, Panama, Ecuador, Peru, Bolivia and Brazil. Countries whose fauna is particularly poorly known include most Cenfral American coimtries between Mexico and Costa Rica, the Guianas and Uruguay. There is an obvious latitudinal gradient in taxa diversity, which peaks in the countries of Colombia and Ecuador. Although Ecuador has the highest recorded species diversity of any country (56 species), this is due to extensive sampling there during the course of my research by myself and Jason Hall. Both Colombia and Ecuador are predicted to contain 59 species, or 69% of the entire genus, and it is possible that an

PAGE 93

86 additional species (A. fessonia) may be found in the isolated dry forests of southwestern Ecuador, which support other dry forest species with disjunct ranges (Willmott & Hall, unpubl.). By contrast, Peru and Brazil, often cited as the most species-rich countries in the world for butterflies (Brown, 1991), are significantly less diverse, with 46 and 42 species respectively, and reasons for this are discussed further below. Since Adelpha species occur in a wide variety of habitats at a broad range of elevations, and show much variation in ecological traits such as hostplant specificity and microhabitat preference, they are probably reasonably accurate indicators of diversity in other groups of butterflies (Brown, 1996). However, certain more sedentary groups, such as the Satyrinae and Ithomiinae, are likely to exhibit higher levels of endemism, probably more similar to the patterns evident from an analysis of the distribution of Adelpha taxa diversity. Again, Colombia proves to be clearly the most diverse country in terms of taxa, with a predicted total of 110 (53% of the genus), followed by Ecuador with a predicted total of 81 (39% of the genus). Of interest is the high number predicted for Venezuela, seldom mentioned in considerations of the distribution of biodiversity (e.g. Brown, 1991), which ranks third with 75 (36% of the genus). An explanation for these patterns of diversity may be gleaned from an examination of the distribution of species by biogeographic region (Table 5). I use the biogeographic regions as defmed by Brown (1982a,b), with several of the subdivisions of Brown (1979) and others that appear useful to understand the distribution Adelpha diversity. The reason for the high species and taxa diversity of Venezuela, Colombia and Ecuador is clear; all three countries contain faunas derived from three of the four principal biogeographic regions (Transandean, Andes and Amazon), while countries such as Peru and Brazil span only two regions (Andes and Amazon, and Amazon and Atlantic, respectively). Despite the high endemism of the Brazilian Atlantic region (11 species), this does not compensate for the great diversity of Andean and Transandean faunas.

PAGE 94

87 Table 4. Distribution of Adelpha taxa by country. Taxon D20mw3CZUcuO>Mo;cQ/.M XX?X??XXXXXXX X gorfmam XX?X??XXX X egregia X seriphia X p/o«e X X a9«r7/ja XXX therasia X X barcanti X 7XXXXXXX hewitsoni ? X ? viracocha X X X X X X flS/fl XXXXXX a/a/a X X completa X X negra X X X X X ar/cia ? X X portunus ? X serenita X ancia X X corcjTa X X X X corcyra X X

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88 Table 4. Continued. salazari X collina X X aretina X X dognini X X tracta XXX pithys XX ? X donysa XX XX donysa X X albifdum X X fessonia XXXXXXXXXXX ?X fessonia XXXXXXXXX ernestoi X cestus ? X lapitha ? X gelania XXX gelania X X arecosa X calliphane XXX? poltius X mythra XXX? basUoides XXXXXXXXXXXX />/«fl«re XXXXX X XXXX pseudomalea X X pAtoifl XXXXX X XXX plesaure X i^/woifl X ? ? X X gavina X falcipennis XXX? /Aoflsa XXXXXXXX ?X? man/Via X XXX thoasa XX X X gerona XXX calliphiclea ? X ? A/c/«s XXX?XXXXXXXXX?XX XXXX iphiclus XXX7XXXXXXXXX XX XXXX estrecha ? X epAeja ? X X iphicleola XXXXXXXXXXXX XXX ???X X iphicleola XXXXXXXX gortyna ? X X X Ihessalita ? XX leucales X XXX ???X iphimedia j( flAy/a X "•e/ona ? XXXXXXXXX XXXX ne;W( ? X deborah ? X X X leucocoma XXXXX X XXXX melona X pseudarete XXX er/teWa X? ???XXXXX ??X ssp. nov. X ? ? ? sophax ? X X eponina X

PAGE 96

89 Table 4. Continued. SwDwjOyouja.m<;a.mDOc/ ipL,H^3:;QfiiiJ zalmona X X ethelda X X galbao X ? ? X X?XXX? ?X agiV/a X ? X X X epione ? ? X syma X X X X cytherea X X X 9 X X X X X X X X X X 9 X X X X X marcia X X X ? X X X daguana X X X X X nahua X olbia X X X cytherea X X X X X X X X X aea ? ? X insularis X viola X X X X 9 X X X X pseudococala X X X X ? X X X X riola X salmoneus X X X 9 X X X X X X X X X salmonides X X X 9 X X X X emilia X X salmoneus X X colada X X X X demialba 9 X X ? epizygis X X X X epizygis X X X X epona X fabricia X X X X X X X 9 9 capucinus X X X X X 9 9 X X X X gutierrezi ? X capucinus X X X X X X X X X velia X ? ? X bamesia X X X X X 9 X X X X X X X X X leucas X X X X X ? X X X X X bamesia ? ? ? X X X trinita X diazi X hesterbergi X 9 9 X abia X X X 9 naxia X X X 9 X 9 X X X X X X X X 9 X 9 X naxia X X X 9 X ? X X X X X X X X 7 X 7 oteroi ? X hieronica X heraclea ? X 9 9 X X X X X X X 9 X heraclea ? X ? 7 ? X X X X ? X X ? X makkeda ? X antonii ? X sp. nov. X malea X X "} ? X X X X X X X X X X X X X fundania X X ? ? X X X X X X ? ixia X malea ? X aethalia X ? X X X X X goyama X X X X boeotia X X 9 9 X 9 X X X X X X X 9 X X 9 X oberthurii X X 9 ? X ? X X boeotia X X X X X X X X 7 X

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90 Table 4. Continued. Taxon DSOcQLLi3:ZOa.U>ULia.m<:a.mDOc/itx.l-^Sa:QeLU fidicula '! X altamazona ? ? ? X ? X X ? ? ximena XXXXX X ??X ximena XXXXX mossi ?X?XX X ??X delinUa XX? ?X?XXXXXXX utina XX??X?XX delinita XXXXX polling ?XXXX X ??X erotia X????XXXXXXXX X X?X erotia X????XXXXXXXX X X?X caphira X X phylaca XXXXX ? XXXXX phylaca XXXXX pseudaethalia XX?XX?XXXXX jojfrei ? X messana ?X????XXXXXXX X 11X1 messana ?X? ? ? 7XXXXX delphicola XXXXX X??X? thesprotia XXXXXXXX XXX mesentina XXXXX XX XXX fycorias XXXXXXXXXXXXXXXX? X melanthe XXXXXXXX wallisii X spruceana X X melanippe X fcra X X X X X X X fycorias XXX? ttttica 1 X X X X X X X hemileuca X X attica X X X X X X X leuceria XXIXXXXXX X leuceria XXI XXXXX juanna X X leucerioides X X erymanthis XI 1 X X X X X ssp. nov. X erymanthis X ? ? X X X adstricta X ssp. nov. X sichaeus XXXXX rothschUdi X X stUesiana X X boreas IXXXXXXX 1 XIX boreas 7XXXXXXX kayei X ? X ? X cocala XXllXXXXXXXXX X XXX lorzae XX? ?XXXXXXX orellanai ? X cocala XXXXX XXXX didia X caninia X fdderi XXX X leucophthalma XXXX X leucophthalma XXXX smalli X irminella ? X

PAGE 98

91 Table 4. Continued. Taxon DSomSlZUQ-0>Ma. m<:Q.eQDDc/iLi.^^S 3:Qa.O irmina -ST A. -5?-57V irmina A v A Y A tumida v A A A V A saundersu V X V V A V A frontina X X saundersii X X X helepecki X X lamasi X X salus X ? ? 1 X X X X ssp. nov. X ? ? ? X X salus X emmeli I X shuara X X f argentea X X X coryneta X X jordani 9 X X X zina X X X X X X X X lacina X X X zina X X X X inachia X desousai X pyrczi X enope X irma X X X X tnilleri X ? justina X X X X X justina X X maira ? justinella X inesae X valentina X X X X olynthia X 7 X X X levona X X ^ W D W D S 0 CD C/3 J w HON z COR PAN 0 z > ECU OS PQ Pn BOL <; Pi <: m > D Oi. D .W D 2 H JAM < a: 0 fi a! D p^ |CUB Taxa Total recorded 4 36 28 14 13 21 18 35 40 86 66 74 52 44 20 19 58 5 18 12 21 11 1 1 2 1 Total expected 4 38 34 24 28 31 32 38 43 no 75 81 57 50 24 24 61 10 27 26 26 11 1 2 2 1 1 Recorded as % 100 95 82 58 46 68 56 92 93 78 88 91 91 88 83 79 95 50 67 46 81 100 100 50 100 100 100 of expected Expected as % 1.9 18 16 12 14 15 15 18 21 53 36 39 28 24 12 12 29 4.8 13 13 13 5.3 0.5 1 1 0.5 0.5 of genus Species Total recorded 3 32 27 14 12 20 18 34 36 53 40 56 46 39 18 18 42 5 18 12 20 11 1 1 2 1 1 Total expected 3 34 33 23 27 30 32 37 37 59 45 59 48 45 22 23 43 10 26 25 25 11 1 2 2 1 1 Recorded as % 100 94 82 61 44 67 56 92 97 90 89 95 96 87 82 78 97.7 50 69 48 80 100 100 50 100 100 too of expected Expected as % 3.5 40 39 27 32 35 38 44 44 69 53 69 55 53 26 27 51 12 31 29 29 13 1.2 2.4 2.4 1.2 1.2 of genus

PAGE 99

92 § > « ^ ^ o o ^^ 3 c ^ o 81 = -S a gs p w si ai _ U U W W X X X tU X X ^ 3 Q 5j y l-l -fi . 5|~ = -S S o S E 5 S g -Si S UBOIUjUIOQ soieuiBf .is sapuyg sapuv M 'H AVIOO M eamiuv S AV o 1^ t Endemic/: fM)EMl Total/subi TOTAL i i usaiuiuioci Bqno sspnv 3 sspuv M eousiuv S M 1 X X X X C' e^ X X X X X X X X XXX XX 3 ^. a o C ^ S 3 ^ s. 5 ^ S -f 2 a 3 ?2 2. I ii g S4 t I 3 i £ 5.-§ S s UUDU xucuxxxx Itliiltiliif gas II ^ s> ^ c 3 P n ii sr^a, -3 -c

PAGE 100

93 An analysis of the distribution of diversity in Adelpha with elevation (Fig. 5) shows the expected pattern of a decrease in diversity with altitude, and a peak at around 7001000m. However, this peak in diversity does not necessarily reflect the local diversity within particular regions, but may be due to the greater endemism of species in low elevation premontane forests, and the extension of many lowland species into the lower elevation zone. 70-1 6050u « 40^ 306 I 20101 I I I I I I I I I I r 0 500 1000 lU n I I I I I I I I I 1 1 I I I I 1500 2000 2500 3000 Elevation above sea level/m Fig. 5. Variation in species diversity with elevation throughout the range of Adelpha. To examine the effect of elevation on local species diversity I tabulated data for Ecuadorian species on either slope of the Andes, thus eliminating the effects of changing latitude and mean diurnal temperature on elevational ranges. I have included only data collected by myself and Jason Hall, and by other reliable workers. While it is clear that several species, particularly on the western slope, have wider ranges than the known

PAGE 101

94 Ecuadorian range, in the majority of cases the Ecuadorian range is the broadest known for the species throughout its geographic distribution, indicating that the data are probably the best available. The distribution of species diversity with ahitude is illustrated in Fig. 6. Noteworthy is the peak in diversity at 600m elevation in the eastern Andean foothills, where a total of 35 species may be found sympatrically (a further three are predicted). Diversity on the western slope is much lower at this elevation, but slope diversities decrease and converge with increasing altitude. "I 1 1 1 1 1 1 I I 1— I— T— I 1 1 1 1 I I 1— I 1 1— I 1 1— I 1 °8888S88888888888888S8888888 ___._,„_„_^__r^r-)r)fNr)fN(NfN Elevation above sea level/m Fig. 6. Variation in Adelpha species diversity with elevation on both Andean slopes of Ecuador. The high diversity in the eastern foothills is due to the combination of two faunas, the lower montane Andean fauna and the Amazonian fauna. I analysed the upper and lower elevational limits of species from both slopes (Figs. 7, 8) to examine whether

PAGE 102

95 faunal changes occur at particular elevations or are continuously distributed, and the results suggest the former. On the east the pattern is less clear, partly due to the absence of lower limit records from very low elevations since the eastern lowlands do not extend below 200m. Nevertheless, there is a peak in lower elevational range at 2-300m, corresponding to the Amazon fauna, which has its upper limits peaking between 8001200m. Lower montane species have a lower elevational limit of 400-600m and an upper limit of 16001900m, while higher montane species have a lower limit of 12001300m and an upper limit of 2000-2700m. Although some of these patterns may be explained by varying sampling intensity at different elevations, the emergence of similar patterns on the western slope provides support that they are not artificial. 25 Fig. 7. Upper and lower limits of elevational ranges for Ecuadorian Adelpha on the east Andean slope. Values are pooled for pairs of adjacent elevations to compensate for sampling errors.

PAGE 103

96 ^ 5 I' 3 2 1 m 8 S S 3 ^ r*l Q Lower limit of range I Upper limit of range 8 S S S I \r> 0\ i ^ i i (N VO 00 Elevation above sea level/m 8 i i i 8 8 8 8 ^ m »r> (S
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97 proportion of shared species occurs at 1000m, the altitude at which most Choco endemics occur (e.g. A. lamasi, A. rothschildi, A. levona). Most of the species occurring in lowland western Ecuador, and indeed the entire Transandean region, are widespread, and this biogeographic region lacks the high endemism of the Amazon. East Andes West Andes I I I I I I I I ogggggooggoggoogogggggggggog Elevation above sea level/m
PAGE 105

98 (Brown, 1982a,b), this is not the case for montane species, with the exception of Adams (1985). I suggest that the relatively low endemism in montane faunas between Andean slopes above 1500m, and the preservation in habitats above this altitude of primitive species in the genus (e.g. A. diodes, A. alala group), might be explained by the long term stability of such habitats. If this hypothesis proves to be widely applicable across other plant and animal taxa it may be significant in terms of the conservation importance of montane areas, since primitive species are often more genetically isolated from the most closely related species and therefore might be considered of higher conservation value (VaneWright et al, 1991). Review of the genus Adelpha The species are arranged here in an order jfrom most primitive to most derived, following the phylogeny derived in Chapter 3 (Fig. 250) and an analysis of wing pattern characters that vary too much throughout the genus to be coded for cladistic analysis. Such characters are discussed under each species, in addition to the characters that I consider important in their identification and which unite the taxa included in each; this information is presented separately fi-om the constituent taxa in the case of polytypic species. In a very few cases there is no single character that defines the species, but taxa are grouped on the basis of shared characters between pairs of allopatric taxa. I use the terms DFW, VFW, DHW and VHW to refer the dorsal and ventral fore and hindwing respectively. For each subspecies 1 discuss how it is distinguished fi-om other subspecies, variation, and the nomenclature and associated synonymy. The dates of publication of names are those given in Lamas et al. (1995), except those for the plates published by Fruhstorfer in Seitz ([1907]1925). In this case, plates 106-109 were published in 1913, plate 110 in 1916 and plate llOA in 1920, and I am informed by Lamas (pers. comm.) that the dates for these plates given in Lamas et al.

PAGE 106

99 (1995) are erroneous. Treatment of the names which appeared on plates 106-109, two years before the text (Fruhstorfer, 1915), is a complicated nomenclatural problem. These names appeared individually beneath figures, with the name "Adelpha" at the top of the plate, and the taxonomic status (species, form, subspecies) that has been accorded them by all subsequent authors, as far as I am aware, is that of Fruhstorfer' s (1915) text. However, Gerardo Lamas (pers. comm.) is of the opinion that such names should be regarded as species with the spelling as on the plate, and Philip Tubbs (pers. comm.), of the ICZN at the BMNH, also confirms this to be the procedure resulting fi-om strict application of the Code. Although the author of the names is not apparent from examination of the plate alone (as pointed out to me by L. Miller, pers. comm.), and the specific name is not explicitly combined with a generic name, such a description is still considered to be valid by the Code. The publication date for these problematic names is therefore 1913 and all are regarded as of specific status, and since the type locality in all cases has been determined fi"om an "outside" source (Fruhstorfer, 1915), I have enclosed such localities in square parentheses. Since there is no information on the plates as to the number of specimens on which the name is based, I have regarded all type material for such names as syntypic, regardless of indications to the contrary in the text (Fruhstorfer, 1915). After the type specimen data, I list the first author to use alternative taxonomic combinations, misidentifications, and the most recent taxonomic combination, which I use to determine whether or not the combination presented here is new or revised. Ideally, new taxonomic combinations should be accompanied by some indication that the author was aware of the fact, but since virtually no previous authors ever explicitly stated that they were making new or revised taxonomic combinations, I have been somewhat arbitrary in my choice of whether or not to regard a combination as valid. I have tended to treat new combinations as valid if proposed in works concentrating either specifically on the genus, or provide fiirther information about taxa, on the basis that the author might

PAGE 107

100 have made more effort in establishing the correct taxonomy. Taxonomic combinations in simple lists of species from coimtries or localities containing no further discussion about the taxon in question have been typically ignored, since many taxonomic combinations in such lists result from an uncritical literature survey or from misidentifications of taxa, and it is often impossible to ascertain which taxon is being referred to. However, even in the case of the latter, I have generally given credit to the authors for new taxonomic combinations that I beheve to be vaUd, even if the taxonomic change was not specified, but have not regarded taxonomic changes that I beheve incorrect as valid unless accompanied by a specific statement of the author's intent. I have not attempted to list the numerous misspellings of names in the literature (e.g. Martin et al., [1923]), and have corrected trivial misspellings in citing previous taxonomic combinations and misidentifications. I use the term syn. nov. to denote a name regarded for the first time as infrasubspecific, stat nov. for any name formerly considered to be of different taxonomic rank or, if a subspecific name, combined with a different specific epithet, and stat. rest for a taxonomic rank or combination reverted to that of a previous author. I have also listed unavailable names, proposed as quadrinomials, under the appropriate taxon, with the bibliographic and type data, since there is the possibility that such names may have been made available by the action of subsequent authors (see under A. seriphia godmani). I list nomina nuda and species erroneously placed in Adelpha in Appendix B. Ecological information on the adults and immature stages is presented for each taxon from my own field work and published sources. Hostplants are listed in Table 2 and are usually not mentioned in the species account, unless there is some problem in identification or extra ecological information that cannot be included in the Table. Specimen data are hsted as detailed above under Materials and Methods.

PAGE 108

101 Adelpha bredorvii Geyer, 1837 Figs. 10; 95; 178 Identification, taxonomy and variation: Adelpha bredowii is readily distinguished from all other species in the genus by its large size, broad wing shape and contrasting ventral colors of white, blue-gray, dark redbrown and orange bands on a black background. There is variation in the extent of the orange DFW subapical marking, the width of the white DFW postdiscal spots at the costa, the configuration of the white DFW postdiscal band, the presence or absence of a second orange band crossing the end of the VHW discal cell and the extent of the bluegray line distal to the white postdiscal band on the VHW, and I recognise three subspecies. Carpenter and Hobby (1945) give an exhaustive discussion of wing pattern variation in this species and only major variation will be noted here. The generic status of this species has had a rather unstable history, for although Geyer (1837) correctly placed it in Adelpha in his original description, being the first author to use Hiibner's recently erected genus, A. bredowii has since oscillated almost continuously between Adelpha, the junior synonym Heterochroa, and Limenitis. This is partly due to its large size and ventral colors which are unusually bright for Adelpha, superficially recalling the North American species of Basilarchia (usually placed in Limenitis) with which A. bredowii conmionly occurs (Seitz, 1915). In addition, the eyes of A. bredowii are smooth, rather than setose as in almost all Adelpha, a character shared with the remaining North American limenitidines (Godman and Salvin, 1884). The lack of other obvious, extemal structural characters by which to separate Basilarchia and Adelpha exacerbated the problem, and the generic placement of the species seems to have been based largely on personal preference. In fact, A. bredowii is closely related to A. diodes, as evidenced by several characters of the male and female genitalia, and probably A. herbita. Adelpha bredowii and A. herbita share the entire loss of the outer postdiscal series on the VHW except in cell Cu2, unique in the Adelpha serpa group. The brownish

PAGE 109

102 shading in the postdiscal area of the VHW of A. bredowii appears to represent the ground color, as indicated by the fact that it differs in color and surrounds the brighter orange marking that represents the outer postdiscal series in cell Cu2. A number of other wing pattern and genitalic characters place A. bredowii firmly within the A. serpa species group, as one of its most primitive members, and one of the most primitive members of the genus (see Chapter 3). Synapomorphies of the Adelpha serpa group possessed by A. bredowii include the reduction of the basal streak in the DFW discal cell, which lies close to the costa and borders an area of dense reddish scales, the absence of a basal streak in the VFW and VHW discal cells, the well-defined black line that extends fi-om the base of the VHW to the tip of the humeral vein, the loss of the clunicula in the male genitalia and gain of a sclerotised, spiny pad in the aedeagus, and the coarsely spined, sclerotised bands on the corpus bursae of the female genitalia. Adelpha bredowii is the only member of the A. serpa group in which the iimer postdiscal series is visible on the VHW, with this marking probably being lost in remaining species, suggesting it may be the most primitive species in the group. Range and status: Western U.S.A. throughout Mexico to Guatemala. Scott's (1986) report of Honduras is based on mislabelled specimens. Common, fi"om near sea level to 2000m. Specimens examined: 1318 (903m, 41 5f) Adelpha bredowii bredowii Geyer, 1837 Figs. 10a,b Adelpha bredowii Geyer (1837: 10, n. 413) TL: Mexico. Types: ZMHU: STf: Mexico//Syntype//Origin//Coll. Sommer [examined] Heterochroa bredowii Geyer, Westwood (1850); Limenitis bredowii HObn. [sic], Edwards (1870), Seitz (1915); Adelpha bredowii bredowii Geyer, Emmel et al. (1 998)

PAGE 110

103 Identification, taxonomy and variation: This subspecies differs from A. b. eulalia in having a narrower, less friangular subapical orange marking on the DFW which forms a spot in cell M3 and extends as orange scaling into cell Cul; in other subspecies the orange marking is much broader at the costa and seldom extends into cell M3, and if so, only as sparse orange scaling. The white postdiscal spots at the costa are much smaller, approximately half the width of the white postdiscal spot in cell M2 instead of being of approximately equal width, and the hindwings are less rounded. Adelpha b. califomica is easily distinguished by having an exfra orange band across the end of the VHW discal cell, the blue-gray band distal to the white VHW postdiscal band broken in the middle, by its smaller size and by a more distally displaced white postdiscal spot in cell M2 of the DFW. hi the small series of specimens examined of this subspecies there is slight variation in the size and extent of the DFW orange subapical marking, and it is possible that more extensive collecting will show this character to vary clinally to that of A. b. eulalia. Both surfaces of this species were accurately figured by Geyer ([1832]-[1833]: pi. 142, fig. 825, 826) several years before he introduced the name based on an unspecified number of male specimens which he had received from Herr Sommer (Geyer, 1837), and there has never been any question as to its identity. I have examined a syntype female in the ZMHU which differs from the original figure in having a more irregular white DHW postdiscal band, the veins in the dark red-brown VHW postdiscal area not lined with silver-blue, and no tiny white postdiscal spot in cell M2 on the DFW, but it is otherwise similar and has all of the characters which distinguish this subspecies from others. It is also set with the forewings distinctly far forward, uncommon for specimens of that time and unlike the original figure. It seems unlikely that this was the figured specimen, but since it has data indicating it to be from the collection of Sommer it is almost certainly a syntype, notwithstanding Geyer's statement that the description was based only on males.

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104 Range: Pacific slopes of central western Mexico, in the states of Guerrero, Jalisco, Oaxaca, Michoacan and Morelos. Vargas et al. (1996) list A. b. eulalia as occurring in Jalisco, but also list de la Maza's (1987) record of A. b. bredowii from that state as A. b. eulalia. Since de la Maza (1987) figured both of these subspecies correctly I have followed his identification here, having seen no specimens fi-om that state myself I have seen three specimens labeled "Guatemala", two of which, in the BMNH, were commented on by Carpenter and Hobby (1945), who suggested, probably correctly, that they had been mislabeled. Habitat and adult ecology: De la Maza (1987) reports that this subspecies has been found in Mexico in pine-oak forests in March. It has been recorded up to 2400m, and its relative rarity in collections probably reflects its limited geographic range. Specimens examined (23m, 3f): t trans, to A. b. eulalia MEXICO (20m, 30: Guerrero: Omiltene 8000* Jul. Nov. 7m BMNH, 2m AME; Rio Balsas Apr. Im USNM, Im, If AMNH; Xucumanatlan 7000' Jul. Im BMNH; no specific locality 2m BMNH; Michoacan: Uruapan Im BMNH; Morelos: Cuemavaca Sep. Im BMNH; Oaxaca: Oaxaca Im BMNH; Not located: no specific locality 2m, If BMNH, Im, If ZMHU, 2m MNHN. GUATEMALA (3m): no specific locality 2m(lt) BMNH, Im BMB. Additional locality data: MEXICO: Guerrero: Cruz de Ocote; Filo de Caballo; Jalisco: Autlan; Mexico: Valle de Bravo (de la Maza, 1987); Jalisco: km. 90 rd. to Autlin Sep. (Field, 1940b); Bosque Escuela Jan. Mar. Apr. Aug. Sep.; Est. Cientifica Las Joyas La Ordefiita Mar. (Vargas et al., 1996: as A. b. eulalia); Michoacan: Carapa Sep. (Field, 1940b); Oaxaca: 12 mi. S. of Oaxaca (Carpenter and Hobby, 1945). Adelpha bredowii eulalia (Doubleday, [1848]) Figs. 10c,d; 95a Limenitis eulalia Doubleday ([1848]: pi. 36, fig. 1) TL; [Venezuela]. Types: BMNH(M): STf "? The Type specimen figured L. bredowi Hbn. s.sp. eulalia Dbl. Det. G.D. Hale Carpenter 1943//2. Heter. ? Bredowii Hb. Zutr. 825-6 Men. p.l Ig-Dbd. p.279 No. 12 Eulalia Dbd. Dium. pi. 36 fig. 1 Mexique donne por M. Doubleday Dbd. a reconnu la Bredowii apres qu'il a fait figure son eulalia//124//Syntype//Type//Ex. Musaeo Ach. Guen6e//Ex. Oberthar Coll. Brit. Mus. 1927-3" [examined] =Limenitis bredowii guatemalensis Carpenter and Hobby (1945: 318, pi. 1, figs. 7-9, pi. 2, fig 12) syn. nov. TL: San Geronimo, Guatemala. Types: BMNH(T): HTm: "Genitalia C//Holotype L. bredowi Hbn. f n. guatemalensis TYPE G. D. Hale Carpenter 1943 M.S. Photo fig. 7 I7//B.C.A. Lep. Rhop. Adelpha bredowi Hbn. Godman-Salvin Coll. 1916.-4.//San Geronimo Guatemala F.D.G. and O.S.//nV/Type H.T."; ATf Polochic Valley; BMNH(M): PTf: San Geronimo; 3PTm: Polochic Valley; PTm: Rio Polochic; 2PTm: Guatemala city; PTm: Verapaz; PTf: Central Valleys; PTm: Mexico [all examined] =Heterochroa bredowii Geyer, Westwood (1850); =Limenitis bredowii HUbn. [sic], Edwards (1870); =Adelpha bredowii HObn. [sic], Kirby (1871); =Limenitis bredowii bredowii Geyer, Seitz (1915); Limenitis bredowii eulalia Dbld., Carpenter and Hobby (1945); Adelpha bredowii eulalia Dbld. and Hew. [sic], Emmel et al. (1998) Identification, taxonomy and variation: Adelpha bredowii eulalia is distinguished fi-om the nominate subspecies in the account of that taxon, and fi-om A. b. californica by lacking a second orange bar at the end

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105 of the VHW discal cell, by having a complete blue-gray line distal to the white postdiscal band on the VHW, and by having the v^hite postdiscal blocks on the DFW in cells Cu2M3 more aligned. The extent of the DFW orange subapical area is slightly variable, sometimes extending as orange scaling into cell M3, and the white DFW postdiscal band also exhibits minor variation in width. A white postdiscal marking is variably present in cell M2 on the DFW, as discussed below. Doubleday ([1848]) accurately figured the dorsal surface of both wings of this subspecies but gave no indication of the specimen on which the figure was based, while the list of plates in Vol. 1, p. viii, gives the certainly erroneous locality of "Venezuela". A syntype female in the BMNH corresponds well with the original description, however, and originated in Mexico. Westwood (1850: 278) placed L. eulalia as a synonym of Adelpha bredowii, a taxon of which it appears Doubleday was unaware when he made his original illustration, and gave Mexico as a locality for both A. bredowii and L. eulalia. Although several subsequent authors (Seitz, 1915) followed Westwood, I discuss my reasons for regarding A. b. eulalia and A. b. bredowii as distinct under the latter taxon. After a very thorough discussion of variation in A. bredowii eulalia. Carpenter and Hobby (1945) described a new subspecies, guatemalensis, for Guatemalan specimens. Their type series is in the BMNH. They state that the subspecies differs from A. b. eulalia in having a narrower DFW orange subapical marking, a broader white dorsal postdiscal band, and a better developed white postdiscal spot on the DFW in cell M2. While this latter spot is often absent in A. b. eulalia, it is present in a number of specimens throughout the range of the subspecies and occasionally as developed as in certain Guatemalan specimens. In addition, although generally the orange DFW subapical marking is slightly narrower and the white dorsal postdiscal band broader in Guatemalan specimens than in typical A. b. eulalia, both of these characters are variable and do not enable consistent recognition of a distinct taxon, and I therefore synonymise guatemalensis with A. b. eulalia (syn. nov.).

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106 Range: Southeastern U.S.A. in the states of Arizona, Utah, Texas, New Mexico, Colorado and Nevada, through Mexico to Guatemala, along the Atlantic slopes of the latter countries. I have examined a few specimens labeled from areas in California, and although these particular records are probably dubious, the subspecies does occur in the extreme southwest of that state (Emmel and Emmel, 1973; Ferris and Brown, 1981). Field (1940a) reported a single female from Scott City in Kansas, and Ferris and Brown (1981) also report it as strays from that state and provide a map of collection localities for Rocky Mountain states in the U.S.A.. Immature stages: Harry (1994) provided the first (and last) fully illustrated color photographic life history of any Adelpha for this subspecies, reared in Utah, U.S.A.. The egg is green and laid singly on the upper surface of the leaf at the margin, at the tip of a leaf "tooth". It appears to be typical of the genus in morphology. The first instar is pale greenish with a brown head and dark spots marking the positions of the ftiture subsdorsal scoli. The second instar has slight projections in place of fixture scoli and has lateral dark brown, pale yellow green and green oblique stripes on each segment, typical of many Adelpha. The third instar has developing scoli on segments T2, A2, A4, A7 and A8 and a similar pattern to the second instar, except with the markings less well defined. The head capsule is pale brown with vertical dark brown stripes. The fourth instar has well developed, orange brown subdorsal scoli on T2, T3, A2, A4, A7 and A8, each with black lateral spines, small, pale suprzispiracular scoli, and the head capsule appears to have a relatively smooth face with two especially long, black tipped dorsal spines. It is pale greenish with paler bumps and dorsally is darker green anterior of Al, between the scoli bases on A7 and A8, and on A9/10. The final instar is a uniform bright green, slightly darker dorsally, with a brown venfral surface, long, relatively thick scoli on T2, T3, A2, A4, A7 and A8 (longest on T2, A2 and A8), each orange brown with fine, thin pale whitish or black lateral spines. All scoli are straight, the thoracic pair anteriorly inclined, those on T3 also orientated slightly laterally. There are short subdorsal scoli on segments

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107 A3, A5 and A6, lacking lateral spines, while all other scoli, notably also the supraspiracular scolus on T2, appear to be absent. Just prior to pupation the larva becomes a yellowish brown. The pupa is dark brown on the wing pads and Tl, but otherwise appears to be silver, with a moderate dorsal projection on A2 and a smaller dorsal hump on T2. The head horns are short but their shape is not visible. Other authors have also described various stages, and it appears that mature larvae may also be mottled green (Ferris and Brown, 1981). Scott's (1986) description may apply either to this subspecies or A. b. califomica. He states that the larva occurs in a second form, which is yellow orange with a brown lateral line bordering the paler brown ventral surface. The pupa may also be straw colored with finer brown marks and gold streaks. Early instars construct typical perches from leaf veins, on which they rest in the Front Curved position (of Aiello, 1984). Harry (1994: figs. 5, 7) also illustrates third and fifth instar larvae resting in the FrontArched-Rear-Up position (of Aiello, 1984), with the face parallel to and closely pressed against the leaf Larvae are in diapause over the winter, according to Ferris and Brown (1981) and Scott (1986). Habitat and adult ecology: In the U.S.A., this subspecies is locally common in moist lowland areas down to 300m. It frequents riparian canyons and forest areas along streams, where both sexes rest frequently on the tips of branches to sun themselves (Ferris and Brown, 1981). Males often congregate at puddles and moist sandy patches along streams (Ferris and Brown, 1981). Bailowitz and Brock (1991) report that this subspecies is most common in mountainous areas, usually above 1300m, though it also has been rarely sighted some distance from natural habitats, in agricultural areas and even in tovms. Adults perch high in oak trees and may occasionally nectar at flowers of Chrysothamnns and Baccharis. Scott (1986) also states that adults of this subspecies, or .4. b. califomica, may be found feeding on fruits (presumably rotting) and aphid honeydew. There are several broods throughout most of the range in the U.S.A., though Emmel and Emmel (1973) report it to have only a single brood, with adults flying from May to July, in the

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108 desert mountain ranges of southern California. Gary (1994) reports that the species flies in southwestern New Mexico from May to June, then August to October, while there are records of adults from elsewhere in the U.S.A. from as early as April and as late as December (Ferris and Brown, 1981; Bailowitz and Brock, 1991). In Mexico the subspecies has been reported from pine-oak forests from February to October, with a peak of abundance in the middle of this period (de la Maza, 1987; de la Maza and de la Maza, 1988), and it has been recorded from 700-2 100m in Chiapas in hot to temperate, humid to semi-humid forests (Beutelspacher, 1983; de la Maza and de la Maza, 1993: as L. bredowii guatemalensis). In Guatemala it has been reported from montane areas above 1200m (Godman and Salvin, 1884). " " ' ' Specimens examined (386m, 1 900: t trans, on dorsal surface loA.b. califomica; i trans, to b. bredowii. U.S.A. (304in, 1400: Arizona: Alpine Jul. Im, If AMNH; Arcenta Canyon Jun. Im USNM; Baboquivari Mts. Jun. Oct. 2m USNM, If AMNH; Chiricahua MU. May Jun. Jul. Sep. Im, If USNM, 8m, 5f AMNH, Im AME, 7m, 9f FSCA; Cochise Co. Im AMNH, 2m MCZ; Coyote Mts. 1 f AMNH; Dinnebito Wash Aug. Im USNM; Ragstaff May 2f USNM; Fort Grant Jul. lfUSNM;Gila Co. Jun. Im MCZ; Globe May 5m AME, 3m AMNH; Globe, 12 mi. N.E., 4000' May Im, If AME; Graham Mts. Aug. Sep. If USNM, Im AMNH; Grand Canyon Aug. Im FSCA; Heber-Pine Sep. Im, If AMNH; Herb Martyr Dam Sep. 2m FSCA; Hereford Jul. Aug. 2m MCZ; Horsethief Basin Jul. 2m FSCA; Huachuca Mts. May Jul.-Sep. Um BMNH, 8m, 5f USNM, 10m, If AMNH, 2m, 3f FSCA, Im MCZ; Jerome, 2 mi. S., 1000' May 3m USNM, If FSCA; Madera Canyon Pima Co. Santa Rita Mts. Jun.-Sep. 18m, 2f USNM, 4m AMNH, 5m, If AME, 3m FSCA; Madua Canyon Santa Cruz Co. 4800-5400' May Jul. Sep. 3m, 2f AME; middle Pioneer Camp Pinal Mte. Aug. 31m, 15f AMNH; Oak Creek Canyon Jun. 2m, 2f AMNH; Onion Saddle Sep. Im, If FSCA; Palmerlee Aug. Im, If BMNH, 9m, 4f USNM; Paradise Jun. Aug. Sep. Im USNM, 9m, If AMNH, Im FSCA; Patagonia Mts. Jun. Sep. 3m AMNH; Payson, 10 mi. S., Aug. Im AMNH; Pine Creek Jun. Aug. Sep. 2m FSCA, Im, If USNM; Portal 5400' May-Jul. Sep. Oct. If USNM, 11m, 15f AMNH, Im, If MCZ; Prescott ImBMNH, ImUSNM; Ramsey Canyon (Huachuca Mts.) Jun.-Sep. 8m, 7f AMNH, 3m, If AME; Rose Creek Jun. 5m, If USNM; Santa Catalina Mts. May 6m, 2f AMNH; Santa Rita Mts. Sep. Im FSCA, If AMNH; Senator Jun. 6m, 6f BMNH; Silver Creek Jun. Im, 2f AMNH; Skeleton Canyon Oct. Im FSCA; Sonoita Jul. If USNM; Water Canyon 5000' Im BMNH; White Mts. May Jun. 6f AMNH; WiUiams Im, If USNM; no specific locality Jun. Jul. Im, If BMNH, Im, 2f USNM, 2m AMNH; California: Redlands, San Bemadino Mts. Jul. 5 1930 Im AME-error; Los Angeles Jun. 12 1923 Im AME-error; Fresno Jun. 18 1928 Im AME-error; Colorado: Durango Aug. Im AMNH; Nevada: Charleston Mts. May Nov. 3mt, I ft USNM, 3m AMNH; Clark Co. Jun. 3mt USNM; New Mexico: Brooks Ranch Sep. 3m AME; Cherry Creek, Pinos Altos Mts. Jun. Jul. Sep. 5m, If USNM, Im FSCA; Central New Mexico Jun. 2m, 4f USNM; Fort Wingate Sep. Im AMNH; Hell Canyon Oct. Im USNM; Jemez Springs Sep. Im AMNH; McMillan Camp Jul. 24m, 8f AMNH; MogoUon 7200' 2m, If AMNH; Organ Mts. 5600' Jun. Im, If AMNH; Pinos Altos Sep. Im AME; San Mateo Mts. 7400' Jul. Im AMNH; Tajaque Canyon Jun. Im FSCA; Texas: Alpine Jul. Im, 2f AMNH, Im FSCA; Big Bend N.P. 3-4000' Sep. If AMNH; Browne Im AMNH; Edwards Mts. Jul. Im USNM; Green Gulch Big Bend N.P. Aug. 2m FSCA; Kerrville Apr. Oct. Im, If USNM, If FSCA; New Braunfels Oct. 2m FSCA; Sunny Glen Ranch Jul. If AMNH; The Basin Big Bend N.P. Aug. Im FSCA; no specific locality 2m BMNH; Utah: Hellhole Pass Denver Dam Mts. Aug. If AME; Leeds Creek Pine Valley Mts. Washington Co. May Jun. Sep. 5m, 2f AME; Oquirrh Mts. Tuoelb Co. Aug. Im AME, Im FSCA; Stockton Aug. Sep. 2f USNM; Tooele Aug. Im AMNH; Utah Im USNM; Not located: "California" Im MCZ; no specific locaHty If BMNH. MEXICO (59m, 370: Chiapas: Campet Aug. Im AME; Chiapas 2m AMNH; Ugos de Montebello I500-I800m Feb. If AME; Montibe Sep. 3m, 3f AME; Santa Rosa 4m, 2f AME; Chihuahua: Catarinas Jun. If AMNH; Chihuahua Sep. Im AMNH; Cidnegas Jun. If AME; La Mesa de la Avema Im AME; Madera Jun. 6m, lOf AME; Pinos Altos If BMNH; Primavera Jun. 2f AMNH; Distrito Federal: Pedregal Apr. Aug.-Oct. 14m, 4f AME, If AMNH; Durango: nr. Durango City If BMNH; Encino Jun. If AMNH; Guerrero: Acahuizotla Oct. 4m AME; Apuico Nov. Im AME; no specific locality 2m BMNH; Hidalgo: Cuesta Colorada 8500' Aug. Sep. 3m AMNH; Puerto de Zorro Apr. If FSCA; Zimapan, 5 mi. N.E., 2100m Aug. If AME; Mexico: Amecameca Apr. If BMNH; Chalma Jul. 3m AME; Nuevo Leon: Hda. Vista Hermoso Villa Santiago 1500' Jun. Im, If AME; Oaxaca: Santa Rosa, 27 km. S.E. Sep. Im BMB; no specific locality 2m(lt), If BMNH; Puebla: Puebla 7000' Apr. If BMNH, Im, If MNHN; Tehuacan 5400' Nov. Im BMB; Sonora: Bacarac Sep. Im AMNH; Cananea Aug. 3m AMNH; Not located: Bolanos-Guadalajara rd. Jul. If USNM; S.W. Colonia Juarez Sep. Im AME; Yautepec Aug. Im AME; no specific locality Im, If BMNH, Im AMNH. GUATEMALA (16m, 60: Alta Verapaz: Polochic Valley 3m, If BMNH, Rio Polochic Im BMNH; Baja Verapaz: San Jeronimo Im, If BMNH; El Quiche: Colotenango Im USNM; Llano del Coyote Sacapulas Oct. Im AME; Guatemala: Guatemala city 2m BMNH, 3m USNM; Santa Rosa: Verapaz Im BMNH; Zacapa: no specific locality Im, If USNM; Not located: Central Valleys If BMNH; no specific locality Im BMNH, Im ZMHU, If MNHN, If MCZ. COUNTRY UNKNOWN (7m, 70: no specific locality 5m, 2f USNM, 2m, 3f AMNH, 2f MCZ. Additional locality data: U.S.A.: California: New York Mts.; Providence Mts.; Granite Mts. N. of Amboy (Emmel and Emmel, 1973); Kansas: Scott City (Field, 1940b); New Mexico (all localities are counties precise data in Toliver et al. (1994)); Bernalillo;

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109 Catron; Chaves; Cibola; Colfax; Dofla Ana; Eddy; Grant; Hidalgo; Lincoln; Los Alamos; Luna; McKinley; Mora; Otero; Rio Arriba; Roosevelt; Sandoval; San Juan; San Miguel; Santa Fe; Sierra; Socorro; Taos; Torrance; Valencia; Apache; Culberson; El Paso; Jeff Davis; Mun. Batopilas; Mun. Casas Grandes; Mun. Janos; Mun. Urique (Toliver et al, 1994); Texas: Concan; Leaky Jul. (Field, 1940b). MEXICO: Chiapas: El Chorreadero 700ni Jan. Aug. Nov. Dec. (Beutelspacher, 1983); Depresidn Central; Macizo Central (de la Maza and de la Maza, 1993); Distrilo Federal: Pedregal de San Angel (Beutelspacher, 1980), Padiema; Tlalpan; Chihuahua: Madera; Durango: Michilia; Hidalgo: La Encamacion; Morelos: Tepoztlan; Nuevo Leon: Monterey; Oaxaca: Ixtlan (de la Maza, 1987); San Luis Potosi: Sierra de Alvarez 1200m Feb.-Nov. (de la Maza and de la Maza, 1988). Adelpha bredowii californica (Butler, 1 865) Figs. 10e,f; 95b; 178a,b Heterochroa californica Butler (1865a: 485) TL: California [U.S.A.]. Types: BMNH(T): STf: "Califor. 56 48//B.M. TYPE No. Rh. 9766 Heterochroa californica f Butl.//16//Type AGB//Syntype//Type" [examined] =Limenitis bredowii Hubn. [sic], Edwards (1870); Adelpha californica Butl., Kirby (1871); Adelpha bredowii californica Bull., EmmeleJo/. (1998) Identification, taxonomy and variation: This subspecies is distinguished from the nominate and A. b. eulalia by having a second orange cell band at the end of the VHW discal cell (this is very occasionally absent (Carpenter and Hobby, 1945)), by the blue-gray band distal to the white postdiscal band being broken or weaker in cells M2 and Ml, its smaller size, more rounded wings and narrower white dorsal postdiscal bands. Carpenter and Hobby (1945) give an extensive discussion of variation in the subspecies. Butler (1865a) described this taxon based on an unspecified number of specimens in the British Museum, which he compared with A. bredowii bredowii, and mentioned the orange bar at the end of the VHW discal cell. I have examined a syntype female in the BMNH. The presence of the orange band at the distal end of the VHW discal cell is unique within the A. serpa group, and strong indication that the continuous orange band basal to the white postdiscal band on the VHW in more derived members {A. nea, A. serpa etc.) is a result of the widening of the area between cell bars one and two. However, the absence of the cell band in occasional specimens of A. b. californica, as well as the similar male and female genitalia and wing pattern, lead me to treat this taxon as conspecific with A. b. bredowii and A. b. eulalia.

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110 Range: Western U.S.A. in the states of California and Oregon, with a very few records in western Nevada (Carpenter and Hobby, 1945), Arizona and north central Utah (Carpenter and Hobby, 1945), to extreme northwestern Mexico in the peninsula of Baja California. Domfeld (1980) provides a map of collecting localities within Oregon. There are several clearly mislabeled specimens from Honduras (see list of specimens, and Carpenter and Hobby (1945)) which are probably responsible for the inclusion of this country in the range of A. bredowii by Scott (1986). Immature stages: The most frequently reported hostplant of this subspecies is Quercus chrysolepis (Canyon Live Oak), in California (Comstock, 1927; Howe, 1975; Orsak, 1977; Domfeld, 1980; Christensen, 1981), while Comstock (1927) also implied that it fed on other, unidentified species of Quercus. In the peninsula of Baja California (Mexico), the subspecies feeds on Quercus agrifolia in the lowlands and Quercus chrysolepis at higher elevations (Brown et al, 1992). Scott (1986) cited a number of species of Fagaceae as hostplants, listed in Table 2 under A. b. eulalia, without stating whether they applied to that taxon or to A. b. californica. The earliest published notes on the immature stages of this taxon are also the first for any Adelpha species, comprising a description of the pupa by Edwards (1873). Subsequently both Dyar (1891) and Comstock and Dammers (1932) described all the immature stages in great detail, though Dyar mentions only four instars, presumably missing the first, and their observations are summarised here. The egg is green, morphologically typical of the genus and laid singly at the tip of a leaf on the upperside. The first instar is olive green with paler flecks, the head capsule mottled light and dark brown, with bumps marking later scoli. The second instar is green (Dyar) or various shades of brown (Comstock and Dammers) with darker spots marking the subdorsal scoli on T2, T3, A2 and A8, a dark brown supraspiracular line and a pale brown head capsule. The third instar is pale brownish yellow and densely covered with yellowish conical tubercles, with developing scoli on T2, T3, A2 and A8, each terminating with 4-5 black

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Ill spines. The head is pale greenish and also densely convered with conical tubercles, with four vertical brown stripes on the front and sides of the head capsule, long black dorsal spines and black-tipped lateral spines. The fourth instar is light reddish-brown with a lateral greenish tinge and the tubercles are yellowish, while the scoli are brown with black tipped spines. Scoli are well developed on the segments as in previous instars, and also there appear small scoli on A4 and A7. The chalazae and spines of the head capsule are now yellowish white with black tips. The fifth instar is olive green and densely covered with conical yellowish tubercles which become more dense vantrally, and the legs and ventral surface are brown. Small supraspiracular scoli are present on all segments except Tl and A9, and that on T2 has lateral spines. Subdorsal scoli are brown and largest on T2, T3, A2 and A8, bearing many white, black-tipped thick conical spines, while remaining scoli are absent on Tl, tiny on Al, larger on A3, A5, A8 and A 10, and largest on A4 and A7. The spiracles on Tl are faintly black rimmed, others are concolorous with the body. The head capsule is flattened and not bilobed, unlike Basilarchia which has a deep dorsal groove at the median sutiore. The face is almost smooth with only a few granulations, but has a lateral double row of long spines, most prominent dorsally, which are yellowish white with the exception of a pair of black dorsal spines. The color is brown on the face, paler laterally and ventrally, with a tapering whitish band over the ocelli and another faded band on each side of the clypeus. The labrum is white and the ocelli black. Just before pupation the larva fades to a uniform straw-yellow. The pupa is pale brown with large metallic areas laterally on the thorax and posterior to the eyes, with long, pointed, triangular head horns. This description was repeated in abbreviated form by several subsquent authors (Comstock, 1927; Howe, 1975; Domfeld, 1980; Scott, 1986, possibly also referring to A. b. eulalia), most of whom remarked on the camouflage of the larva against the hostplant leaves. Comstock (1927) figured the pupa, which is morphologically similar to A. b. eulalia, but with longer head horns and less pronounced dorsal projections on T2 and A2. The pupa is dark fawn in color or pale straw yellow, sometimes with gold

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112 streaks and darker brown, and is attached to tree trunks by a large silken web, "not less than half an inch in diameter" (Edwards, 1973; Comstock and Dammers, 1932). The first instar builds a leaf perch, presumably fi-om the mid-rib, similar to other Adelpha. Second to fifth instars rest in the FrontArched-Rear-Up position (Dyar, 1891; Comstock and Dammers, 1932: pi. 26c). The duration of the various stages imder laboratory condifions was given by Comstock and Dammers (1932) as follows: egg, 11 days; 1*' instar, 9 days; 2"^ instar, 8 days; 3'"'' instar, 4 days; 4"' instar, 9 days; 5"' instar, 14 days; pupa, 10 days; total development time, 65 days. Habitat and adult ecology: Comstock (1927) reports that this subspecies is fi-equently seen gliding high among the branches of the oak tree hostplant. Males often gather in small groups to feed at moist sand along streams, typically in the middle of the morning, and rare individuals have been recorded feeding on flowers. They have also been observed feeding on grape squeezings (Emmel and Emmel, 1973). In California, there are two broods, with adults flying fi-om April to June and again from August to September (Emmel and Emmel, 1973; Orsak, 1977), while Comstock (1927) reports that the flight season varies somewhat with altitude. According to Christensen (1981) this subspecies frequents oak, pine and Douglas-fir woodlands of western Oregon from midJune to mid August. Domfeld (1980) reports it flying from May to October, but that it is most numerous in mid to late summer. Several of the comments of Scott (1986) may apply to either this subspecies or A. b. eulalia, and are mentioned under the latter taxon. Carpenter and Hobby (1945) list additional older works in which there are observations of the adult natural history. Poulton (1908, 1909) suggested that this subspecies is involved in mimicry with Basilarchia lorquini, with which it flies throughout its range. This is indeed possible; of the three subspecies of A. bredowii, the nominate appears to be the most primitive, through comparison with the most closely related species A. diodes, in that the hindwing is still relatively elongate and angular, the postdiscal series on the VFW are more

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113 extensive and not so broadened as in the remaining two subspecies and the inner postdiscal series on the VHW is still present (also true of A. b. eulalia). Although A. b. californica has a second VHW orange discal cell band, a primitive character, I interpret this as a reversal since it is otherwise absent in all members of the A. serpa group, and also occurs occasionally m A. b. eulalia. It is therefore clear that several features that enhance the resemblance of A. b. californica to A. lorquini, such as the more rounded wings and displaced white DFW postdiscal spots, are derived characters, as would be predicted if A. b. californica and B. lorquini are involved in mimicry. Specimens examined (494m, 222f): U.S.A. (490m, 221f); Arizona: Prescott If USNM-error?; no specific locality (Escalante) Jun. If AME, If NlNHN-error?; California: Altascadero Apr. 3m AMNH; Angeles Forest 2500' Jun. Sep. 7m, If AMNH, Im, If MUSM; Auburn May Im AME; Bald Mt. Rd. Humboldt Co. Jul. 2m FSCA; Bangor Sep. If USNM; Barton Flats San Bemadino Co. Jun. Jul. 2m, If FSCA, Im AME; Bear Valley May 2m USNM; Berkeley Aug. Im USNM, 3m, If AMNH; Big Basin State Park Jul. Im, If FSCA; Bonneville Jun. Im AMNH; Boulder Creek Apr. If AMNH; Bronville Jun. If AMNH; Camp Baldy Im MCZ; Camp Nelson Jul. Im AMNH; Capitan Res. S. Diego Co. Aug. If FSCA; Carrista Creek S. Diego Co. If AME; Carrville Im AMNH; Carsona Pass Alpine Co. Jun. Im AME; Cedar Grove Im AMNH; Cedar Pine Aug. Im USNM, 2m AMNH; Chews Ridge Monterey Jun. 2m, 4f AMNH; Coal Creek Canyon Jun. If BMNH; Collins Ranch Aug. If AMNH; Cottonwood Camp Im USNM; Crestline If USNM; Cuyamaca, Rancho State Park Jul. 2m USNM, 2m, If AMNH, 3m FSCA; Davis Creek If USNM; Descanso May Im USNM, 2m AMNH; Devil's Gulch Jun. 7m, 5f AMNH; nr. Downeyville Oct. If AME; Dulzura Oct. If AMNH; Dunsmuir May If AMNH; Eaton Canyon Jul. If BMNH; Echo Mts. Jul. Im BMNH; EI Cajon S. Diego Co. Jul. If FSCA; Elizabeth Uke Canyon Aug. Im FSCA; El Monte S. Diego Co. Aug. 2f FSCA; El Portal May Im, If AMNH; Elstmore Im AMNH; Fairfax Jun. Jul. 3m AME; Figueroa Mt. If AMNH; Frazier Park Ventura Co. 2m AME, Im AMNH; French Gulch Jun. Jul. 4m USNM; Geysers Jun. Im AMNH; Glendale Mar. May Oct. 2f BMNH, 2m FSCA, Im AMNH; Glendora May 3m USNM; Gold Creek May 2m AME; Horn Mts. Kem Co. 1 100' Apr. Jun. Im, If AME; Humboldt 12 mi. E. Orick Jul. Im AME; Idlewild Aug. Im FSCA, Im AMNH; Indian Flat Mariposa Co. May Im AME; Jelachapi Jul. Im AME; Jerseydale 3500' Jul. Aug. 17m, 6f AMNH, 2m, If AME, Im FSCA; Julian Im AMNH; Kings Mt. Jul. If AME; Laguna Mts. Jul. Im AME; Uke Hobergs Im USNM; E. Lakeside S. Diego Co. Mar. 3f FSCA; U Tuna Canyon Mar. -May Aug. -Oct. 5m, 5f AME; Laytonville Aug. If AMNH; Lime Kiln Apr. 3f AME; Livermore, 8 mi. S., Jun. Im USNM; Livermore, 20 mi. S., Jun. Im AME; Lopez Canyon Jun. If AMNH; Los Angeles Apr.-Jun. Sep. 5m BMNH, 2m, If USNM, 8m, 5f AMNH, 2m AME; Los Banos Jun. 2m, 2f AMNH; Macho Canyon May Im, If AMNH; Marin Co. Jun. 2m, If USNM; Mariposa Co. May Aug. Im, 3f AMNH; Marton Peak San Bemadino 3300' Jul. 4m AME; Marysville Jun. Im USNM; nr. McCloud Mt. Shasta 3500' Jun. 3f AME; Mendocino Mts. May Jun. If USNM, 2m AMNH; middle Cahfomia 4m, 3f USNM; middle Fork Butte Jun. 2m, If USNM; Middleton Jun. Im USNM; Mill Creek Canyon San Bemadino Mts. Jun. Im AME, Im FSCA; Mill Valley May If AMNH; Mineral King Jun. Im USNM; Mint Canyon May Im AME; Miramar Feb. Im USNM; Miranda Aug. If AMNH; Mirror Lake Jan. Jul. Im USNM, If AME; Monterey Jun. Aug. 4m FSCA, Im USNM; Morbon Park Jul. Im AME; Mt. Diablo May Im AME, Im AMNH; Mt. Laguna San Diego Co. Im FSCA, Im, 3f AMNH; Mt. Lowe Aug. 9m, 4f BMNH; Mt. Tamalpais Jun. If AMNH; Mt. Wilson Jun. Aug. If BMNH, 2m USNM; Nevada Co. Jul. Im USNM; Newhall Hills Jun. If BMNH; Orange Co. Jun. If AME; Oso Canyon San Rafael Mts. Mar. Im AME; Palmar 3m, If AMNH; Palo Alto Jun. Im, If USNM, Im AMNH; Palomar 5-5300' Jun. Jul. 9m AME, If AME, If FSCA; Jul. 2m, If AMNH; Panther Creek Salt Springs Amador Co. Jun. Im FSCA; Patterson, 22 mi. W., Stanislaus Co. May 2m AME; Pasadena Aug. Im USNM; Pepperwood Humboldt Co. Jul. 2f FSCA; Petaluma Im AMNH; Pfeiffer State Park Monterey If FSCA; Placerville Feb. Nov. Im BMNH, If AMNH; Plumas Co. Jul. If USNM, If AMNH; Pinnacles Nat. Mon. Apr. Im AMNH; Pom Mts. Sep. Im MCZ; Potrero, 2 mi. N., S. Diego Co. Jul. 2m FSCA; nr. Qjal, Lion Creek May 2m FSCA; Riverside Co. May Im AMNH; Rubio Canyon May Im USNM; San Bemadino Mts. May Jun. Sep. 3m, If BMNH, 4m USNM, 2f AMNH. Im MCZ; SanBerdoCo. Im AMNH; San Diego MayAug. 2m MCZ, ImBMNH, 3m, 4f AMNH, Im AME; San Francisco 3m AME, If USNM; San Gabriel Mts. Jun. 22m, 7f AMNH, 2m AME; San Isabel Creek Volcan Mts. San Diego Co. 3000' Jun. 2m AME; San Josi Im AME; San Mateo Jun. Im AME, Im AMNH; San Rafael May Im BMNH; Santa Barbara Co. 3800' Jun.-Aug. 6m AME; Santa Clara Co. Aug. If FSCA; Santa Cniz May Jun. Aug. If BMNH, Im, 2f USNM; Santa Elena 2fAMNH; Santa Monica Mts. May Im FSCA, Jul. 2m AMNH; Saratoga Im AMNH; S. California Im USNM; Seven Oaks San Bemadino Jul. Im, If AME; ShasU 3500' Jun. Aug. 69m, 6f BMNH, 2f USNM, 2m AMNH, Im AME; Shirley Meadows summit Green Horn Mts. 6700' Jun. If AME; Sprinaville Jun. Im AMNH; Sierra City 4000' Jun. Jul. Im, If AME; Silocredo Canyon Grange Co. Jun. Im AME; Snow Creek Mar. If AME; Stunt Canyon LA Co. Apr. Im, If FSCA; Tahachapi Mt. Kem Co. 5600' Jun. Jul. If AME, 4m BMNH, 2m USNM; Tanbark Co. LA Co. Jul. If FSCA; Thompson Canyon Yolo Co. Vaca Mts. May 3m, 7f AME; Trinity Butte 3m AMNH; Trinity Co. Jun. Im FSCA; Trobuco Canyon Orange Co. Jun. 2m FSCA; Tuolumne Co. Jun. Aug. 3m AMNH; Two Rock Im AMNH; Watsonville Im BMB; W. of St. Helena Jul. If FSCA; Yucaima Jul. 4f AME; Ukiah May Im, If USNM; upper Santa Ana River Im AMNH; upper Yuba Res. May Im USNM; Wahwonga May If BMNH; Walsingham Im BMNH; Waterman Canyon San Bemadino Mts. May Im AME; Willits Aug. Im AMNH; Winchester Jul. 2m BMNH; Woodside Jul. Im USNM; Yosemite 4000' May-Aug. 6m, 4f BMNH, 3m, 6f USNM, 8m, If AMNH, 5m, If AME; Yucalpa Jul. Im AME; no specific locality Jun. Jul. 8m, 9f BMNH, Im BMB, Im, 7f MNHN, 4m, 4f USNM, 8m, 6f AMNH, Im MCZ; Oregon: Brooking Vulcan Mts. Jul. Im AME; Butte Falls Jul. 3m

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114 USNM; Central Point Apr. If BMNH; Corvallis Jul. Inn USNM; Eugene Sep. 4m AME; Gold Hill Jackson Co. Jul. Sep. Oct. 5m BMNH, Im FSCA; Goose Lake Jul. Im USNM; Jacksonville Aug. 3m, 2f AME; McMinnville Sep. Oct. Im BMNH, 6m USNM; Mt. Ashland Im AME; Oregon 2m AMNH; Roseburg 2m MCZ; Salem Jul. If USNM; San Mateo Jun. Im AME; Sissons May Aug. 2m, 2f BMNH; Wolf Creek Aug. Im AME; Utah: Utah Im, If USNM; Not located. Abigua Creek Jul. 2m USNM; Laurel Canyon Jun. 2m BMNH; N. America 2f BMNH; Quincy 3400' Jun.-Sep. 7m, 3f BMNH; no specific locality 7m, 6f MCZ, 3m, 4f BMNH, Im, If MNHN, 7m, 4f USNM. COUNTRY UNKNOWN (3m, 10: no specific locality 2m AME, Im, If FSCA. "HONDURAS" (Im): no specific locality Im MCZ-error. Additional locality data: U.S.A.: see Carpenter and Hobby (1945). MEXICO: Baja California None: Siena de Juirez; Sierra San Pedro Martir, numerous localities and map (Brown et al., 1992). Adelpha diodes Godman and Salvin, 1878 Figs. 11; 96; 179 Identification, taxonomy and variation: This is a very distinctive species, characterised by the short "tail" on the hindwing at vein Cu2, the evenly edged orange DFW subapical marking and the even, almost continuous silvery white iimer and outer submarginal series on the VFW. The ventral ground color, length of the hindwing "tail" and shape of the DFW orange subapical marking vary, and two subspecies are recognised. This species is the only member of the Adelpha serpa group not to have a third cell bar in the VFW discal cell, but it has all of the defining synapomorphies of the group. It is closely related to A. bredowii, as indicated by characters of the male and female genitalia. Range and status: Mexico to western Panama in montane areas. From 1400 to 3000m. Very local but not imcommon. Specimens examined: 37 (26m, llf) Adelpha diodes diodes Godman and Salvin, 1878 Figs. lla,b; 96a Adelpha diodes Godman and Salvin (1878a: 270) TL: Volcan Chiriqui [Panama]. Types: BMNH(T): STm: "B.M. TYPE No. Rh. 9851 Adelpha diocles m G. and S.//4 93//B.C.A. Lep. Rhop. Adelpha diocles G. and S. Godman-Salvin Coll. 19l6.-4.//Type. sp. figured//V. de Chiriqui Arce/ZChiriqui Panama ArceZ/m.Type H.T." [examined]

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115 Identification, taxonomy and variation: The nominate subspecies is distinguished from A. d. creton by having shorter hindwing "tails", a narrower DFW orange subapical marking of more even width, a white DFW postdiscal band which does not extend into cell M2, ventral ground color in the distal half of the wings which is reddish brown instead of dark brown, and a white VHW postdiscal band of more even width which does not taper in from cell Rs to the costa. There is little variation in specimens examined. Godman and Salvin (1878a) described this species from an unspecified number of males in their collection collected by Arce on Volcan Chiriquf, and a syntype male is in the BMNH. It was subsequently figured by Godman and Salvin (1884), but the species appears to have been unknown to Fruhstorfer (1915). Range: To date known only from the Chiriqui massif in eastern Costa Rica and western Panama, although possibly also extending into Nicaragua, given the range of A. tracta, which occurs in similar montane habitats. Habitat and adult ecology: The only published observations of this subspecies in nature are those of DeVries (1987) in Costa Rica, and they are summarised here. He reports that it occurs from 2000-3000m in cloud forest, while museum specimen data indicate that it also occurs as low as 1400m. Adults appear to be very local and seasonal, though they may be abundant during the main dry season (February to March), and generally fly at the canopy level. Two male specimens in the ANMH were apparently attracted to lights at night. Specimens examined (Mm, \T): COSTA RICA (4m): Atajuela: Cascajal Jan. Im BMNH; Volcin Poas Im USNM; Puntarenas: Las Alturas 4700' Mar. at light 2m AMNH. PANAMA (10m, 10: Chiriqui. Chiriqui Im BMNH; Volcdn Baru 1700-1800m Dec.-Feb. Im FSCA, 8m, If USNM. Adelpha diodes creton Godman, 1901 Figs. llc,d; 96b; 179a-c

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116 Adelpha creton Godman (1901: 692) TL: Jalapa, Mexico. Types; USMVl: HTf: "A. creton Godm. f type//Type No. 850 [?]//sp. figured/ZCollection W. SchausZ/Jalapa, Mex." [examined] Helerochroa creton G. and S., Hoffmann (1940); Adelpha diodes Godm., Steinhauser (1974), D'Abrera (1987); Adelpha diodes creton Godm., de la Maza et al. (1989), de la Maza and de la Maza (1993) Identification, taxonomy and variation: This subspecies is distinguished from the nominate as detailed in the account of that taxon. There is little variation in the specimens examined except in the extent to which the white DFW postdiscal band extends into the posterior half of cell M2, where it may solidly fill the area or consist of diffuse scaling. This subspecies was described by Godman (1901) in a supplement to the Biologia Centrali Americana from an unspecified number of females collected by Schaus in Mexico. A syntype female is in the USNM. It was initially described as a species distinct from A. d. diodes on the basis of several of the characters listed above as distinguishing the two taxa, and Miller and Miller (1970) retained the two as distinct species stating that "the genitalia suggest that the two species should be kept separate". However, they provided no evidence in support of this assertion, and since I have been unable to find any consistent differences in the genitalia and the two taxa do not differ substantially in wing pattern, morphology or habitat, in addition to being allopatric, I follow de la Maza et al. (1989) and de la Maza and de la Maza (1993) in regarding creton as a subspecies of A. diodes. Range: Adelpha d. creton has been recorded in montane areas from Mexico to Guatemala and El Salvador. It is almost certainly more widespread, extending into Honduras, and possibly Nicaragua. Habitat and adult ecology: Miller and Miller (1970) provide an interesting account of the behavior and habitat of this species in Mexico. They captured 15 specimens and observed many others over several months in mixed pine-oak habitat in the vicinity of La Encamacion, between 2400m and 2450m. In agreement with the observations of the nominate subspecies by DeVries (1987) in Costa Rica, adults typically remained high

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117 among the crowns of trees, especially oak, only occasionally descending to puddle at damp soil, and then remaining very wary. The close association of the adults with oak trees led them to speculate that the hostplant might be in this family, and indeed this would be expected from the close relationship of A. diodes and A. bredowii. In Mexico, de la Maza and de la Maza (1993) report the species occurring in Chiapas from 18002700m in humid temperate forests in the Macizo Cenfral and Sierra Madre areas, while de la Maza (1987) reports records from August to March. Specimens examined (12ni, 100^ MEXICO (12m, 9f): Chiapas: Mt. Huitepec Apr. If AME; San Crist6bal de las Casas Feb. Mar. 6m FSCA, If AME; Hidalgo. La Encamacion 2400-2450m oak-pine forest Jan.-Mar. 2m BMNH, 3m, 5f AME; Zimapan Feb. Mar. Im, If AME; Veracruz: Jalapa If USNM. GUATEMALA (If): Quezaltenango: Volcan Santa Maria Oct. If USNM. EL SALVADOR (If): Melapdn: Metapan 2300m Dec. If AME. Additional locality data: MEXICO: Chiapas: Macizo Central; Sierra Madre (de la Maza and de la Maza, 1993); Huitepec; Guerrero: Filo de Caballo; Teotepec; Oaxaca: La Esperanza (de la Maza, 1987). Adelpha herbita Weymer, 1907 Fig. 12a,b Adelpha herbita Weymer (1907: 21, pi. H, fig. 7) TL: St. Catharina, Brazil. Types: Collection unknown: HTf not located. Identiflcation, taxonomy and variation: Adelpha herbita is a very distinctive, primitive and enigmatic species, to date only known to my knowledge from the holotype female. It is distinguished from A. zea, which it closely resembles on the dorsal surface, by having a complete and distinct white postdiscal band extending to the costa on the VFW (this band is fiised entirely with the subapical marking in A. zea), complete inner and outer submarginal series on the VFW (these are reduced or absent in cell M2 in A. zea), the subapical marking remaining distinct from the white postdiscal band in cells Cul and M3 of the WW, and no reddish bands distal or basal of the white postdiscal band on the VHW. The combination of the latter two characters also distinguishes the species from all other A. serpa group members.

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118 Weymer (1907) described this species carefully and at great length from a single female in his own collection, and the original illustration of the dorsal and ventral surface of half wings shows great attention to detail, notable in the shape of the second VFW discal cell bar which exactly matches other A. serpa group members. However, to date the whereabouts of the holotype female specimen is unknown, and neither myself nor Gerardo Lamas (pers. conrni.) have seen further specimens in any collections. Fruhstorfer (1915) also knew only of the holotype, and I am informed by Olaf Mielke (pers. comm.) that there are no specimens in the collections of the Universidade Federal do Parana, Curitiba, Brazil, which has extensive Adelpha material from Santa Catharina, the type locality. It has been suggested to me that the holotype is actually an aberration, but I do not believe this to be the case. In fact, A. bredowii also shares the loss of the outer postdiscal series on the VHW except for cell Cu2, exactly as depicted in the original illustration oi A. herbita, while A. diodes has much reduced reddish coloration basal of the VHW white postdiscal band, similar to A. herbita. I caimot believe that the number and combination of distinctive wing pattern characters in evidence in the original illustration could have arisen from any other known Adelpha simply as an aberration. Rather, these characters suggest that this is a very primitive species closely related to A. bredowii and A. diodes. If this is the case, then its presence in southeastern Brazil, far removed from these two Central to North American species, is of great evolutionary interest. Range: Currently the range of this species is not known with any certainty, but it probably occurs in the states of Santa Catharina and Parana in southeastern Brazil. Habitat and adult ecology: There is no published information on the natural history of this primitive and extremely rare species. From what is known of the ecology of related species, I would expect it to be confined to montane areas, some of which extend above 1800m, in the state of Santa Catharina and possibly Parana. Given that A. herbita appears to have been unreported since the capture of the holotype, any information on additional

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119 specimens or field observations would be of great value, since its conservation status must be considered vulnerable if the species is even still extant. Specimens examined: NO SPECIMENS KNOWN. Adelpha zea (Hewitson, 1 850) Figs. 13a,b; 97a,b; 180a-c Heterochroa zea Hewitson (1850: 435, pi. IX, figs. 1,2) TL: Rio de Janeiro [Brazil]. Types: BMNH(T): STf: "Heterochroa zea Hewitson Syntype det. R.I. VaneWright 1983//B.M. TYPE No. Rh. 9850 Heterochroa zea m Hew./ZBrazil Hewitson Coll. 79-69 Heterochroa zea. 2.//zea//Type//Syntype"; BMNH(M): ST?f: "Braz./ZBrazil Hewitson Coll. 79-69 Heterochroa zea. 1 ." [both examined] =Adelpha zea serpentina Fruhstorfer (1915: 532) TL: Santa Catharina [Brazil]. Types: BMNH(T): STm: "Fruhstorfer Coll. B.M. 1 937-285//zea serpentina Fruhst.//Brasilien St. Cath. Fruhstorfer//TYPE//Type//Syntype"; STf: "Fruhstorfer Coll. B.M. 1937-285//Brasilien St. Cath. Fruhstorfer//TYPE//Type"; BMNH(M): STf: "serpentina Fruhst.//Brasilien St. Catharina FruhstorferZ/Fruhstorfer Coll. B.M. 1937-285" [all examined] =Adelpha zea tarpeia Fruhstorfer (1915: 532) TL: Paraguay, Rio Grande do Sul [Brazil]. Types: BMNH(T): STf "Fruhstorfer Coll. B.M. 1937-285//zea tarpeia Fruhst.//Rio Grande Brasil ex. Coll. H. Fruhstorfer//TYPE//Type//Syntype"; BMNH(M): 2STf: "Rio Grande Brasil FruhstorferZ/Fruhstorfer Coll. B.M. 1937-285"; BMNH(R): STm: "TYPE/ZSapucay, Paraguay, 19.Vin.06 (W. FosteryZzea tarpeia Fruhst." [all examined] Adelpha zea Hew., Kirby (1 871 ) Identification, taxonomy and variation: Adelpha zea is most similar in wing pattern to A. paroeca, but is distinguished by the orange postdiscal marking touching the white postdiscal spot in cell M3, with no dividing thin dark brown line, and on the VHW by the dark red band basal of the white postdiscal band extending only into the anterior half of cell Cu2, instead of to vein 3A, and the dark line along vein 3A extending to the wing base instead of to the costal margin. Adelpha hyas hyas is similar on the dorsal surface but the ventral surface is markedly distinct. Characters that differentiate this species fi-om A. herbita are discussed under the latter species. There is very minor individual variation in the width of the white dorsal postdiscal bands. The name zea was first introduced as a nomen nudum by Westwood (1850) in his list of Adelpha, and illustrated and described shortly after in characteristic detail by Hewitson (1850) based on an unspecified number of specimens in his own collection. I have examined a syntype female in the BMNH(T) and a fiirther possible syntype in the BMNH(M), which differs slightly fi-om the former only in having a few orange scales at

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120 the DFW discocellulars. Fruhstorfer (1915) described two further subspecies, serpentina, from an unspecified number of specimens from Santa Catharina, and tarpeia (later illusfrated and misspelt tarpeja (Fruhstorfer, 1920: pi. 1 10A,b)), based on two males from Paraguay in the Tring Museum and three females from Rio Grande do Sul in his own collection. The putative differences in the width of the white dorsal postdiscal band, orange DFW subapical marking and brightness of the red bands on the VHW are barely apparent in any of the specimens of the syntypic series examined for either taxa, or indeed in any specimens oi A. zea, and Hall (1938) correctly synonymised both names with A. zea. Although Fruhstorfer (1915) and several subsequent authors freated A. paroeca as a subspecies of A. zea, the two are clearly distinct species as discussed xmder the former species. Range: Southeastern Brazil to Paraguay, northeastern Argentina and Uruguay. Habitat and adult ecology: Ebert (1969) found this species to be not uncommon in Minas Gerais in low canopy forest in hilly areas, up to 1400m, while Hoffinann (1936) observed the species in Santa Catharina feeding fairly commonly at flowers of various Compositae in March and April. Biezanko (1949) reported it (as Adelpha tarpeia) to be uncommon in Rio Grande do Sul from January to April. Otherwise nothing has been published on the ecology of this species and judging from museum specimens it is generally uncommon in the field. Specimens examined: 63 (26m, 37f) "COLOMBIA" (10: no specific locality 1 f BMNH-error. BRAZIL (22m, 320: Espirito Santo: If BMNH; Parana: Castro 4m, 2f BMNH; Femandes Pinheiro 2600' Apr. If BMNH, If AME; Igua9u Oct. Im BMNH; N. Parana 4m, If AMNH; Ponta Grossa 3500" Dec. 2m, 1 f BMB; Uniao da Victoria 2400' Apr. If AME; Rio de Janeiro: Teresopolis Im BMNH; Rio Grande do Sul: Pelotas Jan. Mar. Apr. 2m, 5f MCZ; Rio Grande do Sul Im, 3f BMNH, 2f ZMHU; Santa Catharina: Serrinha do Pirai, W. Joinville 950' Mar. If FSCA; Trombudo Alto May Im AME; no specific locality 2m, 3f BMNH, Im ZMHU, Im, If BMB, 6f AMNH; Sao Paulo: Casa Branca Im, If ZMHU; Not located: no specific locality 2f BMNH, Im AMNH. PARAGUAY (4m, 30: Caaguazu: Yhii Sep.-Dec, Im BMNH; Paraguari: Sapucay Sep. 2m, 2f BMNH; Not located: no specific locality Im, If MCZ. COUNTRY UNKNOWN (10: no specific locality 3m, If BMNH, IfMNHN. Additional locality data: BRAZIL: Rio Grande do Sul: Monte Bonito Jan.-Apr. (Biezanko, 1949); Minas Gerais: Po^os de Caldas 1300-1400m May (Ebert, 1969). ARGENTINA: Misiones (Hayward, 1951). URUGUAY: no specific locality (Biezanko et al., 1978). Adelpha paroeca (H. W. Bates, 1864) stat. rest. Figs. 14a-d; 98; 181a,b

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121 Heterochroa paroeca Bates (1864: 127) TL: Guatemala, central valleys. Types: BMNH(T); STf: : "Heterochroa paroeca Bates f Syntype det. R.I. Vane-Wright 1983//B.M. TYPE No. Rh. 9849 Heterochroa paroeca f Bates//B.C.A. Lep. Rhop. Adelpha paraeca Bates Godman-Salvin Coll. 1916.-4.//Heterochroa paroeca Bates//f//Centr. Valleys Guatemala F.D.G. and O.S.//Guatemala Central Valleys/ZType H.T."; BIVINH(M): ST?f: "Centr. Valleys Guatemala F.D.G. and O.S.//f//Central Val. Type sp. figured/ZB.C.A. Lep. Rhop. Adelpha paraeca Bates Godman-Salvin Coll. 1916.-4." [both examined] =Heterochroa emathia Felder (1869: 473) stat. rest. TL: Potrero [Mexico]. Types: BMNH(R): STm: "481//Mexico Terra cal. Milimet//H. emathia Feld.//Omathia [sic] n. //Type" [examined] Adelpha serpa var. paroeca Bates, Kirby (1871); Adelpha serpa var. emathia Fldr., Kirby (1871); Adelpha zea paroeca Bates, Fruhstorfer (1915), de la Maza and de la Maza (1993); Adelpha zea emathia Fldr., Fruhstorfer (1915), de la Maza and de la Maza (1993); Limenitis (Adelpha) paroeca Bates, Ross (1976) (actually A. serpa celerio); Adelpha zea ssp. de la Maza and de la Maza (1993). Identification, taxonomy and variation: Adelpha paroeca is distinguished from the similar A. nea sentia by having an orange postdiscal spot on the DFW in cell M3 distinct from the white postdiscal spot, and the white postdiscal spot in cell Cul on the VFW being slightly narrower than the spot in Cu2, instead of half as wide again. Adelpha zea has the pale orange postdiscal marking on the VFW in cell M3 fiised with the white postdiscal spot, instead of separate, and the reddish band basal of the white VHW postdiscal band has a different configuration, not reaching vein 3A. Adelpha paraena massilia lacks an orange postdiscal marking in DFW cell M3. All other similar A. serpa group members have the white submarginal series on the VFW consisting of paired spots instead of single dashes in each cell space. There is some variation in the width of the white dorsal postdiscal bands and the width of the orange DFW subapical marking, both tending to be narrower in specimens from Nicaragua to Panama. In addition, specimens from Costa Rica and Panama often have more orange-brown coloring in the band basal of the white VHW postdiscal band, orange-brown ventral ground color in the distal half of the wings and the silvery white markings of the venfral submarginal series reduced in size (Fig. 14c,d). However, all of these characters seem to vary within populations and are best ascribed to clinal variation throughout Central America. Adelpha paroeca was described in detail by Bates (1864) in comparison with A. serpa, from a male specimen or specimens from Guatemala. The putative syntype and second possible syntype in the BMNH are, however, females, but otherwise agree with

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122 the original description so it appears that Bates simply misidentified the sex. Felder (1869) described the taxon emathia from an unspecified number of male specimens as a fiiU species, and the syntype is in the BMNH. However, since he failed to mention the recently described A. paroeca it must be assumed that he was unaware of the description. The two taxa were both placed as "varieties" of A serpa by Kirby (1871), while Fruhstorfer (1915) treated both names as subspecies of A. zea. I have not found any consistently different populations within A. paroeca that I believe merit subspecific recognition and so synonymise emathia with A. paroeca (stat rest), as first proposed by Godman and Salvin (1884). Adelpha paroeca has been and still is variously treated as a subspecies of .4. zea (Fruhstorfer, 1915; DeVries, 1987; de la Maza and de la Maza, 1993), despite the great range disjunction between the two taxa, although it has also been accorded specific status by a number of authors (Godman and Salvin, 1884; D'Abrera, 1987; Lamas and Small, 1992). The differences in wing pattern discussed above, and in the female genitalia, show that the species are clearly very distinct, and I restore A. paroeca to fiill species status (stat. rest.), hi fact, the extension of the first cell bar on the VHW to vein 3A is a synapomorphy which groups this species with the more derived species following this account, indicating that A. paroeca is not the sister species of A. zea but is more closely related to A. nea and A. paraena. Range: Mexico to western Panama. Habitat and adult ecology: This species is found in lower montane rain forest habitats where it appears to be uncommon. DeVries (1987) reports it in Costa Rica from 900m to 1500m, while de la Maza and de la Maza (1993) record it in Chiapas, Mexico, from 800m to 2300m, in hot to temperate semi-humid forests. Specimens examined: 77 (40m, 37f) MEXICO (22m, 14f): Chiapas: Comitan Sq). Im, If AME; Las Delicias Oct. 2m AME; Ocozingo Apr. Jun. 2m AME; Pichucaico May If AME; no specific locality 2f AMNH; Guerrero: Acahuizotla Feb. Aug. 2m AME; Nayarit: Compostela Oct. If AMNH; Nuevo Leon: Cola de Caballo Oct. Im AME; Oaxaca: Oaxaca, 192 km. from, Im AMNH; Puebla: Villa Juarez If AME; Veracruz: Cordoba Aug. 3m, If BMNH, If USNM; Jalapa 4600' Nov. Im BMNH, 2m, If USNM, 3m, If BMB; Presidio Im, 2f AME; Vera Cruz If ZMHU; Not located: Terra Caliente 2m BMNH; no specific locality Im AME, If AMNH. EL SALVADOR (2f): San Salvador: Santa Tecla 900m May If AME; Santa Ana: Citala 800m Oct. If AME. GUATEMALA (6m, 130: Alta Verapaz: Baleu

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123 1350m Sep. Nov. Im, 2f AMNH, Im AME; Baja Verapaz: San Jeronimo Im, 3f BMNH; Guatemala. Guatemala City 4-5000' Sep. Im, 4f USNM, Im, If BMB; Retalhuleu: San Sebastian Im USNM; Not located: Central Valleys 2f BMNH; no specific locality If BMNH. NICARAGUA (Im): Jinotega: Jinotega 3400' Nov. Im BMNH. COSTA RICA (Im, 20: Carlago: Tumalba 2000' Jul. Im USNM; San Jose: Rio Sucio If BMNH; Not located: no specific locality If ZMHU. PANAMA (6m, 40: Chiriqui: Boquete Im BMNH; Chiriqui 2m ZMHU; Potrerillos 3600' Jan.-Mar. Im, 4f USNM, Im STRI; Volcan Baru 1300m Feb. Mar. 2m USNM. COUNTRY UNKNOWN (4m, 20: no specific locality 2m BMNH, If USNM, If MNHN, 2m MCZ. Additional locality data: MEXICO: Chiapas: Cuenca de Grijalva; Cuenca de Tulija; Macizo Central; Selva Lacandona; Sierra Madre; Soconusco (de la Maza and de la Maza, 1993); Jalisco: U Calera Nov. (Vargas et al., 1996). Adelpha nea (Hewitson, 1847) Figs. 15; 99; 182 Identification, taxonomy and variation: Adelpha nea is distinguished from all other members of the A. serpa group by having the pale postdiscal marking in cell Cul of the VFW approximately half again as wride as the marking in cell Cu2, due to the fusion of the postdiscal band with the inner, and possibly outer, postdiscal series in this cell. Another useful distinguishing character is the lack of any whitish postdiscal streaks in cells Ml and R5 between the VFW discal cell and the pale subapical marking, also lacking in A. zea. The shape of the male genitalic valva is also unique within the group, being elongated and curved at the distal tip (Fig. 99). Adelpha nea may be easily distinguished from species outside the A. serpa group by the curved dark third discal cell bar on the VFW. Adelpha nea nea is also the only taxon in the A. serpa group in which the DFW postdiscal band is entirely orange in cells Cul and M3. The color of the postdiscal band varies geographically and two subspecies are recognised. Range and status: Mexico to Venezuela, to southern Peru, Amazonian Brazil and the Guianas, in lowland rain forest. Almost certainly more widespread, extending to northwestern Ecuador and Bolivia. Very rare. Specimens examined: 40 (16m, 24f) Adelpha nea nea (Hewitson, 1847) Figs. 15a,b; 99a; 182a-c

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124 Heterochroa nea Hewitson (1847: 257, pi. XX, fig. 1) TLPara [Brazil]. Types: BMNH(T): STf: "Illustrated in The Butterflies of Venezuela A. Neild, 1996//165//Heterochroa nea Hewitson f Lectotype det. R.I. Vane-Wright 1983 see Hall, 1938: 285//nea Hew.//B.M. TYPE No. Rh. 9844 Heterochroa nea f Hew.//Para//Type//Lectotype" [examined] =Adelpha ma campeda Fruhstorfer (1915: 532) XL: Colombia. Types; BMNH(T): STf: "Fruhstorfer Coll. B.M. 1937-285//nea campeda Fruhst7/Columbien ex. Coll. H. Fruhstorfer//TYPE//Type//Syntype"; BMNH(M): ST?f: "Colombie Collection Le MouIt/ZFrrihstorfer Coll. B.M. 1937-285" [both examined] Adelpha nea Hew., Kirby (1871) Identification, taxonomy and variation: Adelpha nea nea is distinguished from A. nea sentia by the DFW postdiscal band being entirely orange in at least cells Cul and M3. It is very similar to and frequently confrised in collections with A. melona leucocoma, but is most easily distinguished by the curving dark third cell bar in the VFW discal cell that divides the pale area between the second and fourth discal cell bars into three. There is a little local variation in the amount of white on the postdiscal band at the anal margin of the DFW, which tends to be more prevalent in Colombian specimens but also occurs in the syntype specimen oiA. nea. The single Costa Rican specimen has the orange DFW subapical marking barely joined to the remainder of the postdiscal band. Hewitson (1847) accurately figured and described A. nea based on material in the British Museum from Para, and made special note of the differences in the configuration of VFW discal cell bars which readily distinguish this species from A. melona. A syntype female is in the BMNH and was figured by Neild (1996). Fruhstorfer (1915) described an unspecified number of Colombian female specimens as a new subspecies, campeda, and a syntype female is in the BMNH. Hall (1938) synonymised campeda with nea, with which I concur, all Fruhstorfer's supposed differences between the taxa being minor individual variation, if apparent at all. Range: To date known from Costa Rica to Venezuela, to southern Peru, Amazonian Brazil and the Guianas. Although as yet unrecorded, it probably also occurs in the Choco region of western Colombia to northwestern Ecuador.

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125 Habitat and adult ecology: Adelpha nea nea is widespread but appears to be very rare throughout its range. Females outnumber males in collections by almost 2:1. There is almost nothing known of its biology, Neild (1996) never encountered the species in the field in Venezuela, while DeVries (1987) omitted it altogether from the Costa Rican fauna. In eastern Ecuador, Jason Hall (pers. comm.) captured a male specimen in a ridge top subcanopy trap baited with rotting fish in primary lowland rainforest in the middle of the wet season, but despite numerous other visits to the same locality we have never again encountered the species. The species is probably both seasonal and usually confined to the canopy. The close resemblance of the dorsal surface to the much commoner, but relatively unrelated species A. melona, also apparently a canopy species, suggests that there may be some mimetic relationship between the two taxa. In Central America in areas where A. melona has yet to be found (Mexico to Belize) the subspecies of ^. nea has a very different dorsal surface wing pattern. Specimens examined (13tn, 23i): COSTA RICA (If): Costa Rica Coll. Frank Johnson If AMNH. PANAMA (2f): Canal Zone: Cocoli Dec. If STRI; Panama: Colon 1500' Feb. If USNM. VENEZUELA (Im, IQ: Barinas: Reserva Forestal Ticoporo 230m Apr. Im MUSM; Delta Amacuro: Rio Acure If AME. COLOMBIA (3m, 3f): Cundinamarca: Bogota 2m BMNH; Vaupes: Mitu Aug. Im ESM; Not located: no specific locality 2f BMNH, If BMB. ECUADOR (Im): Napo: Chichicorrumi 450m Jul. Im KWJH. PERU (2m, 30: Junin: La Merced 2500' Im, If BMNH; Loreto: Iquitos If AMNH; Puno: Chaquimayo 25-3000' Aug.-Oct. Im BMB; Rio Tavara 450m Aug. If MUSM. BRAZIL (2m, 70: Amazonas: Massaury If ZMHU; Para: Obidos Jan. If AME, Im AMNH; Para Im, 3f BMNH; Not located: no specific locality 2f BMB. GUYANA (Im, 20: Potaro/Siparuni: Potaro River Jul. If AME; Not located: Essequibo R. 140 mi. inland Im BMNH; no specific locality If BMNH. FRENCH GUIANA (3m, 40: Cayenne: Cayenne Im BMNH, If ZMHU; Laurent du Maroni: Nouveau Chantier Im BMNH; St. Laurent du Maroni If BMNH, If BMB; Maroni Im BMNH; Not located: no specific locality IfMNHN. Additional locality data: ECUADOR (lm/0: Napo: Rocafuerte (coll. M. Moreno, Quito). BRAZIL: Amazonas: Ilha de Maraci (Mielke and Casagrande, [1992]; possibly misidentification of A. melona leucocoma, not recorded from this site). Adelpha nea sentia Godman and Salvin 1884 stat. nov. Figs. 15c,d; 99b Adelpha sentia Godman and Salvin (1884: 309, Tab. XXIX, figs. 9,10) TL: Corosal, British Honduras [Belize]. Types: BMNH(T): STm: "B.M. TYPE No. Rh. 9848 Adelpha sentia m G. and S.//Corosal British Honduras Roe.// B.C.A. Lep. Rhop. Adelpha sentia G. and S. Godman-Salvin Coll. 1916.-47/m//Type H.T." [examined] Adelpha serpa sentia G. and S., Fruhstorfer (1915); =Adelpha serpa celerio Bates, Hall (1938); Heterochroa serpa sentia O. and S., Hoffmann (1940) (actually A. paraena massilia); Limenitis (Adelpha) sentia G. and S., Ross (1976) (actually A. paraena massilia); Adelpha serpa sentia G. and S., Austin el al. (1996) (actually A. paraena massilia)

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126 Identification, taxonomy and variation: This subspecies is easily distinguished from the nominate by the DFW postdiscal band being almost entirely white. It is very similar to A. paraena massilia, but distinguished by having orange scaling at the distal edge of the white DFW postdiscal spots in cell M3 and M2, the spot in cell M3 being wider than that in cell Cul, and by lacking any whitish dashes in cells Ml and R5 between the end of the discal cell and the pale subapical marking on the VFW (resulting from the fusion of the subapical marking with the upper postdiscal band). Adelpha seriphia godmani and A. serpa celerio may be distinguished by the same characters, as well as by having paired instead of single white submarginal markings in each cell space on the ventral surface. Adelpha paroeca may be additionally distinguished by having an orange postdiscal spot in cell M3 on the DFW. The amount of orange scaling at the distal edge of the white postdiscal markings in cells Cul and M3 is sHghtly variable mA. nea sentia. Adelpha nea sentia was figured and described as a full species by Godman and Salvin (1884) from a single male specimen which is now in the BMNH. However, all subsequent authors have regarded this taxon as synonymous with A. paraena massilia (see discussion under that taxon). The broad postdiscal marking in cell Cul compared with that in cell Cu2 is a character unique among members of the A. serpa group to A. nea, and the fusion of the subapical marking with the upper postdiscal band on the VFW is a character found within the A. serpa group only in A. nea and A. zea. In addition, the distinctive shape of the male genitalic valva (Fig. 99b) confirms beyond doubt that this taxon is conspecific with nominate A. nea. Range: Mexico, in the state of Oaxaca, to Belize, almost certainly occuring in Guatemala and probably extending to Nicaragua. Austin et al. (1996) list "yl. serpa sentia" from Guatemala, but I presume this is the more common A. paraena massilia. Habitat and adult ecology: Nothing is known of the ecology of this subspecies. It is very rare in collections and its range is no doubt much wider than current specimen data

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127 indicate. All records are from lowland forest. The dorsal wing pattern differences between this subspecies and the nominate may well be the result of mimicry between this taxon and a large number of superficially similar sympatric Adelpha, including A. paraena massilia, A. serpa celerio, A. iphiclus and A. iphicleola. Specimens examined (3m, 10: MEXICO (Im, 10: Oaxaca: Chimalapa Sep. Im AME; Quintana Roo. X-can Jul. If AME. BELIZE (Im); Atlamida. Corozal Im BMNH. COUNTRY UNKNOWN (Im): no specific locality Im MNHN. Adelpha paraena (H. W. Bates, 1865) Figs. 16; 100; 183 Identification, taxonomy and variation: Adelpha paraena is similar to several other members of the A. serpa group. It is distinguished from A. serpa, A. radiata, A. seriphia and A. hyas by the VFW submarginal series being composed of single, instead of paired, whitish dashes in each cell space, most notable in cells Cul and M2. It is distinguished from A. nea by having whitish postdiscal sfreaks on the VFW between the discocellulars and the subapical marking in cells M2-R5 and at the costa Adelpha paroeca is also similar, but always has well developed orange and white postdiscal spots in cell M3 on the DFW, in addition to a much broader inner submarginal series on the VHW. There is variation on the DFW in the size and extent of the orange subapical marking and the white postdiscal spot in cell M3, in addition to the width of the white postdiscal bands, and four subspecies are recognised, one of which is described here. The undivided spots of the WW submarginal series suggest A. paraena is probably most closely related to A. nea and A. paroeca, and this is borne out in the cladistic analysis in Chapter 3, although it has often been considered a subspecies of A. serpa.

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128 Range and status: Lowlands from Mexico to Bolivia, Brazil and the Guianas. Very uncommon to rare throughout the range. Specimens examined: 174 (106m, 68f) Adelpha paraena paraena (H. W. Bates, 1865) Figs. 16a,b; 100 Heterochroa paraena Bates (1865: 331) TL: Part [Brazil]. Types: BMNH(T); STf: "Heterochroa paraena Bates f Syntype det. R.I. VaneWright 1983//Illustrated in The Butterflies of Venezuela A. Neild, 1996//172//B.M. TYPE No. Rh. 9847 Heterochroa paraena f Bates/ZAdelpha paraena Bates Godman-Salvin Coll. 1916.-4.//f Para paraena//f//Para L. Amazons H.W. Bates/ZType H.T./ZSyntype"; BMNH(M): STm: "ParS/Zparana [sic-red label]ZZAdelpha paraena Bates Godman-Salvin Coll. 1916.-4.ZZPara L. Amazons H.W. BatesZZm" [both examined] Adelpha serpa var. paraena Bates, ICirby (1871); Adelpha serpa paraena Btlr. [sic], Fruhstorfer (1915); Adelpha paraena Bates, Hall (1938); Limenilis serpa paraena Fruhst., Brown and Mielke (\967); Adelpha paraena paraena Bates, Neild (1996). Identification, taxonomy and variation: Adelpha paraena paraena is distinguished from A. paraena massilia and A. paraena reyi by having an orange postdiscal marking instead of a white postdiscal marking on the DFW in cell M3. Adelpha paraena lecromi lacks any postdiscal marking in this cell, or may have a small white dot. The subspecies is very similar to A. radiata explicator and A. radiata radiata, but is distinguished by having the submarginal series on the VFW composed of single instead of double white spots in each cell space, especially noticeable in cell M2. Adelpha serpa and A. seriphia may be distinguished in a similar manner, and always have a fiilly developed white postdiscal marking in cell M3 on the DFW. The veins crossing the orange DFW subapical marking in females of A. paraena paraena tend to be orange rather than dark brown, while Guianan females occasionally have faint, sparse orange postdiscal scaling in cell Cul on the DFW. Adelpha paraena was described by Bates (1865) based on the female, and I have examined a syntype female, which was figured by Neild (1996), and possible syntype male (Bates appears to have occasionally misidentified the sexes of type specimens of other new Adelpha taxa that he described). The description is accurate and discusses the diagnostic character for this species, the single submarginal spots on the VFW, although

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129 it appears that all subsequent authors, with the exception of Neild (1996), failed to take note of this important character. Fruhstorfer (1915) placed the taxon as a subspecies of ^4. serpa, while Hall (1938) correctly reinstated it as a distinct species based on the rearing experiments of Miles Moss in Para. I have examined several specimens from Meta Province in Colombia which exhibit variation on the DFW, suggesting intergradation with A. paraena reyi, which occurs in Venezuela near the Colombian border at the base of the eastern Merida slopes. The female specimen from Rio Ariari has a small white spot at the base of cell M3, while both females from Remolinos lack orange postdiscal markings in cell M3, the latter (as hsted below) also has a small white spot at the base of cell M3. The former Remolinos specimen closely resembles A. paraena lecromi described below, but has the narrow white DFW postdiscal band and broad orange DFW subapical marking typical of the nominate subspecies. Two specimens of A. paraena paraena from southeastern Brazil in the USNM and the AME differ from typical A. paraena paraena in having broader white dorsal postdiscal bands. However, I have also seen a specimen in the AMNH labeled "Parana" [S.E. Brazil], which appears to be typical, although it is possibly mislabeled. More material may demonstrate the southeast Brazilian population to be distinct and to merit subspecific status, but I freat it for the present as A. paraena paraena. Range: East of the Andes from Venezuela to Bolivia, Brazil and the Guianas. The population from southeastern Brazil may represent a distinct subspecies. Immature stages: There is some confiision regarding the identification of this species and A. serpa diadochus in the paper of Moss (1933), who reared both species in Para, Brazil. While his specimens were identified by Hall (1933), whose discussion of Moss' species clearly indicated that he was aware of the distinguishing characters of both A. serpa and A. paraena. Moss' collection in the BMNH has a series of A. paraena paraena identified as A. serpa, and a series of A. serpa diadochus identified as A. paraena. It is

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130 therefore unclear whether Moss' (1933) published information for each of these species is imder the correct name or not, and I have been unable to ascertain this through examination of the preserved early stage material, which is in uncharacteristically poor condition. For the present I regard the names in the drawer of Moss' collection as applying to the same taxa as in his paper (Moss, 1933), and the following information, and the hostplant records in Table 2, were reported by him under the name A. serpa. Moss figured both the last instar larva (pi. I, figs. 15, 16) and the pupa (pi. H, fig. 8). The last instar is dark green, with an area behind the head and the posterior end orange brown, the head capsule is striped with black and there are well developed scoli on segments T2, T3, A2, A7 and A8. The A2 scolus is dark green, composed of densely overlapping spines and curved posteriorly, the remaining scoli are orange brown. Supraspiracular scoli are short with terminal spines. When at rest in a curved position (shown in Moss' figure) the larva strongly resembles a bird dropping. The pupa has a shining metallic color, the dorsal projections are relatively small and the head horns are long, thin and curved laterally. The larvae are found on bushes or the lower branches of the hostplant. Habitat and adult ecology: The nominate subspecies is widespread, but with the exception of the Guianas and lower Amazon, it is rare throughout its range. In eastern Ecuador it occurs as solitary individuals in both disturbed primary and secondary rain forest habitats up to 850m and is most fi-equently encountered in the wet season. Males may be attracted to rotting fish in subcanopy traps in light gaps, most often in the early afternoon on hot days. Specimens examined (59m, 380: t trans, to A. p. reyi, no orange in DFW cell M3; % trans, to A. p. reyi, trace white spot in DFW cell M3. VENEZUELA (10: no specific locality If BMNH. COLOMBIA (Im, 30: Meta: Rio Ariari Aug. IfJ LMC; San Jose Guaviare Aug. If JFL; Remolinos (Meta) X/89 leg. J. Burgos Imf JFL; Remolinos Meta 25/3/87 leg. J.F. LeCrom Iftt JFL. ECUADOR (4m): Morona-Santiago: Bomboiza 850m Jul. Im KWJH; Santiago 350m Sep. Im KWJH; Napo: Chichicomimi 450m Jul. Im KWJH; Finca San Carlo 600m Sep. Im KWJH. PERU (20m, 30: Hudnuco: Tingo Maria May Im AME; Junin: La Merced 2500' Apr. Aug. 4m, 3f BMNH, Im BMB; Rio Satipo 750-850m Aug. Im MUSM, Im AMNH; Loreto. Boija 1000' Im BMB; Iquitos Aug. Im BMNH; Pebas 2m ZMHU, Im BMNH; San Roque Im AME; Explomapo Camp, nr. Iquitos Mar. Im AME; Madre de Dios: Boca Rio La Torre 300m Sep. 4m MUSM; Not located: Huallaga Im AMNH. BOLIVIA (Im): Santa Cruz: Santa Cruz Im BMNH. BRAZIL (I4m, 170: Amazonas: Humayta Jul.-Sep. Im BMNH; Massaury Im, If ZMHU; Manaus Im ZMHU; S5o Paulo de 01iven9a Im ZMHU; mnas Gerais: km. 142 Curvelo, Leitao May Im USNM; Para: Obidos If BMNH; Para 7m, 13f BMNH, If AMNH, Im BMB; Santarem Nov. Im AME; Parana: "Parana, Brazil//coll. Frank Johnson" 1 f AMNH-error?. GUYANA (Im, 40: East Berbice/Courantyne: New River 750' Jan. -Mar. If BMNH; Upper Takutu/Upper Essequibo: Kuyuwini R. Im AMNH; Not located: Parish If BMB; no specific locality 2f BMNH. FRENCH GUIANA (15m, 60: Cayenne: Cayenne Im ZMHU, If BMNH; Guatimala Im MNHN; Laurent du Maroni: Maroni River Im AME; St. Laurent du Maroni If MNHN; Not located: no specific

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131 locality 12m, 4f BMNH. SURINAM (Im, IQ: Brokopondo: Geldersland, Surinam River If USNM; Not located: no specific locality Im BMNH. COUNTRY UNKNOWN (Im, 20: Onoribo Mar. If BMNH; no specific locality Im ZMHU, If BMB. ssp. nov.? BRAZIL(lm, 10: Dislrito Federal. "Brazil: Parque do Gama, 950m, D.F. Brazil, 14 May 1969, S.S. Nicolay" Im USNM; Minas Gerais: Paracatu, km. 231 Belo Horizonte-Brasilia Apr. If AME. Additional locality data: BRAZIL: Dislrito Federal: Parque do Gama UOO Jun.; Golds: Leopoldo Bulhdes 1000m Dec.; Minas Gerais: km. 485 BR-040 600m Feb. (Brown and Mielke, 1967). Adelpha paraena lecromi Willmott, ssp. nov. Figs. 16c,d Adelpha nr. paraena Bates, Aiello (1 984); Adelpha serpa ssp. nov. Lamas and Small ( 1 992) Description and diagnosis: FW length of HT: 28mm. Both sexes differ from A. paraena massilia and A. paraena reyi in lacking or only having a faint trace of a white spot at the base of DFW cell M3. In A. p. massilia and A. p. reyi this white spot is always well developed and fills the basal area of this cell as far as the distal edge of the white postdiscal spot in cell Cul. Adelpha p. reyi also has a narrower orange subapical marking and the upper postdiscal band usually consists of more pronounced whitish dashes. The nominate subspecies differs as follows on the DFW: the white postdiscal band is narrower, particularly the marking in cell M3, with the basal edge incised with black at vein Cul, and the marking in cell M3 is therefore ahnost isolated from that in cell Cul; the upper postdiscal band is never visible as faint whitish dashes; the orange subapical marking is broader and extends fully into cell M3. On the VFW of A. p. lecromi the pale subapical marking is more isolated from the upper postdiscal band by dark brown, while on the VHW the submarginal series are often cleaner white and more sharply defined. Types: Holotype male: COLOMBIA: Cundinamarca: "Adelpha sentia Bogota"; in the USNM. Etymology: This subspecies is named for Jean Franfois LeCrom, for his generous hospitality during my visit to Colombia and whose collection supplied important information on the distribution of A. paraena subspecies. Taxonomy and variation: hi the Canal Zone in central Panama, specimens of A. paraena exist with every variation in the size of the white postdiscal spot on the DFW in

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132 cell M3, indicating intergradation between A. p. lecromi and A. p. massilia. A specimen similar to A. p. lecromi from Remolinos in Colombia appears to be an intergrade between the nominate subspecies and A. p. reyi (see under nominate subspecies). I have refrained from designating type specimens from among the Panamanian material I have examined since it all originates in the Canal area, an apparent hybrid zone. The holotype was probably collected in the valleys to the northwest of Bogota. Lamas and Small (1992) refer to this subspecies as A. serpa ssp. nov., while Aiello (1984) referred to it as^. vi. paraena. Range: This subspecies has been recorded only from two areas, central Panama and the general labelling locality of "Bogota" in Colombia, the latter specimen probably being collected in the Magdalena valley west of the Cordillera Oriental. As yet it is unrecorded from western Colombia or western Ecuador, but it is expected to occur there. Immature stages: Aiello (1984) reared this subspecies (under the name nr. paraena) on Combretum decandrum (Combretaceae) in Panama. She figured the larva and pupa (figs. 4, 6, nr. PAR), and described the larva as being very similar to A. serpa celerio, but paler above, mottled brown and black, with a white dorsal patch joining A6 and A7. The scoli are brown, with the scolus on A2 the darkest, and the latter is long, straight and slender, with ascending spines, similar to A. serpa celerio; the subspiracular scoli on segments A2-4 are pink. The ventral half of segments Al-Tl is dark, and there are dark, oblique stripes on segments A2-A7. The larva turned yellowish just prior to pupation. The pupal head horns are shorter than in A. serpa celerio, curving outwards at their midpoint. Habitat and adult ecology: Presumably similar to other subspecies. Specimens examined (12m, 30: t slight hint whitish scaling in base DFW cell M3. PANAMA (1 Im, BQ: Canal Zone: Aiello Lot 82-55 no.l Imf, Lot 82-55 no.2 Im, Lot 97-15 no.2 Im, Lot 93-89 no.2 If STRI; Farfan Jun. Jul. 4m, If USNM, Im STRI; Fort Kobbe Jun. Nov. 3m USNM; Paraiso Jan. If AME. COLOMBIA (Im): Cundinamarca: Bogota Im USNM. Adelpha paraena re^/Neild, 1996 Figs. 16e,f

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133 Adelpha paraena reyi Neild (1996: 43, pi. 4, fig. 174) TL: Las Quiguas, Esteban Valley, N. Venezuela. Types (all Venezuela): BMNH(M): HTm: Las Quiguas; PTm: Macuto; PTf: Caracas; AFEN: IPTm, 2PTf: Hda. Panaga [all examined]; AFEN: IPTm, IPTf: Hda. Pinaga; TP: PTf: Tucuco; DLF: PTf: Via Altamira de Caceres; MALUZ: 2PTm El Tucuco; JB: PTf: Isla de Betancourt; RM: PTm: Rio Chucuri; R: PTm: R. S. Domingo; PTf Maracay [not examined]. Identification, taxonomy and variation: This subspecies is most similar to A. p. massilia, from which it is distinguished by the narrower DFW orange subapical marking and by typically having slightly broader white postdiscal bands. There is some variation in the width of the postdiscal bands and the orange DFW subapical marking, the former being broadest and the latter narrowest along the Cordillera de la Costa. The upper postdiscal band is often present on the DFW as whitish rays, and sometimes in females as solid white streaks (see Neild (1996) for illustrations of a wide range of phenotypes). This subspecies was clearly described and figured by Neild (1996) and the holotype is in the BMNH. Range: North and northwestern Venezuela: west of the Cordillera de Merida to the Sierra de Perija, on the eastern slopes of the Cordillera de Merida and along the northern slopes of the Cordillera de la Costa. Although unrecorded to date from Colombia, it should occur there both in Norte de Santander and extreme northern Meta. Habitat and adult ecology: Neild (1996) reports that he never encoimtered this subspecies in the field, but like other subspecies of A. paraena it appears to be much rarer than the sympatric subspecies of A. serpa. Adelpha p. reyi is sympatric along the Cordillera de la Costa with A. erotia caphira (f lerna), A. naxia, A. malea ixia, and A. bamesia trinita, all of which are unrelated species which have forms strikingly similar to A. paraena reyi, and these five taxa are almost certainly involved in mimicry. Specimens examined (3m, 4f): VENEZUELA (3m, 40: Barinas: Rio Caparo Research Station, 32 km. E. El Canton, b-Iight, Feb If USNM; Carabobo. Las Quiguas, Esteban Vail., N. Ven. Nov.-Mar. Im BMNH; Dislrito Federal: Caracas If BMNH; Macuto Om Mar. Im BMNH; Tachira: Hda. Panaga 800m Mar. Oct. Nov. Im, 2f AFEN. Additional locality daU: VENEZUELA: See type data above (Neild, 1996). Adelpha paraena massilia (C. and R. Felder, 1867) Figs. 16g,h; 183a,b

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134 Heterochroa massilia C. and R. Felder (1867: 423) TL: Mexico. Types: BMNH(R): STm: "Syntype//Type//Mexico Salle Type//H. massilia Felder/Zmassilia n." [examined] Adelpha serpa var. massilia Fldr., Kirby (1871); Adelpha serpa senlia G. and S., Frustorfer (1915), audi., misid.; =Adelpha serpa celerio Bates, Hall (1938); Limenilis (Adelpha) sentia G. and S., Ross (1976) misid.; Adelpha serpa massilia Fldr., Lamas and Small (1992); Adelpha paraena massilia Fldr., Neild (1996) Identification, taxonomy and variation: Adelpha paraena massilia is distinguished from the nominate and A. p. lecromi by having a well developed white postdiscal spot in cell M3 on the DFW. Adelpha p. reyi has the DFW subapical orange patch much reduced in width and broader white postdiscal bands. Adelpha p. massilia is similar to sympatric A. serpa and A. seriphia, but may be distinguished by the submarginal series on the ventral surface being composed of single instead of double white spots in each cell space. Adelpha nea sentia is distinguished by having a trace of orange at the distal margin of the white postdiscal spot in cell M3 on the DFW, in addition to several other characters (see imder A. nea sentia). There is some local variation in the size of the spots of the VFW submarginal series, which are usually reduced or absent in cells M3, Ml and the apical area. Felder and Felder (1867) described massilia from an unspecified number of male specimens collected by Salle in Mexico in their own collection, and compared it with A. serpa celerio, stating that the two submarginal white bands on the VHW were composed of undivided spots, a character separating the taxon from both A. serpa celerio and A. seriphia godmani. I have examined a syntype in the BM^fH. Soon after Kirby (1871) placed massilia as a variety of A. serpa, and ever since there has been confiision as to its correct taxonomic status. Godman and Salvin (1884) figured a specimen oi A. seriphia godmani as Adelpha massilia, misled by the original description's mention of narrower white postdiscal bands. Fruhstorfer (1915) placed the name as a subspecies of A. iphiclus and figured a specimen of A. iphicleola iphicleola as massilia (Fruhstorfer, 1913, pi. 107e), referring to true massilia as A. serpa sentia (Fruhstorfer, 1915), both misidentifications that were followed by most subsequent authors (e.g., Beutelspacher, 1976; de la Maza, 1987; DeVries, 1987; Austin, 1992; Austin et al, 1996). Neild (1996)

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135 was the first to correctly treat massilia as a subspecies of ^4. paraena. In central Panama intergrades to A. paraena lecromi occur, with a reduced white postdiscal spot in cell M3 of the DFW, and a specimen was figured by Aiello (1984) as "m. paraena". Range: Mexico to central Panama, with intergrades to A. p. lecromi in the Canal Zone. Habitat and adult ecology: Little has been published on the adult ecology of this subspecies, except for a report by Austin et al. (1996, as serpa sentia) that it is uncommon in the Tikal area of Guatemala within and along the edges of primary forest, in the months of May and December. DeVries (1987) never saw the species in the field. In Chajul, Mexico, I foimd the subspecies in association with disturbed lowland rain forest. Judging fi-om the nimiber of specimens in collections it is imcommon throughout its range, being heavily outnumbered by the similar and common A. serpa celerio. Specimens examined (32m, 230^ t trans, to A. p. lecromi, with white spot in DFW cell M3 half normal size. MEXICO (I2m, lOf): Chiapas: Chajul Rio Lacuntun Jul. If KWJH; Palenque Jan. Im AMNH; Oaxaca. Chimalapa Oct. Im, 2f AME; Tabasco: La Venta 10m Jan. 2m, If AME; Veracruz: Catemaco Jun. Oct. 2f AME; Coatzacoalcos Jan. Im AME; Franja Vieja Aug. If AMNH; Minatitlin Jul. Im AMNH; Moloacan Sep. Im AMNH; Pajaritos Jul. Im AMNH; Popoctepetl, Tuxtla 400m Jul. If MUSM; Tierra Blanca Aug. Im, If AME; Veracruz Im USNM, If AMNH; Not located: Cotontepec Jun. Im AME; no specific locality Im BMNH. GUATEMALA (9m, 30: El Peten: Sayaxchi Aug. Sep. 6m, 2f AMNH; Escuintla: Palin Oct. Im USNM; Retalhuleu: San Sebastidn 2m, If USNM. HONDURAS (Im, 2f): Colon: Puerto Castilla Jul. Im BMB; Cortes: San Pedro Sula 2f BMNH. COSTA RICA (Im, \{): Not located: Finca Taboga Jan. Im FSCA; no specific locality If AMNH. PANAMA (9m, 7f): Bocas del Toro: Bocas del Tore Jan. Iff USNM; Chiriqui: Boquete Dec. If USNM; Bugaba 800-1500' Im BMNH; Chiriqui Im BMNH, Im BMB; Lino If BMNH; Canal Zone: Aiello Lot 93-76 no.2 1ft STRI; Farfan Feb. Jun. Jul. 2m, 2f(lm, Iff) USNM, 2fl(lt) STRI; Fort Kobbe Jun. Imf USNM; Gamboa, Cerro Pelado Oct. Iff USNM; Paraiso Jan. Imt AME; STRI Tupper Centre Jan. If STRI; Panama: El Llano 330m Cord, de San Bias Jul. Iff USNM; Veraguas: no specific locality 2mt BMNH. Additional locality data: MEXICO: Veracruz: 1.75 mi. E. Sontecomapan sea level Aug.; 2 mi. N.E. Catemaco 300m Sep. (Ross, 1976). GUATEMALA: El Peten: Tikal (Austin et al., 1996). BELIZE: Corozal: Fresh Water Creek Forest Preserve (Meerman, 1 999: as serpa sentia). Adelpha radiata FnihstorfeT, 1915 Figs. 17; 101; 184 Identification, taxonomy and variation: The majority of subspecies of .4. radiata may be distinguished fi-om A. serpa, A. seriphia and A. hyas by the great reduction or loss of the white postdiscal spot in cell M3 on the DFW. Adelpha radiata myrlea closely resembles A. serpa serpa but has the spots of the VHW submarginal series divided in each cell space. Adelpha paraena may be similarly distinguished fi-om all A. radiata subspecies by having undivided spots comprising the ventral submarginal series. There is geographic variation in the shape and

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136 size of the orange DFW subapical marking and presence of a white spot in cell M3 on the DFW, and five subspecies are recognised. Adelpha radiata is most closely related to A. serpa, A. seriphia and A. hyas, with which it shares divided spots in the VFW submarginal series. There are no consistent genitalic differences between any of these species. Range and status: Central Panama to northwestern Ecuador, eastern Ecuador, French Guiana and southeastern Brazil. Very rare, in lowland rain forest below 900m, unrecorded throughout the majority of its presumed range, three of the five known subspecies described since 1995. Specimens examined: 28 (14m, 14f) Adelpha radiata radiata Fruhstorfer, 1915 Figs. 17a,b Adelpha serpa form radiata Fruhstorfer (1915: 531) TL: Santa Catharina [Brazil]. Types: BMNH(T): STm: "Fruhstorfer Coll. B.M. 1 937-285//serpa radiata Fruhst./ZBrasilien Blumenau Fruhstorfer//TYPE//Type//Syntype"; STf: "Fruhstorfer Coll. B.M. 1937-285//Brasilien Blumenau Fruhstorfer/ATYPE/ZType"; BMNH(M): STf: "Brasilien Blumenau Fruhstorfer/Zradiata Fruhst.//Fruhstorfer Coll. B.M. 1937-285"; STm and f: "Brasilien Blumenau Fruhstorfer/ZFruhstorfer Coll. B.M. 1937-285" (all examined] Adelpha radiata Fruhst., Hoffmann (1936); =Adelpha serpa serpa Boisd., Hall (1938); Adelpha radiata radiata Fruhst., Willmottand Hall (1999) Identification, taxonomy and variation: Adelpha radiata radiata differs fi-om A. radiata myrlea in having in cell M3 on the DFW only a small white dot, in addition to a large orange postdiscal marking. It is distinguished fi-om other subspecies by enlarged submarginal series on the VHW (and VFW), separated only by a thin, darker brown line. This subspecies is similar to A. paraena paraena, which has undivided spots in each cell forming the ventral submarginal series. Adelpha hyas hyas is smaller and has, on the VFW, deeper intruding dark lines into the pale subapical marking, a yellowish color in the discal cell and no red between the fourth discal cell bar and the postcellular bar, and on the VHW the reddish postdiscal band is almost fiised with the inner submarginal series. Adelpha serpa serpa always has a well developed white postdiscal spot in cell M3

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137 f • of the DFW, the VFW has dark orange postdiscal dashes in cells Cul and M3 (probably representing the outer postdiscal series) which are absent in A. radiata radiata, and the white blocks composing the inner submarginal series on the VHW are not divided by dark lines. The size of the white dot in cell M3 on the DFW is somewhat variable in the few specimens of A. radiata radiata that I have been able to examine. Fruhstorfer (1915) described this taxon as a form of Adelpha serpa, which he stated was intermediate between typical serpa and A. hyas hyas, based on an unspecified number of specimens fi-om Santa Catharina in southeastern Brazil. There are two males and three females in the BMNH fi-om Fruhstorfer's collection labeled Blumenau, which appear to be syntypes. Hoffinann (1936) stated that he regarded radiata as representing a valid species, since despite having reared many individuals of A. serpa serpa, he never found any specimens similar to radiata. Hall (1938), however, placed the taxon as a synonym of nominate A. serpa, an unsurprising decision since he relegated taxa of no less than six distinct A. serpa group species to the status of forms or subspecies of .4. serpa. In fact, A. radiata radiata is sympatric with A. hyas hyas, A. serpa serpa and probably A. paraena paraena, and distinguished by several characters fi-om each (see above). The true systematic status of the taxon radiata, and indeed all A. radiata subspecies, has been greatly obscured by the rarity of all these taxa in collections and the consequent difficulty in establishing the sympatry of phenotypes and the reliability of characters for distinguishing species. The taxonomic arrangement adopted here is the result of fieldwork in eastern Ecuador, which finally demonstrated the broad sympatry of five species (Willmott and Hall, 1999), and the subsequent tracing of each of these species throughout the Neotropics. While there is no single character which defines A. radiata throughout its range, allopatric taxa in neighboring regions are associated through a consilience of characters and thus may be grouped as a single species. Such characters will be discussed under each subspecies.

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138 Range: The nominate subspecies has been recorded from southeastern Brazil, from Rio de Janeiro to the state of Santa Catharina. It possibly also extends to northern Argentina (Misiones), since Hayward (1973) reports specimens "intermediate" between A. serpa serpa and A. hyas hyas from there. Habitat and adult ecology: Little has been reported on the behavior or habitat of this rare subspecies. Tom Emmel (pers. comm.) captured males hill topping on an open summit in Santa Catharina. Specimens examined (8m, 50: t white spot in cell M3 larger than typical. BRAZU. (9m, 50: Guanabara: Rio Im BMNH; Santa Catharina: Serrinha do Pirai, W. Joinville 950' Mar. 2m FSCA; Blumenau 2m, 3f BMNH; Joinville 20-200m Feb. Dec. 2m MUSM; Rio Natal, Sao Bento do Sul 550m Feb. Im MUSM; Sao Paulo. S5o Paulo 2500" Nov. Dec. 2ft BMB. COUNTRY UNKNOWN (Im): no specific locality Im BMNH. Adelpha radiata myrlea Fruhstorfer, 1915 Figs. 17c,d Adelpha serpa form myrlea Fruhstorfer (1915: 532) TL: Espiritu Santo [Brazil]. Types: BMNH(T): HTm: "Fruhstorfer Coll. B.M. 1937-285//serpa myrlea Fruhst.//Espirito Santo Brazil A. Heyne, Berlin-Wilm. V//TYPE//Type//Syntype" [examined] =Adelpha serpa serpa Boisd., Hall (1938); Adelpha radiata myrlea Fruhst., Willmott and Hall (1999) Identiflcation, taxonomy and variation: Adelpha radiata myrlea differs from the nominate subspecies in having a well developed postdiscal white spot in cell M3 on the DFW, and no orange postdiscal spot in this cell. It is very similar on the dorsal surface to A. serpa serpa, but on the VFW lacks the dark orange postdiscal dashes in cells Cul and M3 and has the spots of the VHW submarginal series divided in each cell. Adelpha hyas hyas has a clearly different dorsal and ventral surface, the latter differing particularly in the deeply intruding dark lines in the pale VFW subapical area and the expanded submarginal series which extend to merge with the postdiscal series in cells Cul and M3; in A. r. myrlea, the postdiscal series are absent in these cells. Fruhstorfer (1915) described myrlea from a single specimen in his collection on the basis of the reduced DFW orange subapical marking, and the holotype is in the BMNH. There is little variation in the three specimens examined. This taxon is placed as

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139 a subspecies of A. radiata on the basis of the divided submarginal series (also shared with A. hyas hyas) and the absence of the postdiscal series in cell Cul on the VHW. Range: Southeastern Brazil, from Espirito Santo to Rio de Janeiro, though possibly extending as far north as Bahia. Habitat and adult ecology: This subspecies is very rare in collections and nothing has been reported on its habitats or behavior. Specimens examined (Im, 21): BRAZIL (Im, 2f): Rio de Janeiro: San Antonio dos Brotos If BMNH; Espirito Santo: no specific locality Im BMNH; Not located: no specific locality If BMNH. Adelpha radiata explicator Willmott and Hall, 1999 Figs. 17e,f; 101 Adelpha radiata explicator Willmott and Hall (1999: in press) TL: Finca San Carlo, km. 13 Puyo-Tena rd., E. Ecuador. Types: KWJH; HTm: Finca San Carlo [examined] Identification, taxonomy and variation: Adelpha r. explicator is distinguished from other A. radiata subspecies in their respective accounts. It is very similar to A. paraena paraena, but may be distinguished by the divided spots in each cell space of the venfral submarginal series. Adelpha r. explicator is sympatric with A. paraena paraena, A. serpa diadochus and A. hyas hewitsoni, while A. seriphia aquillia also occurs in eastern Ecuador but at higher altitudes. It shares with nominate A. radiata the reduction of the DFW white postdiscal spot in cell M3 and the lack of any dark orange postdiscal markings (representing the outer postdiscal series) on the VFW in cells Cul and M3, and with A. r. gilletella and A. r. aiellae, the reduced VHW submarginal series. Range: To date only known from a single site in eastern Ecuador, but presumably occurring at least throughout the upper Amazon basin. Habitat and adult ecology: This subspecies is extremely rare, only known to date from a single specimen from eastern Ecuador, which was captured in a large light gap created by a tree fall along a ridge top in primary forest. This individual was flying about the

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140 clearing in bright sun in the middle of the morning, with the characteristic flight of all 5er/?a-group members, periods of alternating rapid wing beats and gliding. It perched in between flights on bushes 1-5 m above the ground around the edge of the arena, in the company of A. paraena paraena. The great similarity between the dorsal surfaces of these two taxa suggests there may well be a mimetic relationship. The recent discoveries of ^. r. explicator, A. r. gilletella and A. r. aiellae suggest that A. radiata is a very widespread species, and since all known male specimens outside of southeastern Brazil have been captured on hilltops, further collecting in this microhabitat will hopefully extend the known ranges of A. radiata subspecies and possibly reveal further undescribed taxa. Specimens examined (Im): ECUADOR (Im): Napo: Finca San Carlo 600m Sep. Im KWJH. Adelpha radiata aiellae Willmott and Hall, 1999 Figs. 17g,h; 184 Adelpha radiata aiellae Willmott and Hall (1999: in press) TL: nr. Lita, Rio Baboso, ridge to east, W. Ecuador. Types: KWJH: HTm: nr. Lita, Rio Baboso, ridge to east; ATf: km. 40 LitaSan Lorenzo rd., Rio Durango, W. Ecuador; USNM: PTf: Gatun, Panama; STRI: PTf: Gatun, Panama; FSCA: 2PTf: Piila, Panama; LMC: PTf: Yatacue, Colombia [all examined] Identification, taxonomy and variation: Adelpha r. aiellae is superficially most similar on the dorsal surface to A. r. explicator, fi-om which it differs in having a smaller orange spot in cell M3 of the DFW, the orange DFW subapical marking broader in cells M2-R5 with the veins crossing it less heavily lined with black, and a narrower white postdiscal band on the DHW. On the VFW, A. r. aiellae has dark orange dashes in cells Cul and M3 representing the outer postdiscal series, which are absent in A. r. explicator. In fact, A. r. aiellae is probably more closely related to A. r. gilletella, which shares all of the previously mentioned diagnostic characters, and differs from A. r. aiellae only in the presence of an orange spot on the DFW in cell Cul and in having a narrower white DFW postdiscal band. There is some variation in the size of the orange spot in cell M3 on the dorsal forewing, and Panamanian specimens have the veins crossing the orange subapical marking on the

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141 dorsal forewing slightly less strongly lined with dark brown than the two known Ecuadorian specimens. A tiny white dot is sometimes present at the base of cell M3 on the DFW. This taxon is sympatric with A. serpa duiliae and A. seriphia godmani, and associated with A. radiata on the basis of its similarity to A. r. gilletella. Range: Central Panama to northwestern Ecuador, west of the Andes, but possibly more widespread in Central America, probably extending at least into Costa Rica. Immature stages: Constantino (1998) listed the hostplant and described the early stages of this Adelpha taxon under serpa", but an examination of the voucher specimen in the LMC shows it to be A. radiata. The last instar is dark brown with black markings and green spots, resembling lichen, the head is dark brown with black marks. The color and form of the scoli were unfortunately not specified. The pupa has long head horns and is chrome colored, with the sutures lined in black, becoming bright golden just before eclosion. Aiello (1984) reports rearing a species in the A. serpa group (her Group I) with a larva resembling A. serpa celerio but a pupa resembling A. paraena. Unfortunately the pupa died, but this species might well also have been A. radiata aiellae. Habitat and adult ecology: This subspecies is very rare in collections and to date has been recorded from near sea level to 900m in the vicinity of disturbed primary rain forest habitats. The only male specimen known to date was found perching on top of a 4m high bush at the edge of a very wide path along the top of a steep-sided forested ridge. The single female I have seen was flying 5m above a wide river through secondary growth in the early afternoon. In western Ecuador the species has only been encountered towards the end of the wet season. Specimens examined (Im, 6f): PANAMA (4f): Canal Zone: Gatun 350' Jan. May If USNM, If STRI; Colon: Pifla 200m Jul. 5, Feb. 7 2f FSCA. COLOMBIA (10: Valle del Cauca: Alto Anchicaya Yatacue 600m Sq). If LMC. ECUADOR (Im, If): Carchi: Lita, ridge east of Rio Baboso Jul. Im KWJH; Esmeraldas: Rio Durango 250m Jul. If KWJH.

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142 Adelpha radiata gilletella Brevignon, 1995 Figs. 17iJ Adelpha gilletella Brevignon (1995: 17, fig. 27, 28, 29, 30) TL: Galion, Roura, French Guiana. Types (all French Guiana): LCB: HTm: Gallon; ATf: pk 35 RN2, Roura; [original figures examined]; PTm: Galion Jan.; PTm: pk 21 piste de Kaw, Roura Jul.; JYG: PTm and f: pk 21 piste de Kaw, Roura in copulo Oct. [not examined] Adelpha radiata gilletella Brev., Willmott and Hall (1999) Identification, taxonomy and variation: This subspecies differs from all other subspecies in having a well developed orange postdiscal spot in cell Cul on the DFW, and from A. radiata radiata and A. r. myrlea by the much narrower ventral submarginal series. Adelpha r. explicator has a narrower orange block in cell M3, a narrower orange subapical marking with the veins that cross it strongly lined with black, and the white postdiscal band is broader, especially the spot in cell Cul. On the VFW of A. r. explicator the pale subapical marking and postdiscal spot in cell M3 are of uniform color and do not have the distal edge dark orange. Adelpha r. aeillae has a broader white DFW postdiscal band, particularly in cell M3. Brevignon and Brevignon (1997) figure a specimen of A. r. gilletella which has a large, round postdiscal orange marking in cell Cu2 distal of the white band, and thick orange postdiscal dashes on the DHW along the outer postdiscal series in cells M3, M2 and Ml. Brevignon (1995) clearly illustrated dorsal and venfral surfaces of the male and female of this taxon, which he described as a species on the basis of its sympatry with Adelpha serpa and Adelpha paraena. He was, however, unaware of the existence of the taxon radiata, with which this subspecies shares the reduction of the white postdiscal spot in cell M3 on the DFW, only occurring in related species in the sympatric A. paraena paraena. Adelpha r. gilletella and A. r. radiata are also of similar size and wing shape, and the orange DFW subapical marking is of similar shape and lacks the dark intruding lines typical of A. hyas on the ventral surface.

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143 Range: This subspecies is only known to date from northern French Guiana, but it is undoubtedly more widespread, probably extending throughout the Guianas to northern Brazil (Amapa), and perhaps to the lower Amazon. Habitat and adult ecology: Brevignon (1995) reports that the subspecies was encountered in forested hilltop clearings at the end of the morning, and data indicate that it probably occurs throughout the year. Clearly it is very rare and I have seen no specimens in any of the collections that I have examined. Specimens examined (Im, If): FRENCH GUIANA (Im, If): Cayenne: Galion Roura Dec. Im(photograph) LCB; pk 35 RN2 Roura Nov. Ifl^jhotograph) LCB. Additional locality data: See type data above (Brevignon, 1995). Adelpha serpa (Boisduval, 1 836) Figs. 18; 102; 185 Identification, taxonomy and variation: Adelpha serpa is very similar to A. seriphia, but is consistently distinguished (from sympatric subspecies) by having the blocks comprising the DFW postdiscal band arranged more vertically, the DFW orange subapical marking more horizontal, and (from all subspecies except A. seriphia barcanti) by having the spots of the irmer submarginal series approximately parallel to the orange postdiscal band on the VHW, rather than having those in cells Rs and Sc+Rl much closer than the remainder. Adelpha hyas is smaller, has dark brown lines intruding into the pale subapical marking on the VFW, and lacks orange postdiscal dashes on the VFW in cells Cul and M3. Most subspecies of A. radiata lack a well developed white spot in cell M3 on the DFW, and when this spot is developed, the spots comprising the VHW submarginal series are poorly defined and coalesce. All other similar A. serpa group species may be distinguished by having single instead of divided spots in each cell space comprising the submarginal series on the VFW. There is geographic variation in the width and color of the postdiscal bands, the

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144 size of the orange DFW subapical marking and the width of the ventral submarginal series, and four subspecies are recognised. Adelpha serpa forms a clade with three closely related species, A. radiata, A. hyas and A. seriphia (see Chapter 3). Range and status: Mexico to western Ecuador, Venezuela to Bolivia, Brazil, Guianas and Paraguay. In disturbed forest areas, common west of the Andes in lowland and lower montane forest, rarer east of the Andes in lowland forest up to 1000m. Specimens examined: 697 (359m, 3381) Adelpha serpa serpa (Boisduval, 1836) Figs. 18a,b; 102; 185a-c Heterochroa serpa Boisduval (1836: pi. 8, fig 4) TL: Brazil. Types: BMNH(M): STm: "LECTOTYPE m Heterochroa serpa Boisduval 1836 G. Lamas det. 1997//Seipa B.d. iphicla God.Cr. 188 BrasV/Heterochroa Serpa, Bdv. (species gel. pi. 8 fig. 4//Ex. Musaeo Dris. Boisduval//Ex. Oberthiir Coll. Brit. Mus. 1927-3" [examined] =Adelpha damon Fruhstorfer (1913: pi. 107c; 1915: 531) TL: [Paraguay; Santa Catharina]. Types: BMNH(T): STm: "Fruhstorfer Coll. B.M. 1937-285//serpa damon Fruhst.//Brasilien Blumenau Fruhstorfer//TYPE//Type"; STf: "Fruhstorfer Coll. B.M. 1937-285//Brasilien St. Cath. Fnihstorfer//TYPE/A"ype"; BMNH(R): STf: "serpa damon Fruhst.//Holotype//Type//Sapucay Paraguay 16.8.04 (W. Foster)//107 C4" [all examined] =Adelpha serpa form omamenta Fruhstorfer (1915: 531) TL: "presumably Bahia" [Brazil]. Types: MHNG: HTm: data not recorded [examined] Adelpha serpa Boisd., Kirby (1 87 1 ) Identiflcation, taxonomy and variation: The nominate subspecies is distinguished from A. serpa diadochus by the DFW white postdiscal band being narrower and composed of more disjointed spots, by the pale subapical marking on the VFW being almost connected to the white dashes of the upper postdiscal band by pale shading (in A. s. diadochus the two are separated by a brown line), by the more angular hindwing which has a straighter distal margin, by the sfraighter orange postdiscal band on the VHW, by the enlarged ventral submarginal series which are separated only by a thin dark brown line and by not having the spots of the VHW submarginal series divided in each cell. Other subspecies are easily distinguished by the reduced VHW submarginal series, the spots of which are more distinctly paired in each

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145 cell. Adelpha serpa serpa is similar to A. radiata myrlea, which differs by having the VHW submarginal spots divided in each cell. There is some variation in the width of the white postdiscal bands and the extent of the DFW subapical orange marking, which usually enters cell M3 and may extend as far as vein Cul. In the case of the latter, the DHW may have orange dashes in each cell, marking the outer postdiscal series. Specimens with extensive DFW orange occur from Rio de Janeiro to Sao Paulo, but since typical specimens also occur throughout the same range I regard the variation as infrasubspecific. Boisduval (1836) appears to have provided no text to accompany his original figure of Adelpha serpa, which was named on the plate where the butterfly was figured in a natural pose with the wings closed, showing only the ventral surface. From the figure there is, however, no question as to the identity of the taxon, the broad VHW submarginal series being clearly evident. On page three of the "Explications des Planches" Boisduval gives the type locality of Brazil. Recently Lamas (pers. comm.) discovered a specimen among duplicate accession material from Oberthiir's collection at the BMNH which appears to be a syntype. This specimen has slight orange spotting in the middle of cells M3-Sc+Rl along the inner postdiscal series on the DHW, and the DFW subapical orange marking extends to the middle of cell Cul, matching female specimens in the BMNH from Rio de Janeiro. Fruhstorfer (1913) figured his new taxon damon based on a female specimen from Paraguay, which he later placed as a subspecies of A. serpa (Fruhstorfer, 1915), stating that it differed from typical specimens of A. serpa in having a reduced DFW subapical orange marking and a narrower white postdiscal band. The figured female is in the BMNH(R), as are fiirther syntypes from Blumenau; both differences represent no more than individual variation and Hall (1938) correctly synonymised damon with serpa. Fruhstorfer (1915) described a specimen of the form with orange dorsal postdiscal markings as omamenta, based on a single specimen in the Geneva Museum, and this name was also synonymised with A. serpa by Hall (1938). Dr. Lobl, of

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146 the MHNG, kindly supplied me with a photograph of the presumed holotype of ornamenta, which closely resembles the syntype of serpa. The nominate subspecies of A. serpa is actually rather different in several respects from remaining A. serpa subspecies, which have traditionally been treated as a distinct species, A. celerio (Fruhstorfer, 1915; Neild, 1996). In particular, it shares with A. seriphia the disjointed white postdiscal spots on the DFW, a more vertical orange DFW subapical marking, and the tendency to have dorsal orange dashes along the outer postdiscal series. However, the Bolivian A. seriphia therasia, geographically the closest subspecies, does not closely resemble A. serpa, while in the Amazonian A. serpa diadochiis the VHW submarginal spots are enlarged and occasionally become entire, as in nominate A. serpa. Furthermore, Paraguayan specimens of A. serpa, particularly the female syntype of damon, are clearly intermediate in wing pattern between A. s. diadochus and A. s. serpa. These specimens have the hindwing distal margin and orange postdiscal band on the VHW straighter than typical A. s. diadochus, but the black separating the submarginal series is heavier than in typical A. s. serpa, as in A. s. diadochus. There is reduced white scaling in the VFW apex, as in typical A. s. diadochus, and the pale VFW subapical marking is more isolated from the upper postdiscal band. Range: The nominate subspecies occurs from southeastern Brazil to Paraguay and northeastern Argentina. Immature stages: Both Muller (1886) and Hoffmann (1936) reported rearing "Adelpha serpa" in Santa Catharina (Brazil), but while Muller's record is similar to many others of this species, in that it could also apply to A. radiata or A. hyas, Hoffmann clearly was aware of how to distinguish the three species and his hostplant and early stage records are therefore reliable. Both reported that the eggs are typical of the genus and are laid on the upperside of a leaf at the tip. Early instars are greenish with a matt, rust-red head capsule with paler marks, and the body is marked with light green spots, each with a fine gray seta (Hoffmann). Later instars develop distinctive, short, broad scoli on A2 in which the

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147 spines are enlarged, flattened and merged (figured by Miiller on Taf 2, fig. 7). The head capsule (Miiller, 1886: Taf 2, fig. 21) has pronounced spines around the edge, and two black, vertical stripes at the outside edges of the labrum, in addition to vertical stripes at the lateral edges of the face. The pupa is a brilliant golden color with small dorsal projections in the same position as the last instar scoli on segments T2, T3, A2, A3, A4, with those on A2 and A3 the most pronounced (Miiller, 1 886). The development times given by Hoffinann (1937) are as follows: egg, 4 days; pupa, 12 days in February, 16 days in April to May, possibly due to the cooler temperature. The larva extends the leaf midrib and rests on it in early instars (Hoffinann, 1937). Habitat and adult ecology: Ebert (1969) found this subspecies to be uncommon up to 1400m in Minas Gerais in low canopy forest. Brown (1992) reports that this subspecies occurs in humid forest in the Serra do Japi, Sao Paulo, up to 1000m. Judging from the number of specimens in collections it may be common in the field. Hoffinarm (1936) found the larva at Santa Catharina in May. Specimens examined (100m, 780: t orange on DFW in cell Cul ; J trans, to A. s. diadochus. "MEXICO" (Im): no specific locality Im MCZ-error. "GUATEMALA" (If): no specific locality 1 f BMNH-error. "HONDURAS" (Im): Cortes: San Pedro Sula Im BMNH-error. "PERU" (10: Junin: Chanchamayo If ZMHU-error. BRAZIL (86m, 680: Espirito Santo: no specific locality 2m BMNH; Guanabara: Corcovado 2m, If MCZ; Rio May If BMNH, 2m ZMHU, 6m, 13f BMNH; Rio Janeiro Jul. Nov. 2f USNM, 5ft MNHN, 3m MCZ, Im, If USNM, If AMNH; Rio de Janeiro N.P. Dec. Im BMNH; Rio Tijuca Iff MNHN; Tijuca Im, If USNM, 4m, 2f BMNH; Minas Gerais: Leopoldina Im ZMHU; Marumba R. Feb. If BMNH; Nova Lima Apr. Aug. Im, If AME; Rio das Velhas, Sabara-Belo Horiz. If BMNH; Serra do Espinha90, km. 344 Rio-Belo Horizonte Apr. Im AME; no specific locality Im BMNH; Parana: Castro Im, 2f USNM, Im, If BMNH; Caviuna Jul. 2m, If AMNH; Femandes Pinheiro 22600' Apr. If BMNH, Im AME; N. Parana 5m, 8f AMNH; Ponta Grossa 900m Mar. Im USNM; Rio de Janeiro: Organ Mts. If BMNH; Petropolis 1500m May If BMNH, Im, 3f AME, Im, If USNM; Santa Catharina: Rio Natal, nr. Rio Vermelho 900-1450' Mar. Im FSCA; Blumenau Im, 2f BMNH, Im, If ZMHU, 3m MCZ, If AMNH; Campo Nola Im AME; Corupa May Im USNM, Im AMNH; Joinville 10m Feb. Mar. If FSCA, 2m AME; Rio Natal, Sao Bento do Sul 550m Feb. Im MUSM; Rio Vermelho Im, 1 f AMNH; sao Bento do Sul 1750' Mar. Im FSCA; no specific locality 2m, 3f AMNH, Im, If BMNH, Im, If ZMHU, Im BMB, Im MCZ; Sao Paulo: Alto da Serra 500' Nov. Dec. Im BMB; Mendes 2m, If AME; Santos Aug. Imt MNHN; Sao Paulo Im BMNH, 5m, 2f BMB(3mt); Not located: S. Brazil If USNM, Im BMNH; no specific locality Oct. Nov. If ZMHU, 2m BMNH, 2m BMB, Im, 2f MNHN, Im MCZ. PARAGUAY (Im, 20: Paraguari: Sapucay Dec. 2ft BMNH; Not located: S. Paraguay Imt BMNH. "FRENCH GUIANA" (2m): Cayenne: Cayenne Im MNHN-error; Not located: no specific locality Imt MNHN. COUNTRY UNKNOWN (9m, 60: Henrietta ? If ZMHU; no specific locality Im USNM, 5m, 2f BMNH, Im, If MNHN, 2m, 2f MCZ. Additional locality data: BRAZIL: Minas Gerais: nr. Pofos de Caldas 1300-1400m Feb. Mar. May Dec. (Ebert, 1969); Santa Catharina: Jaragui Apr. Dec. (Hoffmann, 1936). ARGENTINA: Misiones (Hayward, 1951). Adelpha serpa diadochus Fruhstorfer, 1915 Figs. 18c,d

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148 Adelpha celerio diadochus Fruhstorfer (1915: 531) TL: Tarapoto, Huallaga, Peru. Types: BMNH{T): ST?ni: "Illustrated in The Butterflies of Venezuela A. Neild, 1996//183//Fruhstorfer Coll. B.M. 1937-285//celerio diadochus FruhstV/Marcapata Peru//Peru H. Fruhstorfer//TYPE//Type//Syntype?" (examined] =Adelpha serpa form timehri Hall (1938: 285) TL: Guiana and Para [Brazil]. Types: BMNH(M): STm: "Illustrated in the Butterflies of Venezuela, Neild 1996//185//Syntype m Adelpha serpa f timehri Hall G. Lamas det. 1987//B.C.A. Lep. Rhop. Adelpha celerio Bates Godman-Salvin Coll. 1916.4.//m//Cariamang River B. Guiana H. Whitely/ZSyntype"; BMB: STm: Para Jan.-Mar. [both examined] =Adelpha celerio florea Brevignon (1995: 17, fig. 31, 32) TL: pk 27 Piste de Kaw, Roura, French Guiana. Types (all French Guiana): LCB: HTm: pk 27 Piste de Kaw [original illustration examined]; PTm: Montsinery, Mar.; PTm: Cacao, Roura, Apr.; PTm: Maripasoula, Jul.; JYG: ATf: Saul, Nov.; Coll. B. Hermier, French Guiana: PTm: Montagne des Chevauz, Roura, Nov.; PTm: Petits Saut, Sinnamary, Jan. [not examined] Papilio iphicla [sic] Linn., Cramer (1777: 139, pi. 188, fig. E, F) misid.; Papilio iphiclus Linn., Herbst (1793: 138, n. 79, tab. 148, fig. 3, 4) misid.; Adelpha iphicla Linn., Hubner ([1819]: 42) misid.; Papilio iphicla [sic] Linn, var.?, Godart ([1824]) misid.; =Heterochroa serpa Boisd., Hewitson (1850); =Adelpha serpa celerio Bates, Hall (1938); Adelpha serpa diadochus Fruhst., Willmott and Hall (1999) Identification, taxonomy and variation: Adelpha serpa diadochus is distinguished from A. s. celerio and A. s. duiliae by the ahnost pure white dorsal postdiscal band which tapers anteriorly on the DFW, typically leaving a relatively small spot in cell M3, by the thicker inner submarginal series on the VHW which has undivided spots in each cell near the costa, and by the smoothly curving orange VHW postdiscal band. Further characters are discussed under A. s. celerio and A. s. serpa. It is also similar to A. radiata myrlea (with which it is not known to be sympatric) and A. hyas viracocha, but has the spots of the VHW submarginal series sharply defined. Adelpha hyas hewitsoni differs by having dark lines intruding into the pale subapical marking on the VFW and by lacking dark orange postdiscal dashes in cells M3 and M3 on the VFW. Adelpha seriphia may be distinguished in the same way as A. serpa celerio from A. seriphia godmani. The DFW orange subapical marking is slightly variable in size, and particularly in Guianan specimens may extend into cell M3. The width of the DFW postdiscal band is also slightly variable, being narrowest in specimens from the base of the Andes. This subspecies was figured for the first time by Cramer (1777) on plate 188, figs. E and F, based on a specimen from Surinam, but misidentified as Papilio iphicla Linnaeus. Hewitson (1850) and Westwood (1850) both stated that Cramer's figure represented A. s. serpa, a view followed by some subsequent workers (Butler, [1870];

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149 Kirby; 1871), but not all, Bates (1865) and Godman and Salvin (1884) regarding it as representing Amazonian A. s. celerio. Fruhstorfer (1915), introducing the name diadochus, avoided the controversy by failing to mention Cramer's figure at all, and provided the briefest description of an unspecified number of specimens from Tarapoto. He stated merely that the size of the DFW orange subapical marking corresponded with that of A. seriphia godmani, while the postdiscal bands were similar to those of A. s. celerio. I have seen no specimens of A. s. diadochus, or indeed any A. jer/7a-group species, in any collections from Tarapoto, and the whereabouts of the specimen(s) on which Fruhstorfer based his description is unknown. The specimen listed above as a possible syntype came from Fruhstorfer' s collection and is labeled "celerio diadochus Fruhst." in Fruhstorfer's hand, and it might possibly have lost an original Tarapoto label during later relabelling; the type of A. thoasa zalma was also supposedly from Tarapoto, but the only syntype known simply has the locality data "Peru". Nevertheless, the possible syntype of diadochus and the type locality leave little doubt in my mind as to the identity of the taxon. Hall (1938) synonymised diadochus with A. s. celerio, but went on to describe Guianan and Para specimens as a form of A. serpa, timehri, of which I have examined two syntypes. The syntype specimen in the BMNH, from Guyana, has the orange subapical marking of the DFW extending into cell M3, a character typically, but not always, present in Guianan specimens, and usually not apparent elsewhere in the range of A. s. diadochus. On the basis of this character Neild (1996) retained timehri as distinct from A. s. diadochus, and placed timehri as a subspecies of A. celerio. However, the second syntype of timehri, in the BMB, has only a hint of orange scaling on the DFW in cell M3, and since this character seems to be variable in the Guianas and lower Amazon, and orange is present in cell M3 on the DFW in a specimen from Ega in the BMNH, I do not regard it as sufficiently constant to retain timehri as a separate subspecies. Brevignon (1995), apparently ignorant of both of the namestimehri and diadochus, described specimens of A. s. diadochus from French Guiana as a new

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150 subspecies, A. celerio florea. The holotype lacks orange in cell M3 on the DFW, while this character is apparently present in three paratypes and the allotype (Brevignon, 1995). Neild (1996) synonymised florea with timehri, while Willmott and Hall (1999) placed diadochus as a subspecies of A. serpa with both florea and timehri as synonyms, the taxonomy adopted here. Range: Eastern Colombia to Bolivia, Amazonian Brazil and the Guianas. Immature stages: Moss (1933) reared this species in Brazil (Para), but there is some confusion over his identification of the species and A. paraena paraena (see discussion under Immature stages of A. paraena paraena). The following information, and the hostplant records in Table 2, were reported by him under the name A. paraena. The egg is laid singly at the tip of the leaf and the brown first instar hatches approximately one week later. The last instar is bright green and differs fi-om the sympatric A. paraena paraena in not having orange brown scoU. Also, the lateral figure of the larva (pi. I, fig. 17) shows the scolus on A2 to be long, slender and straight, as described by Aiello (1984) for A. serpa celerio. The larvae typically occur on low saplings less than a metre above the ground and as early instars expose and extend the midrib in the manner typical of the genus, resting on the upper surface of the leaf in the final instar. Early instars apparently suffer heavy predation by Hymenoptera. Habitat and adult ecology: Adelpha serpa diadochus is very local and uncommon, and is particularly poorly represented in collections fi-om the central Amazon. Its is restricted to lowland rain forest up to 1 000m. Males may be attracted to rotting fish in subcanopy traps in old selectively logged forest light gaps. In eastern Ecuador, I have observed males perching with the wings open or closed in the early to middle afternoon on bushes 3-5m high, in large (20m wide), old clearings surrounded by primary or secondary forest along rivers, patrolling the entire clearing with a gliding flight, with few wing beats. Such microhabitats are also fi-equented by A. thoasa manilia, A. hyas hewitsoni and A.

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151 iphicleola thessalita, and these species are probably involved in mimicry. Andrew Neild (pers. comm.) captured a female flying with a male at Bomboiza in eastern Ecuador, in an area of secondary growth. Brevignon (1995) reports that the species is active in French Guiana from late morning to early afternoon, and frequents both hilly and flat areas. Specimens examined (42tn, 2%f): COLOMBIA (10: Caqueta: Rio Orteguaza 1000' Jan. If USNM. ECUADOR (6m, 30: Morona-Santiago: Bomboiza 850m Nov. 2m, If KWJH; Napo. Pimpilala 600m Aug. Im KWJH; Rio Shandia 550m Sep. Im KWJH; Pastaza: Canelos Jan. If ZMHU; Puyo 1000m Dec. Im USNM; Rio Bobonaza 1000m Oct. Im MJP; Sucumbios: Garzacocha, La Selva, Rio Napo 250m If PJD. PERU (11m, 100: Cuzco: Marcapata Im BMNH; Huanuco: Pozuzo 800' Apr. Im BMB; Tingo Maria 800m Mar. Jun. Im FSCA, Im USNM, If AME; Junin: Chanchamayo 2f ZMHU, If USNM; Ipokiari May If MUSM; La Merced 2500' Im BMB, If MUSM; Loreto: Iquitos 2f AMNH; Pebas 120m Sep. Im MUSM, Im ZMHU; Madre de Dios: Castafla 150m Oct. If MUSM; San Martin: Jepelacio 2m, If AMNH; Not located: E. Peru Im BMB; upper Rio Maraildn Jan. Im AMNH. BOLIVIA (2m, 20: La Paz: Rio Songo Im ZMHU; Santa Cruz: Buenavista 750m Apr. -Aug. If BMNH; Prov. del Sara Im BMNH; Not located: no specific locality If BMNH. BRAZIL (10m, 60: Amazonas: Ega Im BMNH; SSo Paulo de Olivenpa Im ZMHU; Tonantins If BMNH; Maranhao: Mts. Aiireos If BMNH; Para: Para Jan.-Mar. Im BMB, 5m, 4f BMNH, Im MCZ; Villa Nova Im BMNH. GUYANA (2m): Cuyuni/Mazaruni: Cariamang River Im BMNH; Not located: no specific locality Im USNM. FRENCH GUIANA (11m, 60: Cayenne: Cayenne 2m BMNH; pk 27 Piste de Kaw, Roura Aug. Im LCB; Laurent du Maroni: Maroni Im BMNH; Maroni River Im AME; St. Jean du Maroni 2m, If BMNH; Not located: no specific locality 3m, 5f BMNH, 2m MNHN. Additional locality data: VENEZUELA; Panlepui; Ventuari; Imeri (Neild, 1996). FRENCH GUIANA: See type data for florea (from Brevignon, 1 995). Adelpha serpa celerio (H. W. Bates, 1864) Figs. 18e,f Heterochroa celerio Bates (1864: 127) TL: Polochic Valley, Guatemala. OTL: Guatemala; Upper and Lower Amazons. Types: BMNH(T); LTm: "Heterochroa celerio Bates m Syntype det. R.I. VaneWright 1983//lllustrated in The Butterflies of Venezuela A. Neild, 1996//179//B.M. TYPE No. Rh. 9845 Heterochroa celerio m Bates// B.C.A. Lep. Rhop. Adelpha celerio Bates Godman-Salvin Coll. 19I6.-4.//m//Polochic Valley F.D.G. and O.S.//Type H.T.//Syntype"; PLTf: "Heterochroa celerio Bates f Syntype det. R.I. Vane-Wright 1 983//Illustrated in The Butterflies of Venezuela A. Neild, 1996//I82//B.M. TYPE No. Rh. 9846 Heterochroa celerio f Bates// B.C.A. Lep. Rhop. Adelpha celerio Bates Godman-Salvin Coll. 1916.-4.//f//Polochic Valley F.D.G. and O.S.//Type H.T.//Heterochroa celerio n.s"; BMNH(M): PLTm: "m Ega//celerio[red label]//B.C.A. Lep. Rhop. Adelpha celerio Bates Godman-Salvin Coll. 1916.-4.//m//Ega U. Amazons H.W. BaXes"=diadochus; PLTf: "Para//celerio[red label]//B.C.A. Lep. Rhop. Adelpha celerio Bates Godman-Salvin Coll. 1916.-4.//f//Para L. Amazons H.W. Bales"=diadochus [all examined] =Adelpha diademeta Fruhstorfer (1913: pi. 107d; 1915: 530) TL: [Western Mexico and Orizaba]. Types: BMNH(T): STm: "Fruhstorfer Coll. B.M. 1937-285//celerio diademata Fruhst.//W. Mexico H. Fuhstorfer//TYPE//Type//Syntype"; STf: : "Fruhstorfer Coll. B.M. 1937-285//W. Mexico H. Fuhstorfer//TYPE//Type"; BMNH(R): STf: "serpa diademata Fruhst.//Orizaba//R" [all examined] =Adelpha phintias Fruhstorfer (1913: pi. 107c; 1915: 530) TL: [Merida, Venezuela]. Types: BIVINH(R): STf: "serpa phintias Fruhst.//Syntype//TYPE//Montan. Sierra Merida 3000m 8.7.98 (Briceno)//l 8 1 //Illustrated in The Butterflies of Venezuela A. Neild, 1996" [examined] Adelpha serpa var. celerio Bates, Kirby (1871); Adelpha cestus Hew., Boisduval (1870) ? misid. ?; Limenitis (Adelpha) paroeca Bates, Ross (1976) misid.; Adelpha celerio diademata Fruhst., Fruhstorfer (1915), de la Maza and de la Maza (1993), Vargas et al. (1996), Warren et al. (1998); Adelpha serpa celerio Bates, Willmott and Hall (1999) Identification, taxonomy and variation: Adelpha serpa celerio differs from A. serpa diadochus in having broader postdiscal bands which usually have a pale greenish tint and do not taper so much anteriorly, the VHW outer postdiscal series is roughly parallel to the postdiscal band and a darker reddish color, and the spots of the submarginal series are reduced in size, the

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152 inner series being almost equal in width to the outer, and clearly divided in each cell. Adelpha serpa duiliae differs by having narrower, even more bluish green postdiscai bands, which have more disjointed spots on the DFW, and a smaller orange subapical marking on the DFW. Adelpha seriphia godmani is very similar but has the blocks of the white postdiscai band on the DFW displaced more diagonally, the orange DFW subapical marking is orientated vertically rather than horizontally, the orange postdiscai band on the VHW is straighter and the inner submarginal series is not roughly parallel to the postdiscai orange band on the VHW but is much closer in cells Rs and Sc+Rl than throughout the remainder of the wing. Adelpha paroeca, A. paraena massilia and A. nea sentia all have the spots of the ventral submarginal series undivided in each cell. There is some individual variation in the width and color of the pale postdiscai bands, which vary from almost white to a greenish blue. The DFW orange subapical marking also varies slightly in width. Bates (1864) described celerio based on an unspecified number of male and female specimens from Guatemala, and stated that it also occurred on the upper and lower Amazon. Although he compared it to A. serpa, he sfrangely failed to mention the diagnostic undivided spots of the VHW submarginal series in that taxon. I have located four specimens in the BMNH that appear to be syntypes, and since two of these are from Brazil and represent a different taxon (diadochus) to the Guatemalan specimens, I believe a lectotype designation is necessary. Since the taxon was based first and foremost on Guatemalan specimens (later stated by Bates (1865) to have been captured by Salvin), and the name has been applied ever since to the Central American population, I designate the specimen in the BMNH(T) with the following label data as lectotype of Heterochroa celerio: "Heterochroa celerio Bates m Syntype det. R.I. VaneWright 1983//Illustrated in The Butterflies of Venezuela A. Neild, 1996//179//B.M. TYPE No. Rh. 9845 Heterochroa celerio m Bates// B.C.A. Lep. Rhop. Adelpha celerio Bates Godman-Salvin Coll. 1916.4.//m//Polochic Valley F.D.G. and O.S.//Type H.T.//Syntype". Fruhstorfer (1913)

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153 introduced the name phintias for a single Venezuelan specimen in the Tring Museum (now the BMNH(R)), later placing it as a subspecies of celerio" and stating that it differed from typical A. s. celerio in having a narrower DFW orange subapical marking (Fruhstorfer, 1915). However, this falls within the range of variation of A. s. celerio and phintias was synonymised with that taxon by Neild (1996). The two syntypes of Fruhstorfer's (1913) name diademeta (misspelt diademata by Fruhstorfer (1915) and placed as a subspecies of "A. celerio") are in the BMNH, and show little evidence of the broader postdiscal bands which Fruhstorfer (1915) claimed distinguished them from typical A. s. celerio. The name was synonymised with A. s. celerio by Hall (1938), but has been used by recent authors for Mexican A. serpa (de la Maza and de la Maza, 1993; Vargas et al, 1996; Warren et ah, 1998). However, there appear to be no consistent differences between west Mexican and remaining Mexican A. serpa and the name was syonymised with celerio by Willmott and Hall (1999). Adelpha s. celerio was first placed as a subspecies ("variety") of A. serpa by Kirby (1871), but has been treated usually as a frill species (Fruhstorfer, 1915; DeVries, 1987; Lamas and Small, 1992). However, the principal characters that distinguish the two taxa, the DFW postdiscal band and ventral submarginal series, are intermediate in form in the intervening A. s. diadochus, with which intergrades to A. s. celerio occasionally occur in Colombia. Range: Mexico to northwestern Venezuela, extending to the eastern foothills of the Cordillera de Merida. Immature stages: The eggs are deposited singly up to several times per plant in Costa Rica (DeVries, 1986). The larva and pupa of this taxon were figured and described by Comstock and Vazquez (1961), while Aiello (1984) figured the head capsule (fig. 2) and pupa (fig. 6) from Panama. DeVries (1986, 1987) also gives early stage information on this species from Costa Rica and figures the pupa (Fig. 24, F). The first and last of these records may also apply to several other members of the A. serpa group, especially A.

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154 seriphia godmani, but A. serpa celerio is the most plausible species. The following description is therefore taken from Aiello unless noted. The larvae are mostly black dorsally and green laterally, with pinkish lateral spots on A2-4, A7 and A8. The scoli on the thorax and A7 and A8 are pale, and that of A2 is black. The scolus on A2 is long, slender and sfraight, inclined slightly posteriorly, with sparse ascending spines. The subspiracular scoli on A2-4 are pale lime and bright green, and a black subspiracular stripe crosses the thorax, bends sharply in Al and joins with the scolus on A2. A few days before pupation the scoli on T3, A7 and A8, and the pinkish lateral marks, become green, and just prior to pupation the larva fades to a yellowish brown (lemon yellow with black scoli and a dark spot on the dorsum (DeVries, 1987)). The head capsule has a corona of long spines and vertical black stripes on the face. The pupa is shimmering silver with the sutures lined with black, the dorsal projection on A2 is small, and the head horns are long and only slightly curved near the tip. In Costa Rica, the larvae are solitary and feed on all leaves, and early instars make frass chains, while later instars rest on the upper surface of the leaf, on plants in direct sunlight in swampy areas near river or forest edges, or in forest clearings (DeVries, 1986). The larva rests in the curled position with the posterior end elevated, and resemble a piece of moss and lichen (Aiello, 1984) or a moss-covered twig (DeVries, 1987). Specimen label data indicate that a pupa in the PJD took 8 days to develop. Habitat and adult ecology: Adelpha serpa celerio is common in a wide variety of habitats, from wet to moist rain forest and dry forest (DeVries, 1987; de la Maza and de la Maza, 1993). It has been recorded from sea level to 1700m throughout its range (specimen data; de la Maza and de la Maza, 1993), and specimen label data indicate that it flies throughout the year. With Jason Hall 1 have observed females flying along forest edges in Mexico, while Ross (1976) reports it to be common to abundant in the Sierra de Tuxtla, Mexico, along forest margins and in pastures. DeVries (1987) notes that the

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155 species is attracted to rotting finits but does not specify the sex. Srygley and Chai (1990) found this subspecies to be palatable to jacamars in Costa Rica. Specimens examined (210m, 225f); t trans, to A. s. diadochus. MEXICO (83m, 95f): Chiapas: Campet Im AME; Cuauhtemoc 700m Feb. Im AME; La Granja If AMNH; Las Delicias Sep. If AME; Mapastepec Aug. If AME; Tapachula Jan. Im FSCA; San Jeronimo Mar. Sep. Nov. 3m AMNH, Im, If AME; San Jose del Camnen Apr. Im FSCA; no specific locality 3m, If AMNH; Colima: Colima 2f AMNH; Guerrero: Acahuizotla Aug. Sep. 2m, 7f AME; El Treinte Sep. If AMNH; Rincon 2800' If BMNH; no specific locality 7m, 5f BMNH; Hidalgo: Pisaflores Aug. If AMNH; Jalisco: San Sebastian If BMNH; Michoacan: Playa Azul Aug. Im FSCA; Nayarit: Jalisco Sep. If AMNH; Oaxaca: Candelaria Loxicha Aug. -Oct. Im, 2f AME, 4f AMNH; Chiltepec Aug. Im, If AME; Chimalapa Oct. Im AME; El Naranjal-Chiltepec May If AMNH; Mixtepec Dec. If AMNH; Oaxaca Jun. Aug. 4m, 2f AME, Im BMNH; nr. Puerto Eligio 700m Jul. Im KWJH; Tuxtepec Aug. If AMNH; Soyolapan el Bajo Oct. 4m, 6f AMNH; Valle Nacional 800m Jul. Im MUSM; Puebla: Tequelquitla 625m Aug. Sep. 2m, If MUSM; Xicotepec de initcz If MNHN; San Luis Potosi: Arroyo Seco Oct. If AMNH; Ciudad Valles 2m AMNH, If FSCA; El Salto Falls 860m Feb. Aug. Nov. If USNM, Im FSCA, 3m, If AME; Palitla Dec. 3f AMNH; Picolco May If AMNH; San Mateo Yetla May Im USNM; Tamazunchale Aug. Dec. 2f AMNH, If FSCA; Xilitla If AMNH; Tabasco: Tepescuintla Aug. 3m, 3f AMNH; Tamaulipas: Mante, 30 mi. N., Nov. Im USNM; Veracruz: Catemaco Jul.-Sep. Im, If FSCA, Im, 2f AME; Cordoba Sep. 2m, If AMNH; Coatepec If USNM; Cordoba Jun. Jul. Nov. 2m, 5f BMNH, 2f BMB, If MUSM, If MCZ; Dos Amates Aug. Im, 4f AME; Los Tuxtlas 200m Jul. Im KWJH; Fortin Im ZMHU, Im FSCA; Jalapa Im, If USNM, 2m AMNH; Minatitlan Jul. Im AMNH; Misantla Im, If BMNH, If BMB; Nanchita sea level Feb. Im AME; Popoctepetl, Tuxtla 400m Oct. 3m MUSM; Presidio Jul.-Sep. 5m, 2f AME, 4m, If AMNH; nr. Sontecomapan 300m Jul. 2f KWJH; Tezonapa Jun. Nov. If AMNH, Im AME; Not located: W. Mexico Im, If BMNH; no specific locality Im, 3f USNM, 2m BMNH, If BMB, 2m, 2f MNHN, 2f AMNH. GUATEMALA (31m, 200: Alta Verapaz: Baleu 1350m Jun. Aug. 2m, 2f AMNH, Im, 2f AME, Im MUSM; Chiacam 2m, If BMNH; Choctun Im BMNH; La Vega del Cuajilote 750m Aug. If AME; Polochic Valley 3m BMNH; Tamahu 1 100m Nov. 2m AMNH; Baja Verapaz: Chuaciis 1 m BMNH; Panima 1 f BMNH; San Jer6nimo 1 f BMNH; Chiquimula: no specific locality 600m 1 f MNHN; El Peten: Sayaxche Sep. Oct. 2m, 2f AMNH; Escuintla: Escuintla Jul. Aug. 2m, If BMB; Palin Oct. If USNM, 2m BMB; Izabal: Cayuga Im USNM; Puerto Barrios Dec. Im, If BMB; Quirigua 4-5000' Feb. Im BMB; Tenedores Nov. Im AME; Retalhuleu: Retalhuleu Im BMNH; San Sebastian 2f USNM; Santa Rosa: Barbarena Im AME; Guazacapan 4m BMNH, If BMB; Suchitepequez: Mazatenango Jul. Aug. If BMB; Not located: Central valleys If BMNH; no specific locality If ZMHU, Im BMNH, Im MNHN. BELIZE (3m, 3f): Stann Creek: Stann Creek 125m Sep. Oct. 2m, 2f FSCA; Not located: Hummingbird Hwy. Nov. Im AME; Rio Grande If AMNH. EL SALVADOR (3m, 6f): San Salvador: Ilopango 500m Feb. ex larva Melastomaceae If AME; San Salvador Jan. 2f AME; Santa Tecla Oct. If USNM; Santa Ana: Cerro San Jacinto 800m Dec. 2m AME; Citala 800m Oct. If AME; Usulutan: San Augustfn 3500' Sep. Oct. If BMNH, Im BMB. HONDURAS (3m, 90: Comayagua: Comayagua Jun. 2f MCZ; Cortes: San Pedro Sula Dec. 4f BMNH, If USNM; Not located: La Cambre Feb. Im BMNH; Petuc If AMNH; no specific locality Im, If ZMHU, Im BMNH. NICARAGUA (3m, 40: Chontales: Chontales 2m, 2f BMNH; Zelaya: San Ramon 375' Jul. If BMNH; Not located: no specific locality Im, If BMNH. COSTA RICA (12m, 190: Alajuela: Cariblanco If BMNH; Esperanza Im USNM; San Mateo 1-2000' Sep. Nov. Dec. 2m, If USNM, 2f BMNH; Cartago: Juan Villas 2500-3000' Nov. Jan. If USNM, If BMB; Volcan Irazii 6-7000' If BMNH; Heredia: Chilamate 37m Dec. 2f FSCA; Limon: Gu4piles Jun. If FSCA; Puntarenas: Esquinas Aug. If USNM; Palmar Norte Aug. Im USNM; Caldera 250' Oct. Im BMB; San Jose: Carrillo Sep. Oct. 2m BMNH, 2f BMB; San Jose If BMNH; Not located: no specific locality 3m, 2f BMNH, 2m BMB, Im, 2f AMNH, Im, If MNHN. PANAMA (26m, 250: Canal Zone: Aiello Lot 82-41 Im, Lot 84-10 no.2 If, Lot 93-90 If STRl; Barro Colorado Island Feb. Im AMNH; Empire Feb. Im BMB; Lion Hill Im BMNH; Pipeline rd. If STRI; Chiriqui: Bugaba 800-1500" 2m, 2f BMNH; Chiriqui Im, If ZMHU, 2m BMNH, 3m BMB; David If BMNH; V. de Chiriqui 25-4000' 2m BMNH; Potrerillos 3600' Feb. Mar. Im, 4f USNM; RioToIe 120m Jan. 2f USNM; Santa Clara 1200m Jun. Im USNM; Code: El Valle Dec. If BMNH; Colon: Pifla 100m Apr. If FSCA; Panama: Cerro Campana 2500' Jul. Aug. Nov. Im, 2f USNM, If STRI; Cerro Jefe 900m Mar. Im USNM; Rio Bayano Oct. Nov. 2m USNM; Vique Pt. Feb. If BMNH; Veraguas: Santa Fe 230m Oct. Im USNM; no specific locality Im, 2f BMNH; Not located: Isthmus of Panama 3m, If BMNH; no specific locality lm,lf BMNH, 2f AMNH. COLOMBIA (37m, 220: Antioquia: Rio Cocoma 1000m Aug. Im AME; Boyaca: Muzo Jun. Jul. 2m BMB; Otanche Feb. Aug. Dec. 2m JFL, 2f ESM; Caldas: Quebrada El Aguila, Manizales 1700m Aug. If MHNM; Victoria Mar. Im ESM; Cauca: Pescador 1450m Feb. Dec. Im, 2f AME; Cundinamarca: Bogota If USNM, 4m(lt) BMNH, If BMB, 2f MCZ; env. Bogota 2m, 3ft BMNH; El Baldio 5400' Sep. Imf BMB; Veragua 2m MNHN; El Cesar: Manaure Imf BMNH; Risaralda: Hda. Bacori, Pblo. Rico 450m Jul. Im MHNM; Santander: Landaruri Jan. Im AMNH; La Borroscosa Nov. Im AMNH; Tolima: Honda If BMNH; Valle del Cauca: Cali 1000m Dec. If MUSM; Cali, Club Campestre 1000-1 100m Apr. Jul. Im, If LMC; El Cerrito Apr. Im LMC; Pance 1200m May Jun. 2m LMC; Yatacue, Alto Anchicaya Sep. If LMC; Zacarias Apr. Im LMC; Not located: east Colombia If USNM; no specific locality 8m, 2f AMNH, Im, If ZMHU, 2f BMNH, 2mt BMB, Im MCZ. VENEZUELA (4m, XA^): Aragua: Portochuelo 1 100m Aug. If MUSM; Puerto Cabello If ZMHU; Merida: Merida ImBMNH, Im ZMHU; Montan. Sierra 3000m Aug. If BMNH; Mucutin 2600m 2mt BMNH, 2f BMB; Tachira: Hda. Panaga 800m Feb. 7f AFEN; no specific locality If AFEN; Not located: no specific locality If USNM. "BRAZIL" (10: Santa Catharina: no specific locality If BMB-error. COUNTRY UNKNOWN (5m, 70: Central America Im AMNH; no specific locality If BMNH, If BMB, If MNHN, 4m, 3fMCZ, If AMNH. Additional locality data: MEXICO: Chiapas: throughout see de la Maza and de la Maza (1993); Colima: Cofradia de Suchitlan 130m Nov. (Warren et at., 1998); Jalisco: nr. Barra de Navidad Dec; La Calera Apr.; Est. Cientifica Las Joyas Sep.; Mismaloya Jan. Dec; Pihuamo Sep.; Puerto Vallarta Oct.; 53 km S. Puerto Vallarta Jan. (Vargas et ai, 1996). BELIZE: Cayo: Caracol, Slate Creek Preserve (Meerman, 1999).

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156 Adelpha serpa duiliae Fruhstorfer, 1913 Figs. 18g,h Adelpha rfwi/iae Fruhstorfer (1913: pi. 107d; 1915: 531) TL: [Chimbo and Paramba, Ecuador]. Types: BMNH(T): STm: "Fruhstorfer Coll. B.M. 1937-285//celerio duiliae Fruhst.//Chimbo 1000' VII 97 (Rosenberg)//TYPE//Type//Syntype"; BIVINH(R): STm: "celerio duiliae Fruhst.//Paramba 3500" 01. 97 dry season (Rosenberg)//Syiitype//TYPE" [both examined] Adelpha celerio duiliae Fruhst., Fruhstorfer (1915); Adelpha serpa duiliae Fruhst., Hall, 1938 Identification, taxonomy and variation: Adelpha serpa duiliae is distinguished from A. s. celerio by having postdiscal bands which are narrower, composed of more isolated spots on the DFW and typically more deeply tinted blue-green, in addition to a narrower DFW orange subapical marking which is dissected by more heavily marked veins. It is very similar to the sympatric A. seriphia godmani, but that species is distinguished by having even more displaced spots forming the DFW postdiscal band, the orange DFW subapical marking orientated more vertically (notably the basal edge), and the inner submarginal series on the VHW is not parallel to the orange postdiscal band but much closer in cells Rs and Sc+Rl. Fruhstorfer (1913) clearly figured duiliae and later described it as a subspecies of "A. celerio" based on specimens from Chimbo and Paramba (Fruhstorfer, 1915), and there are two syntypes in the BMNH. Hall (1938) placed it as a subspecies of A. serpa. Range: Western Ecuador, as far south as Bolivar Province, possibly also occurring in extreme southwestern Colombia. Habitat and adult ecology: This subspecies is local but not uncommon from near sea level to 1400m in large clearings or secondary growth areas in the vicinity of primary or secondary forest. Males may be encountered in the afternoon perching singly with the wings closed on small bushes, 2-3m high, in open areas along rivers, or from 4-8m high on ridgetops. Males are very wary when perching, and if disturbed will take flight to another perch, often up to 30m away. Females are usually found in areas of secondary growth along forest edges or even roadsides throughout the middle of the day. The thinning of the dorsal postdiscal band, its greenish coloration and the reduction of orange

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157 subapical DFW patch in this subspecies parallels similar features in the sympatric A. iphiclus estrecha, suggesting that the two may be involved in a mimetic relationship. Specimens examined (7m, 70: ECUADOR (7m, 7f): Bolivar. Chimbo 1000' Aug. Im BMNH; Carchi: Las Juntas, nr. La Carolina 1400m Nov. Im K.WJH; Lita, ridge east of Rio Baboso 900m Jul. 2m, If KWJH; Esmeraldas: Est. Ex. U Chiquita 50m Dec. Im KWJH; Imbabura: Paramba 3500' Mar. Im BMNH; Los Rios. U Chima Jun. Jul. If BMNH; Pichincha: Quito-Sto. Domingo old rd. km. 85, 4400' Jul. If FSCA; Rio Sucio, nr. San Miguel de los Bancos Apr. Im KWJH; Salto de Napac, km. 35 Sto. Domingo-Quito rd. 1000m Jul. If KWJH; Tinalandia Jun. If FSCA; Not located. Oriente If AMNH-CTror; Rio Ayada vie. Dec. If USNM-error?. Adelpha seriphia (C. and R. Felder, 1867) Figs. 19; 103; 186 Identification, taxonomy and variation: Adelpha seriphia is distinguished from all related species by the VHW inner submarginal series not being parallel to the orange postdiscal band, but noticeably closer in cells Rs and Sc+Rl, in all subspecies except for A. s. barcanti. In comparison with sympatric subspecies of ^. serpa, the blocks that compose the DFW postdiscal band are always more displaced diagonally, and the DFW orange subapical marking is orientated more vertically. Certain subspecies of A. hyas are very similar, but are smaller, and have the base of the VFW costa white rather than orange. There is much variation in the width of the postdiscal bands, the form of the orange dorsal postdiscal markings and the ventral submarginal series, and seven subspecies are recognised, one of which is described here. The systematic status of this species has always been imcertain, largely due to the rarity of specimens in collections, and many authors have regarded the various taxa as forms of A. serpa. The two are sometimes similar in sympatric populations, although A. seriphia tends to be found at higher altitudes and therefore sometimes develops a completely distinct dorsal pattern (e.g. Adelpha s pione) due to mimicry with other locally occurring species. Fruhstorfer (1915) recognised A. seriphia as a distinct species almost as treated here, with the exception of then undescribed taxa and the Central American A. s. godmani, which was finally correctly placed by Neild (1996) (as godmani). The shape of the VFW subapical marking and its tendency to have darker intruding lines, in addition

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158 to the displaced blocks of the DFW postdiscal band, suggest that A. seriphia is probably most closely related to A. hyas, which occurs at lower altitudes in the eastern Andes. Most subspecies of A. seriphia are still poorly represented in collections, and, particularly in Colombia, accurate locality data are often lacking. Further collecting may well demonstrate certain subspecies recognised here to be composed of more than one taxon, or conversely, to be merely part of clinal variation. Range and status: Mexico to western Ecuador, Venezuela to Bolivia, Trinidad. Usually in montane habitats, everywhere local and imcommon to rare, several subspecies with restricted and isolated ranges. Specimens examined: 199 (96m, 103f) Adelpha seriphia seriphia (C. and R. Felder, 1867) Figs. 19a,b Heterochroa seriphia C. and R. Felder (1867: 423) TL: Venezuela. OTL: Caracas, Venezuela; Bogoti, Nova Granada. Types: BMNH(R): LTf: "Lectotype//type//Felder Colln.//Venezuela Moritz/Zseriphia Feld.//seriphia Felder/ZIllustrated in The Butterflies of Venezuela A. Neild, 1996//168//Heterochroa seriphia C. and R. Felder, lectotype f det. R.l. VaneWright 1983 see Hall, 1938: 284"; PLT?f: "Heterochroa seriphia C. and R. Felder, Tparalectotype f det. R.l. Vane-Wright I983/A'enezuela//Felder Colin." Px)th examined] Adelpha serpa var. seriphia Fldr., Kirby (1 871); Adelpha seriphia seriphia Fldr., Neild (1996) Identification, taxonomy and variation: The nominate subspecies is similar to A. seriphia pione, from which it differs in having broader postdiscal bands and reduced (occasionally absent) orange postdiscal dashes on the dorsal surface. Adelpha s. aquillia also has narrower postdiscal bands and typically lacks any orange dorsal postdiscal dashes. Adelpha seriphia seriphia is similar to A. serpa celerio and A. serpa diadochus, but may be distinguished as detailed under the account for the species as a whole. The orange postdiscal dashes on the DHW are somewhat variable, and may be absent altogether, particularly in males. Felder and Felder (1867) described this species as being similar to A. serpa, except for an orange postdiscal band on the dorsal surface extending to the anal margin. They did not specify the nimiber or sex of specimens on which the description was based,

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159 only that the specimens came from Caracas, collected by D. Moritz, and Bogota, collected by Lindig, and were in their collection. Since specimens from Caracas and Bogota are referable to separate subspecies, I designate a female specimen in the BMNH(R) with the following label data as the lectotype of Heterochroa seriphia: "Lectotype//type//Felder Colln./A^enezuela Moritz/Zseriphia Feld.//seriphia Felder/ZIUustrated in The Butterflies of Venezuela A. Neild, 1996//168//Heterochroa seriphia C. and R. Felder, lectotype f det. R.I. VaneWright 1983 see Hall, 1938: 284". This specimen is figured by Neild (1996: pi. 4, fig. 169) and labeled "Lectotype", but this was due to a label attached to the specimen indicating it to be the lectotype, whereas in fact no such designation has been published previously. The nominate subspecies is very similar, and very occasionally indistinguishable from, A. s. aquillia, but the latter is retained here since it is usually readily identified and is separated from the nominate by the distinctive and geographically intervening subspecies A. s. pione. Range: The nominate subspecies occurs in the Cordillera de la Costa in northern Venezuela, and may also be found in the Sierra de Turimiquire. Habitat and adult ecology: Little has been reported on the habitat or behavior of this subspecies, which is rare in collections and represented predominantly by females. The available data indicate that it occurs from 900-1 100m and has been collected in January, June, September and October. Specimens examined (3m, 9f): VENEZUELA (3m, 60: Aragua: Rancho Grande llOOm Jan. Jun. If AMNH, Im USNM; Dislrito Federal: Caracas 3000' Sep. Oct. Im AME, If BMB; Not located: no specific locality Im, 4f BMNH. COUNTRY UNKNOWN (30: no specific locality 3f BMNH. Adelpha seriphia pione Godman and Salvin, 1 884 Figs. 19c,d Adelpha pione Godman and Salvin (1884: 309, Tab. XXIX, figs. 16, 17) TL: Chiriqui, Panama. Types: ZMHU: HTf: "Chiriqui7/Origin//AdeIpha pione type/Zholotype" [examined] Adelpha seriphia pione G. and S., Fruhstorfer (1915); Adelpha serpa pione G. and S., Hall (1938); Adelpha seriphia pione G. and S., Lamas and Small (1992)

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160 Identification, taxonomy and variation: Adelpha seriphia pione is distinguished from the nominate subspecies by the broader orange postdiscal markings on the dorsal surface and the narrower white postdiscal band. It is also very similar to A. s. therasia, but differs in having the spots of the inner submarginal series on the VHW less clearly divided by dark rays in each cell. There is some variation in the thickness of the dorsal orange postdiscal markings, particularly in specimens labeled "Bogota", which are probably transitional to A. s. auillia. See also the discussion under A. s. aquillia concerning the paralectotype of aquillia from Ocana, Colombia. Godman and Salvin (1884) described and clearly figured pione based on a single female in Staudinger's collection from "Chiriqui" in Panama. I have examined the holotype in the ZMHU. The specimen is clearly, however, very similar to certain specimens from Colombia and Venezuela, and I conclude it is mislabeled and actually originated in one of those countries. This belief is founded on the occurrence in Panama of A. seriphia godmani, the lack of any other specimens of pione in collections from Panama, and the lack of Adelpha in Panama with similar dorsal patterns to pione, the pattern of which is probably mimetic on that of the Andean A. corcyra corcyra and A. alala completa. Fruhstorfer (1915) first placed pione as a subspecies of A. seriphia, but Hall (1938) regarded it as a subspecies of A. serpa, in which he was followed by D'Abrera (1987), until Lamas and Small (1992) restored it to Fruhstorfer's original combination. Range: Adelpha seriphia pione occurs in Venezuela in the Cordilleras de Merida and Tama, to the Colombian Cordillera Oriental north of Bogota. Neild (1996) reports the subspecies in Venezuela from the Sierra de Perija, based on a single specimen collected at El Tucuco, at 400m. This is a most unlikely altitude for this subspecies, suggesting that the specimen may be mislabeled, although it could also have been collected higher up from this site. At the present I regard its presence in the Sierra de Perija as requiring

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161 confirmation, especially in view of the paralectotype specimen of A. s. aquillia (see discussion imder that taxon). Habitat and adult ecology: This subspecies is not uncommon, and has been recorded from 900-3000m in montane forest habitats. Neild (1996) reports that it might occur down to near sea level, though he may be referring to the nominate subspecies. Pyrcz reports (in Neild, 1996) that the female is attracted to rotting fruit baits, while I have observed a male puddling at damp sand along a stream below the town of Merida. The subspecies seems quite tolerant of habitat disturbance, Pyrcz's observations (in Neild, 1996) of a number of individuals at a flowering tree were made in the grounds of the Facultad de Ciencias Naturales in Merida. The range of this subspecies closely corresponds with that of A. alala completa and A. corcyra corcyra, which also occur in similar montane habitats, and the development of orange dorsal postdiscal bands in each of these three subspecies of relatively unrelated species is strong evidence of a mimetic relationship. Specimens examined (29m, 261): "HONDURAS" (Im): Cortes: San Pedro Sula Im BMB-error. "PANAMA" (Im): Chiriqui: Chiriqui Im ZMHU-error. COLOMBIA (14m, 120: Boyaca: Otanche 900m Aug. Im MHNM; Cundinamarca: Bogota Im USNM, 7m, 6f BMNH; Santander: Ocafla Im ZMHU; Pamplona-Sarare 2200m Mar. Im ESM; Virolin 2300m Mar. 2m JFL; Not located: Interior If BMB; no specific locality If USNM, 3f BMNH, 2m BMB, If AMNH. VENEZUELA (13m, 130: Merida: Culata Im BMNH; Escorial Jan. Im BMNH; Merida 1400m Aug. Im KWJH, 3m, If BMNH; Montan. Japi 2-3000m Sep. If BMNH; Montan. Sierra 3000m Jul. Im BMNH; Mucutin 2600m Im BMNH, Im BMB; Pedregosa 3000m Oct. 3f BMNH; Quebrada Campada, km. 39 Barinitas-Las Uvas Jan. Im MUSM; Quebrada Las Uvas, km. 34 Barinitas-Apartaderos Jan. If MUSM; Tachira: Barinitas-Sto. Domingo If AFEN; San Vicente de la Revancha 2350m 3f AFEN; A^or /ocaterf: no specific locality Im, 2f BMNH, 2m, If BMB. COUNTRY UNKNOWN (10: no specific locality If MCZ. Adelpha seriphia aquillia Fruhstorfer, 1915 Figs. 19e,f Adelpha seriphia aquillia Fruhstorfer (1915: 531) TL: Bogota, Colombia. OTL: No locality stated; Ocafia, Colombia. Types: BMNH(T): LTm: "Fruhstorfer Coll. B.M. 1937285//seriphia aquillia Fruhst./ZBogota Colombia//TYPE//Type//Syntype?"; ZMHU: PLTm: "Ocafia Columb. Kalbr.//Syntype//SYNTYPE m Adelpha seriphia aquillia Fruhstorfer G. Lamas det. 87// Adelpha spec. S. and G. [?illegible] ign." = pione [examined] =Adelpha seriphia naryce Fruhstorfer (1915: 531) TL: Chanchamayo, Peru. Types: ZMHU: STf "Chanchamayo Thamni//Syntype//Seriphia var.//SYNTYPE m Adelpha seriphia naryce Fruhstorfer G. Lamas det. '87" [examined] =Adelpha serpa celerio Bates, Hall (1938); Adelpha seriphia aquillia Fruhst., Neild (1996)

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162 Identification, taxonomy and variation: Adelpha s. aquillia is distinguished from A. s. pione and A. s. therasia by the lack of orange postdiscal markings on the DHW, and only faint or absent orange postdiscal dashes on the DFW in cells Cu2-M3. It is very similar to A. s. godmani, but has more red coloration in the DFW discal cell and thinner postdiscal bands on the dorsal surface which tend to be of even width; in A. s godmani the blocks of the postdiscal band on the DFW decrease in width from the anal margin to cell M3. The VFW pale subapical marking in A. s. aquillia has dark red-brown lines intruding in each cell, or expanding to color most of the marking red-brown, while in A. s. godmani the marking lacks the intruding lines and is therefore paler. Adelpha s. aquillia is also very similar to A. hyas hewitsoni, but has thinner white postdiscal bands on the dorsal surface, the base of the VFW costa is orange instead of white, and on the VHW the orange band of the outer postdiscal series is sfraight rather than convex and the markings of the inner submarginal series are wider. Adelpha serpa diadochus may be distinguished by the much broader dorsal postdiscal bands and by not having dark lines intruding into the VFW pale subapical marking, in addition to other characters which distinguish A. serpa from A. seriphia. The width of the white postdiscal bands may vary slightly, as does the shape of the hindwing band, and the width of the DFW orange subapical marking is particularly variable, being very narrow in certain Colombia specimens from the Cordillera Cenfral. I have been unable to discern any consistent geographic variation in the limited series examined, but it may well be the case that fiiture collecting will demonsfrate such variation to exist. Fruhstorfer (1915) described aquillia based on a male in his collection "with reduced reddish-brown band of the forewings" in comparison with A. s. seriphia, but did not state where the specimen was collected. He also stated that the name applied to "some more specimens from Ocana (Colombia) of the coll. Staudinger in the Berlin Museum". A male specimen in the BMNH(T) from Fruhstorfer' s collection bears the handwritten

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' 163 label "seriphia aquillia Fruhst." and is from "Bogota". This supposed syntype differs from the only specimen of A. seriphia that I have been able to locate in Staudinger's collection at the ZMHU from Ocana in lacking any orange postdiscal dashes on the dorsal surface between cell M3 of the DFW to cell Cul of the DHW; the ZMHU syntype has thin, isolated orange dashes in each cell similar to nominate A. seriphia. I believe that Fruhstorfer was referring to the entire postdiscal series on the DFW as his "reddish-brown band", in which case both the BMNH specimen, which lacks the markings in cells M3 to Cu2, and the ZMHU specimen, which has slight orange markings in these cells, fit this description. I therefore regard both of these specimens as valid syntypes. Two lines later Fruhstorfer (1915) described an unspecified number of specimens from Chanchamayo as A. seriphia naryce, which he stated resembled aquillia on the dorsal surface, but were similar on the ventral surface to a third new subspecies, therasia (see below). A female specimen of ^4. seriphia in the ZMHU from Chanchamayo is probably the specimen on which Fruhstorfer based his description of naryce, and closely matches the original description. It differs from A. s. aquillia and A. s. pione, and is similar to A. s therasia, by having on the ventral surface broader submarginal series, the spots of which are less sharply defined, paler, and merge with the forewing subapical marking. This specimen has slight orange postdiscal scaling on the DFW in cells M3-Cu2, but otherwise closely resembles females from northern Peru to southeastern Colombia. Since I have seen no other specimens of A. seriphia from central Peru, for the present I regard the putative syntype female of naryce as representing the same taxon as specimens from northern Peru to southeastern Colombia, although it may prove to be a distinct taxon. The remaining problem is whether to regard naryce or aquillia as the name which should be applied to this population. The Ocana syntype of aquillia is something of a taxonomic mystery, since it originates from the northern tip of the Cordillera Oriental at the northern edge of the range of ^. seriphia pione. It may represent either a specimen of, or an intergrade to, an undescribed subspecies fiirther north in the Sierra de Perija, since no subspecies oiA.

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164 seriphia has definitely been recorded there to date. Possibly it may also be an aberration of typical A. s. pione. Since the Bogota syntype of aquillia occurs at the southern Umit of the range of .4. s. pione, it must represent a different taxon to the Ocana syntype. Given that the Bogota syntype is clearly referable to the subspecies that occurs throughout central Colombia to the eastern Andes, and the lack of information concerning the taxonomic status of the Ocaiia specimen, I regard the latter for the present as a specimen of A. pione and designate the former as the lectotype of Adelpha seriphia aquillia, with the following label data: "Fruhstorfer Coll. B.M. 1937-285//seriphia aquiUia Fruhst.//Bogota Colombia//TYPE//Type//Syntype?". Since specimens from north Peru to southeastern Colombia clearly predominate in collections I choose to apply the name aquillia to this population in the interests of nomenclatural stability, and regard naryce as a synonym for the present, as did Willmott and Hall (1999); should naryce apply to a distinct subspecies the name is still available for use. Finally, on a historical note, it must be mentioned that Hall (1938) synonymised aquillia with A. serpa celerio, and naryce with A. serpa duiliae, while Neild (1996) resurrected aquillia as a subspecies of A. seriphia. Range: Adelpha seriphia aquillia appears to occur in Colombia in the Cordillera Oriental south of Bogota, probably on both slopes, on both slopes of the Cordillera Central and the eastern slope of the Cordillera Occidental, to central Peru on the east Andean slopes. More specimens are needed to accurately determine the range within Colombia. Habitat and adult ecology: This subspecies is rare in collections and in the field it is very local, but although females are typically found as solitary individuals in areas of secondary growth, when males are encountered they are usually found perching in small groups. In eastern Ecuador I have observed males on a wide path cleared along the crest of a forested ridge, perching at a small rise in periods of bright sun on the tops of 5-6m high bushes with their wings shut. From this perch they made frequent flights to investigate passing butterflies, and the same perch was utilised over a period of several

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165 months. Males are also attracted to traps baited with rotting fish. The subspecies is restricted to relatively intact cloud forest habitats from 1000-2000m, where it probably flies throughout the year. Specimens examined (22m, 1 7f): t trans. toA. s. godmani; t trans, to A. s. pione. VENEZUELA (20: Merida: Merida 2ft ZMHU-error?. COLOMBIA (16m, lOf): Boyacd: Otanche Apr. If JFL; Caldas. El Aguila 1700m Oct. Imt, 1ft KWJH; Pensilvania 1000m May Im JFL; Cundinamarca. Aguaditas, Fusagasuga Mar. 2f ESM; Bogoti 3m BMNH, Im USNM, Im BMB; Nariiio: Monopamba 1750m Feb. If ESM(resembles nominate, thin orange thread DHW mislabeled?); Risaralda: Pereira Im BMNH; Tolima: San Antonio 2000m Im BMNH; Valle del Cauca: Rio Aguacatal 2000m Im, If BMNH; Not located, no specific locality 2m, If AMNH, Im, 2f(lt) ZMHU, 2m, If BMNH. ECUADOR (4m, 3f): MoronaSantiago. Rio Abanico 1600m Dec. Im KWJH; Sucumbios: Rio Sucio, nr. U Bonita 1800m Nov. If KWJH; Tungurahua: Chinchin 1800m Nov. Im KWJH; Rio Verde 1600m Jun. If AMNH; Not located: Oriente 2m AMNH; "Tena" purchased If DAT. PERU (Im, 21): Amazonas: Alfonso Ugarte 1000-1200m Jul. Im MUSM; Cajamarca: Charape, N. Peru If BMNH; Jmin. Chanchamayo If ZMHU. COUNTRY UNKNOWN (Im): no specific locality Im BMNH. Additional locality data: ECUADOR: Morona-Santiago: Rio Abanico Oct. Nov. (Willmott and Hall, unpubl. sight records). Adelpha seriphia godmani Fruhstorfer, 1915 Figs. 19g,h; 103; 186a,b Adelpha godmani VrvihsioxfQr {\9\?,: pi. 107d; 1915: 530) TL: [Orizaba]. Types: BMNH(R): STm: "celerio fa. godmani Fruhst.//Syntype//TYPE//R//Orizaba" [examined] =Adelpha celerio celerio form syma Fruhstorfer (1915: 530; fig. as A. massilia. In: Godman and Salvin (1884: Tab. XXDC, fig. 7, 8)) TL: not stated. Types: BMNH(M); STm: "Syntype m Adelpha celerio f. syma Fruhstorfer G. Lamas det. 1987//Fig. 14 G.D.H.C.//2//Genita!ia slide No. 16-1 12 godmani Fruhst.//B.C.A. Lep. Rhop. Adelpha massilia Feld. Godman-Salvin Coll. 1916.4.//m//C6rdova Mexico H. Rumeli/ZSyntype" [examined] Adelpha massilia Fldr., Godman and Salvin (1884) misid.; Adelpha celerio diademala form godmani Fruhst., Fruhstorfer (1915); =Adelpha serpa celerio Bates, Hall (1938); Adelpha syma Fruhst., Steinhauser (1974); Adelpha serpa godmani Fruhst., D'Abrera (1987), Salazar (1996); Adelpha seriphia godmani Fruhst., Neild (1996) Identification, taxonomy and variation: Adelpha seriphia godmani is most similar to A. s. aquillia, and is distinguished under that subspecies. It is distinguished fi^om A. serpa celerio and A. serpa duiliae through the configuration of the VHW inner submarginal series that is diagnostic of the species (see under species account), the more diagonally displaced blocks of the DFW postdiscal band, and the more vertical orange subapical marking on the DFW. There is a little variation in the width of the orange DFW subapical marking, and in one female fi-om Cordova, Mexico, there is a slight orange spot in cell M3 of the DFW. The width of the postdiscal bands is quite constant. Four female specimens in the AME fi-om Malinalco, Valle de Bravo and Chalma in Mexico differ fi-om typical A. s godmani in having pure white postdiscal bands which are slightly broader and less disjointed between

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166 cells Cu2 and Cul, the DFW orange subapical marking is entirely orange, without being dissected by darker veins, and extends slightly into cell M3. These specimens may well represent an undescribed subspecies, as they are reasonably isolated from remaining A. seriphia godmani, but I have seen insufficient specimens given the slight differences to warrant their description at the present. Specimens from westem Ecuador have, on the DFW, the postdiscal band and orange subapical marking notably narrower than typical A. s. godmani, but these differences do not seem sufficiently stable to justify separate subspecific status with the material available. Fruhstorfer (1913) figured godmani, and subsequently described it (Fruhstorfer, 1915: 530), as a form of "A. celerio diademata" (=A. serpa celerid), based on a single male from Orizaba in the Tring Museum. The figure corresponds well with the unique syntype (note that no indication was given of number of specimens in the original description by Fruhstorfer (1913)) in the BMNH(R), and even though the name as proposed in the text (1915) is a quadrinomial, since it appeared singly on the plate two years earlier in 1913 it should be freated as a specific and therefore available name. Shortly after, Fruhstorfer (1915) introduced the name syma for "a melanotic form" of celerio celerid" which was figured as A. massilia by Godman and Salvin (1884) on Tab. XXIX, figs. 7 and 8, supposedly differing from godmani in having a narrower orange DFW subapical marking. Godman and Salvin (1884) state that their specimens were obtained by Hoge and Riimeli in southern Mexico, and a syntype specimen matching their figure with appropriate label data is in the BMNH. In fact, the orange subapical marking in the original figure is represented as narrower than reality, and since the differences between the types ofsyrna and godmani fall within the normal bounds of variation in the subspecies, I follow Willmott and Hall (1999) in regarding the names as synonymous. Although the name syma was described as a quadrinomial and therefore would unavailable, Steinhauser (1974) used it as a species name and therefore made the name available with the original author and date of publication.

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167 This subspecies is very similar to A. serpa celerio, and indeed was synonymised with that taxon by Hall (1938). It is the only subspecies of A. seriphia known at that time which Fruhstorfer (1915) failed to associate with other ^. seriphia taxa. Nevertheless, it has all the diagnostic characters of the species (see species discussion) and was first placed correctly as a subspecies of A. seriphia, under the name godmani, by Neild (1996). Range: This subspecies occurs from Mexico to western Ecuador, apparently extending into the Cauca Valley of Colombia and the northern tip of the Cordillera Central. Habitat and adult ecology: Adelpha seriphia godmani is apparently not uncommon in Mexico, but elsewhere in its range it is local and rare. Although it has been recorded down to sea level in Panama and as high as 2400m in Mexico (de la Maza and de la Maza, 1993) and El Salvador, it is typically encountered from around 700-1 500m, in cloud forest habitats. Males are usually encountered perching in a variety of microhabitats, typically along ridge and hill tops where the vegetation exists in various degrees of disturbance. I have observed males in western Ecuador along a wide ridge top path, perching on bushes with their wings shut at the forest edge, around 3-6 m above the ground, while Raguso and Gloster (1996) reported males (as A. serpa) perching from 81 5m above the ground along forest edges and trails. Perching males are very wary, and at the slightest disturbance they quickly take flight along the ridgetop to land on another suitable perch tens of mefres away. Salazar (1996) also found males hill topping on Cerro Ingruma in Colombia. Females are found in heavily disturbed forest habitats along forest edges, but also in free fall light gaps in primary forest. The species is sympatric in western Ecuador with A. serpa duiliae and A. iphiclus estrecha, and since all three of these species exhibit narrower dorsal postdiscal bands and orange subapical markings, they are probably involved in mimicry. Specimens examined (33ni, 39f): MEXICO (23ni, 28f): Chiapas: Las Casas Mar. Im FSCA; no specific locality 2m, 5f AMNH; Guerrero: Iguala Im AMNH; no specific locality Im USNM; Mexico: Chalma Jul. If AME; Malinalco Aug. Nov. 2f AME; Valle de Bravo Nov. If AME; Oaxaca: Chimalapa Jul. Sep. Im, 2f AME; Comitan Sep. IfAME; Veracruz: Catemaco Sep. ImAME; Cordoba 5m, 2f BMNH; Fortin Im, 2f FSCA; Jalapa 4600' Nov. 2f USNM, Im, If BMB; Orizaba Im BMNH; Presidio May Jul. Im, IfAME; Santiago Tuxtla Sep. If AME; Selva Negra 1700m Aug. If AME; Sierra Mojarra Jul. IfAME; Tezonapa Jun. Im AME; Veracruz Im, If AMNH; Xalapa Nov. Im AME; Not located: no specific locality Im USNM, Im, If BMNH, 2f MNHN, Im MCZ, Im, If AMNH. GUATEMALA

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168 (Im 10Alia Verapaz: Baleu Sep. Im AMNH; Santa Rosa. Guazacapan If BMNH. EL SALVADOR (Im, IQ: Melapan: Metapan 2418m Dec. Im AME; San Salvador. Santa Tecla 900m Oct. If AME. COSTA RICA (30: no specific locality 2f BMNH, If BMB. PANAMA (Im, If): Canal Zone. Gatiin May If STRl; Henera. Cerro Alto Higo 1000m Dec. Im USNM. COLOMBIA (4m, 10: Caldas: Bocatoma, Manizales 2250m Feb. If MHNM; Paramo de Letras 2200m Dec. 2m JFL; Quindio: Cerro Aguacatal 1600m May Im JFL; Valle del Cauca: Queremal Jul. 1 m LMC. ECUADOR (3m, 30: Carchi: Lita, ridge east of Rio Baboso 900m Jul. Aug. 2m KWJH;'A/fl/iaW: Cerro Pata de Pajaro, nr. Pedemales 750m Aug. If KWJH; Pichincha. Tandapi 1550m Mar. Aug. If KWJH, If USNm'; Not located: no specific locality Im BMNH. COUNTRY UNKNOWN (10: no specific locality If ZMHU. Additional locality data: MEXICO: Chiapas. Cuenca de Grijalva; Cuenca de Tulija; Macizo Central; Selva Lacandona; Soconusco; Sierra Madre (de la Maza and de la Maza, 1993). COLOMBIA: Caldas. Cerro Ingruma 2300m Jul. Sep. Dec. (Salazar, 1996). ECUADOR: Carchi: Lita, ridge east of Rio Baboso Sep.; Pichincha: Palmito Pamba 1550m Oct. (Willmott and Hall, unpubl. sight records); Reserva Maquipucuna 1550-1 600m Sep. (Raguso and Gloster, 1996). Adelpha seriphia therasia Fruhstorfer, 1915 Figs. 19i,j Adelpha seriphia therasia Fruhstorfer (1915: 531) TL: Bolivia. Types: ZMHU: ST?f: "Coroico, ca. 2000m, Bolivia, (Yungas) 1895 (2-5) Garlepp//Syntype//SYNTYPE m Adelpha seriphia therasia Fruhstorfer G. Lamas det. '87"; ST?m: Rio Songo (1200m), Bolivia (Yungas), 1896, Garlepp [both examined] Identification, taxonomy and variation: This subspecies is similar to A. s. pione, but may be distinguished by having the ventral submarginal series paler, broader, the spots in each cell more clearly divided by dark rays on the hindwing and extended to almost merge with the pale subapical marking on the forewing. The ventral surface is very similar to the Peruvian syntype specimen of naryce, placed as a synonym of aquillia in this work, which differs as in all A. s. aquillia in lacking the well developed postdiscal series of orange dashes on the DHW. I have seen only a few specimens of A. s. therasia, in some of which the inner and outer ventral submarginal series are almost completely merged. Fruhstorfer (1915) provided no information on the sex, number, depository or exact locality of the type specimens of therasia, except that they were collected by Fassl in Bolivia. The description of therasia is also brief, stating only that the specimens differed from "Colombia forms" in being smaller with narrower white postdiscal bands. I am not certain that the ZMHU specimens are indeed valid syntypes, as they appear to have been collected by Garlepp rather than Fassl, but I have no doubt that the taxon which Fruhstorfer described as therasia is indeed the same as that treated here. Range: Adelpha s. therasia is to date known only from southern Peru to northeastern Bolivia.

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169 Habitat and adult ecology: There are no published observations on this subspecies, which, according to specimen label data, occurs from 1200-2 150m. The only recorded month of collection is August, the transition from the dry to the wet season. Specimens examined (4m, 30^ PERU (Im, If): Cu2co: Marcapata 4500' If BMNH; Quebrada Morro Leguia, Cosflipata 2150m Aug. Im MUSM. BOLIVIA (3m, 2f); La Par. Coroico If ZMHU, Im MCZ; Rio Songo 1200m Im ZMHU; Rio Tanampaya If ZMHU; San Antonio, Yungas 1800m ImZMHU. Adelpha seriphia egregia Rober, 1927 Figs. 19k,l Adelpha egregia Rober (1927: 420, fig. 4) TL: Sierra Nevada de Santa Marta, Colombia. Types: BMNH(T): HTf: "Adelpha egregia Rober, Holotype f (illustrated "Type") det. R.I. Vane-Wright 1983//Brit. Mus. 1928-151//B.M. TYPE No. Rh. 307 Adelpha egregia f R6ber//Adelpha egregia R6b. Type f//Sierra Nevada de Sta. Maria Columbian S. Am. 1500m Juni 1925 Coll. Dr. Kniger W. Niepelt Zirlau//TYPE//TypeSyntype"; PTm; "Adelpha egregia Rober m Paratype det. R.I. Vane-Wright 1983//Brit. Mus. 1928-151//B.M. TYPE No. Rh. 306 Adelpha egregia m R6ber//Adelpha egregia Rob. Type nV/Sierra Nevada de Sta. Marta Columbien S. Am. 1500m Juni 1925 Coll. Dr. Kruger W. Niepelt Zirlau//Type//1 16//T.G.H. 1962 551" [both examined] =Adelpha serpa pione G. and S., Hall (1938) (form); Adelpha seriphia egregia R6b., Willmott and Hall (1999) Identiflcation, taxonomy and variation: As the name suggests, this subspecies cannot be mistaken for any other Adelpha taxon. Rober (1927) described this distinctive subspecies as a separate species based on two males in Niepelt 's collection, one of which he figured and named the Type. This specimen however proves to be a female and is in the BMNH with the male paratype. Hall (1938) synonymised the name as a form of A. s. pione, which it clearly is not given the geographic isolation of the Sierra Nevada de Santa Marta and lack of variation observed in the small series examined. Willmott and Hall (1999) followed an impublished checklist of G. Lamas in treating egregia as a subspecies of A. seriphia, with which it shares the diagnostic characters for this species. Range: This subspecies is endemic to the Sierra Nevada de Santa Marta in northern Colombia. Habitat and adult ecology: Nothing has been reported on the habitat and behavior of this subspecies, which, judging from the number of specimens in collections, appears to

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170 be locally not uncommon. It has been recorded from 1500-2400m with the majority of records in the latter half of the year. Specimens examined (3m, Sf): COLOMBIA (3m, 80: El Cesar. Rio Meoguachucua 1900m Aug. If BMNH; Magdalena: Minca 2400m Oct. 2f JFL; San Lorenzo Mines Dec. If ESM; E. above San Pedro de la Sierra, Sta. Marta, 2000m Aug. Im BMNH; San Sebastian-El Mamon 2200m Mar. Sierra Nevada de Sta. Marta If MUSM; Sarachui 1750m Jul. Im, If BMNH; above Vista Nieva 1600m Jun. If BMNH; Not located: Sierra Nevada de Santa Marta 1500m Jun. Im, If BMNH. Adelpha seriphia barcanti Willmott, ssp. nov. Figs. 19m,n Adelpha seriphia Fldr., Kaye (1 940), Baix:ant (1 970: 1 3 1 , pi. 5, fig. 3) Description and diagnosis: FW length of HT: 28.5mm. Male differs from that of A. seriphia seriphia in entirely lacking the orange subapical marking on the DFW in cells M2, Ml and R5, with the result that the VFW subapical marking is white instead of pale orange, and in having slightly narrower dorsal postdiscal bands. The ventral submarginal markings are also slightly obscured by darker scaling, while the inner submarginal series on the VHW is almost parallel with the orange-brown postdiscal line due to the spots being more elongate and diffuse. Since, however, every other wing pattern character of this taxon is typical of the nominate subspecies, there is little doubt that the two are conspecific. I have not examined the female. Types: Holotype male: TRINIDAD: El Tucuche summit, 3072 ft., 11 Aug. 1979 (M. J. W. Cock); in the BMNH(M). Paratype : 1 male: same data as holotype; in coll. M. J.W. Cock, Reading, England. Etymology: This subspecies is named for Malcolm Barcant, who first figured this new subspecies in his field guide to Trinidadian butterflies (Barcant, 1970). Taxonomy and variation: I have examined only two specimens of this subspecies, that figured by Barcant (1970), which was collected by R. Dick at the type locality in 1927 (now in the Angostura-Barcant collection in Trinidad Cock, pers. comm.) and the holotype, and

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171 there is little variation. Both Kaye (1940) and Barcant (1970) simply refer to it as Adelpha seriphia. Range: Adelpha seriphia barcanti is known only from the island of Trinidad, and has been collected only at the type locality, the summit of El Tucuche. Habitat and adult ecology: Barcant (1970) reports that this subspecies could be regularly found on the summit of El Tucuche, perching in small groups, although Cock (pers. comm.) is of the opinion that he never captured any specimens. I am grateful to Matthew Cock for his comments on this subspecies, which 1 reproduce here: "In Trinidad, A. seriphia appears to be restricted to the one locality: the summit of El Tucuche (3, 072 ft.), a disturbed mountain top in primary forest. Between 1979 and 1982 I visited the summit of El Tucuche five times (once overnight), saw this species on two visits (several specimens on each occasion) and collected it once. At that time ... the summit itself was reasonably clear. . . . Trees growing just below the summit to the south and west overlooked the peak slightly, and the male of A. seriphia settled on these, only rarely coming within reach of a long-handled net. On one occasion I climbed the adjacent, slightly lower peak. El Naranja, but it was forested and collecting was not practical. Apart from my two males, as far as I am aware the only other Trinidad specimens known are those which Barcant refers to . . . and five males and a unique female in the collection of Scott Alston-Smith. . . . Since climbing El Tucuche is normally a dawn to dusk frek ... I would say with some confidence that all specimens were collected between 12.30 and 14.00". Specimens examined (2m): TRINIDAD (2m): El Tucuche summit 3072 ft. Aug. Im BMNH, Im figured by Barcant (1970). Adelpha hyas (Doyere, [1840]) Figs. 20; 104; 187

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172 Identification, taxonomy and variation: Adelpha hyas is distinguished from other members of the A. serpa group by its small size, the lack of orange postdiscal dashes on the VFW in cells Cu2-M3 (occurring in A. seriphia and A. serpa), the dark, deeply intruding lines in the pale VFW subapical marking (also occurring in some subspecies of A. seriphia) and the pale base of the VFW costa (orange in A. seriphia). There is geographic variation in the color of the ventral surface, arrangement of the DFW postdiscal band and ventral submarginal series, and three distinctive subspecies are recognised. Adelpha hyas has rarely been accorded full species status, due to confusion surrounding the systematics of the A. serpa group in general. However, there are several characters which link its respective taxa, and it is clearly sympatric in several populations with A. serpa, whilst occurring at lower altitudes in the same areas as its probable closest relative, A. seriphia (see that species account). Range and status: Eastern Ecuador along the base of the Andes to Bolivia, northern Argentina to southeastern Brazil. Probably extends into southern Colombia. Amazonian subspecies very local and rare, in wet rain forest in Andean foothills, southern subspecies locally common. Specimens examined: 116 (62m, 54f) Adelpha hyas hyas (Doyere, [1840]) Figs. 20a,b; 187a-c Heterochroa hyas Doyere ([1840]: pi. 138, fig. 1, Ibis) TL: not stated. Types: BMNH?: STf?: not located. Adelpha hyas Boisd. [sic], Kirby (1871); Adelpha serpa form hyas Boisd. [sic], Fruhstorfer (1915); =Adelpha serpa serpa Boisd., Hall (1938); Adelpha hyas Boisd.[sic], Brown (1992) Identiflcation, taxonomy and variation: Adelpha hyas hyas is a distinctive taxon, and is easily distinguished from other subspecies by the yellowish-brown (instead of reddish brown) bands basal to and distal of

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173 the white postdiscal band on the VHW. Other characters unique to A. hyas hyas are the bluish lines which fill the interior half of the area between the first and second discal cell bars and line the discocellulars on the VFW, the orange rather than red coloration at the base of cell Cu2 on the VFW, and the configuration of the white postdiscal spots in cells Cul and M3 on the DFW, which have a straight, diagonal distal edge. There is slight individual variation in the width of the dorsal postdiscal bands and extent of the DFW orange subapical marking. The author of the name hyas has been historically regarded as Boisduval, probably because Doyere ([1840]) credited the name to Boisduval in the original description. Nevertheless, since Doyere was the sole author on the work (Lamas et al, 1995), the name should be attributed to him. The original description contains no text to diagnose the species, but since both dorsal and ventral surfaces were accurately figured there is no doubt as to the taxon to which the name applies. Doyere provided no information on the origin of the specimen on which the illustration was based, nor did he indicate the sex, although the illustration suggests a female, and he stated that the specimens figured came fi-om the collection of Boisduval. I have been unable to locate any potential type specimens in the BMNH, where the Adelpha fi"om Boisduval's collection reside, either in the main collection or the separately housed duplicates fi-om Oberthur's collection, where a syntype of Adelpha serpa Boisduval was recently discovered by Lamas (see under A. serpa). However, it is by no means certain that the type is lost and since there is no confiision surrounding the name, 1 deem a neotype designation unnecessary. Adelpha hyas hyas has often been regarded as a form of A. serpa (Hall, 1938; Hayward, 1973), largely on the basis of supposed intermediate specimens which actually prove to belong to a third species, A. radiata. Adelpha hyas hyas is actually sympatric with A. serpa serpa throughout its range and I have seen no specimens intermediate between the two taxa, a fact noted by Hoffmann (1937), who believed it to be a distinct species since he observed little to no variation in other Brazilian Adelpha and, despite

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174 rearing many /i. serpa serpa from eggs, never found a specimen of ^. hyas. Brown (1992) also accorded A. hyas specific status. Range: The nominate subspecies occurs from southeastern Brazil to northern Argentina and Uruguay, and although unreported from Paraguay to date, it probably occurs there. Immature stages: Hoffmann (1937) states that the egg is green. Habitat and adult ecology: Biezanko (1949) reports that this taxon is rare in Rio Grande do Sul and flies in the months of April to June. Brown (1992) also reported it to be rare in the Serra do Japi, Sao Paulo, where it apparently can be encoimtered on hilltops near forest, and it occurs there in a variety of habitats, including gardens, areas with heavy secondary grovv1:h and even open fields. It is found typically above 1 100m, though it may descend to 900m in valleys. Ebert (1969) found the species to be not uncommon to common in Minas Gerais in primary to secondary forest habitats. Specimen data indicate that elsewhere in its range it flies from near sea level to 1500m, from September to at least May. Specimens examined (SOm, 50f): BRAZIL (49m, 520: Guanabara. Rio Im MCZ; Minas Gerais: Passa Quatro Jan. Im BMNH; Parana: Castro Apr. 2f USNM, 3m, 8f BMNH; Femandes Pinheiro 2600' Apr. Im, If AME, 2m BMNH; Ponta Grossa Sep. 2m AME; Uniao da Vitoria 2000' Dec. Jan. 2m BMB; no specific locality Im, If BMNH; Rio de Janeiro: Nova Friburgo If ZMHU; Petropolis 1500m Jan. May 2m AME, Im AMNH, 1 m USNM; Rio Grande do Sul: Pelotas Feb. Apr. 3m, 3f MCZ, 1 m, 4f AMNH, I f AME, 3f USNM; Rio Grande Jan. 1 m, 3f BMNH, If MCZ; Santa Maria 1400' Jan. Im, If BMB; no specific locality Im USNM; Sania Catharina: Blumenau Im MCZ; Cauna 15m, 1 If AMNH; Joinville 100-200m Dec. Im MUSM; Rio Vermelho Im AMNH; Sao Bento do Sul 1750' Mar. If FSCA; Theresopolis Im, 2f BMNH; Villa de Lages 2m BMNH; no specific locality 2m, 6f AMNH, 3f ZMHU; Sao Paulo: SSo Paulo Im BMNH; Not located: no specific locality Im, If BMNH. COUNTRY UNKNOWN (Im): Im ZMHU. Additional locality data: BRAZIL: Minas Gerais: nr. P090S de Caldas 1000-1400m Mar. May (Ebert, 1969). Rio Grande do Sul: Monte Benito (Biezanko, 1949). ARGENTINA: Misiones (Hayward, 1951). URUGUAY: no specific locality (Biezanko et al., 1978). Adelpha hyas viracocha Hall, 1938 Figs. 20c,d Adelpha serpa form viracocha Hall (1938: 285) TL: Chanchamayo, Peru. Types: BMNH(M): HTm: "Holotype/ZPerou Chanchamayo Oswald Schunke Recu 1912//Ex. Oberthur Coll. Brit. Mus. 1927-3//Holotype m Adelpha serpa f viracocha Hall G. Lamas det. 1987"; PTm: Bolivia Hewitson Coll. 79-69 Heterochroa hyas. 7.//B.M.(N.H.) Rhopalocera vial number 4412//Boliv."; PTf: Bolivia Hewitson Coll. 79-69 Heterochroa hyas. 2.//Boliv."; BMB: PTm: Chanchamayo [all examined] Adelpha serpa ? subsp. D'Abrera (1987); Adelpha hyas viracocha Hall, Willmott and Hall (1999)

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175 Identification, taxonomy and variation: Adelpha hyas viracocha is distinguished from the nominate subspecies under that account, and from A. hyas hewitsoni by the submarginal series on the venfral surface being merged with one another and almost blending with the postdiscal markings. It is similar to sympatric A. seriphia aquillia, but may be distinguished by the lack of a broad black band distal to the reddish postdiscal band on the VHW, the lack of any orange postdiscal dashes on the VFW in cells Cu2-M3 and by the base of the WW costa being white instead of orange. Adelpha serpa diadochus has much more sharply defined submarginal markings on the venfral surface and no darker lines intruding into the pale VFW subapical marking. There is slight variation in the extent to which the venfral submarginal series are merged, in some specimens they are all but indistinguishable and the distal half of the wing takes on a uniform, pale brownish color. This taxon was described by Hall (1938) in comparison with A. hyas hyas and A. radiata radiata, and I have examined all of his designated type specimens. The characters which justify its placement as a subspecies of A. hyas are discussed under A. hyas hewitsoni. Range: Adelpha hyas viracocha is known from eastern Andean foothills from cenfral Peru to Bolivia, though it possibly extends further north in Peru. Habitat and adult ecology: There are no published observations of this subspecies in nature. It is clearly very rare and local, and restricted to forest along the base of the Andes up to 1050m. Specimens examined (5m, 3f): PERU (3m, If): Junin: Chanchamayo Im BMNH, Im BMB; Puno: Rio Tivara 1050m Aug. Im, If MUSM(/>i copulo). BOLIVIA (2m, 20: La Paz: La Paz If MNHN; Rio Songo Im ZMHU; Not located: no specific locality Im, If BMNH. Adelpha hyas hewitsoni Willmott and Hall, 1999 Figs. 20e,f; 104 Adelpha hyas hewitsoni Willmott and Hall (1999: in press) TL: Pimpilala, nr. Talag, Napo Province, Ecuador. Types (all Ecuador): KWJH: HTm and 4PTm: Pimpilala; ATf: Apuya; PTm: Rio Llandia; MNCN: PTm: Pimpilala [all examined]

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176 Identiflcation, taxonomy and variation: Adelpha hyas hewitsoni is distinguished from the remaining two A. hyas subspecies by the small, sharply defined spots which compose the ventral submarginal series, which are themselves distinct and clearly separate from the ventral postdiscal markings. There is slight local variation in the extent of the DFW orange subapical marking, which may extend into cell M3. Adelpha hyas hewitsoni was clearly described and figured in the original description, and the type series has been examined. Willmott and Hall (1999) placed the taxon with A. hyas viracocha as subspecies of A. hyas on the basis of the following shared characters: small size, lack of dark orange outer postdiscal series dashes on the VFW in cells Cu2-M3, VHW postdiscal band and inner submarginal series parallel, base of the VFW costa white rather than red-orange, deeply intruding dark lines into the pale VFW subapical marking. Adelpha hyas viracocha additionally shares the following characters with A. hyas hyas: faded, merged ventral submarginal series on the VHW that extend to the orange/yellow-brown postdiscal band and merging of the VFW submarginal series with the subapical marking to form a pale area which fills the distal half of the wing and ends basally along a straight, diagonal line extending from tomus to costa between the subapical marking and upper postdiscal band. Range: To date this subspecies is known only from a small area in cenfral eastern Ecuador. Presumably it is much more widespread, probably extending at least to southern Colombia and northern Peru. Habitat and adult ecology: Adelpha hyas hewitsoni has been recorded at three localities at the base of the Andes, from 600-950m. It is sympatric with Adelpha paraena paraena, Adelpha radiata explicator and Adelpha serpa diadochus, while Adelpha seriphia aquillia also occurs in the east Ecuadorian Andes, but at higher elevations (16001800m). Males of A. hyas hewitsoni are typically encountered in groups in large (20-30m wide), old forest clearings along streams and rivers. Usually they perch from 1200-1 330h in

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177 bright sun on the tops of bushes 5-6m high, making sorties out and returning to the same perch. At the type locality, Pimpilala, the species seems to be present throughout the year in a particular coffee orchard, where it is often the most common perching Adelpha. Other species which resemble it on the dorsal surface and with which A. hyas perches include Adelpha iphicleola thessalita, Adelpha thoasa manilia and Adelpha serpa diadochus. The single known female was flying in heavily disturbed ridgetop forest around midday, at a site where males have yet to be recorded. Specimens examined (7m, 10^ ECUADOR (7m, \f): Napo: Apuya 600m Sep. If KWJH; Pimpilala 600m Apr. Aug.-Oct. 5m KWJH; Talag, Napo Ecuador Alt. 800m D.A.Trembath 5.IX.95 No. 14176 Im DAT; Pastaza: Rio Llandia 950m Sep. Im KWJH. Adelpha alala (Hewitson 1847) Figs. 21; 105; 188 Identification, taxonomy and variation: Adelpha alala is distinguished from A. corcyra and A. aricia by the more rounded hindwing tomus and by having two clearly marked, oval or round white postdiscal spots at the DFW costal margin, instead of diffuse white shading. The clunicula of the male genitalia is shorter than A. aricia, but longer and more pointed than in A. corcyra. There is some variation in the presence or absence of an orange postdiscal band on the DHW, and three subspecies are recognised. Adelpha alala is closely related to A. aricia and A. corcyra, sharing almost full expression of all postdiscal and submarginal series on the ventral surface, a distinct, entirely red-brown margin on the VFW and VHW, overall pale ventral coloration, especially in the basal area of the VHW, and similar genitalia, with the costal lobes of the male genitalic valvae reduced or absent, and the valvae bearing only a single, or few, large, terminal spines. Range and status: Western slopes of the Andes from northwestern Venezuela (Cordilleras de la Costa, Merida and Sierra de Perija) to northwestern Ecuador, Sierra

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178 Nevada de Santa Marta, and eastern Andes from Colombia to northern Argentina. From 450m at the foot of the mountains to 2600m in a wide variety of habitats. Very common. Specimens examined: 875 (859m, 160 Adelpha alala alala (Hewitson 1847) Figs. 21a,b Heterochroa alala Hewitson (1847: 261, PI. XXI, fig. 8) TL: Venezuela. Types: BMNH(T); STm: "Illustrated in The Butterflies of Venezuela A. Neild, 1996//B.M. TYPE No. Rh. 9807 Heterochroa alala Hew./A'enezuela//Type//Syntype"; BIVINH(M): ST?m: "Hewitson Coll. 79-69 Heterochroa alala. 1." [both examined]. Adelpha alala Hew., ICirby (1871) Identification, taxonomy and variation: The nominate subspecies is distinguished from A. alala completa by lacking or having only a faint trace of postdiscal orange dashes on the DHW, and from both other subspecies by having the DFW subapical orange markings filling each cell to form a solid band, instead of being isolated spots in each cell. Hewitson (1847) described this species from an unspecified number of specimens in his own collection and the British Museiun, and two syntypes have been located in the BMNH. The original description, which mentions the white postdiscal DFW costal spots, and figure of the dorsal surface of the right hand pair of wings, which show the enlarged orange DFW postdiscal spots, clearly refer to this taxon. Two of the three specimens that I have examined from the Sierra Nevada de Santa Marta in Colombia differ from Venezuelan specimens in having the DFW orange subapical band extending distal of the white postdiscal band almost to the anal margin, while in the remainder it doesn't quite extend into cell Cu2. In view of the isolation of the Sierra Nevada de Santa Marta from the Venezuelan Cordillera de la Costa, the presence of a different subspecies, A. alala completa, in intervening mountain ranges, and the slight wing pattern differences mentioned, the Colombian population may well merit separate subspecific recognition when more specimens are available to assess variation. In addition, I have to seen no

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179 specimens from the Sierra de Perija on the ColombiaVenezuela border, which might help clarify the situation. Range: Typical A. alala alala occurs in the Cordillera de la Costa of northwestern Venezuela, while phenotypically similar specimens are found in the Sierra Nevada de Santa Marta in north Colombia. Habitat and adult ecology: The nominate subspecies is relatively uncommon in collections and nothing has been reported on its behavior in the field. It is probably involved in mimicry with the sympatric A. seriphia seriphia. Specimens examined (17ni, 7f): VENEZUELA (16m, 5f): Aragua: La Lagunita de la Montana, S.W. Colonia Tovar 1750m Oct. 4m BMNH; Distrito Federal: Avila 1000m Jul. Im AFEN; Caracas 8000' Feb. Im, If AME, 2m, If BMB; S. slope of Mt. Avila 6510-7000' Im USNM; Not located, no specific locality 3m, 3f BMNH, 3m AMNH, Im BMB. COLOMBIA (Im, 2f): Magdalena: "Sarachui 1750metres 18.VlI.1972//North Colombia: 1725 Sierra Nevada de Sta. Marta Exped. M.J. Adams and G.l. Bemard//6 1.7 1/72 A/B/B.M. 1973106" Im BMNH(AandB); "Sarachui 1830 1750metres 17.VI1.1972//B.M. 1973-106" 1 f BMNH(AandB); El Cesar. "MaruamaqueChenduca 1450-1750m 15.VII.972//North Colombia: 1697 Sierra Nevada de Sta. Marta Exped. M.J. Adams and G.I. Bemard//B.M. 1973-106" If BMNH(AandB). COUNTRY UNKNOWN (Im): Im BMNH. Adelpha alala negra (C. Felder and R. Felder, 1862) Figs. 21c-f; 105a,b; 188a,b Heterochroa alala var. negra C. Felder and R. Felder (1862: 115) TL: [fluminis Negro superioris in Brasilia septentrionali] [Peru]. Types: BMNH(R): STm: "Syntype//Rio Negro//v. negra Feld.//negra n."; ST?m: "Rio Negro//f. negra Feld." [both examined] =Adelpha ehrhardi Neuburger (1907: 50) syn. nov. TL: Santa Inez, Ecuador. Types: Wilhelm Neuburger collection (current location unknown): HT [not examined] ^Adelpha alala albifida Fruhstorfer (1907: 172) syn. nov. TL; "Unknown, probably Central America or Colombia". Types: BMNH(T): STm: "cora fa. albifida Fruhst.//H. Fruhstorfer/ZFruhstorfer Coll. B.M. 1933-131//TYPE//Type//Syntype" [examined] -Adelpha alala cora Fruhstorfer (1907: 172) syn. nov. TL: Pozzuzo, Peru. Types: BIVINH(T): STm: "alala cora Fruhst.//Peru H. Fruhstorfer/ZFruhstorfer Coll. B.M. 1933131//TYPE//Type//Syntype" [examined] =Adelpha alala fillo Fruhstorfer (1907: 172) syn. nov. TL: "Argentina, Bolivia??". Types: BMNH(T): STm: "fillo Fruhst.//Argentinien ex. coll. FnihstorferZ/Fruhstorfer Coll. B.M. 1933-131/ZTYPE/ZType//Syntype" [examined] =Adelpha negrina Fruhstorfer (1913: pi. 108a; 1915: 515) syn. nov. TL: [Bolivia]. Types: BMNH(T): STm: "alala negrina Fruhst.Z/Yungas Bolivia/ZFruhstorfer Coll. B.M. 1933131ZZTYPEZZTypeZZSyntype"; BMNH(R): STm: "negrina/ZGuanay Mapiri R. 1500' 8 95 StuartZZalala negrina Fnihst.ZZR/Z108 A3" [both examined] =Adelpha alala completa form praecaria Fruhstorfer (1915: 515) unavailable name TL: not given. Types: BMNH(T): STm: "completa fa. praecaria Fruhst./ZColumbien ex. coll. H. FruhstorferZZFruhstorfer Coll. B.M. 1933-131ZZTYPE/ZTypeZZSyntype"; BMNH(R): STm: "R. Dagua Colombia W. RosenbergZZcompleto fa. praecaria Fruhst.ZZR/Zcompleta" [both examined] -Adelpha alala privigna Fruhstorfer (1915: 51 5) syn. nov. TL: "presumably Peru or the Upper Amazon". Types: BMNH(T): STm: "cora fa. privigna Fruhst.//Peru H. Fruhstorfer/ZFruhstorfer Coll. B.M. 1933-131ZZTYPEZZTypeZZSyntype"; BIVINH(M): ST?m: "privigna Fruhst.Z/Peni H. FruhstorferZZFruhstorfer Coll. B.M. 1933-1 31 "[both examined] Adelpha alala var. negra Fldr., Kirby (1871)

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180 Identiflcation, taxonomy and variation: This subspecies is distinguished from A. a. completa by lacking the orange postdiscal band on the DHW, and from the nominate subspecies by the smaller orange DFW subapical spots, which are isolated in each cell space. It is a rather variable subspecies and a nimiber of phenotypes have been described, particularly by Fruhstorfer (1907, 1913, 1915). The DFW orange subapical spots vary in size and may in some specimens, particularly in north Peru (Tabaconas), be entirely white (an atavism), the dorsal postdiscal bands vary in thickness, and specimens from cenfral to southern Peru frequently have the orange DFW subapical spots continued as a postdiscal band to the anal margin of the forewing. The venfral surface varies from being striped with rich reddish and yellowish brown bands to a washed out silvery gray, the brown colors almost absent. After an examination of a large series I conclude that all of these characters are locally variable and do not define geographic populations worthy of subspecific recognition. However, data for most Peruvian specimens that I have examined are vague and it may be that ftirther collecting in isolated Cordilleras will reveal the presence of phenotypically stable populations. It is very possible that the variable possession of the entire orange postdiscal band on the DFW is due to local mimicry with A. aricia, which has an entire DFW band throughout central Peru and Bolivia. The Felder brothers introduced the name negra as a variety of A. alala, stating that it differed from the Venezuelan form in having a wider postdiscal band and less undulate submarginal brown lines, presumably referring to the DHW. The type specimen(s) supposedly originated from the Upper Rio Negro in Brazil (C. and R. Felder, 1862), but it is clear from other species described in the same paper that the specimens actually originated in the Rio Huallaga valley above Yurimaguas, in north Peru (Lamas, 1976). I have examined a syntype male and a fiirther possible syntype male in the BMNH(R). Subsequently Neuburger (1907) described specimens of A. alala from Ecuador (Santa Inez), Venezuela (Merida) and Bolivia as a new species, A. ehrhardi.

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181 which he stated differed from the nominate in lacking the orange DHW submarginal markings. He also stated that he considered the Ecuadorian specimen to represent the typical form, and I therefore regard this specimen as the holotype and the type locality as Ecuador. Although no syntypes have been located of this taxon, since no mention was made of A. alala negra and Ecuadorian specimens are otherwise typical of this subspecies, I synonymise ehrhardi with negra (syn. nov.). Fruhstorfer (1907, 1913, 1915) described six new taxa based on what I consider to be infrasubspecific variation in width of the dorsal postdiscal band, color of DFW subapical spots and color of the venfral surface. Hall (1938) synonymised pmecaha with negra and retained two of the remaining taxa (cora and fillo) as subspecies, with due skepticism, relegating the remainder to the status of forms of these subspecies. The name A. alala albifida was applied to an unknown number of specimens lacking locality data and the syntype male in the BMNH corresponds well with the description (Fruhstorfer, 1907), having a broad dorsal postdiscal band and white DFW subapical spots. Adelpha alala cora was based on 2 males from Pozzuzo in Fruhstorfer' s collection in which the orange DFW subapical markings extend to the anal margin (Fruhstorfer, 1907), and although the single putative syntype in the BMNH bears only the data "Peru", it agrees well with the original description. Adelpha alala fillo was described for an unknown number of specimens which Fruhstorfer (1907) received from Buenos Aires and suggested originated in Argentina or Bolivia; a syntype male is in the BMNH and differs little from typical A. alala negra. Fruhstorfer's (1915) description of A. alala negrina, from Bolivia, is almost negligible, although the figure (Fruhstorfer, 1913) and two syntype males in the BMNH show it to scarcely differ from typical A. alala negra. The taxon privigna was described based on specimen(s) which differed from albifida only in having narrower dorsal postdiscal bands (Fruhstorfer, 1915), and I have examined two potential syntype males in the BMNH which correspond well with this description. I synonymise all of the preceding five taxa {albifida, cora, fillo, negrina and privigna) with A. alala negra (syn. nov.).

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182 Finally the form praecaria was described for Colombian specimen(s) differing from A. alala completa in lacking the DHW orange submarginal band (Fruhstorfer, 1915); the original figure (Fruhstorfer, 1913: pi. 108, a, as completa) and syntype males in the BMNH show it to be a synonym of ^. alala negra, as Hall (1938) concluded, and the name is in any case unavailable, being a quadrinomial. Range: This subspecies occurs from Colombia, in the Cordillera Oriental south of Bogota, to northwestern Ecuador west of the Andes, to northern Argentina on the east Andean slopes. Habitat and adult ecology: Males of A. alala negra are very common in a wide variety of montane habitats up to 2600m, and may even be found in wet lowland forest as low as 400m near the base of the Andes. Typically they are found in the vicinity of forested mountain rivers and are often observed congregating in very large numbers to feed at moist sand or on water seepage over rocks, particularly when these areas are enriched with urine. They are also attracted to rotting fish and dung on the ground or in traps, and very occasionally may feed on flowers. Males perch on secondary growth bushes from 48m above the ground along the edges of rivers, on the tips of leaves with their wings almost closed. Salazar (1996) also reported males hill topping on Cerro Ingruma in Colombia, at 2300m. Females of this subspecies are very rare in collections and nothing is known of their behavior. The species seems to fly throughout the year. In Peru and Bolivia this subspecies is sympatric and probably involved in mimicry with A. aricia, which inhabits similar montane habitats. Both of these species have local forms with orange postdiscal bands on the DFW throughout southern Peru and Bolivia. Specimens examined (691m, 5f): t trans. loA. a. completa. "VENEZUELA" (3m): Distrilo Federal. Avila Dec. Im JFL-error; Not located: no specific locality Im USNM, Im MCZ-error?. COLOMBIA (167m, 20: Antioquia. Frontino Im BMNH; Mesopotamia 8m AMNH; Rio Cocoma Aug. 2m AMNH; Salinas 48000" Jul. 6m, If BMB; Boyacd: Muzo 12m AME, Im BMB; Caldas: Aguacatal Riosucio Jul. Im KWJH; Bocatoma 2400m Aug. Im USNM; El Aguila Manizales 1800m Oct. Im KWJH; Manizales Feb. Sep. 5m BMNH, 2m MUSM; Quebrada El Bohio, Manizales 1750m Nov. Im MHNM; Caqueta: Caqueta Jun. 4m JFL; Cauca: Popayan Im BMB; Cundinamarca: Bogota 12m BMNH, 2m USNM, 2m BMB; env. de Bogota 12m BMNH; Bogota-Pandi Dec. Im BMNH; Honda 5m BMNH; La Vega Jul. Imt JFL; Rio Negro May Im ESM; Susumuco 2m BMNH; Meta: Guayabetal Jul. Im JFL; Peperital-Buenavista 2m BMNH; Villavicencio 10m AMNH; Naririo: El Palmar 2m AMNH; Quindlo: Cerro Aguacatal May Im JFL; Risaralda: Quebrada Rio Negro, Pblo. Rico 1550m Aug. Im MHNM; Tolima. Las Guayabas, Rio Cucuana 1500m Mar. 10m AME; Rio Ambeima 1400m Feb. Im AME; Valle del Cauca. Bitaco 1700m Jan. Im FSCA; Cali Aug. 2m BMNH, Im MUSM; Calima Dam 3000' Jan. Feb. Nov. Im FSCA, 3m AME; Cerro Gorda 2350m Im AME; El Saladito, km. 12 via mar Aug. Im LMC; Jimenez 1600' Mar. W. Col. Im BMNH; Juntas 4m BMNH; Lago Calima Jul. Im ESM; San Antonio 2000m Aug. Im LMC; Pance 4000' Sep. Im LMC, Im FSCA;

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183 Peflas Blancas 1800m Jan. Dec. Im FSCA, Im LMC; Pichinde Im BMNH; Pico de Aguila 1500m Jan. Jun. 3m FSCA, Im LMC; Rio Dagua 4m BMNH, \mZMHV, Not located: Caucathal Im, If ZMHU; no specific locality 19m AMNH, 2m BMNH, 3mZMHU, 2m USNM, 7m MNHN. ECUADOR (172m, 2f): Carchi: Lita 3000' 3m BMNH; Santa Rosa, Rio Golondrinas 1700m Sep. 2m KWJH; Esmeraldas: Esmeraldas 2m BMNH; Loja: Loja Dec. 4m BMNH, Im BMB; Imbabura: Paramba 3500' Mar.-May 15m BMNH; Morona-Santiago. Rio Abanico Oct. Nov. 3m MJP, Im DAT; Napo: Cotundo 800m Oct. Im USNM; El Capricho Oct. Im DAT; Finca San Carlo Apr. Sep. 2m KWJH; Utas 460m Feb. Im AME; Misahualli 650m Mar. Im MUSM; Rio Arajuno 700m Mar. Im AME; Rio Chalayacu Sep. Im DAT; Rio Coca Jul. 2m AME-error?; Rio Hollin Nov. Im DAT; Rio Napo Im MNHN; Satzayacu Sep. Nov! 3m AME, Im DAT; Pastaza: Alpayacu 3600' 3m BMNH; Mera Jun. Im DAT; Puyo Jun. Im DAT; Puyo, 20 km. N.W., Nov. Im AMNH; Puyo-Tena rd. km. 9, Oct. 2m MJP; Shell Oct. Im KWJH; Tungurahua: env. de Ambato 7m BMNH, 3m BMB; Bailos Jun. Oct. Im MJP, Im DAT, Im BMNH, Im BMB; El Rosario 4900' 2m BMNH; U Mascota 1200m Aug. 5m, If AME; La Victoria 3500' Im BMNH; Mirador Im AMNH; Palmera Im AMNH; Rio Blanco Apr. Aug. 3m AME; Rio Chinchin Grande 1400m Dec. 6m AME; Rio Mapoto Dec. Im AMNH; Rio Negro Apr. Jul. Dec. 2m, If AME; Rio Tigre 1200m Aug. 13m AME; Rio Topo 15-1 700m Jan. Jun. Aug. Sep. Oct. 4m MJP, Im BMNH, 12m USNM; Rio Verde 5000' Jan. Jul. Im BMNH, Im AME; Rio Zuilac 1300m Jul. 3m KWJH; Topo Mar. Dec. 3m AME; Zamora-Chinchipe: Rio Numbala Im BMB; north of Valladolid 2000m May Im KWJH; Zamora 3-4000' 19m BMNH; Zumba Jul. 2m BMB; Not located: middle Ecuador 3m AMNH; Oriente 3m AMNH; Solacama Im BMNH; no specific locality 7m AMNH, 4m BMNH, Im USNM, 2m BMB. PERU (232m, If): Amazonas: Chachapoyas 3m BMNH; Cord, del Condor 10-1200m Jul. 2m MUSM; Huambo Oct. 2m MUSM; Huayabamba 3500' Im BMNH, Im ZMHU; Mendoza, 14 km. W., 1800m Mar. Im MUSM; Ayacucho: Candalosa 1300m Jun. Im AME; Rio Piene 11m AMNH; Cajamarca: Charape 4000' Sep.-Oct. 3m BMNH; Huayabamba 5m BMNH; La Balsa-El Chaupe 1650-1850m Jun. Im MUSM; Rio Chinchipe 6000' 2m BMNH; Rio Tabaconas 6000' 3m BMNH; Cuzco: Buenos Aires, Cosfiipata 2-2300m Dec. 8m MUSM; Callanga 1500m Im ZMHU; Caradoc 4000' Feb. 4m BMNH; Cosfiipata 2m BMNH; Cuzco 4m MNHN; Machu Picchu Sep. Im MUSM; Marcapata 4500' 4m BMNH; Quebrada Morro Leguia 2150m Aug. 6m USNM; Quebrada Quitacalzon, Cosilipata 1050m May Im MUSM; Rio Pampaconas Aug. Im USNM; Santa Ana Im BMNH; Santa Isabel, Cosfiipata Feb. Jun. 3m MUSM; Yanamayo, Cosilipata 2000m Feb. 3m MUSM; Huanuco: Carpish Acomayo Sep. Im BMB; Cushi 1820m Im BMNH; Las Palmas 450m Aug. Im AME; Pozuzo 800-lOOOm 11m BMNH, Im BMB, Im MUSM; Tingo Maria Mar. Apr.-Aug. Im FSCA, 1 Im AME, Im USNM, 3m MUSM; no specific locality 1400m Apr. Im USNM; Junin: Chanchamayo Jun. Jul. 25m BMNH, 2m ZMHU, 2m AME, Im USNM, 2m BMB, Im MCZ; Hda. Naranjal, Mina Pichita 1550m Oct. Im MUSM; U Merced 2500' Mar. Aug.-Nov. 5m, If BMNH, 7m BMB, 2m MUSM; Rio Colorado Im BMNH, Im MCZ; Rio Perene Mar. Im BMNH, Im BMB; Rio Tulumayo, nr. Vitoc 1200m May Im MUSM; San Ramon 2m MUSM; Utcuyacu 5000' Dec-Jan. Im BMNH; "Lima": env. Lima Im MNHN-error; "Loreto": Iquitos Im AMNH-error; "Rio Negro" 2m BMNH; Madre de Dios: Madre de Dios Im MCZ; Pasco: Alto Yurinaqui, Enefias 1400m Apr. Im MUSM; Huagaruncho, nr. Cerro Pasco Im BMNH; Huancabamba Im BMNH; Oxapampa May 2100m 2m MUSM; Pozuzo Im BMNH; Puno: Carabaya May 4m BMNH, 2m AMNH; Inca Mines Oct. Im AMNH; Oroya 3-3500' Apr. Nov. 7m BMNH; Rio Inambari to Limbani Mar. Im BMNH; Sto. Domingo 4500' 4m BMNH; Uruhuasi 7000' Apr. May 2m BMNH; San Martin: Jepelacio 1 100m Jan. May Im BMNH, Im AME, Im AMNH; Juanjui Im MUSM; Rioja Jun. Im AME; Venceremos 1500m Dec. 3m MUSM; Not located: Opara 1600m Im AME; Rio Huallaga 5m AMNH, 2m USNM; no specific locality 10m BMNH, Im USNM, 3m BMB, 3m MNHN, Im MUSM, Im MCZ, Im JFL. BOLIVIA (79m): Cochabamba: Charapaya 1300m Apr. 3m BMNH; Cochabamba 9m BMNH, Im BMB; Incachaca 2220m Apr. 4m AME; San Jacinto 6-8000' Im BMNH, Im ZMHU; La Paz: Chaco 2-3000m Im ZMHU; Chulumani 5200' May Dec. 3m BMNH, Im FSCA, Im MCZ; Coroico Im ZMHU, Im AME; Guanay Im BMNH; U Paz Im BMNH, 3m MNHN; La Paz-Coroico rd. 2000m Sep. Im KWJH; 20 mi. above Mapiri 4000' 2m BMNH; Puenta Villa May 3m FSCA; Rio Puni 900-1400m Dec. Im AME; Rio Songo 1600m Apr. Im MUSM, Im ZMHU; Rio Suapi Im ZMHU; Rio Unduavi 2000m Feb. Im BMNH; San Augustin Sep. Im BMNH; Yolosa 1200m Sep. Im KWJH; Yungas 1200m Nov. 3m BMNH; Potosi: Tupiza Im AME; Santa Cruz: Bueyes 2m BMNH, Im ZMHU; Rio Negro Im AME; Not located: Okara Apr. Im AMNH; Pitiguaya May 1 Im AMNH; no specific locality Aug.6m BMNH, 5m AMNH, Im BMB, 2m MCZ. ARGENTINA (3m): Salta: Rio Arrayazal 600m Sep. 2m AME; Not located: no specific locality Im BMNH. "BRAZIL" (2): Parana: no specific locality 2m AMNH-error. COUNTRY UNKNOWN (33m): U. Amazon Im BMNH; no specific locality 4m BMNH, Im AME, 7m USNM. 6m BMB, Im MNHN, 12m MCZ, Im AMNH. Additional locality data: COLOMBIA: Caldas: Cerro Ingruma 2300m Jul. Sep. Dec. (Salazar, 1996). ECUADOR: Carchi: Las Juntas Nov.; Nariz del Diablo Nov.; Santa Rosa Sep. Nov.; Morona-Santiago: Limon-Gualaceo rd., km. 14, Sep.; Rio Abanico Oct-Dec.; Napo: El Capricho Oct.; Finca San Carlo Apr. Aug. Sep.; Pimpilala 600m Aug. Sep.; Tena-Loreto rd., km. 49, Aug. Oct.; Pastaza: Puyo-Tena rd., km. 35, Oct.; Rio Llandia Aug. Dec; Rio Puyo Oct.; Shell Feb. Apr. Oct.; Sucumbios: La Bonita-Rosa Florida rd., km. 12, Mar.; Quebrada El Copal Nov.; Quebrada El Garrapatal Nov.; Rio Palmar Nov.; Rio Sucio Nov.; Tungurahua: Chinchin Oct.; Rio Zuflac Jul. Oct.; Rio Machay Feb. Apr. Aug. Nov.; Zamora-Chinchipe: Chachacoma Nov.; Chinapintza Sep.; Quebrada Chorillos Apr. Nov.; Quebrada Las Dantas Oct.; Quebrada San Ramon Oct.; Zumba-Loja rd., km. 35, Jul. (Willmott and Hall, unpubl. sight records). ARGENTINA: Misiones (Hayward, 1951)-error?, presumably Salta. Adelpha alala completa Fruhstorfer 1907 stat. rest. Figs. 21g,h Adelpha alala completa Fruhstorfer (1907: 172) TL: Columbien, Muzo. Types: BMNH(T): HTm: "alala completa Fruhst.//Columbien Rochld[?] H. SticheV/Fruhstorfer Coll. B.M. 1933-13l//TYPE//Type//Syntype" [examined]

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184 =Adelpha alala form titia Fruhstorfer (1915: 515) syn. nov. TL [Venezuela]. Types: BMNH(T): STm: "Illustrated in The Butterflies of Venezuela A. Neild, 1996//33//alala titia Fruhst.//Merida Venezuela 1630m//Fruhstorfer Coll. B.M. 1933-1 3 1//TYPE//Type//Syntype"; BMNH(M): 2ST?m: "Fruhstorfer Coll. B.M. 1933-131/A'enezuela ex. Coll. H. Fruhstorfer" [all examined] Adelpha alala var., Staudinger (1886); =Adelpha alala negra Fldr., Hall (1938); Adelpha alala Hew., D'Abrera (1987); Adelpha alala titia Fruhst., Neild (1996) Identification, taxonomy and variation: This subspecies is distinguished by having a postdiscal line of orange dashes on the DHW distal of the white postdiscal band. Although there is some variation in the intensity of the dashes, this appears to be a sufficiently constant feature in the large series examined to warrant retention of this subspecies. Hall (1938) synonymised completa as a form oiA. alala negra, as he believed that the two phenotypes were sympatric. However, an analysis of more accurate locality data than Hall had available suggests that completa is not a local form but occupies a defined geographic area, and so I reinstate it as a good subspecies (stat. rest). As in A. alala negra, there is variation in the width of the white postdiscal band on both wings and size of the DFW subapical spots, but no variation that appears to be geographically correlated. Specimens from the Bogota area show intergradation to A. alala negra through loss of the orange DHW dashes. Fruhstorfer (1907) described A. alala completa from a single male specimen, collected by H. Stichel in 1896 in Colombia, which he stated differed from A. alala alala on the dorsal surface in having a narrower orange postdiscal band and broader white postdiscal band on both wings. The holotype is in the BMNH. Neither of these differences constitute anything more than individual variation, and it is unclear why he failed to mention the diagnostic orange DHW postdiscal markings which are lacking in the nominate subspecies. Fruhstorfer (1915) subsequently described the form titia from Venezuela, without specifying number, sex, location or accurate locality of type specimens, as differing from the nominate in having a narrower white dorsal postdiscal band. Again, his description and discussion are so brief as to be almost worthless, he fails to comment on the DHW postdiscal orange bands, and there is no mention of why he believed his form titia to be distinct from completa. I have examined a single syntype and

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185 two potential syntype males of titia in the BMNH and they do not differ from typical completa. Neild (1996) referred to Venezuelan specimens, treated here as A. alala completa, as A. alala titia, without reference to the former name. Since I can find no consistent differences between Venezuelan and Colombian A. alala completa I place titia as a junior synonym oi completa (syn. nov.). Range: Both Andean slopes from northwestern Venezuela (Cordillera de Merida) to north Colombia (Cordillera Oriental), as far south as Bogota. Immature stages: The only recorded hostplant is Viburnum tinoides var. venezuelense (Caprifoliaceae), a shrub or small tree commonly known in Venezuela as "Cabo de Hacha", which grows in areas of secondary growth and along forest margins (Otero and Aiello, 1996). Aiello and Otero (1996) describe the early stages, illustrating the last instar and pupa, and their observations are summarised here. The egg is laid singly on the underside of a leaf and is similar in morphology oto other Adelpha, as is the first instar larva. The second to fourth instar has a wide, dorsal greenish yellow stripe divided by a thin median line, and a lateral maroon stripe bordered ventrally by a white subspiracular line. The fifth instar is a more uniform green and differs greatly in appearance from all other known Adelpha in having very reduced scoli, with true scoli only on segments T2, T3 and A8, with those on the latter two segments being minute. The T2 scoli are orange with a single central black spine and five radiating spines at the apex. The pupa is bone white to straw colored, and simple in form, lacking obvious dorsal projections on T2 and A2. The head horns are triangular and squat. The construction of perches from leaf veins by the larvae is a behavior similar to that of other Adelpha (see generic introduction), while the late first instar and second instar go on to construct elaborate leaf shelters not reported for any other Adelpha, but very similar to those of Limenitis Camilla (Linnaeus) and Limenitis reducta Staudinger. Two opposing cuts, perpendicular to the leaf margin, are made from the margin to the midrib, and the leaf flaps pulled down and fastened with silk. The larva

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186 rests in the shelter when not feeding, and there is typically a curtain of dead leaf material left dangling by the feeding larva over the sheUer entrance. The larva abandons the shelter in the late fourth or early fifth instar, after v^hich it rests exposed on the leaf Otero and Aiello (1996) make the interesting observation that this is when the larva changes color to a imiform green, and it also seems likely that the great reduction in scoli is due to the protection gained fi-om the shelter rendering them almost obsolete. Habitat and adult ecology: The habitats and behavior of this subspecies are typical of the species; I have observed males puddling along streams near Merida, in Venezuela. Neild (1996) states that it is found from 1000-2000m in Venezuela. This subspecies occurs in the same montane habitats as A. corcyra corcyra and A. seriphia pione, and the ranges of each of these subspecies closely correspond. Although A. alala and A. corcyra are closely related, A. seriphia belongs in a different group within the genus, and the independent development of an entire orange postdiscal band on the dorsal surface of all three of these taxa is probably due to mimicry between them. Specimens examined (153m, 4f): t trans. ioA. a. negra. "U.S.A." (Im): California: no specific locality Im BMNH-error. VENEZUELA (76ni, 2f): Barinas: Quebrada Las Uvas, km. 34 Barinitas-Apartaderos Jan. Im MUSM; Santa Barbara 2m BMNH; Merida: La Mucuy 2300m May Sep. 2m MUSM; La Pueblita 1450-1550m Aug. 2m BMNH; Jacinto 1500m Im BMNH; Merida 1400-1500m Mar. Nov. Dec. 24m, If BMNH, 5m ZMHU, 5m USNM, 3m AMNH, 3m BMB, 2m AFEN, Im AME, Im MNHN, Im KWJH; Montan. Japi 2-3000m 3m BMNH; Monta. Sierra 3000m Oct. 5m, If BMNH; Mt. Serpa May Im JFL; Pedregosa 3000m Oct. 3m BMNH; Rio Aldaregas 8000' 9/11/39 Im BMNH; Valle 2200m 3m BMNH; West of La Pueblita 1500-1600m Aug. 2m BMNH; Tachira: Mata Mula P.N. El Tama 1850-2020m Mar. Im AFEN; Not located: no specific locality 2m BMNH, Im ZMHU, 2m BMB. COLOMBIA (69m, If): Boyaca: Muzo Im BMNH; Santa Maria Im ESM; Cundinamarca: Bogota 23m BMNH, 2m USNM; env. de Bogota 2m BMNH; Cananche Im BMNH; El Baldio 5400' Sep. 3m BMB; Fusagasuga Imf AMNH; La Vega 1900m Jan. Im BMNH; Mt. Redondo Feb. 2m AME; Quebrada Tasajera Feb. Im AMNH; U. Rio Negro 850m Im USNM; Meta: San Martin, Llanos of Rio Meta Im BMNH; Villavicencio Imf AMNH; Narino: Ricaurte 1600m Sep. Im ESM(error?); Santander: La Carmen, Rio Opon region Nov. Im AMNH; La Lechera Nov. Im AMNH; La Soledad Dec. Im AMNH; Rio Opon Dec. Im AMNH; San Gil Nov. Im, If JFL; Vaupes: Rio Vaupes 1400m Aug. Im AME; Not located: no specific locality 8m AMNH, Im BMNH, Im USNM, 2m BMB, 2m MNHN. COUNTRY UNKNOWN (7m, If): no specific locality 2m USNM, 2m MNHN, 3m, If MCZ. Adelpha aricia (Hewitson 1847) Figs. 22; 106 Identification, taxonomy and variation: Adelpha aricia is very similar to A. corcyra, and the only wing pattern character that I can find which reliably separates all taxa of each species is the shape of the VFW subapical marking, hi the subapical area of the VFW in A. aricia the inner and outer

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187 postdiscal series are straight in each cell space, notably in cell R5, and the basal edge of the inner postdiscal series is also straight. In the subapical area of the VFW of A. corcyra the inner and outer postdiscal series are composed of basally pointing arrow-shaped markings in each cell space and the basal edge of the inner postdiscal series is convex. In practice, however, there is little difficulty in identifying specimens since the two species are not known to be sympatric, and in the only area where sympatry is plausible (southern Ecuador and northern Peru), the taxa of each species are usually readily distinguished by other wing pattern characters. Other distinguishing characters include a generally more falcate forewing and produced hindwing tomus in A. aricia, silvery ventral markings in A. aricia serenita and A. a. portunus, and at the distal edge of the VHW white postdiscal band there is a curved red-brown marking in cell M3, but no brovm markings in cell Cul. Adelpha alala is easily distinguished by the two clearly marked, round or oval white postdiscal spots at the DFW costa. There is some variation in the shape and color of the DFW orange subapical marking and VHW markings in A. aricia, and three subspecies are recognised. Given the similar wing patterns, apparent close parapatry and similar habitats, I had considered that A. aricia and A. corcyra might be conspecific. However, the male genitalia are surprisingly different between the two species, the clunicula of A. aricia being high and pointed, that of A. corcyra low and broad, thus warranting their separation. Nevertheless, the two species are clearly close relatives and A. aricia is the probable sister taxon to the group comprising A. corcyra, A. tracta, A. pithys and A. donysa. Range and status: Eastern Andean slopes from north Peru (Amazonas) to Bolivia. In cloud forest habitats fi-om 1500-3000m. Not uncommon. Specimens examined: 174 (174m)

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188 Adelpha aricia aricia (Hewitson 1 847) Figs. 22a,b Heterochroa aricia Hewitson (1847: 263, pi. XXI, fig. 1 1) TL: Bolivia. Types: BMNH(T); STm; "Bolivia/Zaricia Hew.//B.M. TYPE No. Rh. 9811 Heterochroa aricia m Hew.//Syntype//Type"; BMNH(M): STm: "Bolivia Hewitson Coll. 79-69 Heterochroa aricia. 17/B.M. TYPE No. Rh. 9812 Heterochroa aricia m Hew./ZParatype" [both examined] Adelpha aricia Hew., Kirby (1 871) Identification, taxonomy and variation: The nominate subspecies differs fi-om A. aricia serenita by having slightly duller ventral coloration and by having the distal edge of the VHW white postdiscal band straight and bordered by a straight red-brown line in cells M3-M1; in A. a. serenita this red-brown line is formed of concave elements in each cell and disjointed slightly at each vein. Adelpha a. portunus differs in a similar manner on the VHW, and has reduced orange DFW subapical spots. The thickness and intensity of the orange postdiscal band on the DFW is variable to some extent, especially in cell Cul, while a few specimens have trace orange scaling in cell Cu2 over the inner submarginal series on the DFW. While I believe this variation to be infrasubspecific, for the same reasons given under A. alala negra there may yet prove to be phenotypically distinct and constant populations on isolated cordilleras which merit subspecific recognition. Hewitson (1847) described A. aricia based on Bolivian specimens in his own collection and the British Museum, and his detailed description, figure of the dorsal surface of the left hand wing pair and the two syntypes in the BMNH leave no doubt as to its identity. Range: The nominate subspecies occurs along tastem slopes of the Andes fi-om southern Peru to Bolivia. Habitat and adult ecology: Nothing appears to be published on the ecology of this species; judging fi-om museum specimens males are apparently uncommon, while the female appears to be unknown. The habits and habitat of this species are probably similar

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189 to those of the closely related A. corcyra. The nominate subspecies has been recorded from 1500m to 3000m. Specimens examined (72m): PERU (32m): Cuzco: Buenos Aires, Cosfiipata 23(X)m Dec. 2m MUSM; Callanga 1500m Im ZMHU; Marcapata 4500' 2m BMNH; Pillahuata May 2500m Jan. Im MUSM; Yanamayo 2-2200m Feb. May 2m MUSM; Junin: Chanchamayo 3m BMB-error; Puno: Agualani 10000' Oct. 3m BMNH; Limbani 10000' Nov. 8m BMNH; Oconeque-Agualani 6-9000' Mar. 2m BMNH; Oroya-Limbani Jan. Im BMNH; Quiton Carabaya 7000' Sep. Im BMNH; Rio Inambari-Limbani Mar. 4m BMNH; Not located: no specific locality 2in BMNH. BOLIVIA (39m): La Paz: Apolobamba 3m BMNH; Chaco 2-3000m Im ZMHU; Cillutincara 3000m Im ZMHU; Coroico Im MCZ; Cusilluni May Im MCZ; La Paz Apr. 2m BMNH, Im MCZ; Pararani-Mapiri 5200' Mar. Im AMNH; Unduavi Im BMNH; Cochabamba: Cochabamba 6m BMNH; 5 days N. Cochabamba Im MCZ; Incachaca 2220m Mar. Apr. 3m AME; San Jacinto Im BMNH, Im ZMHU; Not located: Cocopunco Im AMNH; Locotal 2600m 2m ZMHU; no specific locality 11 m BMNH, Im AMNH. COUNTRY UNKNOWN (Im): no specific locality Im BMNH. Adelpha aricia serenita Fruhstorfer 1915 stat. rest. Figs. 22c,d; 106a,b Adelpha aricia serenita Fruhstorfer (1915: 516) TL: Peru. Types: BMNH(T): STm: "aricia serenita Fruhst./ZFruhstorfer Coll. B.M. 1937-285//Fruhstorfer Coll. B.M. 1933131//Peru H. Fruhstorfer//TYPE//Type//Syntype" [examined] =A. aricia aricia Hew., Hall (1938) Identification, taxonomy and variation: This subspecies is distinguished from the nominate subspecies as discussed in that account. Adelpha aricia portunus has the orange DFW postdiscal band reduced to subapical spots in cells M2-R5. Variation in A. a. serenita is similar to that seen in the nominate. Fruhstorfer (1915) described this subspecies based on an unspecified number of specimens from Peru, one of which he figured as the dorsal surface of A. aricia (Fruhstorfer, 1913: pi. 108, a). He stated that it differed from the nominate in the thickness of the DFW orange subapical marking and in the pattern of the ventral reddish brown lines. I have examined a single syntype male in the BMNH. While the first of Fruhstorfer' s two characters appears to be unreliable, the second is not, and as discussed under the nominate subspecies the two taxa are consistently distinct. Hall (1938) failed to note the diagnostic ventral characters and synonymised serenita with A. aricia aricia, and I therefore reinstate serenita as a good subspecies (stat. rest.). Range: This subspecies is known from the eastern Andes in central Peru.

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190 Habitat and adult ecology: Presumably similar to nominate subspecies. The locality data indicating that this subspecies occurs as low as 800-900m are almost certainly erroneous. Specimens examined (67m): ? subspecies identification uncertain. "ECUADOR" (Im): no specific locality Im AME-error. PERU (63m): Ayacucho: Rio Piene, Apurimac 4m AMNH; Cuzco. Cuzco Im MCZ; Huanuco. Cushi 1820-1 900m 8m BMNH; Pozuzo 800m Apr. 4m BMNH, 3m MUSM, Im AME, Im BMB; Tingo Maria Mar. Im AME; Junin: Chanchamayo Jan.-Aug. 2m BMNH, 2m BMB, Im ZMHU; El Porvenir 900ni Apr. Im BMNH; Hda. Naranjal, N.E. Mina Pichita 1550-1650m Nov. Im MUSM; La Merced 2500-3000' Aug.-Nov. 3m BMNH, Im BMB, Im MUSM; Perene R. Mar. Im BMNH; Rio Colorado Im BMNH; San Luis de Shuaro Aug. 2m MUSM; Pasco: Huancabamba 6-10,000' Aug. 10m BMNH, Im BMB, Im MCZ; Not located: PN Puente Yanango Oct. Im MUSM; no specific locality 2m? USNM, 5m BMNH, 2m BMB, Im MUSM, Im MCZ. COUNTRY UNKNOWN (3m): no specific locality Im? USNM, Im BMNH, Im MNHN. Adelpha aricia portunus Hall 1938 Figs. 22e,f Adelpha aricia portunus Hall (1938: 209) TL: "Tambillo, Rio Tabacones etc., Peru". Types: BMNH(M): STm: "Syntype m Adelpha aricia portunus Hall G. Lamas det. 1987//EX. Oberthur Coll. Brit. Mus. 1927-3//Tambillo Peru Cordilieres J. Stolzmann 1878 9-II//Syntype"; 4STm: Chachapoyas; 2STm: Manchara; 2STm: Rio Tabaconas; 7STm: Tambillo; STm: no locality; BMB: STm: Rio Tabaconas; STm: Tambillo [all examined] Identification, taxonomy and variation: This subspecies is distinguished from A. aricia serenita by the reduced orange DFW postdiscal band. The orange subapical spots usually have white scaling at the basal edge, though this is variable, as is the size of the subapical spots, which may almost coalesce to form a band. There is a specimen in the MUSM from Libertad which has particularly broad orange markings in the DFW subapical area, lacking any white at the basal edge of the markings, but since these characters show variation in Hall's type series of portunus, some specimens of which approach the Libertad specimen in wing pattern, and I have seen only a single specimen of the latter I am unable to say whether it represents a distinct subspecies. Hall (1938) described this subspecies based on 17 specimens in the BMNH and 2 specimens in his own collection. I have examined all of these, except that I was only able to locate 16 syntypes in the BMNH. Range: This subspecies is known from the east Andean slopes of northern Peru, almost to the Ecuadorian border, sfrongly suggesting that it also occurs in the Cordillera de Lagunillas in southern Ecuador.

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191 Habitat and adult ecology: Presumably similar to the nominate subspecies. Specimens examined (35m): PERU (35m): Amazonas. Chachapoyas 4m BMNH; Cajamarca: Manchara 7000* Sqj. 2m BMNH; Rio Tabaconas 6000' 2m BMNH, Im BMB; Tambillo 7m BMNH, Im ZMHU, Im BMB; Libertad: Cumpang, Tayabamba-Ongon 2400-2700m Oct. Im MUSM; San Martin. La Playa, P.N. Abiseo 2480-2680m Jul. 7m MUSM; Macedonia, P.N. Abiseo Aug. 2m MUSM; Quebrada El Peligro.'p.N. Abiseo 2045m Aug. 2m MUSM; Rio Montecristo, P.N. Abiseo 2600m Aug. Im MUSM; Not located. Hda. Udima, Quebrada El Palmo 240O-260OTn Nov. 3m MUSM; no specific locality Im BMNH. Adelpha corcyra (Hewitson, 1847) Figs. 23; 107; 189 Identification, taxonomy and variation: Adelpha corcyra is very similar to some subspecies of A. aricia, but typically it has rounder hindwings and a duller brown VHW, and the irmer and outer postdiscal series which compose the pale VFW subapical marking consist of basally pointing arrowshaped markings in each cell, instead of straight dashes. The basal edge of the pale VFW subapical marking is also concave, instead of straight. The clunicula in the male genitalia is much shorter and broader than in A. aricia, confirming the distinctness of these two closely related, apparently parapatric species. Adelpha pithys and A. donysa are also similar but have more rounded hindwing shape, especially the distal margin which is scalloped, and the inner and outer postdiscal series on the VHW are even throughout the wing, not more pronounced in cells Rs and Sc+Rl, with a distinctly reddish brown ground color. Adelpha alala is distinguished by the two well defined white oval postdiscal spots in the DFW near the costa. There is geographic variation in the presence or absence of an orange postdiscal band on the DHW, the shape of the orange DFW subapical marking and width of the white postdiscal bands, and five subspecies are recognised, two of which are described here. The distinctive shape of the clunicula in the male genitalia shows that A. corcyra is most closely related to A. tracta, A. pithys and A. donysa. The first three of these four species are allopatric in montane islands. I regard A. corcyra as a distinct species fi-om A. tracta on the basis of the VHW character mentioned above to distinguish A. corcyra from

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192 A. pithys and A. donysa. This character is probably in its plesiomorphic state in A. corcyra{\i occurs also in this state in A. alald), suggesting A. tracta is more closely related to A. pithys. For a discussion of why I regard A. tracta as distinct from A. pithys, see the account under the former species. Range and status: Venezuela (Sierra de El Tama, possibly Merida) throughout Colombia, on both Andean slopes throughout Ecuador to the western slopes of extreme northern Peru. In intact cloud forest habitats from 1500-3000m, locally not uncommon. Probably Viburnum (Caprifoliaceae). Specimens examined: 290 (283m, 7f) Adelpha corcyra corcyra (Hewitson, 1 847) Figs. 23a,b; 107a,b; 189a,b Heterochroa corcyra Hewitson (1847: 262, pi. XXI, fig. 9) TL: New Grenada. Types: BMNH(T): STm: "Illustrated in The Butterflies of Venezuela A. Neild, 1996//36//corcyra Hewitson//B.M. TYPE No. Rh. 9808 Heterochroa corcyra m Hew.//New Grenada//Type//Syntype"; B!V1NH(M); ST?m: "N. Granada Hewitson Coll. 79-69 Heterochroa corcyra. 1 ."; ST?m: "N. Granada Hewitson Coll. 79-69 Heterochroa corcyra. 2." [all examined] Adelpha corcyra Hew., Kirby (1 871 ) Identification, taxonomy and variation: Adelpha corcyra corcyra is distinguished from all other A. corcyra subspecies except A. c. salazari by the orange postdiscal band on the DHW. It differs from A. c. salazari as described under that subspecies. The width of the orange postdiscal markings on the DHW varies slightly, being especially thick in one male from Minas de Muzo and one male from Bogota, both in the BMNH. Very occasionally the orange of the DFW subapical marking extends just into cell Cul, but never as sfrongly as in all specimens examined of A. c. salazari. Adelpha alala completa is also similar, but may be distinguished by the characters discussed in the species account for A. corcyra. Hewitson (1847) described corcyra based on specimens in the British Museum from New Granada, and his description and figure of the dorsal surface leave no question

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193 as to the identity of the taxon. I have examined 3 probable syntypes in the BMNH, all of which correspond to the original description. Range: Venezuela (Sierra de El Tama, possibly Merida) to Cordillera Oriental in Colombia as far south as Bogota. Although accurate locality data are few, in the Bogota area, this subspecies seems to occur in the mountains to the west of the city, while A. c. collina is found in the hills just behind Bogota. Habitat and adult ecology: Judging from the number of specimens in collections this subspecies is common, although if the record from Valencia in Venezuela is to be believed, it is very rare throughout the Merida range. There are no published observations of the subspecies in the field, but specimen label data indicate that it occurs from 22002700m, throughout the year. Specimens examined (99m, Sf): VENEZUELA (3m): Carabobo. Valencia Im BMNH; Tachira: San Vicente de la Revancha 2350m Mar. Im AFEN; Via [)elicias Tama Nov. ImAFEN. COLOMBIA (91m, 4f): fio^'ocd. Arcabuco 2200m Jan. Im, lfUSNM;Muzo ImBMNH, Im BMB; N.E. of El Arenal 2400-2500m Jul. Im BMNH; Parque Iquaque 2700m Jan. Apr. Im ESM, Im JFL; Santa Rosita Nov. If JFL; Cundinamarca: Barro Blanco, Madrid Nov. Im ESM; Bogota 23m, If BMNH, 4m USNM, Im MCZ; env. de Bogota Im BMNH; El Baldio 5400' Sep. 2m BMB; Pacho Im BMNH; Norte de Santander. P.N. El Tama, N. Santander 2500m Dec. Im JFL; Tolima. Rio Chili Apr. -June Im BMNH; Not located. Cauca Im ZMHU; Interior Im BMNH; Magdalena Valley 2m BMNH; Rio Poroeca Jul. If AMNH; no specific locality 20m AMNH, 8m BMNH, 7m MCZ, 3m ZMHU, 6m BMB, 2m MNHN. "BRAZIL" (Im): no specific locality Im BMB-error. COUNTRY UNKNOWN (4m, 10: 2m ZMHU, If AME, Im USNM, Im MNHN. Adelpha corcyra salazari Willmott, ssp. nov. Figs. 23c,d Description and diagnosis: FW length of HT: 25.5 mm. Males differ from the nominate subspecies in having a broader orange subapical marking on the DFW, which is not divided into separate spots in each cell space by darker veins, and is of an even width, instead of being narrowest in cell Ml. The basal edge of the marking is straight, not incised at each vein, and the marking extends into cell Cul to form a prominent triangular orange spot. Very occasionally this marking extends into cell Cul in the nominate subspecies, but only as indistinct orange scaling. The DHW postdiscal orange band is always relatively heavy, there is no thin pale brown band distal of the white postdiscal

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194 band on the DHW, and on the VHW the white postdiscal band is not lined at its distal edge with an uneven, reddish brown line. The female is similar to the male. Types: Holotype male: COLOMBIA: Magdalena: "Kancnia 2108 2450m 26.viii.l972//B.M. 1973-106"; in the BMNH(AandB). Paratypes: COLOMBIA: El Cesar. 1 male: "San Jose N. Colombia F. Simons//m//Adelpha corcyra Hew. Godman-Salvin Coll. 1916.-4."; in the BMNH(M); 1 male: "above Sogrome Rio Meoguachucua 17001800m 28.Vni.72//North Colombia: 2119 Sierra Nevada de Sta. Marta Exped. M.J. Adams and G.I. Bernard"; Magdalena: 5 males: "Above Kancnia (Tromba) 2500m 25.Vm.l972//North Colombia: 2077 Sierra Nevada de Sta. Marta Exped. M.J. Adams and G.I. Bernard"; 2 males: east above San Pedro de la Sierra 22-2500m Aug.; in the BMNH(AandB); 1 male, 1 female: Minca Magdalena 2400m Oct.; in the JFL; Not located: 1 male: Sta. Marta; in the BMNH(R). Etymology: This subspecies is named for my friend Julian Salazar, for his generous hospitality during my visit to Colombia and for allowing me to view and take notes from his collection. Taxonomy and variation: The Santa Marta population differs in several slight but consistent characters from the nominate subspecies, and given the reasonable series of specimens examined, and its geographic isolation from the nominate subspecies, I believe it justifies subspecific recognition. Range: Sierra Nevada de Santa Marta in northern Colombia. Habitat and adult ecology: This subspecies is known from 1700-2500m and has been collected in August and October. Judging from museum specimens it is locally not uncommon. Specimens examined (12m, If): COLOMBIA (12m, If): El Cesar. San Jose Im BMNH; above Sogrome Rio Meoguachucua S. N. de Santa Marta 17(X)-1800m Aug. Im BMNH; Magdalena: Kancnia 2450m Aug. Im BMNH; above Kancnia (Tromba) S. N. de Santa Marta 2500m Aug. 5m BMNH; Minca 2400m Oct. Im, If JFL; E. above San Pedro de la Sierra S. N. de Santa Marta 22-2500m Aug. 2m BMNH; Not located. Santa Marta ImBMNH.

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195 Adelpha corcyra aretina Fruhstorfer, 1907 Figs. 23 e,f; 107c Adelpha corcyra aretina Fruhstorfer (1907: 172) TL: Ecuador. Types; BMNH(M): STm: "Ecuad. Fruhstorfer/ZFruhstorfer Coll. B.M. 1 937-285//Syntype m Adelpha corcyra aretina Fruhstorfer G. Lamas det. 1987//Syntype"; ST?m: "Fruhstorfer Coll. B.M. 1937-285//Ecuad. Fnihstorfer/Zcorcyra collina Hew."; ST?m: "Ecuador ex. Coll. H. Fruhstorfer// Fruhstorfer Coll. B.M. 1937-285//colIina Hew." [all examined] Adelpha corcyra collina Hew., Fruhstorfer (1915) misid. Identification, taxonomy and variation: Adelpha corcyra aretina is most similar to A. corcyra dognini, and is distinguished under that subspecies. It is distinguished from A. c. collina by the orange DFW subapical marking ending in the anterior half of cell Cul, instead of extending at least to vein Cu2, and closely bordering the distal edge of the white postdiscal band, which is diagonal, sloping basally, instead of vertical.. Adelpha c. collina also has a pinkish cast to the ventral surface. Adelpha alala negra is similar but may be distinguished by the characters discussed under the species account for A. corcyra. Adelpha aricia portunus may also be very similar, and is discussed under A. corcyra dognini. Fruhstorfer (1907) described this subspecies through comparison with the nominate, from three Ecuadorian males in his collection, one collected by Richard Haensch, and two ex. Oberthur's collection. He stated that the orange DFW subapical marking was narrower (though this is not a constant character) as were white dorsal postdiscal bands, but mysteriously failed to mention the lack of the orange postdiscal band on the DHW. Later, however, he confirmed that the DHW lacked the orange postdiscal band (Fruhstorfer, 1915). There are three Ecuadorian specimens in the BMNH from Fruhstorfer's collection, two of which are labeled collina Hew., while all three lack any fiirther indication that they are syntypes of aretina. While these specimens may be true syntypes, which have been subsequently relabeled and thus lost data presented in the original description, I believe it is impossible to be sure. However, I am certain that the name aretina does indeed apply to the subspecies found from southern Colombia to

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196 central Ecuador, since Haensch only visited the eastern Andes in the Pastaza and Baeza areas during his trip to Ecuador (Haensch, 1903). He could have collected specimens of A. c. collina in western Ecuador, but since Fruhstorfer (1915) treated this taxon as a separate species, Adelpha epidamna, a synonym of collina, Fruhstorfer's aretina must apply to east Andean specimens of A. corcyra. Fruhstorfer (1915) misidentified typical aretina as collina (explaining why two potential syntypes listed above are labeled collina), and stated that the former differed from the latter in having narrower bands, a purely fictional difference presumably presented so that he would not have to synonymise his own name aretina. At any event, since collina applies to a completely distinct taxon, aretina remains a valid name. Range: The eastern Andean slopes from extreme southern Colombia to central eastern Ecuador. Habitat and adult ecology: This subspecies occurs from 1500-2700m in association with primary cloud forest habitats, where it is almost always found along rivers and appears to fly throughout the year. Males are uncommon and usually encountered as solitary individuals flying swiftly about rocky playas puddling at damp sand, particularly at urine, and are attracted to rotting fish hung in traps along river banks. I have seen no female specimens. Specimens examined (67m): COLOMBIA (2nn): Narino: El Socorro, area a Monopamba 1500m Feb. Im MHNM; Monopamba 1750m Feb. Im ESM. ECUADOR (64m): Morona-Santiago: Limon-Gualaceo rd., km. 22, 2050m Nov. Im KWJH, Im DAT; Napo: Archidona Apr. Im BMNH-error; Rio Arajuno 700m Mar. Im AME-error; Rio Chonta 2000m Oct. Im KWJH; "Pichincha": Aloag 7m USNM-error; Tungurahua. env. de Ambato 16m BMNH, Im BMB; Bafios 5-7000' 3m BMNH; Mirador Im AMNH; Rio Blanco Apr. May 4m AME; Rio Machay Jul. Aug. 2m KWJH; Rio Topo 1400m Apr. Im AME; Rio Verde 5000' Jan. Im BMNH; San Antonio 2100m Apr. Im AME; Yunguilla, nr. Bafios May Im USNM; Not located: Oriente 4m AMNH; Santa Barbara Aug. Im BMB; no specific locality 3m BMNH, 7m AMNH, 4m MCZ, Im BMB, Im JFL. "PERU" (Im): San Martin: Tarapoto Im BMNH-error. Additional locality data: ECUADOR: Morona-Santiago: Lim6n-Gualaceo rd., km. 22, Sep.; Sucumbios: Rio Palmar Nov.; El Higueron Nov.; Tungurahua: Rio Machay Feb.; Rio Ulba 2700m Nov. (Willmott and Hall, unpubl. sight records). Adelpha corcyra dognini Willmott, ssp. nov. Figs. 23g,h Description and diagnosis: FW length of HT: 27 mm. Males differ from A. corcyra aretina by having a broader orange subapical marking on the DFW, of more even width.

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197 which has the veins colored orange within the marking instead of dark brown, particularly vein Ml. The white postdiscal bands are broader on both wings and that on the DFW tapers from anal margin to cell M2, whereas in A. c. aretina the white block in cell Cul is of the same width as that in cell M3. The DHW is darker in the distal half due to the expansion of the black band distal of the white postdiscal band, so that the pale brown area surroimding the triangular, blackish submarginal spots is restricted to a thin line closely bordering these spots, instead of a pale brown band. I have seen no female specimens. This subspecies is also very similar to certain specimens of A. aricia portunus, but has more rounded wings and can be distinguished as discussed under the species account for A. aricia. Types: Holotype male: ECUADOR: "Ecuador Loja Dec. 1892 (P. Dognin) 1915-209"; in the BMNH(M). Paratypes: ECUADOR (17m): Loja: 1 male: same data as HT; 1 male: "Environs de Loja Equateur 83//32. 21. Ex. Coll. Dognin 1921//Joicey Bequest Brit. Mus. 1934120"; 1 male: "Environs de Loja Equateur 1891//Joicey Bequest Brit. Mus. 1934-120"; in the BMNH(M); 1 male: San Francisco Aug.; 2 males: env. Loja Jul.; in the BMB; 1 male: "Environs de Loja Equateur 1889//Collection W. Schaus"; 1 male: "Environs de Loja Equateur 1889//Edw. T. Owen Collection"; 1 male: "Loja Equateur Fevrier '86//Edw. T. Owen Collection"; 1 male: "Environs de Loja Equateur '87//Collection W. Schaus"; 1 male: "Loja Equateur Fevrier '86//Collection W. Schaus"; in the USNM; Zamora-Chinchipe: 1 male: Quebrada San Ramon 1700m Oct.; in the KWJH; 1 male: "Zamora Ecuador 3-4000' (O.T. Baron)"; in the BMNH(R); 1 male: "Zamora, Equateur 1885 Abbe Gaujon/ZCoUection W. Schaus"; in the USNM; Not located: 1 male: "S. Ecuador//m//Adelpha collina Hew. Godman-Salvin Coll. 1916.-4."; 1 male: "Ecuador Hewitson Coll. 79-69 Heterochroa collina. 4."; 1 male: "Ecuador Hewitson Coll. 79-69 Heterochroa corcyra. 3."; in the BMNH(M). PERU (Im): Piura: 1 male: "Peru, Pi,

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198 Camino Japalache-El Carmen, entre Batan y Machete, 17-VI-80 2150m L.J. Barkley"; in the MUSM. Etymology: This subspecies is named for P. Dognin, who collected several of the type specimens. Taxonomy and variation: This subspecies differs from A. c. aretina in several respects, but nevertheless I deliberated for some time before deciding to describe it. Specimens of A. c. aretina from the Limon-Gualaceo road in eastern Ecuador, midway between the Pastaza and Zamora valleys, have slightly thicker DHW white postdiscal bands than typical A. c. aretina, and are probably showing intergradation with A. c. dognini. The single known Peruvian specimen has a slightly broader white postdiscal band on the DHW than typical A. c. dognini, and it thus might be argued that the differences observed between A. c. aretina and A. c. dognini are all just part of clinal variation from north to south. However, since two characters, the broader orange subapical marking and anteriorly tapering DFW postdiscal band, consistently distinguish the two taxa in the relatively large series examined of both, and since both have relatively extensive geographic ranges in which there is little, if any, clinal variation, I believe the southern population merits subspecific recognition. Range: This subspecies occurs in southern Ecuador on the eastern slopes in the Zamora valley, south to extreme northern Peru on the eastern slopes near the Ecuadorian border. Habitat and adult ecology: This subspecies is locally common in the Zamora valley, but apparently rarer in Peru, and has been reliably recorded from 1700-2 150m. It is confined to cloud forest habitats, where males behave similarly to other subspecies and may be encountered as solitary individuals puddling at urine along rivers. Specimens examined (19m): ECUADOR (18): Loja. Environs de Loja Jul. 2m BMNH, 2m BMB, 3m USNM; Lpja Feb. Dec. 2m BMNH, 2m USNM; San Francisco Aug. Im BMB; Zamora-Chinchipe: Quebrada San Ramon 1700m Oct. Im KWJH; Zamora 3-4000' Im BMNH, Im USNM; Not located. S. Ecuador Im BMNH; no specific locality 2m BMNH. PERU (Im): Piura: Camino Japalache-El Carmen, entre Batan y Machete 2150m Jan. Im MUSM. Additional locality data: ECUADOR: Zamora-Chinchipe: Quebrada San Ramon Oct. (Willmott and Hall, unpubl. sight records).

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199 Adelpha corcyra collina (Hewitson, 1 847) stat. rest. Figs. 23i j Heterochroa collina Hewitson (1847: 262, pi. XXI, fig. 10) TL: Quito [Ecuador], Types: BMNH(T): LTm: "B.M. TYPE No. Rh. 9809 Heterochroa collina m Hew.//Quito//Type//Syntype"; BMNH(M): PLT?m: "Ecuador Hewitson Coll. 79-69 Heterochroa collina. 1."; PLT?m: "Ecuador Hewitson Coll. 79-69 Heterochroa collina. 2//B.M. TYPE No. Rh. 9810 Heterochroa collina m Hew./ZParatype"; PLT?m: "Ecuador Hewitson Coll. 7969 Heterochroa collina. 3."; PLT?in: "Ecuador Hewitson Coll. 79-69 Heterochroa collina. 4." [=dognmi\ [all examined] =Heterochroa epidamna C. and R. Felder (1867: 424) TL: Bogota, Nova Granada [Colombia]. Types: BMNH(R): STm: "Syntype//Type//Bogota Lindig Type/Zepidamna Felder/Zepidamna n." [examined] Another Im in BMNH(R) "Bogota Lindig". Adelpha collina Hew., Kirby (1871); Adelpha collina var. epidamna Fldr., Kirby (1871); Adelpha epidamna Fldr., Fruhstorfer (1915); Adelpha corcyra collina Hew., Fruhstorfer (1915)referring to /I. corcyra aretina; Adelpha collina Hew., Hall (1938) Identification, taxonomy and variation: Adelpha c. collina is distinguished from all other A. corcyra subspecies by having the orange subapical marking on the DFW touching the white postdiscal spots in cells Cul and M3. There is local variation in whether the marking extends into cell Cu2 or terminates at vein Cu2, and the marking may be broken into small, isolated spots in each of cells M2-R5. Some specimens of ^4. aricia may be similar, but the orange subapical marking is always approximately straight in that species. Hewitson (1847) described and clearly figured this subspecies based on specimens in the British Museum and his own collection fi-om Quito. There is only a single specimen in the BMNH which bears the label "Quito", and this came from Hewitson's collection and closely matches the original figure, notably in having isolated orange subapical spots on the DFW. Since Hewitson's description was clearly based on more than one specimen, and one potential syntype in the BMNH actually belongs to a different taxon {A. corcyra dognini), I designate as the lectotype of Heterochoa collina the Quito specimen menfioned above with the following label data: "B.M. TYPE No. Rh. 9809 Heterochroa collina m Hew.//Quito//Type//Syntype". It appears that Felder and Felder (1867) mistakenly believed A. corcyra corcyra to be Hewitson's A. c. collina, since in their original description of the name epidamna they stated that it differed from A. c. collina in lacking orange postdiscal dashes on the DHW, while a single specimen in the BMNH(R) of A. c. corcyra from the Felder collection is identified as A. c. collina. The

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200 description of epidamna was based on an unspecified number of male specimens collected by Lindig at Bogota, and a syntype is in the BMNH. The original description refers to the poorly defined white postdiscal band on the VFW, a distinctive characteristic of this taxon. The syntype of epidamna differs fi^om that of collina in having a broad, continuous orange band as the DFW subapical marking, but since this character is variable within populations I regard the former name as synonym of the latter. A. corcyra collina has historically been treated as a distinct species (Kirby, 1871; Fruhstorfer, 1915; Hall, 1938; D'Abrera, 1987), but since the genitalia do not differ fi-om typical A. corcyra, there are no fimdamental wing pattern differences and it occupies similar habitats to A. corcyra, I treat it as a subspecies. Fruhstorfer (1915) placed the name collina as a subspecies oiA. corcyra, but this was based on a misidentification and he was actually referring to A. c. aretina. To date there are no known cases of sympatry of this taxon with other A. corcyra subspecies, and although there are areas of very close parapatry, particularly Bogota, I have seen no specimens indicating intergradation. Further collecting may just possibly demonsti-ate this taxon to be sympatric with other A. corcyra subspecies and therefore a distinct species. Range: There are few accurate distributional data for this subspecies, and more are needed to determine the Colombian range more precisely. It occurs in Colombia on both slopes of the Cordilleras Occidental and Central, and in the Cordillera Oriental as far north as the mountains just behind the city of Bogota. It seems to be replaced in the mountains north and west of Bogota by the nominate subspecies, and in the exti-eme south on the eastern Andean slopes by A. c. aretina. It also occurs on the western slopes of the Andes throughout Ecuador. Habitat and adult ecology: This subspecies seems slightly more tolerant of disturbed habitats than A. c. aretina, but in its behavior and preference for riverine microhabitats, and in its occurrence in cloud forests fi-om 1500-3000m, it is very similar. In western

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201 Ecuador, I have found males to be locally not uncommon and attracted to rotting fish in riverside traps and horse dung, and to puddle along river banks, especially at urine. Specimens examined (86m, If): "PANAMA" (Im): Chiriqui: Lino 800m Fassl Im BMNH-error. COLOMBIA (72m, 10: Antioquia: Antioquia 2m, If ZMHU; Las Palmas 8000' Jul. Im BMB; Frontino 2m BMNH; Sonson 2300m Aug. Im JFL; Valdivia Im BMB; Caldas: Bocatoma, Manizales 2100-2400m Mar. Jul. Im MUSM, 2m MHNM, Im KWJH; Cerro Guadalupe Manzanares 2400m Nov. 2m KWJH; Manizales 4m BMNH, Im BMB; Caquela: Caquetii May Jun. 4m JFL; rd. to Florencia Mar. Im ESM; Cauca: Cerro Munchique Im AMNH; Paletega 3000m Dec. Im ESM; Silvia 2500m Jul. Im LMC; nr. Tacueyo 5500' Cauca Nov. Im BMNH; Cundinamarca: Bogota 2700-2900m Jan. Mar. Nov. 3m BMNH, 2m ESM, Im AMNH, Im USNM; Bogota paramo 3000m Aug. Im AMNH; San Miguel 3000m May Im AMNH; Usquen, Bogota 3000m Feb. 3m AMNH; Naririo: Pasto 2000' Im MNHN; Piedrancha Dec. 3m AMNH; Putumayo: Pitalito-Mocoa Apr. Im ESM; Quindio: Res. Nat. Acaime, Salento 27(X)m Jan. Im MHNM; Salento, Rio Quindio 2800m Jan. Aug. 3m LMC; Risaralda: Pereira Im BMNH; Tolima: Caflon del Tolima Im MCZ; Rio Ambeima 2000m Jun. Im AME; Not located: Cauca Im ZMHU; Cauca, Tom6 Jan. Feb. Aug. 16m BMNH; Cord. Oriental 3000m Im AMNH; El Chico 3000m Im AMNH; no specific locality Im USNM, Im BMB. ECUADOR (10m): Carchi: Santa Rosa, Rio Golondrinas Sep. Nov. 2m KWJH; Pichincha: Quito Im BMNH; Rio Las Palmeras 1900m Aug. 3m KWJH; Tandapi Aug.lm KWJH; Not located: no specific locality 3m BMNH. "PERU" (2m): Junin: Chanchamayo Im BMNH, Im BMB-error. COUNTRY UNKNOWN (Im); no specific locality ImBMNH. Additional locality data: ECUADOR: Carchi: EI Coraz6n 2700m Sep.; Pichincha: Rio Chisinche May (Willmott and Hall, unpubl. sight records). Adelpha tracta (Butler, 1872) Figs. 24a,b; 108a,b; 190a,b Heterochroa tracta Butler (1872b: 102, pi. 38, fig. 6) TL: Cartago, Costa Rica. Types: BMNH(T): STm: "B.C.A. Lep. Rhop. Adelpha tracta Butl. Godman-Salvin Coll. 1916.-4.//B.M. TYPE No. Rh. 9814 Heterochroa tracta m Butl.//H. tracta Butl. Type//m//Costa Rica Van Patten Druce Coll.//Type//Syntype"; BMNH(M): ST?f: "B.C.A. Lep. Rhop. Adelpha tracta Butl. Godman-Salvin Coll. 19I6.-4.//CosU Rica Van Patten Druce Coll.//r [examined] Adelpha tracta Butl., Kirby (1877) Identification, taxonomy and variation: Adelpha tracta is a distinctive species, similar perhaps only to A. levona and A. rothschildi, which have the DFW orange postdiscal band split into separate spots and lack the third discal cell bar on the VFW. There is little variation in the specimens examined. Butler (1872b) described and clearly figured this species based on specimens of an unspecified number and sex collected by Van Patten in Cartago, Costa Rica, in Druce's collection. I have examined a syntype male and probable syntype female in the BMNH. Adelpha tracta is a member of the A. alala group, all of which have the third discal cell bar visible on the VFW, in addition to other less obvious characters. The broad and short clunicula in the male genitalia place it in a group which includes A. corcyra, A. pithys and A. donysa. Among these species, the reduction or loss of the entire inner postdiscal series on the VHW and reddish ground color of the distal half of the VHW

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202 groups A. tracta, A. pithys and A. donysa. The similar hindwing shape and the shape of the DFW orange subapical marking suggests that A. tracta and A. pithys are sister taxa, and I treat them as distinct species on the basis of the fusion of the inner postdiscal series with the postdiscal band in cells Cul and M3 in A. tracta, a character unique to the A. serpa group and rare among primitive Adelpha. I have been unable to find any consistent genitalic differences between the two species, or indeed between any members of the A. corcyra group. Range: Eastern Nicaragua to western Panama, in montane areas. Hostplant: Unidentified genus of Rubiaceae in Costa Rica (Haber and Chacon, In: DeVries, 1987); Viburnum costaricanum (Caprifoliaceae) in Costa Rica (Haber, pers. comm.). Haber (pers. comm.) informs me that he believes the record of Rubiaceae is probably erroneous, since he has numerous oviposition and feeding records on Viburnum, but none on any rubiaceous plant. Viburnum is the expected hostplant, given the close relationship of this species with A. alala. Immature stages: Haber and Chacon {In: DeVries, 1986) report solitary late instar larvae feeding on all leaves of plants along forest edges. The pupa collected by Haber in the PJD apparently hatched after 16 days. Habitat and adult ecology: DeVries (1987) reports this species in Costa Rica to be widespread and common in all forest habitats, fi-om 800-3000m. Solitary individuals are encountered along road cuts and forest edges during periods of sun, and males perch fi-om l-3m high on vegetation until the early afternoon. Both sexes are attracted to rotting finits of Lauraceae and Melastomaceae, and males feed at wet sand and mammal dimg. Judging from specimen label data these observations no doubt apply throughout the range of the species. A single male in the AMNH was apparently captured at lights at night. Specimens examined: 81 (50m, 3 If) NICARAGUA (8m, 20: Managua. Managua Oct. 2m USNM; Matagalpa: Matagalpa Aug.-Oct. Dec. 5m, If AMNH, If AME; Not located: no specific locality Im BMNH. COSTA RICA (17m, 140: Alajuela: Volcdn Poas Mar. Apr. Im BMNH, Im, If USNM; Cartago: Azaher de Cartago If BMNH; Cerro de la Muerte 5000' Sep. Im AME, Im FSCA; El Alto 5000' Oct.-Dec. 2f BMB; Irazii 6-7000' Im BMNH; Tres Rios 5000' Jan. Im, If USNM; Heredia: Rio Sarapiqui 1300m Jun. Im USNM; Puntarenas: Las Alturas 4700' Mar. coll. at light J.S. Miller Im AMNH; Monteverde 4500' Jul. Im AME; San Jose: Bajo La Hondura 1200m Aug. If AME; Carrillo Sep. Oct. 2m BMB; Rio Sucio Im BMNH; Not healed: Agua Caliente Sep. Im BMNH, If BMNH; no specific locality 2m,

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203 5f BMNH, Im ZMHU, If BMB, Im, If AMNH. PANAMA (25m, 130: Chiriqui: Chiriqui Feb. 5m, 3f BMNH, 4m, If ZMHU, 2m USNM, 2m BMB, 2m AMNH; Potrerillos 3600' Jan.-Mar. 2f FSCA, If AME, 2m, 3f USNM; Potrero Seco, Chiriqui Im, If BMNH; Santa Clara 1200m Sep. Im USNM; Valle de Chiriqui 3-4000' 3m, If BMNH; Volcan Bart 1800m Feb. Apr. 2m, 2f USNM; Veraguas: no specific locality Im BMNH; Not located: no specific locality If MNHN. "COLOMBIA" (10: Cundinamarca. Bogota If BMNH-error. COUNTRY UNKNOWN (10: no specific locality If USNM, Im, 2f MCZ. Adelpha pithys (H.W. Bates, 1864) Figs. 25a,b; 109a-c; 191a,b Heterochroa pithys Bates (1864: 128) TL: Guatemala. Types: BMNH(T): STm: "pithys Bates//B.C.A. Lep. Rhop. Adelpha pithys Bates Godinan-Salvin Coll. 1916.4.//B.M. TYPE No. Rh. 9822 Heterochroa pithys m Bates//m Guatemala tableland/ZCentr. Valleys Guatemala F.D.G. and O.S.//Type. sp. figured/ZType H.T."[examined] =Adelpha pithys vodena Fruhstorfer (1915: 524) stat. nov. TL: Guerrero, Mexico. Types: BMNH(M): 2ST?m: "Guerrero Mexico (O.T. Baron)//Adelpha "pithys pithys" det. R.I. VaneWright I989//ex. Rothschild ColI.//Rothschild Bequest B.M. I939-I" [both examined] Adelpha pithys Bates, Kirby (1 87 1) Identification, taxonomy and variation: Adelpha pithys is very similar to A. donysa, from which it is distinguished by its smaller size, by having the area between the first and second VFW and VHW discal cell bars only slightly paler or the same color as the discal cell ground color (in A. donysa this area is reddish brown, as is the discocellular area), and by the pale postdiscal shading which represents the upper postdiscal band on the VFW being cormected to the lower postdiscal band, so that there is a continuous thin brovra line from the costa to vein M3 bordered on both sides by pale shading. In A. donysa the upper postdiscal band is typically present as pale shading only in cells Ml and R5, sometimes slightly extending into cell M2, but almost never so as to form a continuous brown line within the pale area as in A. pithys. The inner and outer submarginal series on the VHW of A. pithys are separated only by a thin reddish brown line, whereas in A. donysa this line is thicker. Other characters which distinguish the two species outside of Guatemala (see under A. donysa for dicussion of Guatemalan specimens) include in A. pithys the more pointed hindwing, the broader hindwing band and the narrower DFW orange subapical marking which does not have so smooth a basal edge. Adelpha corcyra lacks the rich reddish coloration of the ventral surface, in addition to having several other distinguishing characters which are discussed under that species. The venfral surface of A. pithys varies

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204 slightly in the amount of silvery coloration, otherwise this species is quite phenotypically stable. Bates (1864) described this species from a male specimen or specimens, in comparison with A. iphiclus, and his description clearly mentions several distinguishing features such as the third cell bar in the VFW discal cell. Godman and Salvin (1884) later stated that they were figuring the type specimen which they had collected near Duenas, and their illustration corresponds closely with the syntype in the BMNH(T). Fruhstorfer (1915) described vodena as a subspecies of A. pithys, stating that it was larger than typical specimens, with a "considerably narrower" hindwing postdiscal band, darker ventral colors and "prominent reddish-brown delimitation of the white discal band" on the VHW. However, he gave no information on the number, sex or whereabouts of the type specimens, only that they were from Guerrero in Mexico. Miller and Miller (1970) concluded that Fruhstorfer was unaware of true A. donysa and redescribed it as vodena, since A. donysa generally differs from A. pithys in also having a darker ventral surface and thinner white postdiscal bands, and synonymised vodena with donysa. However, I have examined two specimens in the BMNH from Guerrero which are indeed slightly larger than the type of A. pithys, with slightly narrower hindwing postdiscal bands and a conspicuous red-brown coloration distal of the VHW postdiscal band. These are the only specimens, except for a single male of typical A. donysa in the BMNH, that I have seen from Guerrero in a collection which could have been examined by Fruhstorfer, and I conclude that they may well represent the specimens on which vodena was based. Fruhstorfer was prone to exaggeration, particularly when describing new taxa, and it seems that he was aware of true A. donysa, which had been figured by Hewitson (1847), as he discusses accurately its distinguishing features (Fruhstorfer, 1915). I therefore conclude that vodena is a synonym of pithys (stat. nov.), since Guerrero specimens do not differ in any consistent manner from specimens from elsewhere in the range.

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205 Adelpha pithys is probably the sister species to A. tracta, and my reasons for retaining these two allopatric taxa as distinct species are discussed under the latter species. Range: Adelpha pithys is known from montane areas from Mexico to Guatemala, with a single record from Honduras which is plausible but requires confirmation. Habitat and adult ecology: De la Maza (1987) reports this species from cloud forest and occasionally pine-oak forest in Mexico, and de la Maza and de la Maza (1993) give a range of 1300-2800m in Chiapas. Godman and Salvin (1884) reported the species from as low as 700m in Guatemala. Specimen label data indicate that it flies from March to December at least, and is locally common. Specimens examined: 188 (lS2m, 36f) MEXICO (33m, 210: Chiapas. Comitin Mar.-Jun. Sep. 4m, 9f AME; Jaltenango 1000m Apr. Im AME; Lagos de Montebello 13001500m Feb. 4m AME; Mt. Huitepec 8000' Mar. Im AME; Ocozingo 2m AME; Pichucalco May 2m, If AME; San Carlos Jul. 4m AME; San Jeronimo Dec. 2m, If AME; no specific locality Jul. 2m, 6f AMNH, If AME, Im MCZ; Guerrero: Acahuizotla May Oct. 2m AME; El Faisanal, Paraiso Dec. Im AME; no specific locality 2m BMNH; Oaxaca. Candelaria Loxicha Nov. Im AME, Im AMNH; Oaxaca Im, If AMNH; Oaxaca,192 km. from, Dec. If AMNH; Portillo del Rayon 1500m Aug. Nov. 2m AME; Veracruz: El Vigia Aug. If AME. GUATEMALA (112m, 15f): Aha Verapaz: Baleu 1350m Jun. Aug.-Oct. 29m, 6f AMNH, 33m, If AME, Im MUSM, Im MHNM; Polochic Valley Im, If BMNH; Rio Sacaya Sep. Im USNM; San Cristobal 4000' May 3m BMNH, Im BMB; Tactic Im USNM; Tamahu Oct. 8m, 2f AMNH; Antigua: Antigua Jul. Aug. If BMB; Chimallenango: Quisache Jul. Im AME; Yepocapa 2m AMNH; Guatemala: Guatemala city Im BMNH; Quezaltenango: Acatenango Aug. Oct. 2m, If AMNH; Volcan Santa Maria 4-6000' Apr. Oct. Nov. 5m, If BMNH, 9m, 2f USNM, 5m BMB; Retalhuleu: San Felipe 2000' Aug. 3m BMB; Solola: vie. L. Atitlin Apr. Im MCZ; Solala Im BMNH; Zacapa: Zacapa Vail. Oct. Im BMB; Not located: Central Valleys Im BMNH; Chuchumatanes Mts. 1500m Mar. Im AME. HONDURAS (Im): no specific locality Im ZMHU. "PANAMA" (Im): Chiriqui: Chiriqui Im BMNH-error. COUNTRY UNKNOWN (5m): no specific locality 5m USNM. Additional locality data: MEXICO: Chiapas: Selva Lacandona (de la Maza and de la Maza, 1993); Guerrero: Nueva Delhi; Oaxaca: Jalatengo (de la Maza, 1 987). Adelpha donysa (Hewitson, 1 847) Figs. 26; 110; 192 Identification, taxonomy and variation: Adelpha donysa is closely related and can be very similar to A. pithys, and its diagnostic characters are discussed under that species. There is geographic variation in the width of the white DHW postdiscal band and DFW orange subapical marking, and two subspecies are recognised.

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206 Wing pattern and male genitalia indicate that A. donysa belongs to a group of closely related, montane species, which includes A. pithys, A. tracta, A. corcyra, A. aricia and A. alala. This group is one of the most primitive of the genus. Range and status: Mexico to western Honduras, in montane areas from 1800-2700m. Rare in collections but apparently locally common in the field. Specimens examined: 68 (52m, 16f) Adelpha donysa donysa (Hewitson, 1847) Figs. 26a,b,e,f; 110c; 192a,b Heterochra donysa Hewitson (1847: 260, pi. XX, fig. 5) TL: Mexico. Types: BMNH(M): LTm: "B.M. TYPE No. Rh. 9821 Heterochroa donysa m Hew.//Mexico Hewitson Coll. 79-69 Heterochroa donysa. 1. //Donysa W.H.//Syntype"; PLTm: "Mexico 44. 13.//Syntype"; PLTm: "Mexico 44. 13.//Syntype//T.G.H. 1962-541"; PLTf: "Syntype//type//Mexico 44. 13.//B.M. TYPE No. Rh. 9820 Heterochroa donysa f Hew.//Donysa Hewitson" [all examined] =Heterochroa roela Boisduval (1870: 46) XL: Mexico. OTL: Honduras; Mexico. Types: BMNH(T): LTm: "Ex. Musaeo Dris. Boisduval//Ex. Oberthflr Coll. Brit. Mus. 1927-3//=Heterochroa pithys Bates//Vu par Godman en Janvier 1883//Roela Bd. Mexiq.//Heterochroa Roela, Bdv. Mexique//Mexique//Typicum Specinien//Type//Syntype" [examined] Adelpha roela Boisd., Kirby (1871); Adelpha donysa Hew., Kirby (1871) Identification, taxonomy and variation: Adelpha donysa donysa is easily distinguished from A. d. albifilum by the wider DHW white postdiscal band which extends almost to touch vein 2A. It is very similar to A. pithys, and distinguishing characters are discussed under that species. There is a reasonable amount of variation in Mexican specimens in the width of the white dorsal postdiscal bands, the width of the orange subapical marking and the overall size. Miller and Miller (1970) figure a particularly extreme example from Hidalgo, in which the orange DFW subapical marking is almost equal in width to the white DFW postdiscal band. However, there is a specimen in the syntypic series of donysa with the white band almost as thick, in addition to a specimen from Calderas, Guatemala, and since the variation appears to be largely within rather than between populations, I do not recognise distinct subspecies within Mexico.

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207 » Hewitson (1847) described this species based on specimens in his collection and the British Museum from Mexico, and his figure of the dorsal surface of the left hand wing pair establishes beyond doubt the identity of the taxon. There are several syntype specimens in the BMNH, none of which have accurate locality data. The width of the white postdiscal band is slightly variable in this series, and since fiiture more extensive collecting may demonstrate the existence of several consistently distinct populations within Mexico, I designate as lectotype of Heterochroa donysa the following specimen in the BMNH(M), which closely matches Hewitson's original figure: "B.M. TYPE No. Rh. 9821 Heterochroa donysa m Hew .//Mexico Hewitson Coll. 79-69 Heterochroa donysa. 1. //Donysa W.H.//Syntype". Boisduval (1870) introduced the name roela based on a single specimen from Honduras and a single specimen from Mexico, both in the collection of M. de I'Orza in Paris. The only specimen I have seen to date o{ A. donysa from Honduras is clearly the same taxon as albifilum, but this specimen is not from Oberthur's collection, where Boisduval 's collection passed, and therefore cannot be a syntype of roela. Moreover, the original description is sufficiently detailed to ascertain that Boisduval w£is describing a specimen which belonged to the nominate subspecies. He stated that the DHW had a very narrow white discoidal ray which terminated towards the tomus (in albifilum it terminates in the middle of the wing), that this ray was not exactly aligned with that on the DFW (similarly to nominate donysa), and that the DFW orange subapical marking terminated "brusquement" at vein Cul (in albifilum it continues across the vein well into the middle of cell Cul). 1 therefore conclude that the Honduras specimen was either mislabeled or represents a population in Honduras which has remained uncollected ever since. Godman and Salvin (1884) examined the second, Mexican, syntype and synonymised roela with donysa, in which they were followed by subsequent authors (Fruhstorfer, 1915; Miller and Miller, 1970), and to avoid any possible fiiture confiision I designate this specimen, with the following label data, as the lectotype of Heterochroa roela: "Ex. Musaeo Dris. Boisduval//Ex. Oberthur Coll. Brit.

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208 Mus. 1927-3//=Heterochroa pithys BatesWu par Godman en Janvier 1883//Roela Bd. Mexiq.//Heterochroa Roela, Bdv. Mexique//Mexique//Typicum Specimen//Type//Syntype". In the BMNH(M) are two male specimens from the Polochic Valley in Guatemala, which Godman and Salvin (1884) referred to as A. donysa. These specimens bear the following data: "Polochic Valley F.D.G. and O.S.//m//B.C.A. Lep. Rhop. Adelpha donysa Hew. Godman-Salvin Coll. 1916.-4.". They differ from typical A. donysa in having thicker white postdiscal bands, a more pointed hindwing shape similar to A. pithys, a much thinner orange DFW subapical marking, again similar to A. pithys, and the VHW submarginal series are closer together, but not as close as in typical A. pithys. They do however have the characters typical of A. donysa which are discussed under A. pithys, and since typical A. pithys is also known from the Polochic valley I believe the specimens may well represent a new subspecies of A. donysa. Since I have also examined specimens of typical A. donysa from the same region, it may be that subspecies in this region are confined to small montane islands, but the specimens I have examined have insufficiently precise locality data to examine this possibility. Range: Montane areas from Mexico to Guatemala. Habitat and adult ecology: This species is rare in collections, and de la Maza (1987) reports it from November to May in montane semi-deciduous forest and occasionally pine-oak forests, from 1800-2700m (de la Maza and de la Maza, 1993). Miller and Miller (1970) report the species to be locally common in Hidalgo, in Mexico, in association with deciduous, scrub oak-juniper habitat, and suggested that due to the sfrong dry season there that A. donysa might be univoltine. They state that individuals sunned themselves in the morning up to 10.00 a.m., either on leaves or the ground with the wings open, and also perched on prominent leaves with the wings held almost closed, whence they made brief sorties to investigate passing butterflies. Later in the day the butterflies were only seen near the tops of the trees, and due to a close association with oaks by the adults, they

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209 suggested that this might be the hostplant. This may be the case, but possibly a more likely hostplant is Viburnum, which is utilised by the closely related species A. alala and A. tracta. Specimens examined (24m, 1 If): MEXICO (20m, 80: Chiapas: Mt. Huitepec 8000' Mar. Im AME; Rancho Belem Mar. If AME; San Cristobal de las Casas Mar. Sep. 2m FSCA, If AME; Guerrero: Omiltene, wooded mountains 1700-2000m Jul. Im BMNH; Hidalgo: Zimapan, 5 mi. N.W., 2140m pinon-oak-juniper scrub Jan.-Mar. Im BMNH, 1 Im, 4f AME; Mexico: Valle de Bravo Nov. Im AME; Oaxaca: La Esperanza 1750m Mar. If AME; Not located: no specific locality 4m, If BMNH, Im MNHN. GUATEMALA (2m, 30: Alta Verapaz: Baleu If AMNH; Tactic Jul. If USNM; Jutiapa: Calderas Im BMNH; Quezallenango: Volcan Santa Maria Apr. If USNM; Not located: Chicnamac Im AMNH. Additional locality data: MEXICO: Chiapas: Macizo Central; Sierra Madre (de la Maza and de la Maza, 1993); Guerrero: Filo de Caballo; Teotepec; Hidalgo: La Encamacion; Oaxaca: Portillo del Rayo (de la Maza, 1987). ssp. nov.? GUATEMALA: (2m) Alta Verapaz: "Polochic Valley F.D.G. and O.S.//m//B.C.A. Lep. Rhop. Adelpha donysa Hew. Godman-Salvin Coll. 1916.-4." 2m BMNH(M). Adelpha donysa albifilum Steinhauser, 1974 stat nov. Figs. 26c,d; 110a,b Adelpha albifilum Steinhauser (1974: 2, figs. 1-4, 57, 58) TL: Hda. Montecristo, Cerro Miramundo Cloud Forest, El Salvador. Types: AME: HTm: Hda. Montecristo 2418m Mar.; 26PTm, 4PTf; Cerro Miramundo Cloud Forest 2300-241 8m Dec. -Apr. [all examined) See Steinhauser (1974) for list of remaining type specimens. Identification, taxonomy and variation: This subspecies is easily distinguished from other A. donysa subspecies by the narrow DHW postdiscal band, which terminates in the middle of the wing, in addition to the relatively broad DFW orange subapical marking which extends into cell Cul and the dark ventral surface, particularly the reduced or absent hindwing submarginal series. The width, and therefore the length, of the DHW postdiscal band is slightly variable, and in some specimens it may be entirely absent. Steinhauser (1974) described this taxon as a full species from 42 male and 4 female specimens, all captured at the type locality, and I have examined the holotype and a long series of paratypes at the AME. The principal justifications for considering the taxon a distinct species were the wing pattern differences and putative genitalic differences, Steinhauser stating that the climicula in A. albifilum was prominent, while that in A. donysa was absent, and that in A. pithys was only "indicated as a serrate edge". I have examined specimens of all three of these taxa and can find no consistent differences

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210 in the genitalia, and I conclude that Steinhauser's comments were based on the genitalic figures in Miller and Miller (1970), which he references. In those figures the clunicula is omitted in A. donysa, while in A. pithys it is "indicated as a serrate edge". The width of the DHW postdiscal band and DFW orange subapical band is variable both in A. d. albifilum and in the nominate subspecies, and since A. d. albifilum exhibits no fundamental wing pattern differences from nominate A. donysa, and occurs allopatrically in similar habitats, I regard the two taxa as conspecific (stat. nov.). Range: This subspecies is known from a small area in El Salvador, with a single record from Honduras, presumably in the extreme west and probably on the Pacific slope. Habitat and adult ecology: Steinhauser (1974) reports that males of this subspecies were frequently seen at the type locality hilltopping at the summit of Cerro Montecristo, where the cloud forest is replaced by a heath scrub. When hilltopping, males remained on low sunny perches beneath the top of the scrub, while in the forest they could be observed flying in simny areas in the understorey or visiting wet sand, mud and dog dung in clearings. Similarly to the nominate subspecies, males were sometimes seen perching on the road. The subspecies is evidently locally common and has been recorded from November to May, from 2300-2400m. Specimens examined (28m, 50^ EL SALVADOR (28m, 40: Metapan: Cerro Miramundo Cloud Forest 2300-241 8m Dec.-Apr. 28m, 4f AME. HONDURAS (IQ: Noi healed: Montagne de la Sapote, Dept. de Copay If BMNH. Adelpha fessonia (Hewitson, 1847) Figs. 27; 111; 193 Identification, taxonomy and variation: Adelpha fessonia is the only Adelpha species in which the DFW postdiscal band is white and continues unbroken from the costa to the anal margin. There is variation in A. fessonia in the width of the dorsal postdiscal bands and DFW subapical marking, the

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211 extent of reddish-brown scaling in the DFW discal cell, and the color of the ventral surface, and four subspecies are recognised, one of which is described here. Adelpha fessonia is one of the most primitive members of the genus, and despite the differences in dorsal wing pattern, it may well be the sister species to A. gelania. The valvae of the male genitalia in both of these species are of a similar shape and lack spines, an uncommon character amongst other primitive Adelpha, although the female genitalia of A. gelania, which differ from most primitive Adelpha in lacking sclerotised bands on the corpus bursae, suggest that A. gelania and A. fessonia have been isolated for a relatively long time. « Range and status: Extreme southwestern U.S.A. to northeastern Venezuela, and the island of Hispaniola, in deciduous forest and cloud forest habitats, from sea level to 1950m. Common in western parts of its range, rarer in the east. Most populations appear to be disjunct. Not known to date from apparently suitable deciduous forest habitats in central and southwestern Ecuador, but may well occur there. Specimens examined: 307 (173m, 134f) Adelpha fessonia fessonia (Hewitson, 1847) Figs. 27a,b; llla,b; 193a,b Heterochroa fessonia Hewitson (1847: 260, pi. XX, fig. 6) TL: Honduras. Types: BMNH(T): STf: "B.M. TYPE No. Rh. 9817 Heterochroa fessonia f Hew.//fessonia Hewitson//Honduras//Type//Syntype" [examined] Adelpha fessonia Hew., Kirby (1871); Limenitis fessonia Hew., Scott (1986); Adelpha fessonia Hew., DeVries (1987) Identification, taxonomy and variation: Adelpha fessonia fessonia is distinguished from A. f. emestoi under that subspecies, and from A. f. cestus by the much narrower white postdiscal bands, the broader orange DFW subapical marking, and by lacking any reddish-brown scaling in the DFW or DHW discal cell distal of the basal streak. Several other Adelpha species are also superficially similar, but A. fessoniamay be distinguished by the unbroken white DFW

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212 postdiscal band. There is substantial individual, or possibly seasonally induced, variation in the width of the white postdiscal band, which may be as little as half the width of the syntype specimen. All of the specimens reared by Bosese in Texas, U.S.A, have a much darker ventral surface than typical, but it is not clear whether this is due to seasonal effects or not (see comments under A. f. cestus and A. f. lapithd). Hewitson (1847) described this species from an unspecified number of specimens from Honduras in the British Museum, and the syntype female in the BMNH(T) closely matches his original figure. Range: Exfreme southwestern U.S.A. (Texas) to western Panama. Although most authors report this species to be a "stray" in Texas, it clearly breeds there (see Hostplant below). Immature stages: Scott (1986) reports this species in Texas on Celtis lindheimeri (Ulmaceae), and Neck (1996) repeated this record and added Celtis laevigata. However, both state that the early stages are unknown, suggesting that these are oviposition records. Since Celtis is the foodplant of the superficially similar Doxocopa pavon and Doxocopa laure, which both also occur in Texas, and neither Celtis nor Ulmacaeae have otherwise been recorded for A. fessonia throughout the remainder of its range, I regard these records as highly dubious. Late instar larvae in Costa Rica are solitary and feed on all leaves of plants along forest or river edges and in forest light gaps (Janzen, In: DeVries, 1986). There is a series of 6 males and 9 females in the FSCA, reared by R. W. Bosese in Texas, U.S.A., all with the pupal case and some with the head capsule of the last instar larva. All were reared from eggs collected on November 12 1994, and the total development times in days (to adult eclosion) are as follows: 43 (Im, If), 44 (Im), 46 (Im), 47 (If), 50 (If), 52 (Im, 2f), 55 (Im, 2f), 56 (Im), 58 (2f). Habitat and adult ecology: This is a common species throughout most of its range in deciduous forest habitats, occurring only rarely in rain forest at the western edge of its range (DeVries, 1987; de la Maza and de la Maza, 1993). The species has been reported in Chiapas, Mexico, from sea level to 1500m in deciduous forest (de la Maza and de la

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213 Maza (1993), and from premontane forest further north in San Luis Potosi at 1200m, where it is rare and has been recorded only in Ocotober (de la Maza and de la Maza, 1988). DeVries (1987) reports that the species occurs in Costa Rica from sea level to 900m on the Pacific slope of the country, particularly in the lowlands of Guanacaste. Although it flies throughout the year, it is most common in the wet season and in reproductive diapause during the dry season. Both sexes feed at flowers of Cordia and Croton, and may be attracted to rotting finits. Specimens examined (160m, 1 180: U.S.A. (10m, 130: New Mexico: no specific locality Im AMNH; Texas: Bentsen-Rio Grande Valley State Park Hidalgo Co. Jun. Im FSCA; Brownsville Aug. Im AME; Granjeno Hidalgo Co. Sep. 2f FSCA; Madero Hidalgo Co. Dec. Jan. 6m, 9f FSCA; Pharr May Im AMNH; McAllen Hidalgo Co. Oct. If FSCA; Santa Ana Res. Hidalgo Co. If FSCA. MEXICO (99m, 560: Campeche: Campeche Im AMNH; Chiapas: Caflon el Sumidero Jun. Im MUSM; Cuauhtemoc Oct. Im AME; Las Delicias Jun. Im AMNH; Malpaso, km. 25 to, Im AMNH; Ocozocoautla Jan. Im AMNH; Rancho Santa Ana Im AMNH; San Quintin Sep. Im AME; Tonalas Jan. Im AMNH; Colima: Colima 2m, If AMNH, Im BMNH; Comala 2100' May If AME; Guerrero: Acapulco Dec. If AMNH; El Playon Sep. If AME; Zihuatenango Apr. 2m, 2f AMNH; no specific locality 2m, If BMNH; Jalisco: Ayutla-Tierra Colorada 350m Jul. Im BMNH; Chamela Bay Mar. Nov. 4m, If AMNH; Guadalajara Nov. 4m BMNH; Puerto Vallarta Oct. Im USNM; Tenacatitla Bay Apr. If AMNH; Michoacan: Coahuayana Jul. 2m, If AME; Playa Azul Dec. If AMNH; Nayarit: Lag. Maria del Oro 670m Aug. Im AME; Playa Noverillos Jun. Im, 2f AMNH; Tepic 210m Aug. Im BMNH, 3m, If AME; Zapata 900m Aug. If AME; Oaxaca: Oaxaca If BMNH; Oaxaca, 192 km. fi-om, Dec. Im AMNH; Tapantepec 150m Feb. If AME; San Luis Potosi: El Pujal If AMNH; Valles Im AMNH; Sinaloa: Concordia 200m Aug. If AME; El Dorado Sep. Im AMNH; Mazatlan 30m Aug.-Oct. 10m, If AME, 2f AMNH; Santa Cruz Im AMNH; no specific locality 4m USNM; San Luis Potosi: 3 mi. N. Ciudad Valles If FSCA; Tamaulipas: Ciudad Victoria Jan. 2m USNM; Col. Victoria Oct. Im AME; El Limon Mar. Im USNM; Mante, 30 mi. N., Nov. Im USNM; San Fernando 60m 2m AME; Tampico Dec. Im AMNH; Victoria, 60 mi. S., Jul. 2m AMNH; Veracruz: Cordoba Jun. Jul. If BMB; Fortin Im ZMHU; Jalapa 4000' Nov. 3f AMNH, Im BMNH, Im, If USNM, If BMB, If MCZ; Nuevo Morelos Nov. Im AME; Orizaba If BMNH; Palmas Reales Jun. If AMNH; Palo Gacho Oct. If AMNH; Paso San Juan If USNM; Presidio Aug. If AMNH, If BMNH; Tezonapa Jun. Im, If AME; Vera Cruz Im ZMHU; Yucatan: Acanceh Jul. Im AMNH; Chichen Itza Jul. 3f AMNH; Chuminopolis Jul. Im AMNH; Halacal Sep. If AME; Mdrida Jul. Aug. If AME, If AMNH; Pist6 Jul.-Sep. lOm, If AMNH, Im BMNH, 9m, 3f AME, 3m, If USNM, If BMB; no specific locality Im, If ZMHU, Im USNM, If BMB; Not healed: W. Mexico If BMNH; no specific locality 2m, 2f BMNH, Im, If USNM, Im MCZ, If AMNH. GUATEMALA (20m, 110: Chiquimula: Chiquimula Im MNHN; El Progreso: Motagua Valley Im BMNH; Escuintla: Escuintla Jul. Aug. If USNM, 3m, If BMB; Palin Jul. Aug. 2m, 2f BMB; Quezaltenango: Volcan Santa Maria Im BMNH; Relalhuleu: San Sebastien Im, 2f USNM; Santa Rosa: Chiquimulilla Jul. 3m BMNH; Guazacapan Jun. Dec. 6m, 2f BMNH, 3f AME, Im BMB; Zacapa: Gualan Jan. Im USNM. EL SALVADOR (6m, 70: La Libertad: La Libertad 10m Jan. Im, 2f AME; San Salvador: Apopa 2000' Sep. Im, If BMNH, Im BMB; San Salvador Jan. Im, If AME, Im MNHN; Santa Tecla 900m Oct. 2f AME; Zaragoza Jan. If AMNH; Not located: Mina San Juan 600m Mar. Im AME. HONDURAS (30: Cortes: San Pedro Sula 2f BMNH; Not located: no specific locality If BMNH. NICARAGUA (4m, 60: Chontales: Chontales If BMNH; Managua: Managua Dec. If AME; Malagalpa: Matagalpa Im BMNH; Not located: no specific locality 3m, 2f BMNH, 2f MNHN. COSTA RICA (13m, 140: Alajuela: San Mateo Oct. If BMNH; Cartage: Irazu 6-7000' Im BMNH; Guanacaste: Avangarez Jul. If BMNH; Cailas, 24 km. N.W., on flwrs. of Cordia sp. Jun. Im AMNH; Hac. La Pacifica 50m Nov. Jan. 2m USNM; Las Caflas Jul. If AME; San Antonio 3000' Mar. Im BMNH, 3m AME; Santa Rosa lOOm Aug. Im, If KWJH; San Jose: Escazii Jan. 2m, If BMNH; Villa Colon 750-800m Aug. Im AME, If USNM; Not located: no specific locality Im, 7f BMNH, If AMNH. PANAMA (5m, 20: Chiriqui: Armuelles If AMNH; David Im BMNH; Potrerillos 3600' Jan. If USNM; Los Santos: Los Santos Jan. Dec. 4m USNM. COUNTRY UNKNOWN (3m, 60: no specific locality 4f MCZ, ImBMNH, Im, 2fUSNM, Im BMB. Additional locality data: MEXICO: Chiapas: El Chorreadero 700m Jan. (Beutelspacher, 1983); Depresion Central; Estribaciones de los CuchumaUnes; Istmo Seco; Macizo Central (de la Maza and de la Maza, 1993); Colima: numerous records (Warren et al., 1998); Quintana Roo: Carrillo Puerto; Chumpon (de la Maza and Bezaury, 1992); Jalisco: numerous records (Vargas et al, 1996); San Luis Potosi: Sierra de Alvarez 1200m Oct. (de la Maza and de la Maza, 1988); Tamaulipas: Antiquo Morelos Jul.; Forion Sep. (Field, 1940b). BELIZE: Corozal: Shipstem Nature Reserve (Meerman, 1999). Adelpha fessonia ernestoi Willmott, ssp. nov. Figs. 27c-e Adelpha cestus Hew., D'Abrera (1987) misid.

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214 Description and diagnosis: FW length of HT: 26iTim. Both sexes of this subspecies differ from the nominate as follows: the DFW orange subapical marking is much narrower and does not extend into cell M3 as a slight "hook" at the outer postdiscal series, there is slight reddish brown scaling in the DFW discal cell between the first and second cell bars (absent in the nominate subspecies), and the DFW white postdiscal band is less angled at vein M3. Adelpha f. cestus is distinguished by having much broader white postdiscal bands, with the postdiscal and submarginal series on the VHW thus closer together. Types: Holotype male: COLOMBIA: El Cesar. "Colombia L. Sapatoza Region, Chiriguana Dist. C. Allen//Brit. Mus. 1925-576"; in the BMNH(M). Allotype female: COLOMBIA: "Nouv. Grenade de Bogota a Buenaventura Dr. O. Thieme 14 Dec. 77 au 22 Fev. 78//Ex. Oberthur Coll. Brit. Mus. 1927-3"; in the BMNH(M). Paratypes: COLOMBIA: Cundinamarca: 1 female: Finca Guanabana, Fusagasuga 1400m; in the ESM; Magdalena: 1 female: "4 km. S.W. of Atanquez 24.Vn.1972 1950metres//North Colombia: 1808 Sierra Nevada de Sta. Marta Exped. M.J. Adams and G.I. Bemard//B.M. 1 973106//6 1.1 7/72 A/B"; in the BMNH(AandB). Etymology: This subspecies is named for Ernesto Schmidt-Mumm, for his kindness and hospitality during my visit to Colombia, allowing me to take notes from his collection and for his capture of a specimen of this subspecies which established a reasonable geographic range. Taxonomy and variation: There is little variation in the few specimens examined. It might be argued that this subspecies represents merely a step in a cline from A.f. cestus to A. f. fessonia, and indeed it is true that it is intermediate in wing pattern. However, it seems to occupy a sufficiently large range over which it is phenotypically stable, and to be separated by a sufficiently large phenotypic gap from each neighboring subspecies, to warrant its subspecific recognition. The variation discussed within A. f. cestus by Neild (1996) is minor compared with the phenotypic differences between that taxon and A. f.

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215 ernestoi, Venezuelan specimens always having distinctly wider postdiscal bands (Neild, pers. comm.). It is apparently isolated from the nominate subspecies by central and eastern Panama, presumably due to an absence of suitable habitat, while it is also isolated from A. f. cestus by the Sierra de Perija. Further, there appear to be possible differences in preferred microhabitat between this subspecies and A. f. cestus. Range: Colombia, on the western slopes of the Cordillera Oriental, from Fusgasuga north to the foothills of the Sierra Nevada de Santa Marta. Habitat and adult ecology: There are no published observations on this subspecies in nature, but judging from the number of specimens in collections it is very rare. In contrast to other subspecies, it is only known from montane areas from 14001950m. Specimens examined (Im, 3f)COLOMBIA (Im, 3f): Cundinamarca: Finca Guanabana, Fusagasug^ 1400m If ESM; El Cesar. L. Sapatoza Region, Chiriguana Dist. Im BMNH; Magdalena. Atanquez, 4 km. S.W., S. N. de Santa Marta 1950m Jul. If BMNH; Not located, de Bogota a Buenaventura Dec.-Fev. If BMNH. Adelpha fessonia cestus (Hewitson, 1847) Figs. 27f-h Heterochroa cestus Hewitson (1847: 261, pi. XXI, fig. 7) TL: Venezuela. Types: BMNH(T): STf: "niustrated in The Butterflies of Venezuela A. Neild, 1996//158//B.M. TYPE No. Rh. 9818 Heterochroa cestus f Hew./Zcestus Hewitson/A'enezuela//Type//Syntype"; BMNH(M): STf "Illustrated in the Butterflies of Venezuela, Neild 1996//159//B.M. TYPE No. Rh. 9819 Heterochroa cestus f Hew.Z/Venezuela Hewitson Coll. Heterochroa cestus. 1 .//Cestus W.H.Z/Paratype" [both examined] Adelpha cestus Hew., Kirby (1871); Adelpha fessonia cestus Hew., Neild (1996) Identification, taxonomy and variation: Adelpha f. cestus is distinguished from other subspecies by the very broad white postdiscal bands and reduced orange DFW subapical marking, and from the nominate by having additional reddish scaling in the DFW and DHW discal cells between cell bars one and two, and four and the postcellular bar. There is some variation in the white postdiscal and submarginal series on the ventral surface, in some specimens these being almost absent on the hindwing, resulting in a much darker appearance similar to A. f. lapitha. A similarly darkened ventral surface occurs in the specimens of the nominate subspecies in the FSCA from Texas, and I suggest it is due to seasonal effects, though

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216 there are insufficient temporal data to test this theory. According to Neild (1996), the width of the white postdiscal bands varies from west to east, specimens with the broadest bands occurring in the east in Sucre state. I have not seen sufficient specimens in collections to determine whether this variation is purely clinal or subspecific; Neild (1996) mentions specimens in the collection of Benito Gutierrez from Altagracia de Orituco (Guarico) which show variation between the exfremes of the eastern and western forms, suggesting a possible blend zone. Hewitson (1847) described this subspecies based on an unspecified number of specimens in his collection and the British Museum from Venezuela. The original figure closely matches the two syntype females in the BMNH, both of which were figured by Neild (1996). A third female specimen of cestus, in the BMNH(R), with the label data "Lyra//cestus Hew.//Type//Venezuela, Moritz/ZFelder Colin.", is not syntype but probably had a type label attached in the belief that it was the type of "lyra", an unpubUshed manuscript name. Adelpha f. cestus has been treated by all authors prior to Neild (1996) as a distinct species. However, it has no important wing pattern differences and the genitalia are essentially the same, and A. f. ernestoi, described below, forms a phenotypically intermediate population to the nominate subspecies, as noted by Neild (1996). Range: Venezuela, east and west of the Merida range, throughout the Cordillera de la Costa to Sucre state. Although Boisduval (1870) refers to this taxon, as Heterochroa cestus, as being present in Honduras, this is either a misidentification of A. f. fessonia, or A. s. celerio, since he refers to Cramer's (1777) figure of Papilio iphicla (which is, in fact, A. serpa diadochus). Habitat and adult ecology: This subspecies is rare in collections but apparently not uncommon in the field, flying along tracks in disturbed, semi-deciduous forest, where individials may be seen visiting flowers (Neild, 1996). It has been recorded from sea level to 900m from September to February.

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217 Specimens examined (9m, 110' VENEZUELA (8m, 1 If): Aragua: El Limon 450m Feb. 2m MUSM; Puerto Cabello 2m ZMHU; Dislrilo Federal: Caracas 31(X)' Sep Oct. Feb. Im AME, Im USNM, 5f BMB; Guarico: Los Morros de Macaira Dec. 700m If AFEN; Trujillo: Trujillo Im AFEN; Not located: no specific locality Im, 3f BMNH, If MNHN, If AMNH. COUNTRY UNKNOWN (Im): no specific locality Im BMB. Additional locality data: VENEZUELA: Apure; Sucre (Neild, 1996). Adelphafessonia lapitha Hall, 1929 stat. nov. Figs. 27iJ; 111c Adelpha cestus lapitha Hall ( 1 929: 1 32, pi. m, fig. 4) TL; Colombia. Types: BMNH(T): HTm: "Ex. Musaeo Oris. BoisduvaI//Ex. OberthQr Coll. Brit. Mus. I927-3//Problt.-la forme Colombienne de Fessonia God. Salv. Biolog. pi. 29 fig. 18.19.//Lapitha BD. Colombie//Columb.//Type//Holotype" [examined] Adelpha lapitha Hall, Schwartz (1989a), Smith et al. (1994) Identification, taxonomy and variation: Compared to other subspecies, A. f. lapitha has a much darker ventral surface (but see discussion below), particularly basal to the postdiscal band, the first cell bar in the VHW discal cell does not continue outside the cell to touch vein Sc+Rl, but is separated fi-om that vein by a white streak, the orange DFW subapical marking is darker and closer to the apex, and there is an orange spot in cell Cu2 distal of the white postdiscal band. Both sexes of A. f. lapitha also have prominent reddish scaling between cell bars one and two, and four and the postcellular bar, in the DHW and DFW discal cells. This character is present but less pronounced in A. f. cestus, even less pronouced in A. f. emestoi, and absent in the nominate subspecies. There is little variation in the few specimens examined. Hall (1929) described this species as a subspecies of A. (fessonia) cestus, presumably on the basis of the similar orange DFW subapical marking and very dark ventral surface which occurs in some individuals of A. f. cestus. He supplied a black and white photograph of the dorsal surface of the holotype male, which bears the erroneous locality data of "Colombia". Schwartz (1989a) speculates that Hall himself may have captured the holotype specimen and subsequently mislabeled it, but this is not the case since it arrived in the BMNH fi-om Boisduval's collection, possibly sent along with a number of Central American specimens from which Boisduval (1870) described several

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218 new Adelpha taxa. The rarity of the species and the false type locality led to its omission from works on the Caribbean fauna (Riley, 1975), and it was thus not until Schwartz (1989a) that the taxon was again reviewed. Schwartz (1989a) placed A. f. lapitha as a species distinct from A. (fessonia) cestus and A. fessonia fessonia, on the basis of several wing pattern and genitalic characters. However, the comparison between the male genitalia of A. f. lapitha, A. f. fessonia, A. abyla and A. iphicliis iphimedia, is ftmdamentally flawed by clearly mislabeled genitalia. Schwartz (1989a) claims that the genitalia of A. abyla do not differ from A. f. lapitha, while those of A. f. fessonia bear seven recurved teeth, and those of A. iphiclus are described as having numerous straight teeth and various spines. The description of the genitalia of A. f. fessonia clearly indicates that these belong to either A. abyla or A. iphiclus, neither of which differ in genitalia, while those assigned to A. abyla clearly belong in fact to A. f. fessonia. I am unable to even guess at what species the genitalia assigned to A. iphiclus belong to, certainly not to that species though. The genitalia of A. f. lapitha and all mainland A. fessonia in fact show no important differences. Schwartz (1989a) went on to discuss distinctive wing pattern characters in A. f. lapitha, such as the narrow postdiscal band, the lack of VHW postdiscal and submarginal series, the distinctive configuration of the VHW discal cell bars, and the orange submarginal spot in cell Cu2 on the DFW. The first two of these characters vary within mainland subspecies, whereas the latter two occur only in this taxon. However, I have examined a specimen of A. f. cestus in the AME which has a very dark venfral surface, almost entirely lacking whitish coloration except for the postdiscal band distal of the second discal cell bars, exactly as in /I. / lapitha. Furthermore, the inner submarginal series on the VFW in cells Ml and R5 are faded and diffuse, extending to the subapical marking, zsmA.f lapitha, and the inner discal cell bar on the VHW is curved distally in cell Sc+Rl to almost run alongside vein Sc+Rl, intermediate between the configuration of this bar in typical A. f. cestus and A.f lapitha. The orange spot in the DFW tomus represents the outer postdiscal series, which is variably present in subspecies

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219 of other primitive Adelpha, such as A. alala and A. corcyra. I therefore consider all of the wing pattern characters that separate A. f. lapitha and mainland A. fessonia to be relatively weak, and since all of these taxa are defined by the the sharing of a continuous white postdiscal band on the DFW, I place lapitha as a subspecies of A. fessonia (stat. nov.). The shape of the orange DFW subapical marking and reddish scaling in the DFW and DHW discal cells, both primitive states relative to the state in A. f. fessonia, suggests that A. f. lapitha is most closely related to either A. f. cestus or A. f. emestoi, or, more likely, the most ancestral taxon of A. fessonia. Range: The island of Hispaniola, to date known only from the Dominican Republic, with several sites near the Haitian border indicating its almost certain occurrence in that country. Habitat and adult ecology: The following comments are condensed from Schwartz (1989a). This subspecies is known from a variety of habitats, including pine and mixed pine-hardwood forest, xeric-mesic transitional forest and xeric scrub and Acacia forest. It has been recorded almost throughout the year from 1801900m, although the majority of records are below 600m in the months of June and August. Most individuals have been seen flying along forest edges or feeding on flowers, which attract both sexes, including Croton barahonensis (Euphorbiaceae) and Ageratum conyzoides (Asteraceae). Specimens examined (3m, 2f): "COLOMBIA" (Im): no specific locality Im BMNH. DOMINICAN REPUBLIC (2m, 20: San Juan: Vallejuelo, 9 km. E., 2000' Aug. Dec. 2m, 2f AME. Additional locality data: DOMINICAN REPUBLIC: Azua: Azua, 2.5 km. W., 6.6 km. N., 183m Jun.; Peralta, 5 km. S., 305m May Jun.; Dajabon: vie. Restauracion 550m Sep.; nr. Villarpando, 30 km. N. junction Hwy. 2 and Hwy. 44, 370m Mar.; Pedemales: Banano, 1 km. S.E., 488m Jun. (sight record); San Juan: vie. Piramide 204 Sierra de Neibe 1900m (Schwartz, 1989a,b). Adelpha gelania (Godart, [1824]) Figs. 28; 112; 194 Identification, taxonomy and variation: This is a distinctive species which can be confused with no other Adelpha. The dorsal surface bears a remarkable resemblance however to that of the Asian limenitidine.

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220 Sumalia daraxa Moore, but the patterns are produced through modification of different wing pattern elements; in A. gelania the white spots in cells Ml and R5 on the DFW are derived from the inner and/or outer postdiscal series, in S. daraxa they represent the upper postdiscal band. Adelpha gelania is one of the few species in the genus which has white postdiscal series on the DFW, and the absence of many typical pattern elements on the VFW, such as the upper postdiscal band and postdiscal series with the exception of the subapical spots, are distinctive characters. There is geographic variation in the pattern in the DFW discal cell, the orientation of white spots on the DFW, the VHW coloration and the presence of a "tail" at vein Cul on the hindwing, and two allopatric subspecies are recognised. Adelpha gelania is a primitive species within the genus, probably most closely related to A. fessonia, with which it shares reddish bands in the DFW discal cell, a third discal cell bar on the VFW, a similar pattern of lines in the cells anterior and posterior of vein 3 A on the VHW, a line joining the base of the VFW to the tip of the humeral vein, and similar male genitalia, which lack distal spines on the valvae. Range and status: The islands of Puerto Rico and Hispaniola. In dry and mesic forests up to 1700m. Locally common. Specimens examined: 51 (24m, 27f) Adelpha gelania gelania (Godart, [1824]) Figs. 28a,b; 112a,b; 194a,b Nymphalis gelania Godart ([1824]: 378) TL: Amerique equinoxiale. Types; MNHN?: ST: not located. Heterochroa gelania Godt., Westwood (1850); Adelpha gelania Godt., Kirby (1871) Identiflcation, taxonomy and variation: Adelpha g. gelania is distinguished from A. g. arecosa by having, on the DFW, reddish scaling which fills only the anterior half of the space between the first and second discal cell bars, reddish scaling over the DFW discocellulars, and more evenly spaced

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221 white subapical spots in cells M2-R5. On the DHW the orange marking in the tomus is less extensive, not extending into cell Cul, there is no pronouced "tail" at vein Cul, on the VHW the outer postdiscal series is completely absent, and on the VFW the whitish scaling in cell R5 is more extensive than the corresponding spot on the DFW. There is little variation. Godart ([1824]) described this species based on an unspecified number of specimens from "Amerique equinoxiale". I have been imable to locate with confidence any syntype specimens in the MNHN, the only specimen in that collection of this taxon is a clearly ancient female specimen with a handwritten label "H. Gelania, God." and a modem label "Museum Paris, Mexique". However, the original description, in Latin and French, is sufficiently detailed to unambiguously recognise this distinctive taxon. In particular, the French description mentions three reddish-brown marks at the base of the DFW, corresponding to the bands in the discal cell, whereas A. g. arecosa has only two such marks. At the time, the Dominican Republic was a French colony, whereas Puerto Rico was Portugese, and therefore the former was the only plausible locality within the range of this species where the type the specimen(s) could have originated (Lamas, pers. comm.). Lucas (1835: pi. 68) figured this taxon shortly after its description and there has never been any doubt as to its identity. Range: The island of Hispaniola. Habitat and adult ecology: Schwartz (1989b) reports that this species is locally not uncommon in the Dominican Republic but rare in Haiti. It is most commonly encountered in mesic deciduous forests at high elevations, but it also occurs in dense xeric forests in the lowlands and occurs in both pine and mixed pine hardwood forests. It has been recorded from near sea level to 1700m and flies throughout the year, though it is most abundant in July and August. Individuals have been observed from 7.30a.m.-4.00pm, and typically rest or perch in frees 3-4m above the ground. Two specimens, presumably males, were recorded puddling on rocks in a stream bed. Smith et al. (1994) report that

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222 individuals typically fly along paths near the forest edge, usually within 2m of the ground, with alternating periods of wing beating and gliding. They suggest that the dorsal wing pattern renders the butterfly inconspicuous when flying in sun dappled areas. The species has apparently never been observed at flowers or on any other food source. Specimens examined (23m, 17f): DOMINICAN REPUBLIC (I3m, 15f): La Vega: Jarabacoa, 19 km. S., 3675' Jun. Sep. 2f FSCA; Pedernales: Aceitillar 1 130m Jul. 9m, 8f AMNH; Las Abejas 1 160-1250m Jun. Jul. Im, 3f AME; Not located: Chacquay Feb. Im AMNH; Isla Saona, San Domingo Jan. If MCZ; Pico Duarte 3800' Apr. Im KWJH; Sto. Domingo Im, If BMNH. HAITI (10m, If): La Visite and vie. La Selle Range Sep. Im MCZ; Port-au-Prince Im BMNH, Im MCZ; no specific locality 6m, If BMNH, Im MCZ. "MEXICO" (IQ: no specific locality 1 f MNHN-eror. Additional locality data: DOMINICAN REPUBLIC: numerous records in Schwartz (1989b); Not located: woodland above and west of Constanza; nr. Playa Bavaro (Smith et al., 1994). HAITI: Not located: Foret de Pins, 4 km. N.W., 1500m (Schwartz, 1989b). Adelpha gelania arecosa (Hewitson, 1847) Figs. 28c,d Heterochroa arecosa Hewitson (1847: 263, pi. XXI, fig. 12) TL: Mexico; West Indies. Types: BMNH(T): STf: "Hewitson Coll. 79-69 Heterochroa gelania. 1.//B.M. TYPE No. Rh. 9834 Heterochroa arecosa f Hew./Zarecosa W.H.//Syntype//Type" [examined] ^Heterochroa gelania Godt., Westwood (1850); =Adelpha gelania Godt., Kirby (1871); Adelpha gelania arecosa Hew., Fruhstorfer (1915); Adelpha arecosa Hew., Brown and Heineman (1972); Adelpha gelania arecosa Smith et al. (1994) Identification, taxonomy and variation: This subspecies is distinguished fi-om the nominate under that account. There is little variation. Hewitson (1847) described this taxon from specimens in his own collection from Mexico and the West Indies, and the syntype female in the BMNH closely matches the original figure. It seems he was unaware of Godart's gelania, as he did not mention the name in the original description. Soon after Westwood (1850) placed the name as a synonym of A. gelania, until Fruhstorfer (1915) resurrected the name and treated it as a subspecies. Ironically, later workers often regarded the taxon as a distinct species (Brown and Heineman, 1972; Schwartz, 1989a,b), unfil Smith et al. (1994) again relegated it to subspecific rank. Despite the differences mentioned in the account above that distinguish the two taxa, I believe they are best regarded as subspecies, although this decision is subjective. They are clearly sister taxa sharing a large number of derived wing pattern and

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223 female genitalic characters, and have no important wing pattern or genitaHc differences. They also occupy similar habitats. Range: The island of Puerto Rico. Habitat and adult ecology: According to Smith et al. (1994), this subspecies is common in the coastal Cambalache Forest Reserve near Arecibo, and in the highlands of Maricao and Toro Negro to over 1000m. hidividuals may be encountered along roadsides and forested paths, and show territorial behavior. It appears to fly throughout the year. Specimens examined (11m, lOf): PUERTO RICO (11m, 80: Cambalache 150m Aug. If AME; Ensefiada Jul. If AMNH; Guajataca 400m Mar. Sep. Nov. 2f FSCA, 3m, If AME, Im AMNH; Maricao 2500' Aug. Im AME, If AMNH; Mt. Brittan Jan. If AMNH; San Juan Jul. If AMNH; Toro Negro 3000' Aug. 4m AME; no specific locality Im BMNH, Im ZMHU. COUNTRY UNKNOWN (It): Valera Ex. Musaeo Oris. Boisduval If BMNH; no specific locality If BMNH. Adelpha calliphane Fruhstorfer, 1915 Figs. 29a,b; 113a,b; 195a,b Adelpha calliphane Fruhstorfer (1915: 526; 1916, pi. 109b, missp. as calliphana) TL: Rio de Janeiro, Brazil. OTL: Rio de Janeiro, Sao Paulo [Brazil]; Paraguay. Types: BMNH(T): LTm: "Fruhstorfer Coll. B.M. 1937-285//calliphane Fruhst.//Brasilien Rio Fruhstorfer//TYPE//Type//Syntype" [examined]; PLTf: "Fruhstorfer Coll. B.M. 1937285//Brasilien Rio Fruhstorfer//TYPE//Type " [examined] = poltius; BMNH(M): PLT?m: "Sao Paulo Fruhstorfer/ZFruhstorfer Coll. B.M. 1937-285"; PLT?m: "Brasilien Sao Paulo Fruhstorfer/ZFruhstorfer Coll. B.M. 1937-285"; BMNH(R): PLTm; "Yhii, Paraguay XII.96 (Ander)//calliphane Fruhst." [all examined] Adelpha gavina Fruhst., Brown (1992) misid. Identification, taxonomy and variation: This species is distinguished from the majority of similar species by having a continuous postdiscal band on the VFW, the lower portion of which is white, the upper portion pale gray. Adelpha poltius also has a similar band, but differs in the orange-brown VFW subapical marking being of more even width, with the basal edge notably curved in cells M2-R5 so that it is much nearer the upper postdiscal band in cell M2 than Ml, in having reduced postdiscal and submarginal series on the ventral surface, and in having a thin, dark bown line immediately bordering the distal edge of the white postdiscal band on the VHW, instead of a broader, red-brown line. There is little variation in the small series examined.

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224 Fruhstorfer (1915) described this species based on the male and a female, the latter from Rio de Janeiro, stated the range to be Rio de Janeiro to Sao Paulo, and also mentioned two males from Paraguay in the Tring Museum (now the BMNH(R)). The description is uncharacteristically detailed and mentions the continuous postdiscal band on the VFW, which characterises this species and A. poltius, and the dorsal surface of the species was figiu-ed on plate 109b (Fruhstorfer, 1916). The syntype male specimen in the BMNH(T) closely matches the figure, which shows the anteriorly broadening DFW orange subapical marking with a straight basal edge which distinguishes the species from A. poltius. Furthermore, Fruhstorfer's (1915) description of the male genitalia clearly refer to those of A. calliphane. However, Fruhstorfer also failed to note the distinctness of A. poltius, since the syntype female mentioned in the original description is actually that species. The only other Fruhstorfer specimens of this species in the BMNH(M) are two males from Sao Paulo, and they are almost certainly syntypes, while only a single male in the BMNH(R) from Paraguay bears a label indicating it to be a probable syntype. Fruhstorfer most probably regarded the syntype male in the BMNH(T) as his "Type", and it appears to be this specimen that was illustrated, but since there is some doubt and the syntype female belongs to a different species, I designate the former specimen as the lectotype of Adelpha calliphane, with the following label data: "Fruhstorfer Coll. B.M. 1937-285//calliphane Fruhst./ZBrasilien Rio Fruhstorfer//TYPE//Type// Syntype". This species belongs to a group which includes several southeast Brazilian species, including A. poltius, A. falcipennis and A. gavina, among which the shared possession of a continuous VFW postdiscal band suggests a close relationship with A. poltius. Range: Southeastern Brazil to northern Argentina and Paraguay, and probably Uruguay. Habitat and adult ecology: This species is relatively uncommon in collections, particularly females, and has been recorded up to 1400m throughout the year. Ebert (1969) reports it to be not uncommon to common in both secondary and primary forest in

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225 Minas Gerais, while Brown (1992, as gavina) reports it as being rare in open forest to areas of extensive secondary growth in the Serra do Japi, Sao Paulo, Brazil. Hoffmann (1936) reported it to feed on flowers of Compositae in Santa Catharina, Brazil. Specimens examined: 53 (50m, 3f) BRAZIL (30m, 3f): Guanabara: Rio Im BMNH; Minas Gerais: Passa Quatro 1000m Dec. Im BMNH, Im BMB, 2m MNHN; Parana. Castro Im BMNH; Caviuna Im AMNH; Igua(;u Jan. Feb. 3m BMNH; N. Parana 2m AMNH; Ponta Grossa 3500' Dec. If BMB; no specific locality Im AMNH; Rio de Janeiro: P.N. Itaitiaia 800m May Im USNM; Rio Grande do Sul: Guarani Im MCZ; Sanla Catharina: Rio Vermelho If AMNH; no specific locality If ZMHU, Im BMB; Sao Paulo: Casa Branca 7m ZMHU; Sao Paulo 2300' 4m BMNH, Im AMNH; Not located: no specific locality Im BMNH, Im BMB. PARAGUAY (19m); Caaguazu: Yhii Dec. Im BMNH; Central: Asuncion Im MNHN; Guaira: Colonia Independencia Aug. Sep. 2m AME; Misiones: no specific locality Oct. Nov. Im BMNH; Paraguari: Sapucay Jul. Sep. Oct. 6m BMNH, 3m BMB; San Pedro: Nueva Germania Im BMB; Not located: Central Paraguay Im BMNH; no specific locality 3m BMNH. ARGENTINA (Im): Misiones: no specific locality Apr. Im BMNH. Additional locality data: BRAZIL: Minas Gerais: nr. P090S de Caldas 1000-1400m Jan. Feb. Apr. May (Ebert, 1969); Santa Catharina: Jaragua Apr. (Hoffmann, 1936). PARAGUAY: Guaira (Kochalka et ai, 1996). Adelpha poltius Hall, 1938 Figs. 30a,b; 114a,b; 196a,b Adelpha poltius Hall (1938: 258) TL: Minas Geraes [Brazil]. Types: BMNH(M): HTm: "Minas Geraes. Brazil. Bates Coll.//m//Minas Geraes m//Holotype//Adelpha abia Hew. Godman-Salvin. Coll. 1916.-4.//Holotype male Adelpha poltius Hall G. Lamas det. 1987"; PTm: "Minas Geraes. Brazil. Bates Coll.//m//Adelpha abia Hew. Godman-Salvin. Coll. 1916.-4.//Minas Geraes m//Paratype"; PTm: "Minas Geraes. Brazil. Bates ColI.//m//Adelpha abia Hew. Godman-Salvin. Coll. I916.^.//Minas Geraes m//Paratype//not in B.M.//ex. coll Saunders"; PTm: "Rio. Hewitson Coll. 79-69 Heterochroa l//Rio//Paratype"; BMB: PTm: Barbacena 4000' Oct. [all examined] Adelpha calliphane Fruhst., Brown (1992) misid. Identification, taxonomy and variation: This species is distinguished from all similar species, with the exception of A. calliphane, by having a continuous postdiscal band on the VFW. Characters distinguishing it from A. calliphane are discussed under that species. There is some variation in the shape of the hindwing, which may be rounded and of similar shape to A. cocala, while the DFW orange subapical area may also be broaden slightly towards the costa, but not so much as in A. calliphane. Hall (1938) described this species based on a holotype male and four syntypes from Minas Geraes and Rio de Janeiro, and I have examined all of the type series. His description clearly applies to this species, mentioning the diagnostic shape of the DFW orange subapical band, at least as it occurs in the type series. Adelpha poltius is probably most closely related to A. calliphane, as discussed under that species. Range: Only known to date from southeastern Brazil, but probably more widespread.

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226 Habitat and adult ecology: This species is rare in collections and most specimens have been collected from 8001200m. It has been recorded from December to May. Ebert (1969) found it to be not uncommon in Minas Gerais in primary to secondary forest habitats, up to 1400m. Brown (1992, as calliphane) also reports that it occurs, in the Serra do Japi, Sao Paulo, in a very wide range of habitats, from humid lowland forest to forest above 1100m, and also may be found in open fields, gardens and areas of extensive secondary growth. Specimens examined: 39 (23m, 160 BRAZIL (23m, 150: "Amazonas": Canuma 2m, If AMNH-error; Guanabara: Rio Im, If BMNH; Minas Gerais: Barbacena 4000' Oct. ImBMB; no specific locality 3m BMNH; Para/io: Castro If BMNH, If USNM; Femandes Pinheiro 2600' Apr. If BMNH, Im, If AME; S5o Luiz do Puruna 90Om Mar. Im FSCA; Rio de Janeiro: Petrdpolis Independencia 900m Im AME; Santa Catharina: Cauna Apr. 9m, 6f AMNH; Rio Vermelho 3350-3550' Mar. If FSCA; no specific locality If MCZ, 4m, If AMNH. COUNTRY UNKNOWN (10: no specific locality If BMNH. Additional locality data: BRAZIL: Minas Gerais: nr. Po90S de Caldas 1000-1400m Jan.-May Dec. (Ebert, 1969). Adelpha mythra (Godart, [1824]) Figs. 31a,b; 115a,b; 197a,b Nymphalis mythra Godart ([1824]: 374) TL: Bresil. Types: MNHN?: HTm: [not located] =Heterochroa zeba Hewitson (1850: 435, pi. IX, figs. 3, 4) TL: Rio de Janeiro [Brazil]. Types: BMNH(T): STm: "B.M. TYPE No. Rh. 9833 Heterochroa zeba m Hew.//Rio Hewitson Coll. 79-69 Heterochroa mithra 2.//zeba//Type//Syntype" [examined]; BMNH(M): ST?m: "Rio R. Hewitson Coll. 79-69 Heterochroa mithra. 1 .//Rio R."; ST?f: "Rio R. Hewitson Coll. 79-69 Heterochroa mithra. 3." [all examined] Heterochroa mythra Godt., Westwood (1850); Adelpha mythra Godt., Kirby (1871) Identification, taxonomy and variation: This species is readily distinguished from all other similar species by the extensive yellow orange coloration suffusing the venfral surface. It is superficially most similar to A. cytherea, but the orange DFW subapical marking does not touch the white postdiscal band, the third discal cell bar is absent on the VFW, the outer submarginal series on the VFW are absent within the broad yellow orange distal marginal border of both wings, and the upper postdiscal band is clearly visible on the VFW as a gray line. There is slight variation in the width of the white postdiscal bands, in particuar the marking in cell M3 on the DFW which may be reduced to a small spot, and in the width of the orange DFW subapical marking.

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227 Godart ([1824]) described this species jfrom a single male from Brazil, the whereabouts of which is unknown. However, the description, which compares the species to A. cytherea, is sufficiently detailed for there to be no doubt as to the identity of the species. Godart mentions that the orange DFW marking is large and triangular and completely isolated from the white DFW postdiscal band, that the postdiscal band on the VFW extends to the costa ("jusqu'en haut") and is accompanied by only two white lines, which I believe refer to the pale bands in the VFW discal cell; in A. cytherea there are three such bands, since the third discal cell bar divides the pale area between the second and fourth cell bars into two. Godart also mentions that the tomus of the VHW has two black spots, which are absent in A. cytherea. The name zeba was first introduced by Westwood (1850) as a nomen nudum, then described shortly after by Hewitson (1850) based on an unspecified number of specimens in his own collection from Rio de Janeiro. Hewitson noted that the species was close to A. mythra, but did not state how it differed, and the detailed and accurate decription, plate showing the dorsal surface and syntype in the BMNH, closely matching the plate, show that zeba is a synonym of mythra, where it was placed by Kirby (1871). The name patricia, listed by Martin et al. ([1923]) as a subspecies of A. mythra and represented in the BMNH(T) by a specimen from Rio Grande in Brazil, is a nomen nudum. Despite a superficial similarity to A. cytherea, this species lacks the third VFW discal cell bar of ^4. cytherea and has the expanded orange DFW subapical marking typical of members of the A. iphiclus group. Within this group, its male genitalia are distinctive in having a relatively broad, short clunicula, and the valvae are short with reduced dorsal and ventral medial lobes. The continuous VFW postdiscal band suggests that the species may be most closely related to A. poltius and A. calliphane. Range: This species occurs in eastern Brazil from Bahia south to Rio Grande do Sul, inland to the state of Minas Gerais, and westwards as far as northeastern Argentina and Paraguay. It probably also occurs in Uruguay.

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228 Immature stages: MuUer (1886) reared this species and reported the hostplant from Santa Catharina (Brazil). The coloring of the larva seems to be similar to that of A. iphiclus, but otherwise little information was given. Muller said that of all the species that he reared, the dorsal projection on segment A2 in the pupa reached a similar scope to A. lycorias lycorias only in this species. This is interesting as no other members outside of the morphologically relatively well characterised A. phylaca group have been reported with such a pronounced dorsal projection. It is unfortunate that the larva was not well described. The pupa also has indistinct dorsal projections in the position of the last instar larval scoli on segments T2, with more pronounced projections on A2-A4. Habitat and adult ecology: Adelpha mythra is very common in collections and has been recorded up to 1500m. It appears to fly throughout the year, but is most abundant in the early months of the year. Ebert (1969) found it to be common in primary to secondary forest in Minas Gerais. Brown (1992) reports that it flies high along paths and within the forest at the Serra do Japi, Sao Paulo, Brazil. Hoffmann (1936) reported that he often found inviduals at flowers of Mikania in Santa Catharina, Brazil. Specimens examined: 163 (121m, 42f) "COLOMBIA" (10; no specific locality If MCZ-error. BRAZIL (1 18m, 41f): Bahia. Bahia If BMNH; Espirito Santo: Santa Cruz Mar. If AME; no specific locality 2m BMNH; Guanabara: Rio Im MCZ, 2m, If BMNH; Minas Gerais: Passa Quatro 1000m Im BMNH, Im MNHN; Parana: Castro Feb. Apr. 9m BMNH, Im USNM; Caviuna May Im, If AMNH; Curitiba Mar. Im FSCA; Igua9u Oct. Nov. 3m BMNH; N. Parana 2m, If AMNH; Ponta Grossa 3500' Dec. Im BMB; S5o Luiz do Puruna 900m Mar. 4m FSCA; Tijucas do Sul 600m Mar. Im, If FSCA, Im USNM; Uni3o da Vitoria 2000' Dec. Jan. 2m, 2f BMB; Rio de Janeiro: Maravalco Jan. Im AMNH; Maromba Jan. Im AMNH; Nova Friburgo Im BMNH, Im ZMHU; P.N. Itaitiaia May Im AMNH; Petr6polis 1000-1500m Jan. Mar. May 7m AME, 5f USNM; Petropolis Independencia 900m Jan. 3m AME; Teresopolis Feb. 2m USNM, Im BMNH, If ZMHU; Rio Grande do Sul: Northeast Im BMNH; Pelotas Jan.-Jun. 8m, 8f AMNH, 7m, 5f MCZ, Im BMNH, Im AME, 3m, If USNM; Rio Grande do Sul Im, 2f BMNH, If ZMHU, Im BMB, Im, If MNHN; Santa Catharina: Serrinha do Pirai, W. Joinville 950' Mar. Im FSCA; Blumenau 2m, 2f MCZ; Brusque Jul. Dec. Im AMNH, If AME; Cauna Mar. Apr. Im, If AMNH; Joinville 20-200m Dec. Im MUSM; Rio Garcia If AMNH; Rio Vermelho 3350-3550' Mar. 7m, If FSCA; Sao Bento do Sul 850m Feb. Mar. Im FSCA, Im AME; no specific locality Oct. 4m, If AMNH, If BMNH, Im ZMHU, Im FSCA, Im AME, Im USNM; Sao Paulo: Alto da Serra Feb. Aug. Dec. 3m BMNH, Im MNHN; Casa Branca Im ZMHU; SSo Paulo 3m BMNH, Im MCZ, Im ZMHU, Im USNM; Ypiranga Dec. Im BMNH; no specific locality 2500' Dec. Im BMB; Not located: Maramba Jan. Im, If MCZ; Rio R. Im BMNH; no specific locality 3m BMNH, Im MNHN, Im AMNH. COUNTRY UNKNOWN (3m): no specific locality 3m BMNH. Additional locality data: BRAZIL: Minas Gerais: nr. Poifos de Caldas 1000-1400m Jan. Feb. Apr. May Nov. Dec. (Ebert, 1969); Rio Grande do Sul: Cap^o do Le3o; Cascata; Monte Bonito; Retire (Biezanko, 1949); Santa Catharina: Jaragua Jan. Mar. Apr. (Hoffmann, 1936). PARAGUAY: Alto Parana (Kochalka et al.. 1996). ARGENTINA: Misiones (Hayward, 1951). Adelpha basiloides (H.W. Bates, 1865) Figs. 32a-d; 116a,b; 198a-c

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229 Heterochroa basiloides Bates (1865: 332) TL: Mexico. Types: BMNH(T): STf: "B.M. TYPE No. Rh. 9837 Heterochroa basiloides f Bates/ZB.C.A. Lep. Rhop. Adelpha basiloides Bates Godman-Salvin Coll. I916.-4.//Type. sp. figured//f//Mexico Salle Bates Coll.//f Mexico Basiloides Bates/ZType H.T." [examined] ^Heterochroa lydia Butler (1865b: 398) stat. rest. TL: Honduras. Types: BMNH(M): STf: "Syntype f Heterochroa lydia Butler G. Lamas det. 1987//Honduras// Syntype" [examined] ^Heterochroa lemnia C. and R. Felder (1867: 417) TL: Mexico. Types: BMNH(R): STm: "Syntype/ZFelder Colln.//Lenmia n.//Mexico Salle Type" [examined] =Adelpha basiloides var. lativittata Staudinger (1886: 142) syn. nov. TL: Puerto Cabello, Venezuela. Types: ZMHU: STm: Puerto Cabello, Hahnel/ZOrigin [examined] =Adelpha basiloides caelia Fruhstorfer (1915: 524) syn. nov. TL: Colombia. Types: BMNH(T): STf: "Fruhstorfer Coll. B.M. 1937-285//basiloides caelia Fruhst.//Columbien ex. Coll. H. Fruhstorfer//TYPE//Type//Syntype" [examined] Adelpha basiloides Bates, Kirby (1871); Adelpha iphicla var. lemnia Fldr., Kirby (1871); Adelpha iphicla var. lydia Butl., Kirby (1871); Adelpha iphicla basiloides Bates., Hall (1938) in part; Limenitis (Adelpha) basiloides Bates, Ross (1976); Adelpha basiloides basiloides Bates, Neild (1996); Adelpha basiloides lativittata Staud., Neild (1996); Adelpha basiloides caelia Fruhst., Neild (1996) Identiflcation, taxonomy and variation: This species is distinguished from all similar species in having the DFW white postdiscal band terminate at vein M2, after which it may or may not be followed by separate white postdiscal dashes. There is much variation in the width of the white postdiscal band, the size and number of postdiscal dashes anterior to it and the shape of the orange DFW subapical marking, and this is discussed below. Bates (1865) described this species based on a female specimen or specimens from Mexico, and correctly noted the character of the white postdiscal band extending to vein M2 which distinguishes this species from similar species. He also noted that there was a white spot on the DFW in cell Ml, and this is the case in the syntype female in the BMNH. In December of the same year, nine months after and apparently ignorant of Bates' (1865) description oiA. basiloides, Butler (1865b) introduced the name lydia for a species from Honduras, based on a specimen or specimens in the BMNH, which he stated differed from A. iphiclus by having the white DFW postdiscal band extending beyond vein M3. The long, detailed description clearly applies to A. basiloides, another character mentioned being the narrower, red-brown discal bands on the VHW. Butler also mentions "two minute and indistinct white spots" anterior of the upper postdiscal band. The putative syntype female in the BMNH bears no data to indicate its type status, but is the

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230 only specimen from Honduras which Butler could have seen. This specimen has a faint, white postdiscal dash in cell Ml on the DFW. hi any case, there is no doubt as to the identity of lydia, but although Godman and Salvin (1884) correctly synonymised it with A. basiloides, Fruhstorfer (1915) resurrected the name for no reason, and I here place it back in synonymy (stat. rest). The Felder brothers, in their description of Heterochroa lemnia, also appeared unaware of Bates' basiloides, although they mentioned Butler's lydia as being the most closely related species (Felder and Felder, 1 867). They compared an imspecified number of female specimens from Mexico to A. iphiclus, and mentioned the more extensive DFW white postdiscal band, the orange DFW subapical spot not extending into cell M3, and the narrower red-brown VHW discal bands. These characters clearly indicate lemnia to be a synonym of A. basiloides, where it was placed by Godman and Salvin (1884), and this is confirmed by the syntype (actually a male) in the BMNH. Staudinger (1886) infroduced the name lativittata (misspelt lativitta by Fruhstorfer (1915)) for specimen(s) from Puerto Cabello in Venezuela, which differed from the nominate in having wider white postdiscal bands. I have examined a syntype male in the ZMHU. Fruhstorfer (1915) subsequently described caelia from Colombia, supposedly differing from the nominate in having a smaller orange DFW subapical marking and broader white postdiscal bands. No information was given on the specimen(s) on which the description is based, but neither of these characters are apparent in any examined Colombian specimens, or in the syntype female in the BMNH. There is substantial variation in this species throughout its range. Most noticeable is the presence or absence of white postdiscal dashes in cells Ml and R5 on the DFW, which occur often in females from Mexico to Honduras, being particularly pronounced in specimens from Tamaulipas (Mexico), and occasionally in Mexican males. Females without these white dashes, corresponding to caelia, occur throughout the range of the species as far north and west as Tabasco in Mexico. The width of the white postdiscal bands is also variable; specimens from Tamaulipas and northwards in Mexico typically

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231 have broader bands, in addition to specimens from northern Venezuela, the latter corresponding to lativittata. Specimens from northwestern Ecuador have the narrowest bands, in certain specimens the width approaches that in A. donysa albifilum, but again the character is variable and some specimens in western Colombia have bands of equal width to Ecuadorian specimens. The orange DFW subapical marking is narrowest in specimens from northwestern Ecuador, and broadest in specimens from Tamaulipas (Mexico), where it also extends into cell M2. Some case can be made ft»r freating lativittata from north Venezuela and caelia from Colombia as subspecies of A. basiloides (see Neild, 1996), but my examination of long series of specimens in museums suggests that all of the characters discussed above are locally shghtly variable and geographically clinal, and do not allow unambiguous identification of distinct subspecies. I therefore regard lativittata and caelia as synonyms of basiloides (syn. nov.). Forbes (unpublished manuscript) intended to describe specimens from Tamaulipas with the DFW orange extending into cell M2 as a new species, occidentis, stating that they differed also in the male genitalia, a claim I have been unable to substantiate. Adelpha basiloides is a member of the A. iphiclus group, as defined by the shape of the DFW subapical marking, but although Hall (1938) placed basiloides as a subspecies of A. iphiclus, the genitalia and wing pattern of the two taxa are clearly distinct and they are sympatric throughout Central America. In fact, A. basiloides is most closely related to A. plesaure, which replaces it east of the Andes. Both A. plesaure and A. basiloides share the DFW postdiscal band extending always to vein M2 (sometimes beyond, in both species), narrower red-brown discal bands on the VHW and similar genitalia and immature stages (Aiello, 1984). However, A. basiloides differs from A. plesaure in having the inner and outer postdiscal series entirely fiised on the VFW, a derived character with respect to other A. iphiclus group members, while A. plesaure is unique in the A. iphiclus group in having the inner postdiscal series fiised with the postdiscal band

PAGE 239

232 on the forewing. These characters suggest that the two species are best retained as distinct. Range: Extreme southwestern U.S.A. (Texas) throughout Central America to northern Venezuela and western Ecuador, west of the Andes. The species appears to have been only recently recorded in the U.S.A. by Neck (1996), since it is not mentioned by either Scott (1986) or Miller and Brown (1981). Neck reports a single specimen from the Rio Grande valley, but he also figures a specimen of this species under the name A. fessonia (pi. 43, fig. 193), and it seems unlikely that the supposedly unique specimen would have been photographed in the wild. I have also seen a single specimen in the FSCA, and it may be that this species is actually represented in more collections but confused with the more common A. fessonia. Immature stages: Aiello (1984) reared this species in Panama and figured the final instar larva and its head capsule (figs. 2, 4) and the pupa (fig. 6). The egg is typical of the genus and is deposited on the upper surface of the leaf, either at the tip or at a damaged portion. Larval color depends on the foodplant, being light brown or reddish brown on Amaioua, and dark brown on Alibertia and Bertiera. The third and fourth instars develop a sharply defined, pale, triangular, dorsal "saddle" marking with its base on the posterior portion of segment A4, and its apex at the middle of A6. Otherwise the larva is mottled black and brown, paler posteriorly, with the boundary between the dark and light areas being an oblique, anteriorly descending line from A7 to A4. Late final instars may be tinged with pink or green, especially at the bases of the scoli, and laterally on A4-A6. The scoli vary substantially in form; that on Tl is moderately thick and long with long lateral spines, those on T3 and A7 are long and thin with very short lateral spines, that on A2 is very thick and curved posteriorly with short, thick lateral spines, those on Al, A3-A6 are short, with only a terminal rosette of thin spines, and that on A8 is short and thick with a broad, clubbed tip armed with long spines. Supraspiracular scoli are also well developed, of similar form to the scoli on A3-A6. The head capsule is patterned with darker lines.

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233 lacks most setae and has well developed chalazae. The pupa is pearly white with black tipped head homs, which vary in length and curvature. There are preserved head capsules and pupal cases in the USNM, collected by Small in Panama. Marquis, Mallet and Janzen (/«: DeVries, 1986: Costa Rica) report that late instar larvae are solitary and feed on all leaves of plants along forest and river edges or in forest light gaps. According to Aiello (1984), the larva of .4. basiloides is almost unique among known larvae in consistently contructing a small, curved mass of frass which strongly resembles a larva on the upper surface of the leaf, away from either the leaf edge or the usual mass of frass accumulated at the leaf base. Presumably this serves some protective function. Development times of the various stages are Hsted by Aiello (1984: Table 2), and the total development time varies from 39-48 days. This is longer than other Panamanian species, and in addition, A. basiloides sometimes passes through a sixth instar. Six instar larvae were reared from the same batch of eggs as five instar larvae, and on several occasions. All six instar larvae developed on the hostplant Amaioua, and although the development time for individuals with six instars is no longer than that for those with five, all individuals were slower to develop on Amaioua than on other hostplants (Alibertia, Bertiera). Habitat and adult ecology: This is one of the commonest species in the northern and western parts of its range, becoming progressively rarer towards and in South America. De la Maza and de la Maza (1993) report it from evergreen and deciduous forest in Mexico, from sea level to 1700m, where it flies from April to December. In the Tikal area of Guatemala it has been recorded from primary to second growth habitats where it is uncommon (Austin et ai, 1996), while in Costa Rica the species occurs up to 1000m, in all forest habitats, being most abundant in the wet season in deciduous forest (DeVries, 1987). Both sexes feed on rotting fioiits of Genipa, Alibertia and Guazuma, and apparently mammal dung (DeVries, 1987), though this is an unlikely food source for females. Orellana (pers. comm.) also reports the species to be attracted to fermenting

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234 fruits of Bunchosia comifolia (Malpighiaceae) in Panama. The species occurs in association with secondary growth habitats in rain forest, and open areas in dry forest. In western Ecuador I have found the species to be rare, and only occurring in wet rain forest below 300m. All the individuals I have seen have been males perching around the edges of ridgetop light gaps in selectively logged primary forest, in the sun, usually around 68m high but as low as 3m on occasion. In western Ecuador, individuals typically have much narrower white postdiscal bands than elsewhere in the range of the species, and the occurrence of narrower bands in a number of other Adelpha species in western Ecuador, with similar dorsal patterns, is strong evidence for mimicry between these species. Other similar species which develop narrow postdiscal bands in western Ecuador include: Adelpha iphiclus, A. iphicleola, A. bamesia, A. erotia f lema, A. serpa and A. seriphia. Specimens examined: 335 (198m, 137f) t with orange on DFW in cell M2 (only applies to specimens in the MCZ, USNM and AMNH). U.S.A. (Im): Texas: Bentsen-Rio Grande Valley State Park, Hidalgo Co. Jun. Im FSCA. MEXICO (97m, 62f): Chiapas. Comitin 2m AME; Mapastepec 2m USNM; Monterrey 2m AMNH; Santa Rosa Aug. Im, If USNM; Guerrero: Acahuizotia Aug.-Oct. 4m AME; Ayutla-Tiena Colorada 350m Jul. Im BMNH; no specific locality Im, If BMNH; Jalsco: Zapotlan Jun. Jul. Im BMB; Nayarit: vie. Compostela Jun. 2mt MCZ; Nueva Leon: Cola de Caballo 610m Sep. Im AME; Hda. Vista Hermosa Im AMNH; Horsetail Falls Aug. Im, If FSCA; Michoacan: Coahuayana Jun. Aug. 3m AME; Oaxaca: Candelaria Loxicha Aug.-Nov. Im FSCA, 5m, 2f AME, Im, If BMB, 2mt, 2ft USNM, 4m, 4f AMNH; Chiltepec Jun.-Sep. 3m AME, 3m, 3f AMNH; Chimalapa Aug. Sep. 3m AME; El Naranjal-Chiltepec May If AMNH; Portillo del Rayon If AME; San Mateo Yetla May 1ft USNM; Tapantepec 150m Im AME; Tuxtepec 2f USNM; Quintana Roo: X-can Aug. 4m AME; San Luis Polosi: Ciudad Valles Aug. Im FSCA; El Barlito Im, 2f AMNH; El Punjal Mar. If AMNH; El Salto Falls Aug. Oct. Nov. 2f FSCA, Im AME, Imt USNM; Palitia Apr. Im AME; Quinta Chilla Aug. Dec. 3f AMNH; Rancho Santa Maria Dec. If AMNH; San Isidro Apr. If FSCA; Santa Rosa 1ft USNM; Tamazunchale Aug. 3m, 3f AMNH; Tamazunchale, 11.7 mi. S., Aug. Ift USNM; Tamazunchale, 30 mi. N., Aug. Imt USNM; Valles Im, If AMNH; Tabasco: Balancan Im AME; La Venta 10m Jan. If AME; Tabasco If BMNH; Teapa Feb. Im AMNH; Tamaulipas: Ciudad Victoria May If FSCA; El Limon Mar. 2mt USNM; Galeana Canyon Oct. Imt USNM; Gomez Farias Mar. Oct. 2m FSCA, 2m, If AME; Mante, 30 mi. N., Nov. Im, If USNM; Rio Sabinas Mar. Im MCZ; Tamaguachale Jul. If AME; Tampico Dec. Im AMNH, Im BMNH; Victoria, 60 mi. S., Jul. 2m AMNH; Veracruz: Allende 3f AMNH; Atoyac If BMNH; Cardel Dec. Im AMNH; Catemaco Im AME; Coatepec Oct. If AMNH; C6rdoba Jun. Jul. 2m, 2f BMNH, 2m AMNH, If BMB; Dos Amates Im AME; Jalapa Im, If BMNH, If USNM, If AMNH; Misantla If BMNH; Ojo de Agua If AMNH; Peso del Toro May If FSCA; Presidio Jun. 3m, If AME; Tecolutla Jul. If AMNH; Vigia Aug. Im AME; Yucatan: Piste Jul. -Sep. 5m AME; Not located: Cotontepec Jun. Im AME; no specific locaHty 2m AMNH, Im, 2f BMNH, Im, If AMNH, If MCZ, Im MNHN. EL SALVADOR (Im, 20: La Libertad: Zaragoza If AMNH; Santa Ana: Citala 800m Oct. Im, If AME. BELIZE (5m): Cayo: Camp Sibiin 200m Jul. 2m AME; Not located: Silk Grass Nov. 2m FSCA; Rio Grande Im AMNH. GUATEMALA (16m, 14f): Alta Verapaz: forests of N. Vera Paz If BMNH; Baja Verapaz: San Jerdnimo If BMNH; El Peten: Sayaxche Sep. Oct. 4m, 2f AMNH; Escuintla: Escuintla 1200' Jul. Aug. 4m BMB; Izabal: Cayuga 2m, 2f USNM; Puerto Barrios Dec. 3m, 2f BMB; Quezaltenango: Volcan Santa Maria 2m, 3f BMNH; Retalhuleu: San Sebastian If USNM; Santa Rosa: Barbarena Im AME; Guazacapan If AME; Suchitapequez: Mazatenango Jul. Aug. If BMB. HONDURAS (Im, 60: Cortes: San Pedro Sula If BMNH; Not located: U Cambre Im BMNH; no specific locality 2f BMNH, 3f ZMHU. NICARAGUA (4m): Chontales: Chontales 2m BMNH; Zelaya: Bluefields Dec. Im AMNH; Nueva Guinea Oct. Im AMNH. COSTA RICA (6m, 5f): Alajuela: San Mateo Im USNM; Guanacaste: Hac. La Pacifica Jul. Im USNM; Limon: Port Limon 2f USNM; San Jose: Puriscal If USNM; Not located: no specific locality 2m BMNH, Im, If ZMHU, Im, If AMNH. PANAMA (24m, 320: Canal Zone: Barro Colorado Island Mar. 4f AMNH; Canal area Mar. Im AME; Cerro Galera Dec. If USNM; CoI6n Jan. Im, If USNM; Farfan Feb. Jun. Jul. Dec. 3m, 4f USNM, Im, If AME; Fort Kobbe Jun. Jul. 3m, 2f USNM; Fort Sherman Mar. If FSCA; Gamboa Oct. Im USNM; Los Rios Mar. Im USNM; Madden Dam 2m FSCA; Matachin Im, If BMNH; Rodman Jan. If USNM; Chiriqui: Bugaba 800-1500' Im BMNH; Chiriqui If BMNH, Im, 2f ZMHU, If MCZ; Rio Tole 120m Jan. Im USNM; Valle de Chiriqui 25-4000' Im, If BMNH; Colon: Piila 200m Jul. Im, 2f AME; Darien: Rio Tuguesa Jul. If USNM; Herrera: Cerro Montuoso 600m Jan. Im USNM; Panama: Arraijan rd. Feb. 2m AMNH; Cerro Jefe 600m Jan. If USNM; Cocoli Oct. Nov. 3f USNM; Veraguas: Calobre Im, If BMNH; Not located: Las Cumbres Oct. Im FSCA; no specific locality 2f ZMHU, If USNM. COLOMBIA (23m, 90: Antioquia: Crystallina 1 100' Jun. Jul. Im BMB; Cundinamarca: Bogota 2m BMNH, 2m USNM; env. Bogoti 2m BMNH; Region du Bogota 6m BMNH; Magdalena: F. Union-F. Mercedes, Rio Guachuca, Santa Marta lOOm Aug. If BMNH; Santander: El Centro Im AMNH; La Borroscosa If AMNH; La Danta If AMNH; La Lechera Im AMNH; U Lindera Im AMNH; Valle del Cauca: El Engafio Aug. If LMC; Not located: no specific locality 3m AMNH, 2m, 3f MCZ, Im, If BMNH, Im

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235 ZMHU, If AMNH. VENEZUELA (7m, 60: Aragua: Puerto Cabello 2m, 2f ZMHU; Carabobo. San Esteban Jul.-Sep. Im BMNH, If BMB; Distrito Federal: Caracas Im BMNH; Merida: Merida If BMNH; Zulia. Mision El Rosario 50m Jan. Im MUSM; Tres Bocas 200m Jun. 1 f AFEN; Not located: Sinistara del Sur 1 m AMNH; no specific locality 1 f BMNH, 1 m MNHN. ECUADOR (7m): Esmeraldas: El Durango 300m Sep. 2m KWJH; U Punta, km. 44 Lita-San Lorenzo rd. 300m Jun. Aug. 2m KWJH; San Lorenzo-Lita rd., km. 20, 100m Aug. Im KWJH; Not located: no specific locality 2m BMNH-error (wide dorsal white bands). "BRAZIL" (Im): no specific locality 1 m AMNH-error. COUNTRY UNKNOWN (2m, If): no specific locality Imt MCZ; If FSCA, Im AME. Additional locality data: MEXICO: Chiapas: throughout see de la Maza and de la Maza (1993); Colima: Agua Dulce 600m Oct.; Cerro de la Media Luna; Coquimatlan Sep.; Colima city 500m Nov. (Warren et al., 1998); Guerrero: Acapulco; La Sabana; Oaxaca: Metates; Chacalapilla; Puebla Patia; Veracruz: El Vigia (de la Maza, 1987); Quintana Roo: Carrillo Puerto; Chumpon; Chunyaxche; Noh-bek; Ramonal; Tres Reyes; Tuliim (de la Maza and Bezaury, 1992); Jalisco: Boca de Tomatlan Dec; Chico's Paradise Dec; La Calera Mar. Nov.; Mismaloya Jan. Mar. Apr. Dec; Puerto los Mazos Mar. May Jun. Nov.; Puerto Vallarta Apr. Dec. (Vargas et al., 1996). BELIZE: Corozal, Toledo (Meerman, 1999). Adelpha plesaure Hiibner, 1823 Figs. 33; 117; 199 Identification, taxonomy and variation: Adelpha plesaure is distinguished from all other species by having the postdiscal series fiised to form broad spots in cells Ml and R5, as in all A. iphiclus group members, and the inner postdiscal series fiised with the postdiscal band, the latter extending at least as far as vein M2, but never extending unbroken to the costa. There is geographis variation in the extent of the postdiscal series and postdiscal band on the DFW, and the presence or absence of an orange postdiscal band distal of the white DHW postdoscal band, and four subspecies are recognised, all of which are connected by intergrades. This is a distinctive species, and except for being inexplicably split into two species by Fruhstorfer (1915), there has never been any confiision over its identity. It is the sister species of A. basiloides, with which it is allopatric, and my reasons for retaining the two as distinct species are discussed under A. basiloides. Range and status: Venezuela to Bolivia, Brazil, the Guianas and Trindad, in lowland rain forest up to 1250m. Common. Specimens examined: 443 (253, 190) Adelpha plesaure plesaure Hubner, 1 823 Figs. 33a,b

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236 Adelpha plesaure Hiibner (1823a: 1 1) TL: Brasilien. Types: ZMHU: STm: "Brasilien//Origin//Coll. SommerZ/Plesaure H.//Syntype" [examined] -Adelpha plesaure heredia Fruhstorfer (1915: 521) TL: Espirito Santo, Brazil. OTL: Bahia, Espirito Santo [Brazil]. Types: BMNH(M): LTm: "plesaure heredia Fruhst.//Espirito Santo Brasil ex. Coll. Fruhstorfer/ZFruhstorfer Coll. B.M. 1937-285" [exainined] =Adelpha plesaure antoniae Fruhstorfer (1915: 521) TL: Santa Catharina [Brazil]. Types: BMNH(T): STm: "plesaure antoniae Fruhst.//Fruhstorfer Coll. B.M. 1933-I31//Brasilien Blumenau Fruhstorfer//TYPE//Type"; STf: "plesaure antoniae Fruhst./ZFruhstorfer Coll. B.M. 1937-285//Brasilien Blumenau Fruhstorfer//TYPE//Type//Syntype" [both examined] Heterochroa plesaure Hiibn., Westwood (1850); Adelpha phliassa plesaure HQbn, Hall (1938); Limenitis plesaure heredia Frahst., Brown and Mieike (1967); Adelpha plesaure Neild (1996) Identification, taxonomy and variation: This subspecies is distinguished from A. p. phliassa by the orange on the DFW only extending as far as the middle of cell Cul, instead of broadly bordering or crossing vein Cu2. Some specimens may have the orange extending as slight scaling to vein Cu2 along the postdiscal series, but not along the distal edge of the white postdiscal band, and I interpret these specimens as showing intergradation to A. p. phliassa, denoting them with an "X" in the list below of specimens examined. The white at the basal edge of the postdiscal band also extends to almost touch vein Cul. Both of these characters are slightly variable, the orange becoming less extensive and the white more so from north to south, with specimens most closely resembling the syntype of plesaure originating from Sao Paulo and Santa Catharina. The ventral surface is also variable, and the basal half may almost entirely lack red-brown coloration in the discal cells, while the outer postdiscal and inner submarginal series may be expanded and almost merged together. These variations however appear to be individual rather than geographic. Hiibner ([1819]) first introduced this name as a nomen nudum in a list of species in the genus Adelpha, then described it briefly in German in comparison with A. iphiclus, based on an unspecified number of specimens collected by Sommer in Brazil (Hiibner, 1823a). The color figure published in the same year by Hiibner (1823b: pi. 41, fig. 231, 232) shows the dorsal and venfral surfaces of a male, which closely match the syntype specimen in the ZMHU, except that the DFW orange extends as slight scaling to touch vein Cu2 along the inner postdiscal series, hi this it closely resembles the syntype male of

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237 antoniae in the BMNH. Fruhstorfer (1915) described antoniae in comparison with another new subspecies, heredia (see below), from an unspecified number of specimens from Santa Catharina. I have examined two syntypes in the BMNH(T), and a fiirther male and two females in the BMNH(M) which may also be syntypes, from "Santa Catharina" and "Blumenau". None of these differ substantially from the syntype or original illustration of A. plesaure, and Hall (1938) synonymised antoniae with the nominate subspecies. Fruhstorfer (1915) stated that heredia, based on specimens of an unspecified number and sex from Bahia and Espirito Santo in Brazil, differed from other subspecies in having the DFW white postdiscal band extending as "far as the wing centre", and referred to the figure named erroneously "A. plesaure" on pi. 107a (Fruhstorfer, 1913) as representing this subspecies. The figure shows a specimen which is almost identical on the dorsal surface to the syntype specimen of antoniae, with the orange terminating in the middle of cell Cul on the DFW, except for slight scaling along the postdiscal series. The specimen in the BMNH(T), which is labeled "Syntype m Adelpha plesaure heredia Fruhstorfer G. Lamas det. 1987//Fruhstorfer Coll. B.M. 1933-131//Brasilien Fruhstorfer//TYPE//Type//Syntype", is probably not a syntype oi heredia, but in fact is A. p. phliassa; the orange on the DFW extends fully to vein Cu2, but the white postdiscal band extends to the middle of cell Cul. One male specimen in the BMNH(M), bearing the following label data: "plesaure heredia Fruhst.//Espirito Santo Brasil ex. Coll. Fruhstorfer/ZFruhstorfer Coll. B.M. 1937-285", more closely matches the original figure referred to by Fruhstorfer (1915), and I designate it the lectotype of Adelpha plesaure heredia. This specimen is slightly intermediate between typical phliassa and plesaure as dicussed above, but I regard it as representing the nominate subspecies. I therefore follow Hall (1938) in treating heredia as synonymous with plesaure. Range: Eastern and southeastern Brazil, from Bahia to Santa Catharina. Immature stages: Muller (1886) reared this subspecies on a rubiaceous plant resembling species of the genus Bathysa, in addition to various other species of Rubiaceae, in Santa

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238 Catharina, Brazil. The eggs are typical of the genus and are deposited on the upperside of a leaf at the tip. The mature larva is dark in overall color with a paler "saddle" mark on A4 and A5. The pupa was figured on Taf 4, fig. 13, and is shining silvery white with long, laterally curving head horns. Habitat and adult ecology: Hoffmann (1936) states that this subspecies may be found on flowers of various Compositae. Brown (1992) reports this subspecies fi-om the Serra do Japi, Sao Paulo (the figure numbers on the plate have been erroneously transposed between this taxon and A. cocala caninia), where it occurs in disturbed forest areas up to 1000m. Judging fi-om specimens in collections it is locally common, and label data indicate that it probably flies throughout the year. Specimens examined (61m, 53f): t trans to A. p. phliassa, DFW white to vein Cul and/or orange to vein Cu2. "HONDURAS" (If): Cortes: San Pedro Sula 1ft BMNH-error. "COLOMBIA" (Im, If): Cundinamarca: Bogota If BMNH, Im MCZ-error. BRAZIL (60m, 50f): Bahia: Bahi'a Imf BMNH; Cachimbo 1ft BMNH; Espirito Santo: Itaguassu Sep. Im AME; Linhares 4m, 4f AME; Santa Teresa Apr. Im AME; no specific locality Imt, 1ft BMNH, Im MCZ; Goias: Goyaz Imt BMNH; Guanabara: Corcovado Im, 2f BMNH; Restinga Jacarepagua Jul. 4m, If AME; Rio de Janeiro Apr. Nov. 6m(lt), 7f(3t) BMNH, If BMB, Im, If AMNH, Im MNHN, Im, 2f MCZ, Im, If ZMHU, Im USNM, 1ft MNHN; Mato Grosso: Burity Jul. If AME; Minas Gerais: Belo Horizonte 3000' Nov. Im BMB; Belo Horizonte-Brasilia, km. 500, Apr. Im AME; Leitao, km. 142 Curvelo May Im USNM; N. Anapolis, Belo Horizonte-Brasilia Apr. 7m AME; Nova Lima If AME; Paracatu Im AME; no specific locality 1ft BMNH; Para: Cuiaba-Santarem, km. 1130, Jul. Im AME; Parana: N. Parana If AMNH; Rio de Janeiro: Itabapoana 1ft BMNH; Laguna de Sacuarestna Aug. Sep. Imt BMNH; Paineiras May 2m USNM; Petropolis Im USNM; Santa Catharina: Blumenau Im, 2f BMNH, 2m MCZ, If AMNH, 3f ZMHU; Corupa 2f AMNH; S3o Bento do Sul 850m Sep. If MUSM; no specific locality Im, If BMNH, Im BMB, If MCZ, 2f ZMHU, Im, If USNM; Sao Paulo: Alto da Serra Santos 800m Feb If BMNH; Anhangahy Dec. Im BMNH; Casa Branca 2m ZMHU; Sao Paulo 2500' Nov. Dec. 2m, 2f MCZ, If BMB; Not located: no specific locality 5m, 5fl[3t) BMNH, Im ZMHU, Im MNHN. COUNTRY UNKNOWN (10: Amazon 1ft BMNH-error. Additional locality data: BRAZIL: Distrito Federal: Sobradinho Woods 1050-1 150m Feb. Aug.; Goids: Rio MaranhSo 700m Jun. Aug.; Minas Gerais: Paraopeba Esta93o Florestal de Experimenta9ao 750m Jun.; Paraopeba Woods 750m Feb. Juti. (Brown and Mielke, 1967); Santa Catharina: Jaragua Mar. Apr. (Hoffmann, 1936). Adelpha plesaure phliassa (Godart, [1824]) Figs. 33c,d; 117a,b Nymphalis phliassa Godart ([1824]: 373) TL: Guyane; Bresil. Types: MNHN?: ST: not located. =Heterochroa euboea C. and R. Felder (1867: 422) TL: Surinam. Types: BMNH(R): STm; "Surinam Cll. V. Lennep Type//Syntype//Type//H. euboea Felder/Zeuboea n." [examined] =Adelpha phliassa implicata Fruhstorfer (1915: 521) TL: Peru. Types: BMNH(T): STm: "phliassa implicata Fruhst.//Fruhstorfer Coll. B.M. 1937-285//Peru H. Fruhstorfer//TYPE//Type//Syntype"; BMNH(M): STm; "Peru H. Fruhstorfer//TYPE//Paratype// Fruhstorfer Coll. B.M. 1937-285" [both examined] =Adelpha phliassa bartolme Fruhstorfer (1915: 521) syn. nov. TL: Mato Grosso [Brazil]. Types: BMNH(T): STm: "bartolme Fruhst.//Fruhstorfer Coll. B.M. 1933-1 31 //Mato Grosso H. Fruhstorfer//TYPE//Type//Syntype" [examined] =Adelpha plesaure cerachates Fruhstorfer (1915: 521) TL: Mato Grosso [Brazil]. Types: BMNH(T): STm: "plesaure cerachates Fruhst.//Fruhstorfer Coll. B.M. 1937-285//Mato Grosso H. Fruhstorfer//TYPE//Type//Syntype" [examined]

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239 =Adelpha plesaure sirona Fruhstorfer (1915: 521) TL: Eastern Bolivia. Types; BMNH(T): STm: "plesaure sirona FruhstV/Fruhstorfer Coll. B.M. 1937-285//Ptov. Sara Dept. S. Cruz de la Sierra February 1904 (J. Steinbach)//TYPE//Type//Syntype" [examined] Heterochroa phliassa Godt., Westwood (1850); =Adelpha plesaure Hubn., Kirby (1871); Adelpha plesaure var. euboea Fldr., Kirby (1871); Adelpha plesaure phliassa Godt,, D'Abrera (1987); Adelpha plesaure phliassa Godt., Neild (1996) Identification, taxonomy and variation: Adelpha p. phliassa is rather weakly distinguished from the nominate subspecies by the orange on the DFW extending broadly to vein Cu2, with no dark brown at the distal edge of the white postdiscal band in this cell. Typically, also, the postdiscal band is only white from the anal margin to vein Cu2, or is colored orange throughout. There is much variation in the color of the posdiscal band in cell Cu2, the width of the DHW white postdiscal band, and the ventral surface postdiscal and submarginal series, which may be sharply defined or almost completely merged. The amount of red-brown scaling between cell bars one and two in the VFW and VHW discal cells is also variable, and may be entirely absent. Godart ([1824]) described phliassa based on specimens from French Guiana and Brazil, and the description corresponds well with this species. Part of the French description, franslated here, reads: "the upperside of the wings is blackish brown, with a white discoidal band, terminating in a sharp point at the anal angle of the hindwings by a yellow marking, losing itself, towards the internal border of the forewings, in an orange band, transverse and larger, of which the external border is sinuous and the internal border deeply indented towards its origin". I have not been able to locate any type specimens in the MNHN, and there remains the slight possibility that the syntype specimens from Brazil may actually be nominate A. plesaure. However, since French Guiana is the first named country in the original description, the description states that the orange band of the DFW begins "towards the internal border of the forewings" as in phliassa as treated here, and certain French Guianan specimens have a sinuate distal border of the DFW orange band, being slightly transitional to A. p. symona, I follow all previous authors (Fruhstorfer, 1915; Hall, 1938; D'Abrera, 1987; Neild, 1996) in

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240 regarding the name as applying to the Guianan population. Felder and Felder (1867) described euboea from a male specimen or specimens from Surinam, ex. coll. Klinkenberg, in their own collection. They compared it to A. plesaure, stating the the DFW postdiscal band was narrower, and the syntype specimen in the BMNH shows it to be a synonym of A. p. phliassa, where it was placed by Hall (1933). There then followed Fruhstorfer's (1915) inexphcable treatment of this species as two distinct species, phliassa and plesaure, for which he provided no justification. As subspecies of the former "species", he described two names, implicata, based on specimens from Peru, and bartolme, based on specimens from Mato Grosso. The characters used to justify these taxa, namely the differing width of the white postdiscal band on the DHW, the overall color of the ventral surface and the slightly increased white shading within the DFW orange band, are all so slight that they would be insignificant even were they not subject to much individual variation. As subspecies of A. plesaure, Fruhstorfer (1915) described cerachates, based on specimens from Mato Grosso, and sirona, based on specimens from Bolivia, each of these supposedly differing from each other and known subspecies only by the characters listed in the previous sentence. Of all of these, cerachates is perhaps the most distinctive, with faded and coalescing postdiscal and submarginal series on the ventral surface, a character often occurring in specimens from drier areas. However, the differences ascribed to all of these taxa, bartolme, implicata, cerachates and sirona, fall well within the usual bounds of variation for this subspecies, and since the last three were synonymised by Hall (1938) with A. p. phliassa, I here synonymise bartolme with that same taxon (syn. nov.). I retain A. p. phliassa as distinct from A. p. plesaure for the present, although the characters which separate the two are weak and variable to some extent. More collecting, or examination of series of specimens with more accurate locality data than those to which I have had access, may well show the two to be connected by continuous clinal variation, and therefore phliassa will need to be synonymised. Certain specimens from

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241 Remolinos, Restrepo and Villavicencio, in Meta, Colombia, have the white postdiscal band on the DHW half the usual width, with more extensive orange on the DFW, and appear to be transitional to A. p. pseudomalea from Venezuela. Range: This subspecies occurs from southeastem Venezuela to Bolivia, throughout Amazonian Brazil and the Guianas. Immature stages: Moss (1933) reared this species in Brazil (Para), and figured both the last instar larva (pi. I, fig. 14) and the pupa (pi. H, fig. 9). In common with the observations of the nominate subspecies by Muller (1886), the larvae are very dark in all instars, the last instar with a few fine white dots, with a lateral oblique stripe of pink and white marking the posterior segments. The scoli are diverse in form, similar to A. basiloides, though Moss' illusfration shows the scoli on T3 and A7 to be shorter and thicker than in A. basiloides. The pupa is pale silvery gray with segments A2 and T2 only weakly dorsally produced, the head horns are long, thin and curved anteriorly. The larva appears to rest in the FrontArched-Rear-Up position (Aiello, 1984). Habitat and adult ecology: This subspecies is common in most lowland forest sites, from primary forest to disturbed forest with much secondary growth, up to 1250m. Males are readily attracted to rotting fish in forest light gaps, and more rarely, banana, hi eastern Ecuador I have observed males perching along a wide ridge top path through secondary forest, from 2-4m above the ground, in the middle of the day. X^VjZT^'^'^^l^^r' ' ' V,*"!^'" * '° PP^-''don,alea; • trans, to A. p. symona. MiKM pf n ^r\ ^wwcw^"° ^""^^ '"^ '^""''^ l'" AMNH; Bolivar. Cuchime, Rio Caura Apr Im 5Se iarioitotn 3^^MNH T''''} ^"^^ '^''^"'^ ^^^Z; Monagas: Barrancas 1" Swk UUcTa Oct 1^ F^M -'/r "^^"'y 2f BMNH. COLOMBIA (13m, IIQ: Amazonas: Florida 2m BMNH Uticia Oct. Im ESM; AnUoquia": "Rio Cocoma, Antioquia, 800-1 100m Aug. L. Richter" Im AMNH-error"'CaauriABogota 2f BMNH; yt/era. Remohnos Apr. Jun. 2f(lt) JFL; Restrepo IfJ JFL; San Jose Guaviare May Jun. Im, 1 f JFL VH avk^do L in ^E(lt), Im, 2fl:it) MCZ, If JFL, Im LMC; No, located: no specific locality Im BMB. ECUADOR ^l^lllZ^T .TJu fr"""" '"^ '"^ ^A'TM^ndez-Santiago rd.,Tm. 40. Nov Im DAT; 2M-S Tg'W 875ni Nov^Dec. Im AMNH; Napo: Apuya Aug. Oct. Im KWJH, Im DAT; Chichicomimi Jul. 2m KWJHCoca 30 km S W AdT m MJP; Coca. 35 km. S.W^, Apr. Im MJP; Finca San Carlo Oct, Im DAT; R,o Latas Im BMB; Pas,aza'. Pu^o Jun If FsS; X, BMNH Im ZMHU, Im USNM, Im BMB; U Merced If BMNH, Im BMB; Satipo 3m AMNH; Rio Colorado 2m BMNHIqu.tos Mar. Jul. Aug^2m, If BMNH, Im ZMHU, Im FSCA, Im USNM; Pebas Jan Oct. Dec. 3m, 1 f BMNH L ZMHt^tm M^^^ ^o Aguas Negras 150m Mar. Im MUSM; Rio Sucusan 140m Sep. 3f MUSM; San Roque Im AME Y^nlguas 1 m zS BOLIVIA (18m. 10. La Paz: Caranav, 1200m Feb. Im MUSM; Coroico 2m MCZ; Rio Songo 1200m Im ZMHU; ^o^" Tup.^

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242 Im AME; Santa Cruz: Buenavista Mar. Aug. Dec. If BMNH, 3m AME; Prov. Sara Feb. Mar. May 4m BMNH, 3m BMB; Santo Cruz Im BMNH; Not located: no specific locality 2m AMNH. BRAZIL (42m, 550: Amazonas: Manaus If BMNH; Manicore If ZMHU; Massaury If ZMHU; S3o Paulo de 01iven9a Im, If ZMHU, Im AME; Serpa Jan.-Mar. If BMB; Tefe Oct. 2m BMNH, Im MCZ; Maranhao: Mts. Aiireos 2f BMNH; Mato Grosso: Burity Jul. Im BMNH; Chapada Im BMNH; Cuiaba Im BMNH; Cui'abSCorumba river system 3m BMNH, Imt BMB; no specific locality 2m BMNH; Para: Itaituba Im ZMHU; Itaituba-Obidos Apr. 2f BMNH; Obidos 2m, 2f BMNH, 3f AMNH; Obidos-Serpa If BMNH; Para Jan.-Mar. Aug. 14m, 29f BMNH, Im* AMNH, If BMB, IfMNHN; Santorem Jan.-Mar. lm,3fBMNH, ImZMHU, U WAB; Ronddnia: Cacaulandia Oct. Nov. IfFSCA, 2m, 2f USNM; A^o/ located: S5o Felippe 2m BMNH; no specific locality 3m, If BMNH, Im, 1ft MNHN. GUYANA (6f): Upper TakuluAJpper Essequibo: Aunai, Essequibo 2f BMNH; Not located: Essequibo R., 140 mi. inland If BMNH; Sabina, Rio Berbice Mar. Apr If BMB; no specific locality 2f BMNH. FRENCH GUIANA (12m, 130. Cayenne: Cayenne 2m, 2f BMNH, If ZMHU, 3m MNHN; Laurent du Maroni: Maroni River 3f AME; St. Laurent du Maroni Jan. May Im, If BMNH; Saul May If MUSM; Not located no specific locality Jun. 6m, 3f BMNH, If BMB, IfMNHN. SURINAM (3m, 60: Para: Bersaba If ZMHU; Para Dist. Nov. If BMNH; Paramaribo: Paramaribo Oct. Nov. If BMNH; Not located: no specific locality 2m, 3f BMNH, Im ZMHU. COUNTRY UNKNOWN (8m, 40: Amazons 2f BMNH, 2m USNM, 4m BMB, Im MCZ; no specific locality 2f BMNH, Im MNHN. Additional locality data: ECUADOR: Morona-Santiago: Bomboiza May; Taisha Jun.; Napo: Apuya Sep.; Capiron Jul.; Chichicorrumi Sep. Dec.; Finca San Carlo Sep. Dec; Rio Tiputini Jun.; Satzayacu Sep.; Yarina Jul.; Pastaza: Pitirishca Jul.; Sucumbios: Pailacocha 250m Oct.; Zamora-Chinchipe: Chachacoma 1250m Nov. (Willmott and Hall, unpubl. sight records)! BRAZIL: Amazonas: Wh& de Maraca (Mielke and Casagrande, [1992]). Adelpha plesaure symona Kaye, 1925 Figs. 33e,f; 199a-c Adelpha phliassa symona Kaye (1925: 414) TL: Northern Hills, Tabaquite, Trinidad. Types: AME: STf: "Type H.TV/Sto. Cruz, Trinidad, 4.XI.21, R. Dick/Zphliassa symona Kaye"; ST?f: Tabaquite [examined] 0996)" "'^'""'^ ' ^'^'"^ ' ^'"""'^ "y"""" l^ye, Neild Identification, taxonomy and variation: This subspecies is distinguished from A. p. phliassa by the scalloped distal margin of the DFW orange postdiscal series, which extend right to the anal margin and the more extensive postdiscal band, which extends as an orange block into cell Ml, with faint orange scaling anterior cell Ml. There is slight variation in both of these characters towards typical A. p. phliassa. Kaye (1925) described this subspecies with great alacrity, stating merely that the forewing band was broader. The description was based on several specimens from the Northern Hills, and Tabaquite, and it is probable that several more of the Trinidadian specimens in the AME are syntypes. This subspecies may occur in coastal areas of the Guianas, or there may be some gene flow, since certain Guianan specimens have the distal edge of the orange DFW postdiscal series scalloped. This scalloping may be what Godart ([1824]) refers to as "sinuate" in his original description oi phliassa, though the deep indentation that is

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243 mentioned of the basal edge of the forewing band near the costa, confirms that Godart's specimens belonged to phliassa as treated here, rather than symona. Range: Extreme eastern Venezuela and Trinidad. Several specimens labeled "Cayenne" in the USNM suggest that this subspecies may also extend along coastal areas of the Guianas, or occasionally disperse there, or it is possible that these are mislabeled. Habitat and adult ecology: According to Barcant (1970), this subspecies is not rare, especially in the wet season, and it occurs up to 500m. Specimens examined (18m, 190: t trans. \oA. p. phliassa. TRINIDAD (17m, HQ: Caroni: Tabaquite Jan. If AME; St. Geroge East: Arima District Jan.-Mar. Oct. Dec. If BMNH, Im, If BMB; Simla Sep. Im FSCA; St. George West: Chancellor's rd. Im AMNH; Fondes Amandes Im AME; Hololo 1000' Oct.-Dec. '2m AME, Im BMB; Macqueripe Bay Apr. If BMNH; Maraval Nov. Dec. Im BMB; St. Anns Oct.-Dec. 5m, 3f BMB; SanU Cruz If AME; St. Patrick: Siparia Oct.-Dec. If BMB; Not located: no specific locality Jan. Im AME, 4f BMNH, Im USNM, 2m BMB If MNHN. VENEZUELA (If): Monagas: U Pica Oct. If MUSM. FRENCH GUIANA (Im, 3f): Cayenne: Cayenne Im. 3fr'2« USNM. COUNTRY UNKNOVm (If): no specific locality If USNM. Additional locality data: TRINIDAD: St. George West: W. of Diego Martin Mome Pierre 1500' Dec.; Mome Catharine 1500' Jan • St. Patrick: Parrylands Oct. (Cock, pers. comm.). Adelpha plesaure pseudomalea Hall, 1938 Figs. 33g,h Adelpha plesaure pseudomalea Hall (1938: 234) TL: Muchuchachi, Venezuela. Types: BMNH(T): HTf: "Illustrated in The Butterflies of Venezuela A Neild 1996//101//pseudomalea Hall i.l.//Joicey Bequest Brit. Mus. 1934-120//44. 20 Muchuchachi Venezuela/ZHolotype" [examined] Identification, taxonomy and variation: This subspecies is easily distinguished from all others by the orange scaling distal of the white DHW postdiscal band, and by the latter being considerably narrower than usual. This subspecies is also similar to A. cytherea nahua, but may be readily distinguished by the ventral surface pattern. The extent of orange on the DHW is slightly variable in the few specimens I have seen. Hall (1938) described this subspecies based on a single female in the BMNH, which I have examined. Range: Western Venezuela, east of the Merida range and west of the llanos, south to Meta province in Colombia.

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244 Habitat and adult ecology: This subspecies is very rare in collections and nothing has been reported on its habitat or behavior. The development of the orange postdiscal band on the DHW produces a remarkable resemblance between this species and the numerous other Venezuelan Adelpha which develop a similar character in populations sympatric with this taxon, and there is ahnost undoubtedly a mimetic relationship between all these taxa. Specimens examined (2m, 1 0: VENEZUELA (Im, \ Merida: Mucuchachi If BMNH; Tachira: La Morita 300tn Aug. Im MUSM. COLOMBIA (Im): Meta: Remolinos Jan. ImJFL. Additional locality data: VENEZUELA: Barinas: Reserva Forestal Caparo (Neild, 1996). Adelpha gavinaVrvihsXoxf&c, 1915 Figs. 34a,b; 118a,b; 200a,b Adelpha gavina Fruhstorfer (1915: 529) TL: Blumenau, Brazil. OTL: Rio de Janeiro, Espirito Santo, Blumenau [Brazil]. Types: BMNH(T): LTm: "Fruhstorfer Coll. B.M. 1937-285//gavina Fruhst.//Brasilen Blumenau Fruhstorfer//TYPE//Type//Syntype"; PLTf: "Fruhstorfer Coll. B.M. 1937285//gavina Fruhst.//Brasilen Blumenau Fruhstorfer//TYPE//Type"; BMNH(M): PLT?m: "Brasilien Fruhstorfer//St. Cath. Sch.//Fruhstorfer Coll. B.M. 1937-285"; PLT?m: "Brasilien Blumenau FruhstorferZ/Fnihstorfer Coll. B.M. 1937-285"; PLT?m: "Brasilien Rio Fruhstorfer/ZFruhstorfer Coll. B.M. 1937-285" [all examined] Adelpha poltius Hall, Brown (1992) misid. Identification, taxonomy and variation: This species is distinguished from the similar A. calliphane and A. poltius by the VFW postdiscal band terminating at vein M3, after which it consists of pale gray shading in cells Ml and R5, instead of the prominent whitish dashes seen in A. thessalia indefecta. It may also be distinguished from the latter species by lacking the outer submarginal series on the venfral surface, instead having a uniform orange-brown submarginal border, and by the slightly darker line which runs through the middle of the pale VFW subapical marking being in the middle of the marking, instead of nearer the distal edge, especially noticeable in cell M3. Adelpha falcipennis is also similar, but has a produced hindwing tomus, the inner submarginal series on the ventral surface is clearly defined, the inner postdiscal series on the VHW consists of a uniform, slightly paler orange-brown line near the distal edge of the white postdiscal band, instead of a pale silvery gray line which is thicker in cells Rs and Sc+Rl, and the VFW orange-brown

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245 marginal border is broader and lacks whitish shading in the apex. Adelpha epizygis and A. abia have the orange DFW subapical marking touching the white postdiscal spot in cell M3 on the DFW, or the orange notably reduced. There is little variation in the small series of specimens examined, except that one male in the AME is an aberrant form in which the usually orange DFW subapical marking is entirely creamy white, and a sinfle male in the FSCA has the white spot in cell M3 on the DFW ahnost completely absent. Fruhstorfer (1915) described this species from two males and four females in his own collection from several BraziUan localities (see TL above). Although the description is rather vague and fails to comment on the most important features of the species, the comparison of the dorsal surface with A. calliphane, and the ventral surface with A. plesaure plesaure, in addition to the statement that the white postdiscal band on the VFW terminates at vein M3, seem to apply only to this species. I have been able to locate only one male and one female syntype, and an additional three possible male syntypes, none of which are from Espirito Santo, one of the type localities. Since it is possible that Espirito Santo specimens may prove to belong to another taxon, I designate the male specimen in the BMNH(T) with the following label data as the lectotype of Adelpha gavina: "Fruhstorfer Coll. B.M. 1937-285//gavina Fruhst./ZBrasilen Blumenau Fruhstorfer//TYPE//Type// Syntype". The precise relationships of this species are not clear, but the VFW white postdiscal band, which is broken at vein M3, suggests it may be more closely related to A. falcipennis, A. thessalia, A. iphiclus, A. iphicleola and A. abyla, than to other A. iphiclus group members. Range: Southeastern Brazil, from Rio de Janeiro to Santa Catharina. Habitat and adult ecology: This species has been recorded from December to March and is rare in collections. However, unless the species was misidentified (very possible), Ebert (1969) reports that it can be locally common in Minas Gerais in high and low canopy primary forest up to 1400m. Brown (1992, as poltius) reports that the species

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246 occurs in similar habitats to A. abia, openings in humid forest from 800m to above 1 100m, in the the Serra do Japi, Sao Paulo, Brazil. Specimens examined: 33 (16m, 170 BRAZIL (15m, 160: Goias: Tower 10 km. N. Geianda 850m Mar. Im USNM; Guanabara. Rio If BMNH; Minas Gerais: no specific locality 2f BMNH; Parana: Uniao da Vitoria 2000' Dec. Jan. If BMB; Rio de Janeiro: Petropolis Independencia 900m Im AME; Santa Catharina: Serrinha do Pirai, W. Joinville 950' Mar. Im FSCA; Blumenau 2m, If BMNH, Im ZMHU; Joinville 80200m Dec. Im MUSM; no specific locality Im, 3f BMNH, 2f ZMHU; Sao Paulo: Alto da Serra Aug. Im MNHN; Casa Branca Im, 2f ZMHU; Cauna Mar. Im, 3f AMNH; Sao Paulo 2m, If BMNH, Im AMNH; Not located: no specific locality Im BMNH COUNTRY UNKNOWN (Im, 10: If BMNH, Im AME. Additional locality data: BRAZIL: Minas Gerais: nr. P090S de Caldas 1000-1400m Apr. May Nov. Dec. (Ebert, 1969); Espirito Santo: no specific locality (Fruhstorfer, 1915). Adelpha falcipennis Fruhstorfer, 1915 Figs. 35a,b; 119a-c; 201a-d Adelpha falcipennis Fruhstorfer (1915: 526) TL: Rio Grande do Sul [Brazil]. Types: BMNH(T): HTm: "Fnihstorfer Coll. B.M. 1937-285//falcipennis Fnihst.//Rio Grande Brasil Fruhstorfer//TYPE//Type//Syntype" [examined]; BMNH(M): PTm: "falcipennis Fruhst.//Rio Grande Brasil Fruhstorfer/ZFruhstorfer Coll. B.M. 1937-285"; 4PT?f: "Rio Grande Brasil Fruhstorfer/ZFruhstorfer Coll. B.M. 1937-285" [all examined] =Adelpha falcipennis perga Fruhstorfer (1915: 527) stat. rest. TL: Santa Cathanna [Brazil]. Types: BMNH(T): STm: "Fruhstorfer Coll. B.M. l937-285//falcipennis pei^ Fruhst.//Brasilen Blumenau Fruhstorfer//TYPE//Type//Syntype" [examined] Adelpha perga Fruhst., D'Abrera (1987) Identification, taxonomy and variation: This species is distinguished from similar species by the elongate hindwing tomus, the broad, orange-brown marginal bprder on the VFW, the clearly defined, silvery gray inner submarginal series, and the VFW white postdiscal band, which is broken at vein M3. There is a little variation in the specimens examined in the color of the brownish areas of the ventral surface, which may be tinted red or orange. Fruhstorfer's (1915) original description of this species is detailed and closely corresponds with the specimen in the BMNH(T). He stated that the Type of this species was in his own collection, and I regard the specimen in the BMNH(T), which bears a red TYPE label, as representing the holotype. He also described a new subspecies, perga, from an unspecified number of specimens from Santa Catharina, which supposedly differed from the nominate in having narrower orange on the DFW and a more 'Variegated" hindwing. In fact the syntype perga in the BMNH is simply a fresher, less faded specimen oi A. falcipennis, and all Fruhstorfer's suggested differences prove to be

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247 individual variation. Hall (1938) synonymised perga with falcipennis, where I also place it (stat. rest.), since D'Abrera (1987) resurrected the name with no explanation. Range: Adelpha falcipennis ranges from southeastern Brazil, from Rio de Janeiro to Rio Grande do Sul, to northeastern Argentina and Paraguay. It probably also occurs in Uruguay. Habitat and adult ecology: This species has been recorded from August to May up to 1400m, and is relatively uncommon in collections. Ebert (1969) reports it to be not uncommon in Minas Gerais in low and high canopy forest. Specimens examined: 62 (45m, 17f) BRAZIL (44m, 17f): Minas Gerais: Nova Lima Sep. Im AME; Parana: Castro 950m 3m BMNH, 2m USNM; Femandes Pinheiro 2600' Apr. 2m AME; UniSo da Vitoria 3m BMB; Sao Paulo: SSo Paulo Im MCZ; Rio de Janeiro: Novo Friburgo 2m ZMHU; Rio Grande do Sul: Catarina Feb. ImAME; Guarani 1 f MCZ; Pelotas Jan. Mar. Apr. 9m, 5f MCZ, 2m BMNH, ImAMNH; Rio Grande 2m, 4f BMNH; Santa Rosa Dec. Im BMNH; Santa Catharina: Serrinha do Pirai, W. Joinville 950' Mar. Im FSCA; Blumenau Im BMNH; Cauna Apr. 5m AMNH; Corupa Im AMNH; no specific locality 5m, 3f AMNH, 4f ZMHU; Sao Paulo: Alto da Serra Aug Im MNHN; Casa Branca Im ZMHU. Additional locality data: BRAZIL: Minas Gerais: nr. Poifos de Caldas 1000-I400m Jan.-May Dec. (Ebert, 1969). PARAGUAY: Alto Parana (Kochalka et at., 1996). ARGENTINA: Misiones (Hayward, 1951). Adelpha thoasa (Hewitson, 1850) Figs. 36; 120; 202 Identification, taxonomy and variation: Adelpha thoasa is distinguished from all other similar species by having orange filling the area between cell bars one and two on the VHW, extending in a band to the costa, in the majority of subspecies being merged with cell bar 3 and the postcellular to form a single orange band bordering the basal edge of the white postdiscal band. In specimens where the VHW cell bars are not merged to form a single band, there appear to be two orange stripes basal of the white postdiscal band which extend and converge from the VFW discal cell onto the hindwing. Often there are two postdiscal dashes on the DFW in cells Ml and R5, which distinguish the species from all similar species. The male genitalia are unique in having large, laterally pointing teeth at the distal edge of the valvae and a very pronounced ventral costal bulge. There is geographic and individual variation in the expression of the white DFW postdiscal dashes in cells Ml and R5, the

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248 width of the white postdiscal band, the shape and size of the orange DFW subapical marking, and the expression of the ventral submarginal series, and four subspecies are recognised. Adelpha thoasa is a distinctve species which has seldom been confused with any other. The wing pattern characters are similar to A. thessalia, and the valvae of the male genitalia, with laterally pointing "teeth" at the distal edge, very possibly indicate an intermediate stage between teeth pointing posteriorly in a plane, as in all previous A. iphiclus group members, and pointing dorsally, as in all following A. iphiclus group members. Range and status: East of the Andes from Venezuela to northern Argentina, Brazil, Paraguay and the Guianas. Uncommon to rare in lowland forest below 1000m, often in drier areas. Specimens examined: 138 (124m, 14f) Adelpha thoasa thoasa (Hewitson, 1850) Figs. 36a,b Heterochroa thoasa Hewitson (1850: 436, pi. DC, fig. 6) TL: river Amazon. Types: BMNH(T): STf?(badly damaged could be m): "Illustrated in The Butterflies of Venezuela A. Neild, 1996//121//B.M. TYPE No. Rh. 9839 Heterochroa thoasa f Hew./ZAmazons Hewitson coll. 79-69 Heterochroa thoasa 2.//thoasa//Paratype"; STm: "Illustrated in The Butterflies of Venezuela A. Neild, 1996//1 19//B.M. TYPE No. Rh. 9838 Heterochroa thoasa m Hew./ZThoasa Hewitson//Type//258//Tapajos//Syntype"; BMNH(M): STf: "Amaz./ZAmazons Hewitson Coll. 79-69 Heterochroa thoasa I."; STm: "Amaz./ZAmazons Hewitson Coll. 79-69 Heterochroa thoasa 3." [all examined] =Adelpha siliaVmhsXoTiQr {\9U:y>\. 107e; 1915: 522) TL: [Encorado, Sa. Cruz de la Sierra, Argentina]. Types: BMNH(R): STm: "thoasa silia Fruhst.//Syntype//TYPE//S. Cruz de la Sierra, E. Bolivia, I905//6 (J. Steinbach)" [examined] =Adelpha thoasa cuyaba Fruhstorfer (1915: 522) TL: Mato Grosso [Brazil]. Types: BMNH(T): STm: "thoasa cuyaba FRihst.//Fruhstorfer Coll. B.M. I937-285//Brasilien Mato Grosso Fruhstorfer//TYPE//Type//Syntype" [examined] Adelpha thoasa Hew., Kirby (1871) Identification, taxonomy and variation: The nominate subspecies is distinguished from remaining subspecies by having two white postdiscal dashes in cells Ml and R5 on the DFW, a narrow orange DFW subapical marking, typically of almost equal width to the white postdiscal dashes, and a broad white postdiscal band. It is most similar to A. t. manilia, which has only a single

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249 white postdiscal spot in cell Ml, with occasionally a few white scales in cell R5, and a broad orange subapical marking. There is much variation in the width of the white DFW postdiscal dashes and the orange subapical marking; in the BMNH(M) there is one male from Villa Nova and one from Sao Paulo de 01iven9a which have a very narrow orange subapical marking and the postdiscal dashes of almost equal width to the postdiscal band, while a single male labeled "Amazon" is intermediate in both these characters to typical A. t. thoasa. The width of the postdiscal band is also slightly variable. This subspecies is comparatively rare in collections and I have seen insufficient specimens with accurate locality data to assess the true nature of variation, but at present it seems largely individual or local. The name thoasa was first introduced as a nomen nudum by Westwood (1850), then described shortly after by Hewitson (1850). Hewitson referred to specimens in the British Museum and his ovra collection from the river Amazon, and I have examined four syntypes in the BMNH. The description and original illustration most closely match the syntype specimen in the BMNH(T) that was illustrated by Neild (1996: pi. 3, fig. 121), and given the variation in this subspecies and the possibility of fiirther stable populations being discovered in the Amazon basin, I designate this specimen as the lectotype of Heterochroa thoasa, with the following label data: "Illustrated in The Butterflies of Venezuela A. Neild, 1996//121//B.M. TYPE No. Rh. 9839 Heterochroa thoasa f Hew.//Amazons Hewitson coll. 79-69 Heterochroa thoasa 2.//thoasa//Paratype". Fruhstorfer (1913) figured, as a new taxon silia, a specimen almost idenfical on the dorsal surface to the lectotype of thoasa, of which he later placed it a subspecies (Fruhstorfer, 1915). Fruhstorfer (1915) refers to the type as being in his own collection, and a fiirther specimen in the Tring Museum (now the BMNH(R)), but I have only been able to locate the latter. Although the type locality is "Encorado, Sa. Cruz de la Sierra, Argentina", this appears to be an error, the specimen actually originating in Santa Cruz de la Sierra in Bolivia. It matches Fruhstorfer's (1915) description in having a darker, more richly

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250 colored ventral surface, but this represents no more than individual variation. The same is true of cuyaba, also described by Fruhstorfer (1915), the syntype of which differs even less from typical A. t. thoasa, and both names were synonymised with A. t. thoasa by Hall (1938). Although cuyaba was described from Mato Grosso, it is possible that the specimen is mislabeled, since all other specimens from that state are A. t. gerona. Range: Southern Venezuela to northeastern Bolivia and Amazonian Brazil. Brown and Mielke (1967) report a specimen of this subspecies in the Museu Nacional in Brazil from "Rio Maranhao" in Goias, where they also recorded A. thoasa gerona, and speculate that these two subspecies meet at the edge of the planalto. It may also be that the specimen of A. t. thoasa is mislabelled. Habitat and adult ecology: There are no published observations on the nominate subspecies, which appears to occur typically in drier regions with mixed cerrado vegetation, such as the Ventuari area of Venezuela, the Meta region of Colombia, the middla and lower Amazon and eastern Bolivia. The subspecies is rare in collections. Specimens examined (31m, 60: t trans, to A. I. manilia. VENEZUELA (Im): no specific locality Im MNHN. COLOMBIA (2m): Cundinamarca. Bogota Im BMNH; Meta: San Josi Guaviare Imt JFL. BRAZIL (15m, 30: Amazonas: Manaus Im BMNH; Maues Im BMNH, 3m ZMHU; Rio Purus If MNHN; 830 Paulo de Oliven9a Im ZMHU, Im BMB, Im BMNH; Serpa Jan.-Mar. If BMB; no specific locality Im ZMHU; Mato Grosso: no specific locality Im BMNH; Para: Itaituba Im, If ZMHU; Para 2m BMNH; Tapajos 2m BMNH. BOLIVIA (5m): Santa Cruz: Buenavista 750m Jan.-Apr. Im BMNH; Prov. del Sara 2m BMNH, Im BMB; Santa Cruz de la Sierra Im BMNH. COUNTRY UNKNOWN (8m, 30: Amazon 3m USNM; Amazons Im, 2f BMNH; Villa Nova Im BMNH; no specific locality 3m, If BMNH. Additional locality data: VENEZUELA: Ventuari (Neild, 1996). BRAZIL: Goias: Rio MaranhSo 700m (Brown and Mielke 1967). Adelpha thoasa manilia Fruhstorfer, 1915 Figs. 36c,d; 120a-c Adelpha thoasa manilia Fruhstorfer (1915: 522) TL: Bolivia. Types: BMNH(T): STm: "thoasa manilia FruhstZ/Fruhstorfer Coll. B.M. 1937-285//Bolivien Frulistorfer/nTPE//Type//Syntype" [examined] =Adelpha thoasa zalma Fruhstorfer (1915: 522) TL: Tarapoto, Peru. Types: BMNH(T): STm: "thoasa zalma Fnisht.//Fruhstorfer Coll. B.M. 1937-285//Peru H Fruhstorfer//TYPE//Type//Syntype" [examined] Identification, taxonomy and variation: This subspecies is distinguished from A. t. thoasa by having only a single, small white postdiscal dot in cell Ml, while that in cell R5 is absent or represented by a few

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251 scales, in addition to having a broader orange DFW subapical marking. Adelpha t. calliphiclea is distinguished under that subspecies, while A. t. gerona has a much more extensive orange DFW subapical marking and darker ventral colors. There is variation in the presence or absence and size of the white postdiscal spot on the DFW in cell Ml and in the width of the orange subapical marking on the DFW. Fruhstorfer (1915) described manilia based on an unspecified number of specimens from Bolivia, stating that it had a broader orange DFW subapical marking and only a small white postdiscal dot on the DFW anterior of the postdiscal band. He went on to describe (Fruhstorfer, 1915) zalma as a subspecies from Tarapoto in Peru, stating that it differed from manilia in having a broader white postdiscal band and the DFW orange subapical marking more vertical than horizontal. Both of these differences are apparent between the syntypes of each of these taxa in the BMNH, and for this reason I regard the zalma syntype as valid, even though it lacks a label indicating it to be from Tarapoto. Probably the original Tarapoto label was lost during subsequent relabelling. Hall (1938), presumably (correctly) regarding the differences as representing merely individual variation, placed zalma as a synonym manilia. Only other Fruhstorfer specimen in BMNH(M) lacks locality data and also white costal spots DFW. This subspecies is rather weakly separated from the nominate subspecies, and fiirther collecting may show the observed differences to be clinal, with characters changing from those typical of A. t. manilia in the foothills of the Andes to A. t. thoasa further east into the Amazon basin. Range: Eastern Colombia to Bolivia, near the base of the Andes. Habitat and adult ecology: This subspecies is generally uncommon throughout its range, and is confined to lowland rain forest in the foothills of the Andes from 2001000m. In eastern Ecuador males are known from three sites, all of which consist of a large clearing with small bushes along a stream or river surrounded by forest in various degrees of

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252 disturbance. In these clearings males perch from 3-5m high on the edges of leaves and patrol the area with a gliding flight with few wing beats, landing on specific bushes up to 20m apart. These sites are also frequented hyA. serpa diadochus, A. iphicleola thessalita, A. thessalia thessalia and A. hyas hewitsoni, and these species are probably involved in mimicry. Males may also be found puddling at damp sand. I have not seen the female in the field and it is rare in collections. Specimens examined (53m, 40: t trans, to A. t. thoasa. COLOMBIA (4m): Cundinamarca: Region du Bogota Im BMNH; Huila: Gigante Aug. Im AME; Mela: San Martin, Llanos of Rio Meta Imt BMNH; Not located: no specific locality Im USNM. ECUADOR (4m, If): Napo: Pimpilala 600m Sep. Oct. If MJP, 2m KWJH; Rjo Shandia 550m Sep. Im KWJH; Yasuni, Est. Cientifica 300m Jul. Im KWJH. PERU (36m, 3f): Cuzco: Quillabamba 950-1050m Mar. Apr. Im MUSM; Hudnuco: Tingo Maria Mar. May Oct. 2m AME, If BMB; Junin: Chanchamayo 1000m 2m BMNH, 2m ZMHU, Im USNM, 2m BMB, Im MUSM; La Merced 2500' Jun. Nov. 2m BMNH, 2m BMB; Rio Colorado 2000' Mar. Apr. 4m BMNH; Satipo Im AMNH; Loreto: Estacion Biol6gica Pithecia, Rio Samiria 180m Sep. Im MUSM; Iquitos Im AMNH; lower Rio Tapiche Jul. Im AMNH; Pebas 2m ZMHU, Im USNM; Sarayacu Im ZMHU; Yurimaguas Im ZMHU; San Martin: Jepelacio 4m, 2f AMNH; Moyobamba 850m Dec. Im MUSM; Rioja Im MUSM; Not located: Huallaga Im AMNH; no specific locality Im BMNH, Im AMNH. BOLFVIA (4m): La Paz: Hotel Rio Selva Resort, Rio Huarinilla 1000m Mar. Im FSCA; Santa Cruz: Juntas 300m Im ZMHU; Not located: no specific locality Im BMNH, Im MCZ. COUNTRY UNKNOWN (5m): Amazon Im BMNH; no specific locality Im ZMHU, 2m BMB, Im BMNH. Adelpha thoasa calliphiclea (Butler, [1870]) stat. nov. Figs. 36e,f Heterochroa calliphiclea Butler ([1870]: 58) TL: Suriname. Types: Coll. unknown: HT: not located Papilio cytherea Linn., Cramer (1781: 170, pi. 376, fig. C, D) misid.; Nymphalis iphicla Linn., Godart ([1824]) misid. as female; Adelpha calliphiclea Bull., Kirby (1 871); =Adelpha thoasa gerona Hayward (1951, 1973) misid. Identification, taxonomy and variation: This subspecies is distinguished from A. t. gerona by the narrower postdiscal bands, the orange DFW subapical marking extending only as orange scaling into cell M3, rather than reaching cell Cul, and on the ventral surface by the submarginal series on both wings being well marked, the pale VFW subapical marking having a dark brown line intruding a short distance from the posterior edge, dividing the two postdiscal series in cell Ml, and the orange stripes in the VHW discal cell being slightly separated by white scaling, instead of fijsed. Adelpha t. thoasa is easily distinguished by having white postdiscal dashes on the DFW in cells Ml and R5, while A. t. manilia has the orange stripes in the basal half of the VHW merged into a single band.

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253 Cramer (1781) figured the dorsal and ventral surfaces of an Adelpha from "Suriname" on plate 376, fig. C and D, which he called Papilio cytherea, stating that he believed it to be that species based on the illustration of Linnaeus' Papilio cytherea published by Clerck (1764). Clearly the specimen is not A. cytherea, since the DFW orange marking is completely isolated fi-om the white postdiscal band, among many other obvious differences. Butler ([1870]) therefore proposed the name calliphiclea for this specimen, and said "There is a species very near to, if not identical with, this insect in the British Museum collection fi-om Bolivia; it may possibly be a race of P. iphicla, which it nearly resembles". Possibly Butler was referring to A. thessalia, which in Bolivian specimens may have an elongated orange DFW subapical marking resembling Cramer's figure. Adelpha thessalia, however, lacks the orange stripes through the discal cell bars in the basal half of the VHW, these being typical of A. thoasa. These stripes appear to originate on the forewing and converge as they cross the hindwing, a character unique to A. thoasa. Both Jorgensen (1922) and Hayward (1973) regarded calliphiclea as representing the same taxon as A. t. gerona, although the specimen figured by Cramer differs in a number of respects (see identification notes above) fi-om typical A. t. gerona, which does not occur in the Guianas. The only explanation seemed to be that Cramer's figure was badly drawn and the specimen mislabeled. However, whilst examining the specimen in the Linnaean collection, courtesy of Martin Honey at the BMNH, referred to by Butler ([1870]) as the type of Linnaeus' Papilio iphiclus, I was surprised to discover that this specimen is a very close match in all important respects to Cramer's figure, and it too was collected in Surinam. I therefore conclude that calliphiclea represents a valid subspecies of A. thoasa (stat. nov.) that is merely very rare (the specimen does not in fact represent Linnaeus' type of Papilio iphiclus, as explained in the notes under A. iphiclus). I have been unable to locate the type of calliphiclea in the BMNH, where several Cramer types are housed (VaneWright, 1975), but it is possible that it is in the RHNL.

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254 Range: To date known only from Surinam, but presumably occurring throughout the Guianas. Habitat and adult ecology: This subspecies is clearly extremely rare in nature, I have seen only a single specimen in all collections examined, collected at least two centuries ago, and it has remained unrecorded in French Guiana, despite an enormous amount of lepidopterological acitivity there in recent years (Brevignon and Brevignon, 1997). Honey (pers. comm.) informs me that the specimen in the Linnaean collection cannot possibly have been the one on which Cramer's figure was based, so I conclude that the subspecies has been collected at least twice! Specimens examined (Im): SURINAM (Im): Not located: "Iphicla 780 Surin. Voght" Im Linnaean Collection, London. Adelpha thoasa gerona (Hewitson, 1 867) stat. rest Figs. 36g-j; 202a,b Heterochroa gerona Hewitson (1867a: pi. Heterochroa, fig. 5, 6) TL: Minas Geraes [Brazil]. Types: BIVINH(T): STm: "Minas Geraes/ZMinas Geraes Hewitson Coll. 79-69 Heterochroa gerona 1.//B.M. TYPE No. Rh. 9841 Heterochroa gerona m Hew.//Type//Syntype" [examined] =Adelpha thoasa cuyaba f brevifascia Talbot (1928: 208) unavailable name TL: Melguira, Mato Grosso, Brazil. Types: BMNH(T): HTm: "39, 27, Melguira, 10 miles S. of Diamantino 2000' 23.v.-3.vi. '27 Matto Grosso C.L. Collenette/presented by J.J. Joicey Esq. Brit . Mus. 193 1-291 /Adelpha thoasa cuyaba f. brevifascia m H.T. Talb. 1928/Type H.T."; BMNH(M): PTm: "39, 27, Melguira, 10 miles S. of Diamantino, 2000', 23.v.-3.vi. Matto Grosso C.L. Collenette/919/Type P.T./Joicey Bequest Brit. Mus. 1934-120" [both examined] Adelpha calliphiclea But!., Kirby (1871) (NOTE: Kjrby listed this as No. la, and iphicla as 1); Adelpha gerona Hew., Kirby (1871); Adelpha thoasa gerona Hew., Fruhstorfer (1915); Adelpha thoasa cuyaba Fruhst., Talbot (1928) misid.; Limenitis thoasa gerona Hew., Brown and Mielke (1967); Adelpha gerona Hew., D'Abreia (1987) Identification, taxonomy and variation: This subspecies is distinguished from the nominate and A. t. manilia by always lacking any white postdiscal dashes on the DFW in cells Ml and R5, by having a broad orange DFW subapical marking which typically extends into cell Cul, by having the pale VFW subapical marking a uniform yellowish brown, with no darker brown scaling, and by usually having the submarginal series on the ventral surface reduced, so that the distal half appears darker in color. Adelpha t. calliphiclea is very similar and is distinguished under that subspecies. Other similar species, such as A. thessalia indefecta, can be distinguished through the diagnostic characters of this species. There is slight variation in

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255 the expression of the VHW inner submarginal series, which may be entire or obscured throughout, and in the extend of the DFW orange subapical marking, which may only just enter cell Cul. All of the specimens I have seen from two localities in Mato Grosso, Melguira and Buriti, differ from typical A. t. gerona in having narrower postdiscal bands, a reduced orange subapical marking that rarely reaches cell Cul and usually only just enters cell M3, and a darker, red-brown VFW subapical marking. These specimens may very possibly represent a distinct subspecies, perhaps inhabiting the watershed of the upper Rio Paraguai, since the specimens I have seen from Nivac (Mato Grosso) and Paraguay are typical A. t. gerona. However, I refrain from describing a new subspecies until more specimens become available, since all of the specimens examined originate from essentially the same site and show variation in the characters that distinguish them from A. t. gerona. There are two male specimens in the BMNH(T) and BMNH(M), from Melguira, that have atypically narrow postdiscal bands, apparently representing aberrations. These specimens are the holotype and paratype of the name brevifascia (see below). Hewitson (1867a) described A. t. gerona based on an unspecified number of specimens in his ovm collection from Minas Geraes, and the syntype in the BMNH(T) very closely matches the original illustration. Talbot (1928) proposed the name brevifascia for two specimens collected by Collenette at Melguira, Mato Grosso, differing from typical A. thoasa thoasa by having narrow white postdiscal bands. Although the narrow width of the band is probably aberrant, these specimens are part of a population that may prove to be subspecifically distinct (see above), but since the name brevifascia was proposed as a form of A. thoasa cuyaba it is a quadrinomial and is therefore unavailable. Range: Eastem and southeastern Brazil to northern Argentina. Habitat and adult ecology: Little is known of the habitats or adult behavior of this subspecies, which is uncommon and has been recorded from 6001000m.

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256 Specimens examined (39m, 4f): BRAZIL (27m, 30: Bahia: San Antonio da Barra Im, If BMNH; Distrito Federal: Parque do Gama 950m May Im USNM; Mato Grosso: Nivac Im BMNH, Im BMB; Minas Gerais: Belo Horizonte 3000' Aug. Nov. 2m BMB, Im USNM, Im AMNH; Nova Lima Apr. Im AME; Passa Quatro 1000m Im MNHN; no specific locality Apr. 2m BMNH; Parana. S. Parana Im BMNH; Sao Paulo: Anhangahy Nov. Dec. 2m BMNH; Casa Branca 3m, If ZMHU, Im BMB, Im MCZ; SSo Paulo 3m, If BMNH, Im ZMHU, 2m USNM; Not located: no specific locality Im BMNH. PARAGUAY (Im): no specific locality Im BMNH. ssp. nov.? BRAZIL (1 Im, 10: Mato Grosso: Buriti 30 mi. N.E. Cuyaba 2250' Jul. 3m BMNH, 4m, If AME; Colegio Buriti 700m May 2m USNM; Melguira, 10 mi. S. Diamantino 2000' May-Jun. 2m BMNH. Additional locality data: BRAZIL: Distrito Federal: Sobradinho Woods 1050-1 150m Jan.; Goias: Anapolis lOOOm Jan.; Cavalcante 900m; Goiania 800m Aug.; Leopoldo BulhSes 1000m Dec; Rio MaranhSo 700m Aug.; Vianopolis 1000m Mar.; Minas Gerais: Paraopeba Esta^ao Florestal de Experimenta^ao 750m Feb. Jun.; Paraopeba Woods 750m Jun.; Uberlandia 800m; km. 222 BR-040 700m Aug. (Brown and Mielke, 1967). ARGENTINA: Misiones (Hayward, 1973). Adelpha thessalia (C. and R. Felder, 1867) Figs. 37; 121; 203 Identification, taxonomy and variation: This species is very similar to A. iphiclus and A. iphicleola, from which it is distinguished by having a red-brown line dividing the postdiscal series in the pale VPW subapical marking in cells M2 and Ml, and the inner and outer postdiscal series on the VHW are straight and parallel, rather than closer at vein M3, though the latter character is sometimes not evident in certain specimens of A. iphicleola. Adelpha iphiclus iphiclus is also always distinguished by having the orange DFW subapical marking only extending into cell M2 as a "hook" at the distal edge of the marking, instead of broadly bordering vein M3. The male genitalia of A. thessalia differ slightly but consistently from those of A. iphiclus, A. iphicleola and A. abyla by having the line of dorsally pointing "teeth" at the distal tip of the valve arising from the middle of the side of the valve, rather than near the top, with the teeth not extending above the dorsal edge of the valve in lateral view, and the valve not tumed upwards at the distal venfral tip. There is local and geographic variation in the size of the orange DFW subapical marking and the expression of the postdiscal and submarginal series on the ventral surface, and three subspecies ae recognised. Adelpha thessalia belongs in a group of species that includes A. iphiclus, A. iphicleola and A. abyla, all of which have a series of dorsally pointing "teeth" arising from the side of the male genitalic valvae near the distal tip. Within this group, A.

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257 thessalia appears to be the most primitive species, with the postdiscal series still visibly distinct in the pale VFW subapical marking. Range and status: Foothills of the eastern Andes from Colombia to northern Argentina, eastern Brazil to Uruguay and Paraguay. In forest up to 2000m. Common. Specimens examined: 476 (436m, 40f) Adelpha thessalia thessalia (C. and R. Felder, 1867) Figs. 37a,b Heterochroa thessalia C. and R. Felder (1867: 417) TL: Rio Negro Superior, Brasilia Septentrional. Types: BMNH(R): STm: "Type//Syntype//Rio Negro Type/ZThessalia FelderZ/Thessalia n." [examined] Heterochroa ephesa ? Men., Felder and Felder (1862) misid.; Adelpha basilea var. thessalia Fldr., Kirby (1871)Adelpha thessalia thessalia Fldr., Fruhstorfer (1915) Identification, taxonomy and variation: This subspecies is distinguished from A. t. cesilas by having a darker ventral surface, with thinner postdiscal and submarginal series and less or no red-brown shading between discal cell bars one and two on the forewing, and by often having the DFW orange subapical marking confined to cells M2-R5, instead of extending into cell M3. Adelpha iphicleola thessalita and A. iphiclus ephesa (not sympatric) are also very similar, but may be distinguished by the characters that define the species. There is individual variation in the size of the orange spot in cell M3 on the DFW, which is usually absent in the north of the range and more prevalent in the south, one male Bolivian specimen has the orange extending along the outer postdiscal series into cell Cul, probably representing intergradation with A. t. cesilas. Felder and Felder (1867) described this species based on an unspecified number of male specimens from the upper Rio Negro in Brazil, hi fact, this locality is erroneous, and all Felder specimens labeled "Rio Negro" actually originated in the Huallaga valley above Yurimaguas in northeastern Peru (Lamas, 1976). The original description compares the species with A. iphiclus ephesa, which this species closely resembles on the dorsal

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258 surface, and comments on the most important diagnostic character for A. thessalia, the red-brown Hne dividing the pale VFW subapical marking, in addition to the less heavily marked ventral postdiscal and submarginal series. I have examined a syntype male in the BMNH. Early authors regarded this species as a variety of A. iphiclus, until Fruhstorfer (1915) correctly treated it again as a distinct species. Range: The eastern slopes of the Andes from Colombia to northern Bolivia. Habitat and adult ecology: This subspecies is common in intact to heavily disturbed primary cloud forest habitats and lowland rain forest along the base of the Andes, from 600-1 800m, with records of 2000m in Bolivia, though it occurs most commonly from 8001500m. It flies throughout the year, at least in eastern Ecuador. Males may be seen puddling singly or attracted to rotting fish along forested rivers, while the female is much rarer, I have only seen a single specimen in collections and have never observed it in the field. Specimens examined (199m, If): t trans, to^. t. cesilas. Z^^M^r p^"™^ " o^if ' '"""J'^y '"^ BMNH-error. COLOMBIA (31m); Boyaca: Muzo 8m AME; Santa Maria Dec. L aLh T; f°P^y^" '"B'^NH. C««rf,„a^«.c«: Bogota Im BMNH; Meta: Rio Negro 2400' Jan. Im USNM; Villavicencio M,T,ff a'mmhT •.u'''''^' I? "° "^^"^ "^NHN. 5m AMNH. ECUADOR (46m): Morona-SantiaZ Macas Im AMNH; Rio Abanico Oct. Im MJP; 2°-4=S, Tg-W 875m Nov. Dec. 2m AMNHNana Satzavacu Dec Im AMp: Pasta.a: ?uyo Dec. 2m AMNH; R-o Puyo 1300m Oct. Im KWJH; Shell 1050m Feb. 2mj;i!w 'Sc/.S .-S^^^ NeZtTooi^rt A^^^^^^^^ M "tu,"'"" ''^'"^ Tunsurahua: Rio Cholo Hay 1300^ Sep. ImTS fSo ?X)'r,^RM>.H ' ^7 '"^ ^"'""'"-Chinchipe: Quebmda Chorillos Nov. 3m DAT; Za^ra ^N^ PERU^8^m m r IT^I' f "^.i"' ^NH; no specific locality 3m AMNH_ PERU (82m lf)^mazo«a5: Falso Paqu.sha 800m Oct. Im MUSM; Huambo 3700' 2m BMNH; Huayabamba Im ZMHUu7 Aus Oct Tm FS?A 7m A^ 1 ^"^ ^MNH, 2m, If BMB, 2m MUSM; Tingo Maria Mar. t x],,cTx c 'J ' Chanchamayo 1000m 14m BMNH, 2m ZMHU, 2m BMB; La Merced 6m BMBRio Per^e Im MUSM; San Ramon 3000' Aug.-Oct. 2m BMNH; Satipo Mar. Sep.-Dec. Im BMNH. 4m AME^rer>Peba^ Im Tm R^NH V ° T ini BMNH; />«.o: Carabaya Jun. Im BMNH; Chaqu.mayo 2500' Apr. Im BMNHTLa uZ 2000N^^ Im BMNH; 5a„ A/«rtm: Huayabamba 3500' Im BMNH; Jepelacio 3m AMNH; Tarapoto Im BMNH, Im ^H^cavMBoqueron Abad Mar. Im AME; No, located: no specific locality 6m BMNH, Im BMB, Im MNHN, Im JF^ bSu (37mV Som Z ZMHrr' '^"fT^^ ^"^^"^ ^^""^^ ^^^^ ^ME; 5 days N. Cochabamba Aug. 2m Lnh San Antonio J"^^, i,? ^""^^ ^'P'"'° ^^"'^ 5"' BMNH; La Paz: below Coroico 1200m Mar Im FSCA Caranav^^ 70^ . m S Anr ^7:.^u ir^^^''''' ^^^^ ^""S^L; ChuVulnSm J^Tm B^^ C^o co (2m): no specific locality Im MNHN, ImMCZ oivid. i,uur«iiKY UNKINOWN

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259 Adelpha thessalia cesilas Frahstorfer, 1915 Figs. 37c,d; 121a,b Adelpha thessalia cesilas Fruhstorfer (1915: 523) TL: Pilcomayo to the Rio Grande [Bolivia]. Types: BMNH(T): STm. "Fnihstorfer Coll. B.M. 1 937-285//thessalia cesilas Fruhst.//R. Pilcomayo-R. Grande December 1903 (J. Steinbach)//TYPE//Type//Syntype" (examined] =Adelpha abia fa. ampla Hayward (1935: 189, pi. XIII, fig. 5) syn. nov. TL: Yuto, Salta, Argentina. Types: FIML: HTm: "Adelpha abia Hew. fa. ampla Holotipo m/A'uto Salta 2I/6/33//Tipus//313r* [photograph examined]; ATf, 3PTm, 2PTf: Tabacal, Salta, 18.VI.33, Kohler; IPTm, 2PTf: Urundel, Salta, 16.V1.33, KOhler [not examined] Adelpha mincia ampla Hay., Hayward (1973) Identification, taxonomy and variation: This subspecies is distinguished from the nominate by the enlarged, more heavily marked postdiscal and submarginal series on the ventral surface, particularly the latter, v^hich almost merge on the hindwing. The area basal of the white VHW postdiscal band is also lighter, the red-brown line bordering the basal edge of the postdiscal band is thinner and does not merge with the third discal cell bar, the discal cell bars on the VFW are thinner and have less red-brown scaling between them, and the silvery white dashes of the upper postdiscal band are much broader, ahnost filling the area between vein M3 and the costa. Characters that distinguish this subspecies from A. t. indefecta are discussed under that subspecies. The certain specimens also have a longer orange DFW subapical marking, which extends into cell M3, and although this is subject to slight individual variation, there appears to be a general tendency for the marking to increase in size from north to south of the range. Specimens from Mato Grosso typically have a slightly narrower orange DFW subapical marking which does not extend into cell M2, and broader white postdiscal bands. Fruhstorfer (1915) described this subspecies based on an unspecified number of specimens collected by Steinbach in December, from the Rio Pilcomayo to the Rio Grande in Bolivia. I have examined a syntype in the BMNH(T), while there are a fiuther two possible syntypes in the BMNH(R) with the same locality data as this syntype. Fruhstorfer mentioned most of the distinguishing characters discussed above, and attributed them to some extent to the subspecies inhabiting a drier zone. He states that

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260 specimens of the nominate subspecies from Coroico, collected in August, also have a paler ventral surface, being the product of the dry season. Jason Hall and I have collected specimens from Coroico and Caranavi in September which have slightly paler venfral surfaces, but it is unclear from the specimens that I have examined in collections from this region whether these differences are seasonally induced or the result of intergradation with A. t. cesilas. Certainly the subspecies seems phenotypically stable for the diagnostic characters over the range given below. Hayward (1935) described ampla as a form of A. abia, following Fruhstorfer's (1915) erroneous application of the name to A. thessalia indefecta, from a type series of ten specimens from Yuto, Tabacal and Urundel in northern Argentina. I have examined a photograph of the holotype, courtesy of Gerardo Lamas, and the original figure of, I presume, one of the paratye males, all of which are in the FIML. These specimens differ from the syntype of cesilas only in having the DFW orange subapical marking extended just past vein Cul, a typical characteristic of southern specimens which also occurs in some Bolivian specimens, and I therefore synonymise ampla with cesilas (syn. nov.). Hayward (1973) lists both A. thessalia cesilas and ""A. mincia ampla" from Salta in Argentina, presumably based on the variation in the DFW orange subapical marking. Range: Andean foothills in southern Bolivia to northern Argentina, northeast to Mato Grosso in Brazil. Habitat and adult ecology: There are no published observations on this subspecies, which is not uncommon in collections. It is not clear whether this subspecies is confined to hilly areas, and thus exists in three disjunct populations, or is continuously distributed throughout the northeastern Bolivian plains. Specimens examined (54m, 2f): t intermediate to ,4. i. indefecta, orange on DFW into anterior half of cell Cul. BOLIVIA (llm): Chuquisaca: Rfo Burmejo-Rio Pilcomayo Dec. 2m BMNH; Rio Grande Dec. 2m BMNH; Rio Pilcomayo-Rio Grande Dec. 3m BMNH; Santa Cruz: Buenavista 400m Im MUSM; Chiquitos 2m MNHN; Santiago de Chiquitos Aug. Im BMNH. ARGENTINA (34m, 2f): Jujuy. Arroyo Apr. 5m AMNH; Calilegua 600m Apr. 5m AMNH; P.N. Calilegua 1600-2500m Feb Apr 9m, 2f AMNH; Salta: Agua Blanca 500m Jun. 2m AME; Colonia Santa Rosa 350m Oct. Im AMNH; Mosconi May Im AMEOran Jun. 3m(lt) AME; Piquerenda Vieja Feb. 550m Im AMNH; Rio Seco, Urundel 350m May Jun. 5m AMNH, Im AMESalta '450m Apr. Im USNM, Im BMB. BRAZIL (8m): Mato Grosso: Buriti Apr. Jul. 4m USNM, Im AME; Chapada Im MCZCuiabaCorumba nver system Im BMNH; Cuiaba Im USNM. COUNTRY UNKNOWN (Im): Puihal, L.C. Apr. Im FSCA(=Pinhal Sta.

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261 Additional locality data: ARGENTINA: Salta: Tabacal (Hayward, 1935). Adelpha thessalia indefecta Fruhstorfer, 1913 stat. nov. Figs. 37e,f; 203a,b Adelpha indefecta Fruhstorfer (1913: pi. 107e; 1915: 523) TL: [Paraguay]. Types: BMNH(R): STm: "Syntype//TYPE//33//R//Sapucay Paraguay 12.X.03 (W. Foster)//iphicIa indefecta Fruhst." [examined] =Adelpha mincia Hall (1938: 257) syn. nov. TL: Paraguay. Types: BMNH(T): LTm: "Fruhstorfer Coll. B.M. 1937-285//abia mincia Fruhst./ZParaguay H. Fruhstorfer//TYPEi//Syntype"; numerous PLT, see below [all examined] Adelpha iphicla indefecta Fruhst., Fruhstorfer (1915); Adelpha abia Hew., Fruhstorfer (1915), Hayward (1931) misid.; Limenitis mincia Hall, Brown and Mielke (1967); Adelpha mincia Hall, D'Abrera (1987) Identification, taxonomy and variation: This subspecies is distinguished from A. t. cesilas by having narrower white postdiscal bands, especially on the DFW where the band usually tapers anteriorly, a broader orange DFW subapical marking that typically extends to cell Cul, and a darker ventral surface similar to the nominate subspecies. It differs from the nominate subspecies in the more extensive orange DFW subapical marking, which extends at least to broadly border vein Cul, and in the inner sumarginal series on the venfral surface typically being broader. Adelpha iphiclus ephesa and A. iphicleola leucates are distinguished by the characters diagnostic for this species, in addition to several mentioned in the discussion below concerning the identity of the holotype specimen of indefecta. There is some variation in whether the orange DFW subapical marking enters cell Cul, and slight variation in the width of the white postdiscal band. Fruhstorfer (1913) figured the dorsal surface of this subspecies based on a single specimen in the Tring Museum from Paraguay, and later (Fruhstorfer, 1915) described it as a subspecies of ^. iphiclus. It should be noted that although the syntype specimen is known to be unique from Fruhstorfer's (1915) comments, it cannot be regarded as a holotype since there was no information on the number of specimens in the original description (Fruhstorfer, 1913). The syntype specimen is in the BMNH(R), but unfortunately lacks an abdomen. It differs from all other specimens o^ A. t. indefecta that

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262 I have examined in the orange DFW subapical marking consisting of an irregular pentagon in cells M2-R5, as in typical A. t. thessalia, with an orange bar in the anterior half of cell M3, and slight orange scaling at the outer postdiscal series at vein Cul. After a great deal of thought and examination of long series of specimens, I conclude that this syntype specimen is actually aberrant in the shape of the reduced orange DFW subapical marking, which causes the resemblance to A. iphiclus that led Fruhstorfer (1915) to place it as a subspecies of that species. The syntype of indefecta differs from specimens transitional between A. iphiclus iphiclus and A. iphiclus ephesa in the BMNH from Sapucay, the type locality, in having orange in cell M3 on the DFW at the basal edge of the subapical marking (absent in all A. i. ephesa and A. i. iphiclus), the basal edge of the orange DFW subapical marking is more vertically orientated, the hindwing is more triangular with a sfraight rather than convex distal margin, the white postdiscal band on the DHW is thinner and has a slightly concave rather than convex distal margin, the postdiscal series on the VHW are parallel rather than closer at vein M3, and the pale VFW subapical marking has a trace of a red-brown line dividing the postdiscal series in cells M2 and Ml, absent in all A. i. ephesa and A. i. iphiclus. All of these differences are characteristic of typical specimens of this taxon, and of A. thessalia in general, and I therefore place indefecta as a subspecies of A. thessalia (stat. nov.). Fruhstorfer (1915) treated A. t. indefecta as A. abia, an error noticed by Hall (1938), who described the taxon as a new species, A. mincia, based on a Fruhstorfer manuscript name published earlier as a nomen nudum by Martin et al. ([1923]). All subsequent authors then followed Hall in referring to this taxon as Adelpha mincia. Hall mentions there being specimens of this taxon in the BMNH from a long series of localities, including Goyaz, Bahia, Minas Geraes, Rio de Janeiro, Sao Paulo, Santa Catharina, Parana, Rio Grande do Sul, Paraguay and N.W. Argentina. He also included specimens from his collection (now the BMB) from Parana and Novo Friburgo. The description is brief but clearly refers to this species, and Hall mentioned that he was introducing the name based on a manuscript name

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263 attached to certain Fruhstorfer specimens and that the types were jfrom Paraguay in the BMNH. As it is not possible to trace all of the specimens designated by Hall as types, since Hall failed to label any of them, and given the remote possibility that some might apply to other taxa, I designate as the lectotype of Adelpha mincia the specimen in the BMNH(T) with the following data: "Fruhstorfer Coll. B.M. 1937-285//abia mincia FruhstV/Paraguay H. Fruhstorfer//TYPE//Syntype". This specimen is typical of A. t. indefecta and I therefore synonymise mincia with indefecta (syn. nov.). Range: Eastern Brazil to Uruguay, west to northeastern Argentina and Paraguay. Habitat and adult ecology: This subspecies is one of the most common species in its area, and has been recorded in every month of the year up to 1400m. Contrary to other subspecies, the female appears to be not uncommon. Ebert (1969) found the subspecies to be not uncommon to common in Minas Gerais in secondary forest habitats. Brown (1992) reports the subspecies from the Serra do Japi, Sao Paulo, in lowland to premontane, semihimiid forest, where it occurs in open areas, as well as in fields, gardens and areas of extensive secondary growth. Specimens examined (1 83m, 37f): t trans, to A. t. cesilas. BRAZIL (128m, 270: Amazonas: Canuma Apr. 4m, If AMNH-error; Bahia: Cachimbo Im BMNH; Goias: Goyaz 2m BMNH; Guanabara: Rio 2m BMNH, Im MCZ; Minas Gerais: Belo Horizonte Jun. Aug. If AMNH, Im USNM; Passa Quatro Im MNHN; no specific locality 2m BMNH; Parana: Castro 950m Feb. Mar. Oct. 8m, 3f BMNH, 3m, If USNM; Caviuna Jun. 2m, 2f AMNH; Femandes Pinheiro 2600' Apr. 3m AME; Igua9u Oct.-Dec. 6m BMNH; N. Parana 5m, If AMNH; Rolandia Nov. If AMNH; Toledo Im AME; Uni3o da Vitoria 2000' Dec. Jan. I8m BMB; no specific locality Im AMNH; Rio de Janeiro: Nova Friborgo 2800' Jan. Im BMB; Petropolis Im USNM; Rio Grande do Sul: Guarani Jan. Dec. 2m MCZ, Im BMNH; Pelotas Nov. Jan. Feb. May 5m, 4f MCZ; Rio Grande do Sul 4m, 9f BMNH, Im ZMHU; Santa Cruz Apr. Im AME; Santa Rosa Dec. Im BMNH, Im MCZSanla Catharina: Blumenau 2m BMNH, Im MCZ; Campos de Palmar Im MCZ; Cauna Apr. 15m, 2f AMNH; Lages Im ZMHU; Pinhal Im AME; Trombudo Alto Jan. Im AME; no specific locality 7m AMNH, 3m(2t) MCZ, 2m USNM, Im ZMHU; Sao PauloAnhangahy Nov. Im BMNH; Aratatuba Oct. Im AMNH; Casa Branca 3m, If ZMHU; SSo Paulo Mar. May 3m, If MCZ, 4m BMNH, Im AMNH; Not located: no specific locality Im MNHN. PARAGUAY (37m, 50: Caaguazu: Yhii Sep.-D^. 7m BMNH Central: Aregua Im AME; Asuncidn Jul.-Sep. Nov. Im JFL, Im AMNH, Imt MNHN; Nueva Italia Sep. Im AMNH; Trinidad Jul 8m, 3f AMNH; Im, If USNM; Paraguari: Sapucay Oct. Dec. 6m BMNH, Im USNM, Im BMB; San Pedro: Jejuy R. Im BMNH Not located: Central Paraguay 3m BMNH; no specific locality Nov. Im JFL, Im, If BMNH, Im USNM, Imt MNHN ARGENTINA {8m, 20: Buenos Aires: Buenos Aires If MNHN; Entre Rios: la Soledad, close to frontier of Uruguay If BMNH Misiones: Puerto Aguirre May 3m BMNH; Rio Parana Nov. Im BMNH; San Ignacio May Sep. Im USNM, Im BMNHno specific locality Nov. Im AMNH; Not located: Chaco de Santiago, Rio Salado Mar. Im BMNH. URUGUAY (20: A/o«/ev;V/eo: Montevideo "at light" If MCZ; Not located: Isla de Paredan Mar. If MCZ. COUNTRY UNKNOWN (10m, 10: no specific locality 6m MCZ 3m USNM, If BMNH, Im MNHN. .iA.-my om iviL,z., Additional locality data: BRAZIL: Goitis: Anapolis 1000m Jul. (Brown and Mielke, 1967); Minas Gerais: nr. Pocos de Caldas I200-1400m Jan.-May Dec. (Ebert, 1969). PARAGUAY: Alto Parana, Canindeyu; Guaird; ItapuaMisiones, Paraguari (Kochallca et al., 1996). URUGUAY: Not located: Artigas (Yacare) Jan. (Biezanko et al., 1957). Adelpha iphiclus (Linnaeus, 1758) Figs. 38; 122; 204

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264 Identification, taxonomy and variation: There is no single character which distinguishes this species from its sister species, A. iphicleola, but taxa that are apparently parapatric or closely allopatric are grouped together on the basis of shared characters between pairs of taxa. In fact, I regard the problem of how the taxa are related between these two species as certainly the most difficult in the genus, and one of the few which I have not been able to solve to my complete satisfaction. In particular, specimens from the biogeographically complex region of northern and northwestern Venezuela, central and northwestern Colombia and eastern Panama are occasionally impossible to identify with certainty using the characters that I have found usefiil to distinguish the two species elsewhere in their ranges. The same is true of specimens from Trinidad and the coastal regions of eastern South America. Part of the problem is the probable influence of seasonality in these peripheral areas, the lack of long series in collections with accurate locality data and the natural variability of these species. Nevertheless, I offer a discussion below of the characters that are stable in certain regions, and hope that workers will concenfrate on solving this problem in their respective areas. In northwestern central America, where this species is sympatric with nominate A. iphicleola, the two species are always readily distinguishable. Adelpha iphiclus has the orange DFW subapical marking only entering cell M2 as a small, posteriorly directed orange dash at the distal edge of the marking, whereas in A. iphicleola the marking extends to broadly border along vein M3. Other characters are discussed under A. iphicleola, and many of these, particularly ventral characters, are apparent and usefiil in separating specimens of ^. iphiclus from A. iphicleola leucates, while the enlarged shape of the DFW orange subapical marking distinguishes A. iphicleola thessalita from A. iphiclus. Problems in identification arise when the orange DFW subapical marking is reduced in width in A. iphicleola, such that it extends into cell M2 only at the distal tip, as in typical A. iphiclus. The identification of these subspecies is discussed under each

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265 taxon. I have dissected a number of specimens from throughout the range of both of these species, and can find no consistent genitahc differences, in contrast to the claim of Austin (1992). Adelpha iphiclus is subject to both individual and geographic variation in the shape of the orange DFW subapical marking, the wing shape and the width of the postdiscal band, and three subspecies are recognised. This species is one of the most taxonomically confiised, and a detailed discussion is given under the nominate subspecies. It is the sister species of A. iphicleola and possibly yl. abyla. Range and status: Mexico to western Ecuador, Venezuela to Bolivia, Brazil, Paraguay, Guianas and Trinidad. Common everywhere in disturbed forest habitats below 1250m. Specimens examined: 1113 (946m, 167f) Adelpha iphiclus iphiclus (Linnaeus, 1758) Figs. 38a-d; 204a,b Papilio iphiclus Linnaeus (1758: 486, n. 172) TL: Cayenne, French Guiana. OTL: Indiis. Types: BMNH(M): NT: "Cayenne//Ex. Oberthur Coll. Brit. Mus. 1927-3." [examined] ^Papilio basilea Cramer (1777: 139, pi. 188, fig. D) stat. rest. TL: Cayenne, French Guiana. OTL: Suriname. Types: BMNH(M): NT: "Cayenne//Ex. Oberthur Coll. Brit. Mus. 1927-3." [examined] ^Adelpha iphicla funalis Fruhstorfer (1915: 523) TL: Rio Dagua, Colombia. Types: BMNH(R): HTm: "Syntype//TYPE//Rio Dagua, Colombia W. Rosenberg/Ziphicla funalis Fruhst." [examined] =Adelpha iphicla exanima Fruhstorfer (1915: 523) TL: Venezuela. Types: BMNH(T): LTm: "Illustrated in The Butterflies of Venezuela A. Neild, 1996//122//Fruhstorfer Coll. B.M. 1937-285//iphicla exanima Fruhst./A'enezuela Fruhstorfer//TYPE//Type//Syntype"; BMNH(R): PLTm: "TYPE/ZSuapure Venez. 13.XI.99 (KJagesy/iphicla exanima Fruhst." [both examined] =Adelpha iphicla pharae Fruhstorfer (1915: 523) TL: Mato Grosso [Brazil], Peru, Bolivia. Types: BMNH(T): STm: "Fruhstorfer Coll. B.M. 1937-285//iphicla pharae Fruhst./ZPeru H. Fruhstorfer//TYPE//Type//Syntype"; BMNH(M): STm; "TYPE//Peru H. Fruhstorfer//Paratype//Fruhstorfer Coll. B.M. 1933131"; STm: "Mato Grosso H. Fruhstorfer//Paratype//iphicla pharae Fruhst./ZFruhstorfer Coll. B.M. 1937-285" [all examined] Papilio iphiclus Linn., Linnaeus (1764: 31 1, n. 129), Drury (1770: 27, pi. 14, fig. 3, 4); Papilio cythereus Linn., Herbst (1793: 137, pi. 148, fig. 1, 2) misid.; Nymphalis iphicla Linn., Godart ([1824]: 374, n. 80) in part, refers to Dairy's and Clerck's figs.; Adelpha basilis Hubn., Hubner ([1819]) repl. name?; Heterochroa iphicla Linn., Westwood (1850) in part, refers to Drury's and Clerck's figs., Hewitson (1850), Bates (1865), C. and R. Felder (1867), Butler ([1870]: 58); Heterochroa basilea Cram Westwood (1850), Menetri^s (1857), Boisduval (1870); Adelpha basilea Cram., Kirby (1871); Adelpha iphiclus Linn Aunvillius (1882), Fruhstorfer (1915), Hall (1938), Hayward (1973), DeVries (1987), Neild (1996); Adelpha iphicla Linn Godman and Salvin (1884); Limenitis iphicla Linn., Ross (1964, 1976) (probably/), iphicleola iphicleola). Brown and Mielke ( 1 967); Adelpha iphicla iphicleola Bates, D'Abrera (1 987), in part, misid.; Adelpha iphiclus iphicleola Bates, Umas and Small (1992) in part, misid. Misapplication of the names iphiclus and basilea: Papilio iphiclus Linn., Clerck (1764: tab. 41 [fig. 3, numbered in register at back]) = A. naxia; Papilio iphicla Linn., Cramer (1777: 139, pi. 188, fig. E, F) = A. serpa diadochus, Papilio iphiclus Linn., Herbst (1793: 138, n 79 tab 148 fig 3 4) = ^ serpa diadochus; Adelpha iphicla Linn., Hubner ([I8I9]: 42) refs. Cramer, 188, E. F = /f. serpa diadochus; Nymphalis iphicla

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266 Godt. [sic], Lucas (1835: 126, pi. 68) = A. iphiclus ephesa; Heterochroa basilea Cram., Bates (1865) = A. naxia; Heterochroa iphiclus Linn., Boisduval (1870) = A. naxia; Adelpha iphicia Linn., Kirby (1871) = A. naxia, A. iphicleola, A. basiloides; Adelpha basilea Cram., Godman and Salvin ( 1 884) = A. naxia Identification, taxonomy and variation: This subspecies is distinguished from A. i. ephesa by having the orange subapical marking on the DFW usually only extending into cell M2 as a "hook" along the outer postdiscal series, rather than extending to form an irregular pentagon with one side broadly bordering along vein M3. Some specimens of A. i. iphiclus have the orange of the marking extending into cell M2 along the length of the marking, but the posterior edge of the marking in cell M2 is curved towards vein M2 in A. i. iphiclus, and away from the vein in A. i. ephesa. Adelpha i. iphiclus typically has broader white postdiscal bands than A. i. ephesa. Adelpha i. estrecha is distinguished from A. i. iphiclus by the very narrow dorsal postdiscal band, tinged very pale blue, which has the spot in cell M2 wider than the rest of the band, in addition to the reduced orange subapical marking on the forewing. The forewings are also distinctly more pointed and falcate than the nominate subspecies. The characters that distinguish A. thessalia from A. iphiclus are discussed under that species. Adelpha iphicleola iphicleola is distinguished as discussed under the general identification section for A. iphiclus. Adelpha iphicleola thessalita may be distinguished from A. i. iphiclus by the same characters of the DFW orange subapical marking as A. i. ephesa, in addition to the slightly tapering white band on the DFW with a slightly concave distal edge, and by its overall smaller size and less rounded, but more distally dentate, hindwing. Adelpha i. iphiclus is very similar, if not sometimes indistinguishable from, A. iphicleola gortyna and A. iphicleola leucates. Adelpha iphicleola gortyna typically has the orange DFW subapical marking shaped as a rough trapezoid, with one sloping side along the costa, the other parallel to the distal margin, such that the orange area in cell M2 forms a posteriorly pointing triangle, the base of which is the same width as the orange area in cell Ml. ha .4. iphiclus iphiclus, the orange area in cell M2 forms a posteriorly pointing, attenuated triangle, the base of which is typically only half the width

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267 or less than that of the orange area in cell Ml. In ^. iphicleola gortyna the distal edges of the orange marking in cells M2-R5 are slightly concave, whereas they are straight in A. iphiclus iphiclus, while the white DFW postdiscal band in the former species does not taper anteriorly and has a slightly concave distal edge. The pale WW subapical marking in A. iphicleola gortyna is typically less obscured with red-brown scaling, and has more sharply defined basal and distal edges. Adelpha iphicleola leucates is distinguished by having the orange DFW subapical marking of similar shape to A. iphicleola gortyna, and has additional ventral characters typical of A. iphicleola, as discussed under that species. The nominate subspecies exhibits slight local and geographic variation, but none in my opinion that is sufficently pronounced or consistent to warrant separate subspecific recognition. However, quite possibly when more material becomes available from localities in areas such as northern Venezuela, there may prove to be consistently distinct populations worthy of subspecific recognition. Central American specimens tend to have slightly narrower postdiscal bands and a less reddish ventral surface, while specimens fi-om Guerrero in Mexico tend to have an orange DFW subapical marking similar to A. iphicleola gortyna (these specimens however are easily distinguished fi-om the sympatric A. iphicleola iphicleola, see under that subspecies). Specimens fi-om Colombia and Venezuela west of the easternmost Andean cordillera typically have a slightly narrower and paler orange DFW subapical marking, and thinner white postdiscal bands, while certain north Venezuelan specimens have broader white bands than usual and a narrower orange DFW subapical marking, more acute forewings, and the VHW postdiscal and submarginal series more straight and parallel (these specimens are marked f in the list of specimens examined). In northern Venezuela and Trinidad occur specimens that appear to be typical A. iphiclus, but intermediates are known to specimens which are much smaller, with wider white bands and a reduced orange DFW subapical marking. It is not clear whether these specimens represent a distinct subspecies of ^. iphiclus, or A. iphiceloa leucates, but I treat them as the latter in this work.

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268 The taxonomic history of this species, one of the most common in the entire genus, is one of the most confused. The difficulties stem fi-om Linnaeus' all too brief original description of Papilio iphiclus, which may be quoted in its entirety (words in square parentheses added by me): "P. [Papilio] B. [Barbarus] alis angulatis: supra fuscis fascia alba, maculaque ferruginea anguli ani. 3. Habitat in Indiis. Macula magna ferruginea intra apicem alae primoris. Fascia alba ab angulo ani ad medium alae primoris ducta". At first glance this would seem to apply to A. iphiclus as treated here (fi-om now on referred to as A. iphiclus), since Linnaeus mentions both a white band traversing the wings fi-om the anal angle (of the hindwing, presumably) to the middle of the forewing and a single, large red-brown spot in the forewing apex. However, Clerck's (1764) figure on plate 41 of the dorsal and ventral surfaces of Linnaeus' Papilio iphiclus unquestionably represents the species A. naxia, which differs most notably on the DFW fi-om A. iphiclus by having an additional orange subapical spot. Martin Honey (pers. comm.), who is completing a detailed examination of the Lepidoptera described by Linnaeus, is of the opinion that since Clerck supposedly figured the specimens on which Linnaeus based his descriptions, his figure should be taken to represent the type specimen. However, as Honey admits (pers. comm.), this view makes no allowance for subsqent usage and nomenclatural stability, the principal aim of the Code. Honey informs me that the type specimen of Papilio iphiclus is certainly lost, as Aurivillius (1882) stated, and that Butler's ([1870]) claim that the type of iphiclus is in the Linnaean collection in London is erroneous. The only specimen similar to A. iphiclus in the Linnaean collection was added at a later date and does not represent the type specimen, or even this species, in fact being the only known specimen of ^. thoasa calliphiclea. Early authors varied as to their treatment of Linnaeus' Papilio iphiclus, and this merely served to compound subsequent conftision. Drury (1770: 27) referenced both Linnaeus' description and Clerck's figure, but figured both surfaces of a specimen fi-om Surinam (pi. 14, fig. 3, 4) which is typical A. iphiclus. Despite the obvious differences between

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269 Drury's and Clerck's figures, both Cramer (1777) and Herbst (1793) referred to them as both representing Papilio iphiclus, but figured yet another unrelated species, A. serpa diadochus, under the name Papilio iphiclus (Cramer, 1777: 139, pi. 188, fig. E, F; Herbst, 1793: 138, pi. 148, fig. 3, 4). Herbst (1793: 137, pi. 148, fig. 1, 2) instead figured A. iphiclus imder the name Papilio cythereus Linnaeus, while Cramer (1777: 139), having mistaken A. serpa diadochus for Linnaeus' Papilio iphiclus, clearly figured the ventral surface (plate 188, fig. D) and redescribed A. iphiclus as Papilio basilea, based on a specimen or specimens fi-om Surinam. Cramer's types should be in either the RNHL or the BMNH (VaneWright, 1975), but they have not been located in either collection (Lamas, pers. comm.; pers. obs.). Hubner ([1819]: 42), in a hst of species in his new genus Adelpha, also referred to Cramer's (1777) figure on plate 188, fig. E, F (=A. serpa diadochus) as representing Linnaeus' iphiclus, and without explanation emended Cramer's Papilio basilea to Adelpha basilis. Other authors referred to the figures of Drury (true A. iphiclus) and Clerck (true A. naxia) as both representing Linnaeus' iphiclus (Godart, [1824]; Westwood, 1850), while Westwood (1850) went on to also Hst Cramer's basilea as a distinct species, though how he thought it differed fi-om Drury's figure is not clear. Authors following Westwood were divided in their treatment of the various names in circulation. Some used the name iphiclus in the sense that it is treated here (Hewitson, 1850; Bates, 1865; Felder and Felder, 1867; Butler, [1870]), while others used basilea to represent the same taxon (Menetries, 1857; Boisduval, 1870; Kirby, 1871) or, conftisingly, to represent A. naxia (Bates, 1865; Godman and Salvin, 1884). Boisduval (1870) and Kirby (1871) both used iphiclus as the name for^. naxia, the species figured by Clerck, while the latter also synonymised two other distinct species with iphiclus, A. basiloides and A. iphicleola. It is clear then that considerable confiision surrounded the names basilea and iphiclus, and only Boisduval (1870) seems to have used iphiclus to apply exclusively to the species figured by Clerck (1764) as Linnaeus' Papilio iphiclus. Aurivillius (1882) finally put a stop to the confiision by designating Drury's (1770) figure

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270 as representing typical A. iphiclus, and listed P. basilea Cramer as also representing that species. Godman and Salvin (1884) appear to have been the last authors to use, or even mention, the name basilea, which they mistakenly applied to A. naxia, and all subsequent authors (Fruhstorfer, 1915; Hall, 1938; D'Abrera, 1987; DeVries, 1987; Neild, 1996; many other more general works) have unanimously referred to the taxon treated here as A. iphiclus under that name. Given the historical confusion discussed above, and absence of type specimens of both Papilio iphiclus and Papilio basilea, I believe it is in the interests of future nomenclatural stability to designate neotypes for both of these names. Although I have not tried to trace every single reference to these two names in the literature, I have tried to examine those during the century following their description that are most widely cited and figure specimens, thus allowing me to ascertain how the two names were treated. Since only Boisduval (1870), among early authors, seems to have unambiguously treatedunder the name A. iphiclus the species figured by Clerck (1764), whereas a number of other authors regarded Drury's figure as representing the Linnaean type, and since the name has been used in the latter sense for at least the last century, I believe that the neotype designation should reflect the more widespread usage of the name. Furthermore, Linnaeus' original description, which mentions only a single red-brown spot in the apex of the DFW, actually corresponds more closely to specimens of A. iphiclus than A. naxia, as treated here. It is also possible that Linnaeus based his description on specimens belonging to both A. iphiclus and A. naxia. Drury's figured specimen of Papilio iphiclus Linnaeus came from Surinam, as did Cramer's figured specimen of Papilio basilea, and I therefore choose to designate the same specimen as neotype for both names. Unfortunately there are no specimens in the BMNH in good condition from Surinam, and I therefore designate a specimen from French Guiana, in the BMNH(M), drawer 16-101, which bears the following label data "Cayenne//Ex. Oberthiir Coll. Brit. Mus. 1927-3.", as the neotype for Papilio iphiclus and Papilio basilea. This specimen is figured (Fig.

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271 38a,b) and corresponds well with Drury's figure, being typical of the species, as currently treated, in all respects. Fruhstorfer (1915) described a new subspecies of A. iphiclus, funalis, from a single specimen fi-om Rio Dagua in western Colombia, which was at the time in the Tring Museum. The holotype male is in the BMNH(R), in addition to four fiirther specimens from the same locality. All of these differ fi-om typical A. i. iphiclus in having a slightly narrower white postdiscal band (Fruhstorfer' s claim that the white postdiscal band of both wings is reduced to half its normal width is an exaggeration) and narrower orange DFW subapical spots. Adelpha iphiclus exanima was described by Fruhstorfer (1915) based on an unspecified number of specimens from Venezuela. The syntype male in the BMNH(T) differs from typical A. i. iphiclus in having a narrower DFW orange subapical marking and slightly broader white postdiscal bands, as stated by Fruhstorfer. This phenotype is typical of specimens from northern and northwestern Venezuela, to which Neild (1996) tentatively applied the name, whilst pointing out the considerable difficulties in assessing variation given the inaccurate data and few specimens available from this biogeographically complex area. There is a second syntype in the BMNH(R) from Suapure, Venezuela, which may actually be a specimen of A. iphicleola leucates. Although it has broad white postdiscal bands and a narrow orange DFW subapical marking as in the first syntype, it originates from an area where one would expect typical A. iphiclus to occur. Since the first syntype was figured by Neild (1996) as A. iphiclus exanima, I designate that specimen as the lectotype of Adelpha iphicla exanima, with the following label data: "Illustrated in The Butterflies of Venezuela A. Neild, 1996//122//Fruhstorfer Coll. B.M. 1937-285//iphicla exanima Fruhst.Wenezuela Fruhstorfer//TYPE//Type//Syntype". Finally Fruhstorfer (1915) described A. iphicla pharae from specimens from Mato Grosso, Peru and Bolivia, as differing from the nominate subspecies in having a larger orange DFW subapical marking and broader white postdiscal bands. These differences are neglible in the several syntype specimens that I

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272 have located. I follow Willmott and Hall (1999) in considering all differences between funalis, exanima, pharae and nominate iphiclus to be too minor and subject to variation to sustain the recognition of any the first three names as valid subspecies, the second two of which were synonymised by Hall (1938), whilst recognising that more extensive collecting in future may well necessitate a review of this taxonomy. To this end, I have added footnotes by certain specimens in the list below that exhibit notable phenotypic differences from typical A. i. iphiclus. Range: This very widespread taxon ranges from Mexico to the western slopes of the Cordillera Occidental in Colombia, to northwestern Venezuela, south to Bolivia, Amazonian Brazil as far south as the central Brazilian plateau in the state of Goias, Paraguay, northeastern Argentina, the Guianas and Trinidad. Immature stages: All of the early stage information known to date, except that of Moss (1933) for which I have identified voucher specimens, could apply to either this species or A. iphicleola, but I summarise it here for convenience and since the life histories of these two species are probably similar. Moss (1933) reared this species in Brazil (Para), and figured both the last instar larva (pi. I, fig. 18) and the pupa (pi. II, fig. 10). Aiello (1984) also reared this species in Panama, and figures the final instar larva and its head capsule (figs. 2, 4) and the pupa (fig. 6). The egg is typical of the genus and may be deposited on the upperside, or, more rarely, the underside of the leaf, typically at the leaf tip but also at its margin. The color of the larvae varies from dark gray, to golden brown, to ahnost black, and is not dependent on the hostplant. The head capsule has a relatively smooth face and long lateral spines, and also varies, from yellow brown with black pits to uniform dark brown or black. Light individuals of the final instar have drak, oblique lateral stripes on A2-A5, and often white lateral marks on A2, A7 and A8. The A2 scolus has a broad base, and all the scoli are all relatively long and thin, with thin, tapering lateral spines, except for that on Al, which is reduced with no lateral spines. The pupa has a moderate projection on A2 and small, triangular head horns. It varies from waxy

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273 white to straw-colored, and is often partly or entirely shining gold or silver, expecially on the dorsal surface of Tl, T3 and Al. Almost always there is a small silver diamond at the base of each mesothoracic leg. There are preserved head capsules and pupal cases in the USNM, collected by Small in Panama. Janzen {In: DeVries, 1986: Costa Rica) reports that late instars are solitary and feed on all leaves of plants along forest or riparian edges or in light gaps. DeVries (1987) states that first to third instars make firass chains, and later instars either rest on these or on damaged portions of the leaves. A single larva observed by Aiello (1984) constructed an additional, curved, larva-form mass of fi-ass on the upper surface of the leaf, as do all A. basiloides. Full development times are given for all stages, and the total development time for three individuals was 29 days (Aiello, 1984: Table 2). Aiello and Orellana (pers. comm.) report that in Panama in August, 1998, there was what appeared to be an outbreak of larvae of A. iphiclus or A. iphicleola. Larvae were abundant on the leaves of Antirrhoea, a roadside weed with broad leaves. Observed mortality agents included an unidentified bacteria/virus, parasitic Hymenoptera and predatory Hemiptera. Habitat and adult ecology: This subspecies is one of the most fi-equently encoimtered Adelpha in South America, although it is less common in Central America than its close relative, A. iphicleola. It occurs in both secondary and primary forest, fi-om sea level to 1250m, throughout the year, although adults are typically most abundant in the wet season. Males may be attracted to rotting fiiiit and excrement (Neild, 1996), and to rotting fish, and are fi-equently found puddling along wide sandy roads through secondary forest and at streams. According to specimen label data, a series of seven males and a female fi-om Costa Rica in the AMNH were collected in June feeding on flowers of Tridex procumbens L., Stachytarpheta jamaicense (L.) and Melanthera sp.. Orellana (pers. comm.) reports that he found this species or A. iphicleola feeding at ripe and fermenting fiaiits hanging on a Bunchosia cornifolia (Malphigiaceae) tree in Panama in August, while

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274 Aiello (pers. comm.) also rqjorts adults to be attracted to fruits of Antirrhoea trichantha, the principal local larval hostplant. DeVries' (1987) observations in Costa Rica are of interest, but they could also apply to A. iphicleola, which he did not recognise as a distinct species. He reports that males perch along 'forest edges in groups, a behavior that I have only observed in A. iphicleola in eastern Ecuador, while both sexes visit fruits of mango, Guazuma and Genipa. DeVries also observed individuals visiting flowers of Vouchysia and Paullinia in the forest canopy. Pinheiro (1996) found this species to be palatable to kingbirds in Brazil. Specimens examined (868m, 125f): t specimens like lectotype exanima (see Neild, 1996, pi. 3, fig. 122), wide postdiscal bands, thin orange DFW subapical marking, acute forewing shape; % DFW orange subapical marking like A. i. iphicleola, ventral surface like A. iphiclus; * trans, to A. i. ephesa; ? identification uncertain, wide postdiscal bands, DFW subapical marking typical A. iphiclus, wing shape like A. iphicleola gortyna. MEXICO (45m, 14f): Chiapas: Cuauhtemoc Oct. Im AME; Las Delicias Sep. Im AME; Musti Jul. 9m AMNH; Ocozingo Im, If AME; San Carlos Jun. If AME; San Jeronimo 450m Aug. Im FSCA; San Quintin Aug. 3m AME; Guerrero: Acahuizotla Jun. Im AME; no specific locality 17m, 2f BMNH; Oaxaca: Chiltepec Aug. Im AME; San Luis Polosi: Tamazunchale Jul. Im FSCA; Xilitla Nov. Im AMNH; Tamaulipas: Mante, 30 mi. N., Nov. Im USNM; Veracruz: Cordoba Jul. If AMNH; Cuesta de Misantla 4f BMNH; Motzorongo Jun. Im USNM; Orizaba Jun. Im AMNH, Im AME; Presidio Aug. Im, 3f AME; Tezonapa If AME; Zongolica Jun. Im BMNH; Not healed: Isthmo de Tehuantepec Im MCZ; no specific locality If BMNH, Im AMNH. BELIZE (2f): Cayo: Camp Sibun 200m Jul. 2f AME. GUATEMALA (11m, 30: Alta Verapaz: Chocnin Im BMNH; Polochic Valley 2m BMNH; Izabal: Cayuga Mar. Im USNM; Quezaltenango: Volcan Santa Maria 4m, If BMNH; Santa Rosa: Guazacapan Im AME; Not located: W. Guatemala Aug. Im MCZ; no specific locality Im, If BMNH, If MNHN. HONDURAS (5m, 20: Comayagua: Comayagua Jun. If MCZ; Cortes: San Pedro Sula Im, If BMNH; Not located: Central Honduras Im AMNH; no specific locality 3m USNM. NICARAGUA (3m, 40: Managua: Managua Oct. If AME; Matagalpa: Matagalpa If BMNH; Chonlales: Chontales If BMNHZelaya: El Bluff Jan. Im BMNH; Not located: no specific locality 2m BMNH, If MCZ. COSTA RICA (24m, 120: Alajuela: San Mateo Jul. Im BMNH; Cartago: Cachi Jul. 3m BMNH; Irazii Im BMNH; Juan Viilas Jun. Im BMNH; Turrialba Jul. 2m, 2f AMEGuanacaste: Cafias, 24 km. N.W., Jun. 7m, If AMNH; Finca Taboja Jun. If AME; Santa Rosa 100m Aug. Im KWJH; Heredia Puerto Viejo, 3 km. S.W., May Oct. Im, 2f USNM; Limon: Port Limon Jan. 2m, If USNM; Santa Clara Valley 1200' If BMNHPuntarenas: Palmar Norte Jul. Im USNM; Rio Tarcoles 50m Sep. Im KWJH; San Jose: San Isidro If MCZ; San Jose If BMNH, Im MNHN; Villa Colon 750m Aug. Im USNM; Not located: no specific locality 2f AMNH, Im BMNH. PANAMA (26m 270' Canal Zone: Barro Colorado Island Feb. Sep. Dec. 2m, 2f AMNH, 2m, If MCZ; Farfan Jun. Dec. 3m(It), 5f(3t) USNM, Im AMNHFort Kobbe Jun. 2m, 3f USNM; Gamboa Jan. 2f USNM; La Pita Jun. Im USNM; no specific locality Mar. If BMB; Chiriquv Bugab'a Im BMNH; Chinqui 2m BMNH, Im, If MCZ, Imt USNM; Code: El Valle If AMNH; Colon: Pifia lOOm Apr. If FSCADarien Cana 900m Jul. Aug. 3m(It), 2f USNM; Los Santos: Rio Oria Jan. 2ft USNM; Pfl/iamo: Arraijan Feb. Sep. 2f AMNHCocoli Nov Im USNM; El Valle 800m Aug. If USNM; Rio Bayano Nov. Im USNM; Rio Trinidad Im, If AMNH; Not located: Las Cumbres Oct Im FSCA; no specific locality Im, If MCZ, Imt USNM, Im, If AMNH. VENEZUELA (35m, 10: AmazonasMt Duida Im AMNH; Puerto Ayacucho lOOm Im AFEN; Rio Mavaca Mar. Im AMNH; Aragua: San Estevan, pres Puerto Cabello 7/1877 Imt BMNH; Bolivar. Caura 3m AFEN; El Dorado Jun. 5m FSCA; Javillal 1 00m Aug. Im AFEN; Suapure Im BMNHCarabobo Us Quiguas Nov.-Mar. Iff BMNH; Distrito Federal: Caracas 3000' Sep. Oct. Im BMB; Lara: Quebrada Negra Yacambu 800m Im' AFEN; Sarare 4m MNHN; Merida: Merida Im BMNH; Sucre: Patao, Guiria Aug. Im BMNH; Tachira: Hda. Panaga 800m Imt AFEN; La Monta 300m Aug. Im MUSM; Trujillo: U Gira Betijoque 550m May Im? (postdiscal band greenish), Dec. Imt AFENZulia: El Tucuco Jun. Im USNM, Imt AFEN; Not located: Alto Orinoco 3m USNM; Morrisca Sep. Im MCZ; Palo Alto Rio Tigre Oct. Im USNM-error; Smistara del Sur Im AMNH; no specific locality 2m BMNH, 3m MNHN COLOMBIA (328m 240' kT'^T^.'^T'^^ ^"^ LMC; Aniioquia: Casabe, R.o Magdalena 13m, 2f AMNHLa Mesa 4000 Jul. 2m BMB; Medellin 3m AME, If USNM; Rio Corcoma Im AMNH; Valdivia Dec. Im AMNH; Boyaca: Minas de ^x"/xm"I. '^"^ 2'"' "^N™Nov. 2m JFL; Caldas: Guamoco Im AMNH; Itaburi Jan 2m AMNH; Manizales 2m BMNH; Rio de la Miel Jun. Im ESM; Victoria Jan. Feb. 2m USNM; Caqueta Montaflita Jan Im USNM Rio w^^^fl"^™ " ^"^^ '"^ ^^S^' Cundinamarca: Bogota 90m, 3f BMNH, I2m BMB 3m MCZ 5m DNXMu' ]^ If BMNH; Cananche Jun. 3m BMNH; c'anire Nov' Im ^^^'^ Im USNM; U Vega 1900m Jan. Im BMNH; head Rio Carare 2m USNM; Cauca: Pescador 1450m Feb 5m If M i^' r^'^".''"'^,'/^^'^ "^""^ ^^""^ •^^N"' ^' S*"*^ ^^"^ 950m Jul. Im BMNH; Onaca Santa RMNH p ?M^""'^ ^'"^ ^"^ ^"1 AMNH; Peperital-Buenavista Jan. Im BMNH, Remolmos Mar. If JFL; Restrepo Jul. Im JFL; Rio Ariari Aug. Im LMC; Rio Negro Jan. Im, If USNM; San Jos^ Guaviare Im JFL; Toquiza Apr May Im BMNH; Villavicencio Feb. Jul. Aug. 3m AMNH. If USNM, Im BMNH, Im, If AME Im MCZV,llav,cenc,a.Mt Redondo Mar. Apr. Im BMNH; Putumayo: Umbria Im AMNH; Santander: Bznzi^c^h^i^. 2m AMNH li Borroscosa Dec. Im, If AMNH; Undaruri Jul. 2f AMNH; La Sevilla Dec. Im AMNH; R,o Qu.rata Dec fTSlNH^ mUna'l^o 5m BMNH-"].^/'TwT^ 'r'l ^ ^^'^^ "'^^^^ ^"""^^ ^auca "m BMnA; ^o Da^ 5m BMNH, Not located: Bogota-Buenaventura Dec.-Feb. Im BMNH; Cauca Im BMNH, Im AMNH; Rio Magdalena Im BMNH,

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275 Im MNHN; San Rafael 3500' Jun. Jul. 2m BMB; no specific locality 52m, 2f AMNH, If BMNH, Im AME, 2m BMB, 3m MNHN, 2m MCZ, 6m USNM. ECUADOR (55m, 20: Morona-Santiago: Bomboiza Nov. 2m DAT; Macas Oct. 3m BMNH, 2m MNHN; Mendez Oct. Im MJP; Napo. Apuya 600m Oct. Im KWJH, Im DAT; Archidona Apr. 3m BMNH; Chichicorrumi 450m Sep. Im KWJH; Coca, 35 km. S.W., Apr. 2m MJP; Cotundo Oct. Im USNM; Finca San Carlo 600m Sep. Im KWJH; Guacamayos Sep. Im DAT; Us Minas de Misahualli 400m Jul. Im KWJH; Napo 2m AME, 7m MNHN; Nushifio Im BMNH; Rio Chalayacu Sep. Im DAT; Rio Coca Jun. 3m AME; Rio Napo Aug. Sep. 2m BMNH; Rio Sliandia 550m Sep. Im KWJH; Rio Tiputini Apr. Im MJP; Santa Rosa Nov. Im DAT; Satzayacu Sep. Oct. 2m DAT; Pastaza. Sarayacu Im BMNH, Im USNM; Pichincha: Quito Im MNHN; Sucumbios: Garzacocha, La Selva, Rio Napo 250m 2m, 2flan. Jul.-Nov. PJD; Rio Aguarico Im MNHN; Tungurahua: Rio Topo Sep. Im USNM; Not located: E. Ecuador Im USNM, 2m KWJH; no specific locality 4m MCZ. PERU (183m, 6f); Amazonas. Falso Paquisha 800m Oct. Im MUSM; Rio Santiago Oct. 4m AMNH; Cuzco: Marcapata Im BMNH; Rio Urubamba Feb. Im MUSM; Huanuco: Cord, del Sira Aug. Sep. 1050m Im, If MUSM; Las Palmas 450m Aug. Im AME; Pozuzo 800-lOOOm 3m BMNH; Tingo Maria May Jul. Dec. Im FSCA, 13m AME; Toumavista 2m AME; Junin: Chanchamayo Im BMNH, Im AME; La Merced 2500' Aug. 2m BMNH; Rio Perene Jun. Im AMNH, Im BMNH, Im BMB, Im MNHN; San Ramon Im MUSM; Satipo Jan. Oct. Nov. 5m AME, Im AMNH; Satipo, 25 km. N., 800m May Im MUSM; Loreto: Arcadia 150m Nov. 3m MUSM; Caballo Cocha May-Jul. 3m BMNH; Castaila 150m Oct. Im MUSM; Curaray 3m MNHN; Guepi May Im MUSM; Iquitos Feb. Jul. Aug. Oct. 7m AMNH, 6m BMNH, Im FSCA, 3m AME; lower Rio Tapiche Jul. Aug. 11m AMNH; Pebas 5m BMNH; Rio Cachiyacu 4m BMNH; Rio Morona Jan. Im AMNH; Rio Sucusari 140m Sep. Um, If MUSM; upper Rio Tapiche Aug. 2m AMNH; Yanamono Jul. Sep. 2m MUSM; Madre de Dios: Boca Rio La Torre 300m Feb. Jul. Oct. Nov. 9m, If MUSM; Pakitza 400m Sep.-Nov. 10m MUSM; Pampas del Heath 220m Jun. Im MUSM; Puerto Maldonado 250m Aug. Oct. Im, If USNM, 2m AME; Puerto Maldonado, 30 km. S., Oct. Im, If USNM; Rio de Las Piedras Aug. Im MUSM; Shintuya 460m Jul. Im AME; Pasco: Alto Yurinaqui 1400m Nov. Im MUSM; Puno: Chaquimayo 2500-3000' Jun.-Aug. Oct. 2m BMNH; U Union 2000' Jan. Im BMNH; San Gaban 2500' 2m BMNH; San Martin: Achinamiza Jun. Sep.-Dec. 6m AMNH; Jepelacio Im AMNH; Rioja Im MUSM; Tarapoto Im BMNH; Yumbatos Sep. If MUSM; Ucayali: Boqueron Abad 600m Aug. Sep. 2m MUSM, Im AME; Contanamo Dec. Im BMNH; Not located: Rio Huallaga 7m AMNH, 2m USNM; upper Rio Marafidn Nov. Dec. 5m AMNH; middle Rio Ucayali Apr. 3m AMNH; Rio Ucayali 2m BMNH; no specific locality 4m BMNH, 2m MCZ, 2m USNM, Im AMNH, Im JFL. BOLIVIA (42m, 20: Cochabamba: Cochabamba Im BMNH; El Palmar 1600m Apr. Im AME; El Beni: Cachuela Esperanza Sep. 5m AME; Reyes Jun. Im BMNH; La Paz: Chimato R. Apr. 2m AMNH; Coroico Mar. 5m AMNH, If MCZ; Potosi: Tupiza Im AME; Santa Cruz: Buenavista Mar. Apr. 2m AME; forest between Santa Cruz and Tapirapuan plains Sep. Im BMNH; Prov. Sara Mar. -Jun. 7m BMNH; Rio Negro Aug. Im AME; Santa Cruz botanical gdns. Dec. Im AMNH; Santa Cruz de la Sierra Jul. Im FSCA; Not located: Yungas Im BMNH; no specific locality 2m AMNH, If MCZ. BRAZIL (70m, 100: Amazonas: Benjamin Constant Oct. 4m AMNH; Caiary-Vaupes Nov. 5m AMNH; Ega Jan. Apr. Jul. Aug. 4m BMNH; Fonte Boa Jul. Im BMNH; Humaita Im BMNH; Labrea, Rio Purus Nov. Im BMNH; lower Rio Madeira Jul.-Sep. Im BMNH; Madeira If AMNH; San Antonio do If a If MCZ; San Joas, Solimdes Im BMNH; Sao Paulo de Olivenfa Aug. 3m BMNH; Tefe Oct. Dec. 2m BMNH, Im AMNH; Golds: Guyabera Nov. Im BMNH; Mato Grosso: Buriti May 4m USNM; Campo Grande Nov. Im* AMNH; Chapada Im BMNH, Im USNM; Cuiaba Im* BMNH; lower Rio Arinos Jun. Im BMNH; Nivac Im* BMNH; no specific locality 4m BMNH; Minas Gerais: Belo Horizonte-Brasilia, km. 500, Im AME; Leitao May Im USNM; Para: Obidos 4m, If AMNH, Im, If BMNH; Para 3m, 3f BMNH; Santarem Im, If BMNH; Tapajos If BMNH; Rondonia: Cacaulandia Jan. Mar. Apr. Oct. Nov. 3m USNM, Im FSCA, 10m AME, Im KWJH; Jaru Aug. Im AME; Not located: Brasilia Im MCZ; no specific locality 2m BMNH, Im MNHN, If AMNH. PARAGUAY (3m): Paraguari: Sapucay, W. Foster 3I/X/04 3m(l*) BMNH. GUYANA (2m, 30: Barima/Waini: Mabaruma Dec. Jan. If BMB; Cuyuni/Mazaruni: Kartabo If AMNH; Upper Takutu/Upper Essequibo: Kuyuwini R. Im AMNH; Not located: Demerara If BMNH; no specific locality Im BMNH. FRENCH GUIANA (6m, 30: Cayenne: Cayenne 4m, 2f BMNH; Not located: no specific locality 2m, If BMNH. SURINAM (40: Brokopondo: Berg-en-Dal Apr. If BMNH; Saramacca River May If BMNH; Not located: no specific locality 2f BMNH. TRINIDAD (9m, 20: Caroni: Tabaquite Oct.-Dec. Im BMB; St. George East: Caura Aug. 2m BMNH; St. George West: Macqueripe Bay If AMNH; St. Patrick: Siparia Oct.-Dec. Im BMB; Not located: Cochrane Sep. Im? FSCA; no specific locality Oct. -Dec. If BMNH, Im BMB, 3m MNHN. COUNTRY UNKNOWN (23m, 40: Amazons Im, If USNM; Mexico? If BMNH; Peru/Bolivia frontier Im MNHN; no specific locality 1 3m MCZ, Im, If BMNH, 3m AME, 4m MNHN, If MCZ. Additional locality data: MEXICO: Chiapas: throughout see de la Maza and de la Maza (1993); Colima: some records in Warren et al. (1998: as A. iphiclus massilides) may apply to this taxon. BELIZE: Cayo; Orange Walk; Toledo (Meerman, 1999). ECUADOR: Morona-Santiago: Bomboiza Oct.; Mendez-Santiago rd., km. 40, Oct.; Rio Shangaime Dec; Santiago Sep.; Napo: Apuya Aug.-Dec; Chichicorrumi Jul. Dec; nr. Coca Oct.; Coca-Loreto rd., km. 21, Mar.; Finca San Carlo Aug. Dec; Pimpila'la Sep.; Rio Tiputini Mar. Jun. Sep.; Rio Yuturi Oct.; Satzayacu Apr.; Yarina Jul.; Yasuni Jul.; Pastaza: Puyo-Canelos rd., km. 30, Oct.; Rio Llandia Dec; Sucumbios: Laguna de Pailacocha Oct.; Paflacocha Oct.; Zamora-Chinchipe: Quebrada Chorillos 1250m Nov. (Willmott and Hall, unpubl. sight records). BRAZIL: Distrito Federal: Parque do Gama 1 100m Jun.; RibeirSo da Contagem 850m Feb.; Sobradinho Woods 1050-1 150m Feb.; Golds: Anapolis 1000m Feb.; Campinas 800m Jan. Dec; Chapada dos Veadeiros 1000m; Leopoldo BulhSes 1000m Mar.; Rio MaranhSo 700m; Vianopolis lOOOm Nov. (Brown and Mielke, 1967). PARAGUAY: Alto Paraguay; Concepcion; Amambay; Paraguari (Kochalka et al., 1 996). Adelpha iphiclus estrecha Willmott and Hall, 1999 Figs. 38e,f; 122a,b Adelpha iphiclus estrecha Willmott and Hall (1999: in press) TL: Ecuador. Types (all Ecuador): BMNH(M): HTm: "Ecuador. Hewitson Coll. 79-69 Heterochroa 1//Ecua "• ATf "Ecuador Hewitson Coll. 79-69 Heterochroa 2//Ecua."; BMNH(R): PTm, PTf: "Quevedo, W. Ecuador. (V. Buchwald)"BMBPTm"St Ana Mana. Quevedo. Ecuador. 1. von Buchwald"; KWJH: PTm: Lita, ridge east of Rio Baboso; PTf: EI DurangoPTm PTf San Lorenzo-Lita rd., km. 18; PTm: San Lorenzo-Lita rd., km. 20; PTm: La Punta, km. 44 Lita-San Loranzo rd.; 6PTmRio Tanti-

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276 USNM: "Alluriquin, 700m, Pichincha, Ecuador 28 Aug. 77 S.S. Nicolay"; AMNH: PTm: "Ecuador/ZColl. Frank Johnson"; PTm: "Oriente Ecuador/ZColl. Frank Johnson". Adelpha iphicla funalis Fruhst., Strand (1918) Identification, taxonomy and variation: This subspecies is distinguished from the nominate by the much thinner postdiscal band that has a pale bluish tint and narrows anteriorly on the DFW, with the spot in cell M3 being the widest, in addition to the more falcate forewing and reduced orange DF subapical marking. There is slight variation in the width of the postdiscal band, but specimens with the broadest bands are still clearly distinct from A. iphiclus iphiclus. A single specimen of this subspecies in the BMB has an Arthur Hall manuscript name, A. tenuivittata, attached to it, but this name was never described. Strand (1918) referred to it as A. iphicla funalis, a synonym of the nominate subspecies, probably based on Fruhstorfer's exaggerated original description of funalis. This taxon is regarded as conspecific with A. iphiclus, rather than A. iphicleola which the wing shape resembles, because of west Colombian specimens of A. iphiclus which have slightly more falcate wings and thinner postdiscal bands than typical A. iphiclus, the shape of the orange DFW subapical marking, and the apparent presence in western Ecuador of A. iphicleola gortyna or a new subspecies of A. iphicleola (see Willmott and Hall, 1999). Range: West of the Andes in Ecuador, from the extreme north to Quevedo area. Although this subspecies has not been recorded to date from southwestern Colombia, it probably occurs there. Habitat and adult ecology: This subspecies is not uncommon in the wet season and early dry season from May to August, in association with disturbed primary and secondary forest, from sea level to 1000m. Both sexes may be found feeding at flowering bushes along forest edges, while males are attracted to rotting fish in ridge top forest light gaps. I have found males perching in ridge top forest light gaps 3m above the ground, and females flying typically along forest edges, or even roadsides. The parallel development of a thinner discal band and reduced orange DFW subapical marking in a number of

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277 western Ecuador Adelpha species {A. serpa duiliae, A. basiloides, A. bamesia, A. iphicleola, A. erotia f. lema) suggests there may be a mimetic relationship between these species. Specimens examined (16tn, 4f): ECUADOR (16m, 4f): Carchi: Lita, ridge east of Rio Baboso 1050m Jul. Im KW iH; Esmeraldas: El Durango 400m Jul. If KWJH; San Lorenzo-Lita rd., km. 18, 100m Jul. Im, If KWJH; San Lorenzo-Lita rd., km. 20, 100m Aug. Im KWJH; La Punta, km. 44 LitaSan Loranzo rd. 300m Aug. Im KWJH; Los Rios: Quevedo Im, If BMNH(R); St. Ana Maria, Quevedo Im BMB; Pichincha: Alluriqurn 700m Aug. Im USNM; RioTanti 750m May Jul.-Sep. 6m KWJH; TVo/ /oca/erf: no specific locality Im, If BMNH(M), Im AMNH; "Oriente" Im AMNH. Additional locality data: ECUADOR: Pichincha: Rio Tanti Jun. (Willmott and Hall, unpubl. sight records).; Chimborazo: Los Llanos 1400m (Strand, 1918). Adelpha iphiclus ephesa (Menetries, 1857) Figs. 38g,h Heterochroa ephesa Menetries (1857: 104, pi. IX, fig. 2) TL: Brazil. Types: ZISP: LTm(?): "Bras./ZLectotypus Heterochroa ephesa M6nitrids, Lectotypus m 1875 design. A. Lvovsky" [photograph examined]; PLTm: "Bras." [not examined]; 2PLT: [not located] =Adelpha iphicla gellia Fruhstorfer (1915: 523) TL: Sa. Catharina, to S3o Paulo [Brazil]. Types: BMNH(T): STm: "Fruhstorfer Coll. B.M. 1937-285//iphicla gellia Fruhst.//Brasilien Blumenau Fruhstorfer//TYPE//Type//Syntype"; STf: "Fruhstorfer Coll. B.M. 1937-285//iphicla gellia Fruhst./ZBrasilien Blumenau Fruhstorfer//TYPE//Type" [both examined] =Adelpha abyla abylina Fruhstorfer (1915: 524) TL: Guyane; Bresil. Types: MNHN?: STf?: [not located] Nymphalis iphicla Godt. [sic], Lucas (1835: 126, pi. 68); Adelpha basilea var. ephesa M6n., Kirby (1871); Adelpha iphicla ephesa Men., Fruhstorfer (1915) Identification, taxonomy and variation: This subspecies is distinguished fi-om the nominate by having a larger orange DFW subapical marking, that typically broadly borders along vein M3 or is at least represented in cell M2 as a broad, triangle, instead of a small, attenuated point at the distal edge of the marking. The white postdiscal band is typically narrower. Adelpha iphicleola leucates is very similar, but it tends to have more pointed forewings, a straighter distal margin on the hindwing, noticeably broader postdiscal bands which are typically of equal width to the orange DFW subapical marking and broadest in cell M3, instead of tapering, the distal edge of the orange DFW subapical marking in cell R5 is typically concave rather than straight, the ventral surface has more heavily marked pale postdiscal and submarginal series and so appears lighter in color, the pale VPW subapical marking is less obscured by brown scaling along the veins and the basal and distal edges

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278 are more clearly defined. The extent of the orange DFW subapical marking is variable in A. i. ephesa, either extending to border vein M3, or just to touch it, while in some specimens orange continues as dashes down the outer postdiscal series in cells Cul and M3, a diagnostic character not found in any A. iphicleola examined, occurring particularly in specimens from Sao Paulo and Santa Catharina. The width of the white postdiscal band varies slightly, generally being thinner in specimens from Sao Paulo and Santa Catharina. Menetries (1855: 33, n. 575) first introduced the name ephesa as a nomen nudum in a catalogue of specimens in the collection of the ZISP, then described and figured it two years later (Menetries, 1857) based on four specimens collected in Brazil. The description compares the subspecies with Cramer's figure of Papilio basilea {=A. iphidus), but fails to note the only constant difference between this and the nominate subspecies, namely the larger orange DFW subapical marking, hi fact, Menetries claims that this marking is smaller. The figure shows both surfaces of a specimen which has characters typical of this subspecies that distinguish it fi-om the similar, possibly sympatric A. iphicleola leucates, including a relatively thin white postdiscal band and an orange DFW subapical marking which extends to the midpoint of cell M2 in its basal half, and to vein M3 in its distal half I have also examined a photograph of a syntype specimen, courtesy of A. Lvovsky, which differs fi-om the original illustration in having a slightly broader orange DFW subapical marking extending to vein M3 along most of its length, a character found in many specimens of this taxon. Since it is just possible that specimens of ^. iphiceloa leucates may be included within the type series, I designate this syntype specimen as the lectotype of Heterochroa ephesa, which bears the following label data: "Bras./ZLectotypus Heterochroa ephesa Menetries, Lectotypus m 1875 [sic] design. A. Lvovsky". According to Lvovsky (pers. comm.), the specimen was placed above the following label in the drawer: "Ephesa nob. Brasil", in addition to a second syntype male specimen, bearing simply the label "Bras.". Fruhstorfer (1915) described A. iphicla gellia fi-om an unspecified number of specimens fi-om Sao Paulo to Santa

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279 Catharina, supposedly differing from ephesa in having a darker ventral surface and narrower postdiscal bands. Both of these differences are apparent in the two syntypes examined, but since they are minor and variable I agree with Hall (1938), who synonymised gellia with ephesa. There are other potential syntypes lacking both handwritten identification and type labels from Fruhstorfer's collection in the BMNH (Blumenau, Im, If; Sao Paulo, Im; Espirito Santo, If), all of which are typical A. i. ephesa. Lucas (1835) figured the dorsal surface of an Adelpha on plate 68, as Nymphalis iphicla, which he claimed was from French Guiana and Brazil. The figure shows a specimen very similar to the syntype female of A. iphiclus gellia, with an enlarged orange DFW subapical marking extending as dashes along the outer postdiscal series into cell Cul. Fruhstorfer (1915) however, without giving ay explicit reason, regarded the figure as representing a new subspecies of A. abyla, which he named abylina, and this name was duely and correctly synonymised by Hall (1938) with ephesa. The type of abylina should be in the MNHN (Horn and Kahle, 1935-37), but I have not been able to locate it. Although this taxon has an orange DFW subapical marking typical of subspecies of A. iphicleola, it otherwise has characters typical of A. iphiclus, such as the rounded wing shape and the pale VFW subapical marking which is more obscured by red-brown scaling and less sharply defined. Furthermore, there are rare specimens of A. i. iphiclus (e.g. one male, Peru, Boqueron Abad in AME) which have the orange DFW subapical marking extending as dashes along the outer postdiscal series into cell Cul, in addition to similar specimens from probable blend zones, such as Paraguay and Mato Grosso. These strongly suggest that ephesa should be regarded as a subspecies of ^. iphiclus. Range: Eastern to southeastern Brazil, to Paraguay. Hayward (1931) figures a sketch of "A. iphicla indefecta" from Alto Parana, and later (Hayward, 1973) reported the taxon from Misiones, Formosa, Argentina; the sketch (Hayward, 1931) shows the typically narrow DFW postdiscal band of A. iphiclus ephesa, compared with A. iphicleola leucates, and I tentatively identify it as the former.

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280 Hostplant: Miiller (1886) reported a species, "/I. iphicla", from Santa Catharina, Brazil, to feed on Bathysa nr. barbinervis; this record might also apply to A. iphicleola leucates, but since A. iphiclus ephesa is clearly much more common in Santa Catharina, it most probably applies to the latter species. Miiller (1886) also reported "/4. basilea" to feed on Calycophyllum (Rubiaceae). Since basilea is a synonym of iphiclus, it is not possible to say to which species Miiller was referring, but if his "iphicla" did not refer to this taxon, then it may be that his "basilea" did. Immature stages: According to MuUer (1886), the egg is deposited on the underside of the leaf at the tip. The larva is in general form similar to A. lycorias lycorias, with a dark transverse band on segment A2 as well as on segments A3-7, a tapering lateral stripe from T1-A2 and dark coloring on the dorsal surface in segment A7. The pupa has indistinct dorsal projections in the position of the scoli on segments T2, with more pronounced projections on A2-A4. D'Almeida's (1933) description of the early stages of "A. iphicla ephesa" probably also apply to this species. He states that the egg is pale yellow and deposited singly on the upper surface of the leaf near the tip. The first instar turns from whitish yellow to gray, with a pale brown head, the second instar is brown with small yellow "warts", and scoli begin to develop as spiny tubercles. The fourth instar is black on the first 4-5 segments and brown on remaining segments, with oblique black stripes and yellowish "warts". The scoli and head capsule are black, the latter with dark yellowish chalazae and spines. The fifth instar is black with yellowish "warts" and scoli of similar or more greenish color, darker towards the base, with those on T2 and T3 more reddish yellow. The head capusle is black with four small chalazae and numerous small pits, ringed with two rows of short yellowish spines. Later the ground color of the larva becomes yellowish brown turning partly reddish, with lateral oblique black markings, which towards pupation becomes a more or less reddish brown with black markings, or green markings on the four posterior segments. Dorsally on the there is a reddish pattern consisting of several indistinct transverse rays extending from T2 to A2. The scoli are

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281 best developed on T2 and T3, with those on A7 and A8 larger than other abdominal scoli, and they are greenish at the base with reddish brown lateral spines. The scoli on AlO are short, broad and black. The supraspiracular and subspiracular scoli are also present but very short, with the latter whitish yellow in color. The pupa has dorsal projections at A2 and T2, the former rounded and the latter pointed, and small conical head horns. It is dark brown marbled with pale brown, paler on the wing pads and ventral surface, with dorsal gold markings and a silver streak ventrally at the base of the antennae. Segments T2, T3 and A2 have small conical lateral projections. The larvae retained the leaf mid-rib where they rested. Development times were as follows: 1'' instar, 3 days; instar, 4 days; 3"* instar, 4 days; 4'*' instar, 5 days; S"' instar, 9 and 7 days; pupa, 8 days (2 individuals). Habitat and adult ecology: Brown (1992) reports that this subspecies is common in disturbed forest areas, particularly at forest and stream margins, in the Serra do Japi, Sao Paulo. It is common in a wide variety of habitats, from humid, undisturbed forest along streams to open fields and gardens, and can also be found hilltopping. D'Almeida (1933) reported the species to be common in hilly areas around Rio de Janeiro, and observed that the female held the wings closed while ovipositing. The subspecies has been recorded up to 1000m (Brown, 1992) and appears to fly throughout the year. Both author's comments may also apply in part to A. iphicleola leucates. Specimens examined (62m, 38f): t trans, to A. i. iphiclus. BRAZIL (60m, 37f): Bahia: Cachimbo Im BMNH; Espirito Santo: Leopoldina If BMNH; Linhares May-Sep. 6m, 16f AME; Baixo Guandu Dec. Im AME; Santa Cruz Feb. 1 f AME; no specific locality If BMNH; Guanabara. Corcovado Im BMNH; P N da Tijuca Feb. If USNM; Rio Janeiro Apr. May Jul. 2m, If AMNH, Im, 3f BMNH, 4m MNHN, Im, If MCZ, Im USNM; Goids: Goyaz 5m BMNH; Serra Dourada Im AME; Maranhao: Montes Aureos Im BMNH; Mato Grosso: Cuiaba-Corumba river system Imf BMNH; Villa Maria-Diamantino Jan. Im BMNH; Minas Gerais: Anapolis Apr. Im AME; Marianna Im AME; San Jacinto Im BMNHno specific locality Im BMNH; Rio de Janeiro: Canto Gallo If MCZ; Itabapoana 2m, If BMNH, If MCZ; Itatiaia Mts 3m MCZUguna de Sacuaresma Im BMNH; Petropolis Im BMNH; Rio das Ostras Im AME; Rio Teresopolis Jun. If AME; Santa Catharina Rio Julio, 30 km. N.W. Joinville 2500' Mar. Im FSCA; Blumenau Im, If BMNH; Brusque Jul. Im AME; Corupa Im AMNHno specific locality Im, 2f MCZ, Im BMNH, 2m BMB, If AMNH; Sao Paulo: Anhangahy Nov. Im BMNH; Ara9atuba 450m Mar '2m BMNH; Mendes Im AME; Rio Pardo 2m BMB; Sao Paulo 2500' Oct.-Dec. Im MCZ, Im BMNH, If BMB Im USNM Not located: no specific locality 2m MNHN, Im, If MCZ, Im BMNH. PARAGUAY (2m): Central: Asuncion Im MNHN; Santisima Tnnidad Jul. Im AMNH. COUNTRY UNKNOWN (1 0: no specific locality If MCZ. Additional locality data: BRAZIL: Santa Catharina: Jaragua Apr. May Jul. Aug. (Hoffmann, 1936).

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282 Adelpha iphicleola (H. W. Bates, 1864) Figs. 39; 123; 205 Identification, taxonomy and variation: This species cannot be distinguished from its sister species A. iphiclus by any single character, but I have, as in A. iphiclus, grouped taxa on the basis of characters shared with neighboring taxa. In Central America, A. iphicleola is distinguished from A. iphiclus by having a more extensive orange DFW subapical marking, which extends to broadly border along vein M3, instead of entering cell M2 just as a narrow dash at the distal edge of the marking. In A. iphicleola the distal edge of the marking in cells M2 and R5 is typically concave, whereas in A. iphiclus the edge is straight or convexly rounded. The forewing and hindwing are typically more triangular than rounded, the forewing more falcate and the hindwing distal margin straighter but more dentate in A. iphicleola than in A. iphiclus. The white DFW postdiscal band on the DFW, in A. iphicleola, often broadens from the anal margin to cell M3, so that the spot in cell M3 is the widest, whereas in A. iphiclus it is usually slightly tapering from the anal margin to cell M3. On the VFW, A. iphicleola has the pale subapical marking less obscured with red-brown scaling along the veins and around the edges of the marking, so that it is more sharply defined against the dark ground color of the wing, while in A. iphiclus the ground color at the basal edge of the marking is usually dark red-brown instead of black. The shape of the distal edges of the marking mirrors that of the forewing but is sometimes more obvious. The pale dashes which compose the VFW postdiscal series are typically more concavely curved in each cell in A. iphicleola than A. iphiclus, while on the VHW the inner and outer postdiscal series converge at vein M3, where the inner series is marked more brightly with white; in A. iphiclus the series are more parallel and evenly marked with white. Many of the ventral characters are slightly variable, but if present are good characters for identification. The enlarged orange DFW subapical marking helps to distinguish this species from A. iphiclus in the Amazon basin, while the ventral characters

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283 typical of the nominate sbspecies also occur in various other populations. I discuss my rationale for considering each taxon as conspecific with the nominate subspecies in each subspecies account. There is variation in the width of the postdiscal band, the size of the orange DFW subapical marking, and the expression of the ventral characters typical of the nominate subspecies, and I have recognised five subspecies. Several of these are only slightly differentiated and future collecting may show them to be merely part of clinal variation. For the present I recognise such subspecies to facilitate future identification and the study of the true nature of variation in this species. This species has typically been regarded as a form or subspecies of A. iphiclus. However, throughout western Central America, and in eastern Ecuador, there occur specimens which are clearly phenotypically stable, sympatric with and consistently distinct from A. iphiclus. Closer examination of series of supposed A. iphiclus revealed the presence of two phenotypes in several other areas of the Nootropics, leading to the preliminary taxonomic arrangment adopted here, which was first proposed by Willmott and Hall (1999). Range and status: Mexico to western Ecuador, eastern Ecuador to northeastern Peru, northwestern Venezuela and coastal areas of Venezuela to eastern Brazil, Paraguay and Trinidad. An isolated subspecies occurs on the island of Cuba. Very common in disturbed forest in Central America, uncommon to rare everywhere else. Specimens examined: 785 (500m, 285f) Adelpha iphicleola iphicleola (H. W. Bates, 1864) Figs. 39a,b; 205a,b Heterochroa iphicleola Bates (1864: 128) TL: Guateniala. Types: BMNH(T): STm: "Illustrated in The Butterflies of Venezuela A. Neild, 1996//131//B C A Leo RhoD .nVH 1 'Pf"f*ir'""; °<^'™"ff'^*" Coll. mt.AJIBM. type No. Rh. 9835 Heterochroa iphicleola m Bates/^Heterochroa iphicleola Bates J^^^/GuM^^^^ Valley F.D.G. and O.S.//nV/Type H.T.//Syntype"; STf: "Illustrated in The Butterflies of Venezuela A. Nerld, 1996//I32//B.C.A. Up. Rhop. Adelpha iphicia Unn. Godman-Salvin Coll. I9I6.-1.//B M TYPE No Rh 9836 Heterochroa iphicleola f Bates/ZHeterochroa iphicleola Bates Type/ZPolochic Valley F DO and 0S//f7/Type HT"'

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284 BIVINH(M): ST?m: "Pacific slope//iphicleola[red label]//B.C.A. Lep. Rhop. Adelpha iphicla Linn. Godman-Salvin Coll. 1916.4.//m//Pacific Coast Guatemala F.D.G. and O.S." [all examined] =Adelpha iphicla massilides Fruhstorfer (1915: 523) TL: Western Mexico. Types: BMNH(T): STf: "Fruhstorfer Coll. B.M. 1937-285//iphicla massilides Fruhst./AV. Mexico Fruhstorfer//TYPE//Type//Syntype" [examined] Adelpha iphicla var. iphicleola Bates, Kirby (1871); Adelpha iphicla massilia Fldr., Fruhstorfer (1915) misid.; Adelpha iphicla basiloides Bates, Hall (1938) in part, misid.; Adelpha massilia Fldr,, Beutelspacher (1976: fig. 4A, 5A), de la Maza (1987), Austin (1992), de la Maza and de la Maza (1993) misid.; Adelpha iphicleola iphicleola Bates, Neild (1996) Identification, taxonomy and variation: This subspecies is distinguished from the neighboring A. iphicleola gortyna by typically having a larger orange subapical marking in cell M2, in addition to the ventral surface characters that distinguish A. iphicleola iphicleola from A. iphiclus (discussed under general identification of A. iphicleola). Other distinguishing characters are discussed under each respective subspecies. There is some individual variation in the width of the white postdiscal band and therefore the spacing of the postdiscal and submarginal series on the ventral surface. Bates (1864) described this species based on an unspecified number of male and female specimens from Guatemala, distinguishing it from A. iphiclus by the shape of the orange DFW subapical marking, in addition to several other characters listed above. He also mentions the presence of a "slender arcuated rufous line" in the VHW discal cell, actually the third cell bar, which may or may not be fiised with the red-brown line at the basal edge of the white postdiscal band, and therefore this is not a constant character. I have examined the syntype male and female in the BMNH and a ftirther possible syntype male, and all are referable to this taxon. Fruhstorfer (1915) described massilides as a subspecies of iphiclus based on a specimen or specimens from western Mexico, and a syntype female is in the BMNH. It differs in no appreciable way from typical A. iphicleola, and the name was synonymised by Hall (1938), but then resurrected to frill species status by Beutelspacher (1976). This was due to a misidentification by Beutelspacher of typical A. naxia, and the name was once again returned to synonymy by Neild (1996).

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285 Adelpha iphicleola has had an ignominious taxonomic history, being usually treated as the Central American subspecies of A. iphiclus (Fruhstorfer, 1915; DeVries, 1987; D'Abrera, 1987; Lamas and Small, 1992), and remaining unrecognised as a distinct species until Austin (1992) figured it alongside typical A. iphiclus from Costa Rica. Fruhstorfer (1913, 1915) treated the taxon as massilia, with the result that the majority of subsequent authors referred to it under this name, including Austin (1992). Neild (1996) was the first author since Bates' original description to treat the species as distinct with the correct name (see Willmott and Hall, 1999). Range: Mexico to eastern Panama, intergrading to A. iphicleola gortyna fi-om Costa Rica to western Panama. Immature stages: Ahnost certainly some of the hostplants records for A. iphiclus actually apply to this species, but given their close relationhsip the two probably share the same hostplants anyway. Similarly, the early stages of the species have probably been reared and reported under the name A. iphiclus. Given the phenotypic similarity of the adults there will probably prove to be no discernible differences in the early stages, and I summarise the available information under A. iphiclus. Habitat and adult ecology: This is possibly the most common Adelpha species in the western parts of its range, extending fi-om sea level to 1500m in humid and subhumid evergreen and semideciduous forest, where it flies throughout the year (de la Maza, 1987; de la Maza and de la Maza, 1993). Austin et al. (1996, as massilia) record it fi-om the complete range of habitats from fields and second growth to primary forest in Guatemala, where it it common throughout the year. In Costa Rica it is most frequently encountered in drier areas on the Pacific slope, where it flies with A. iphiclus, but it always appears to be more abundant than this species (Austin, 1992). DeVries (1987) reports various observations in Costa Rica under the name A. iphiclus, but these may apply to either that species or this (see this section under A. iphiclus). A single male in the AMNH was captured in Costa Rica on flowers of Cordia.

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286 Specimens examined (310m, 1430: t trans, togortyna. MEXICO (215ni, 92f): Campeche. Campeche Jul. 2m AMNH; Chiccna Ruins Im AMNH; China Oct. Im AMNH; Escarcega Aug. Im FSCA; Chiapas. Campet Aug. If AME; Chajul Rio Lacuntun 200m Jul. Im KWJH; Comit^n Aug. 2m AME; Comitan, 10 km. S.E., Cuauhtemoc Sep. 2m AME; El Chorreadero Oct. Im MUSM; El Sumidero Oct. If MUSM; U Chacona Aug. 2m MUSM; Las DeHcias Jun.-Sep. 5m AMNH, 2m, If AME; Mazatan Jul. If AME; Paraiso Oct. Im AME; Muste Jul. 3m AMNH; Ocozocoautla Feb. If AMNH; Pinola Sep. 2m AME; Rancho Santa Ana Jun. 2m AMNH; Rio Chixoy Aug. 2m AME; San Antonio Buenavista 1500m Aug. If MUSM; San Carlos Jul. 3m, 2f AME; San Cristobal de las Casas Jan. Feb. Aug, Im, If AMNH, 2m AME; San Jeronimo 600m Aug. Im USNM; San Quintin Aug. Sep. 4m AME; Tapachula Aug. Im AME; Im, If AMNH; Tonags Jan. Im AMNH; no specific locality Jul. Sep. 4m, 4f AMNH, Im FSCA, 2m AME, If MNHN; Colima. Comala 2100' Oct. Im, If AME; La Salada 1000' Jan. Im AME; Guerrero: Acahuizotia Sep.-Nov. If BMNH, 3m, If AME; Acapulco Sep. 4m AMNH, If AME; Chilpancingo Aug. 1200m 4m AMNH, If BMNH; Colotlipa 1020m Aug. Im AME; Dos Arroyos 1000m Sep. If BMNH; El Treinte Sep. 2m, If AMNH; Iguala 2m AMNH; Tierra Colorada 500m Aug. Im AME; Zihuatenango Dec. Im AMNH; no specific locality 21m, If BMNH, Im VSHM, Jalisco. Cumbre If AMNH; Magdalena 1380m Aug. If AME; Puerto Vallarta Oct. If USNM; San Luis Allende-Ayutla Aug. 2m BMNH; Tenacatitla If AMNH; Michoacan: Coahuayana Aug. 3m, 4f AME; Morelos: Ainacuzac Aug. If AMNH; Nayarit: Compostela Aug. If AME; Tepic Im BMNH, Im MNHN; Zapata 900m Aug. Im, 2f AME; no specific locality Oct. If AMNH; Oaxaca: Candelaria Loxicha 550m Aug. -Nov. 6m, 4f AMNH, Im USNM, If FSCA, 3m, If AME; Chiltepec Aug. Oct. 2m AMNH, 2m AME; Comaltepec Oct. Im AMNH; Espinal Jun. If BMNH; Juchatenango Dec. If AMNH; Mixtepec Dec. Im AMNH; Naranjal-Chiltepec 6m AMNH; Oaxaca Aug. If BMNH, If USNM, Im, 2f AME; Rancho San Carlos Aug. Im AMNH; Salina Cruz Jun. Jul. If BMB; Soyolapan el Bajo May Im, If AMNH; Valle Nacional 800m Jul. Im MUSM, If AMNH; Quintana Roo: X-can May. Jul. Oct. 2f FSCA, Im AMNH; Sinaloa: Mazatlan 30m Aug. Im BMNH, 3m, 3f AME; Papachal Dec. If AMNH; Sinaloa Feb. 2m AMNH; Sinaloa, 24 mi. N.W., 15 Jun. Im AMNH; Tabasco: La Venta 10m Jan. Im AME; Tepescuintla Aug-Oct. 15m, 3f AMNH; Veracruz: Atoyac Im BMNH; Coatepec 2f USNM; Cordoba Mar. Sep. Dec. 2m AMNH, 2m USNM, Im BMNH; Fortin Aug. Im AMNH; Jalapa May Im BMNH, 4m USNM; Motzorongo If USNM; Nanchital sea level Feb. Im AME; Orizaba Im BMNH; Palo Gaucho Aug. Im USNM; Popoctepetl Tuxtia 400m Oct. Im MUSM; Presidio Jul. Aug. Sep. 3m, If AME, 3m USNM, If AMNH; Tezonapa Jul. Im AME, Im AMNH; Villa Ju4rez If AME; Zongolica Jun. 2m BMNH; Yucatan: Dolores Otero Jul. If AMNH; MiridaJul. If AMNH; Piste Jul.-Sep. 18m, 6f AMNH, 7m, 4f AME, 4m, If FSCA, ImBMNH, If USNM; Tekax Sep. Im AMNH; Tzitas If AMNH; Valladolid Im, If BMNH; Not located: W. Mexico If BMNH; no specific locality 4f AMNH, 2f BMNH, Im BMB, 5m, If MNHN, 2f USNM. BELIZE (Im): Corozal: Sarteneja Dec. Im FSCA. GUATEMALA (46m, 230: Alta Verapaz: Cubilguitz Im BMNH; La Vega del Cuajilote 250m Aug. If AME; Polochic Valley 4m, 2f BMNH; Tamahu 2m BMNH; Tucuru Jun. Jul. 4m BMNH; Baja Verapaz: Panima Im, If BMNH; San Jeronimo Im, 2f BMNH; Chiquimula: no specific locality Im MNHN; El Peten: El Ceibal Nov. Im AMNH; Sayaxche Sep. Oct. 1 5m, 9f AMNH; El Progreso: Motagua Valley Im BMNH; Escuintla: Escuintia Jul. If USNM; Palin Jul. Aug. Im BMB; Guatemala: Amatitl4n 3800' Jul.-Sep. 2f BMB; Izabal: Dartmouth Im BMNH; Izabal Im BMNH; Puerto Barrios If BMB; Quezaltenango: Volcan Santa Maria Jul. If USNM; Retalhuleu: San Sebastian If USNM; Santa Rosa: Guazacapan Im BMNH, Im, If AME; Zacapa: Zacapa Aug. If USNM; Not located: El Chila 2200' Sep. 2m BMB; Pacific Coast 2m BMNH; no specific locality 4000' Dec. 2m BMNH, 2m BMB, 2m MNHN. EL SALVADOR (2m, 50: La Libertad: La Libertad 10m Dec. Im, If AME; San Salvador: Apopa 2000' Sep. Im BMNH; Ilopango 1200' Sep. Oct. If BMNH; Santa Tecla 900m Nov. 2f AME; Santa Ana: Citala 800m Oct. If AME. HONDURAS (4m, 30: Colon: Trujillo Dist. Im AMNH; Cortes: San Pedro Sula Jul. If USNM, If BMNH; Francisco Morazan: Tegucigalpa If BMNH; Not located: Las Minas Jul. Im USNM; no specific locality Im BMNH, Im USNM. NICARAGUA (8m, 70: Boaco: Comoapa 2000' Jul. 2m BMNH; Chontales: Chontales 2f BMNH; Managua: Managua Oct. Nov. 3m, If AMNH, 2f USNM; Managua, 12 km. S., Nov. If USNM; Rio San Juan: San Carios Im AMNH; Zelaya: Bluefields Dec. If AMNH; Not located: Rochonil Corozal Jul. Im AMNH; no specific locality 2m AMNH, Im BMNH, Im MNHN. COSTA RICA (16m, 90: Alajuela: San Mateo Dec. Im BMNH; Cartago: Juan Vifias 2500' Jan. Sep. Nov. 4m BMB, 2m USNM; Guanacaste: Cailas Jul. Im, If FSCA; Cailas, 5 km. N.W., Aug. If USNM; Cailas, 24 km. N.W., Im AMNH; Nosara Sep. Im FSCA; Santa Rosa 1 00m Aug. Nov. Im KWJH, If AME; San Antonio Feb. If AME; Heredia: San Antonio de Belen Aug. Im USNM; Puntarenas: Palmar Norte Aug. If USNM; San Jose: Puriscal Nov. If USNM; Villa Colon 600m Jul. Im USNM; Not located: no specific locality Sep. 2f BMNH, 2m MNHN, If USNM, Im AMNH. PANAMA (11m, 40: Canal Zone: Caldena Hydro Plant Jan. Im USNM; Farfan Feb. 3m AME, 4m, 2ft AME; Madden Forest Feb. Im AME; Howard AFB Oct. Im AME; Chiriqui: Valle de Chiriqui 25-4000' If BMNH; Colon: Pifia If AME; Not located: no specific locality Im USNM COUNTRY UNKNOWN (7m) Im AME, 6m MNHN. Additional locality data: MEXICO: Chiapas: throughout see de la Maza and de la Maza (1993); Guerrero: La Sabana; Jalisco: El Tuito; Oaxaca: Metates (de la Maza, 1987); Quintana Roo: Carrillo Puerto; Chumpon; Chunyaxche; Noh-bek; Tres Reyes; Tulum (de la Maza and Bezaury, 1992); numerous records in Beutelspacher (1976: as A. massilia) and Warren et al. (1998: as A. iphiclus massilides) may apply to this species or A. iphiclus iphiclus. BELIZE: Cayo; Orange Walk. Toledo (Meerman, 1999). Adelpha iphicleola leucates Fruhstorfer, 1915 Figs. 39c,d Adelpha iphicla leucates Fruhstorfer (1915: 523) TL: Bahia [Brazil]. Types: BMNH(R): HTf: "Holotype//TYPE//Bahia//Felder ColIn.//ephesa Men./Ziphicla leucates Fruhst." [examined] =Adelpha iphicla daceleia Fruhstorfer (1915: 523) TL: Trinidad. Types: BMNH(T): LTm: "Fruhstorfer Coll. B.M. 1937-285//iphicla daceleia Fruhst.//Tnnidad//TYPE//Type//Syntype"; BMNH(M): PLT?m: "Trinidad/ZFruhstorfer Coll. B.M. 1937-285" [both examined]

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287 Adelpha iphicla Linn., Kaye (1904); Adelpha iphicla phera Fruhst., Hall (1938) misid.; Adelpha iphiclus phera Fruhst., Neild (1996) misid.; Adelpha iphicleola leucates Fruhst., Willmott and Hall (1999) Identification, taxonomy and variation : This subspecies is distinguished from the nominate by having sHghtly rounder wings, a broader white postdiscal band and a narrower orange DFW subapical marking, which usually consists of a triangle in cell M2 that touches vein M3 at its apex. It is distinguished from A. i. iphimedia by that subspecies having an even broader postdiscal band and narrower orange DFW subapical marking, which usually only extends into cell M2 at its distal edge, not reaching vein M3. It is distinguished from both A. i. thessalita and A. i. gortyna by the broader postdiscal bands and characters of the pale VFW subapical marking that distinguish this species from A. iphiclus (see under species account for A. iphicleola). This subspecies is very similar, if not indistinguishable from, A. iphiclus in northern Venezuela and Trinidad. It is generally smaller, and I have used the characters discussed under the identification of A. iphicleola from A. iphiclus in species account of the former, particularly those of the venfral surface, to recognise specimens of ^4. iphicleola leucates from that area, but there are certain specimens which appear to be intermediate. This subspecies is also very similar to A. iphiclus ephesa, and the characters that distinguish it are discussed under the latter subspecies. This subspecies is individually and geographically variable, and it may well be the case that more material will illustrate consistently distinct populations that merit taxonomic recognition. For the present, given the relatively few specimens that I have examined and the individual variation, I treat all specimens of this species from northern Venezuela to Paraguay as a single taxon. Northern Venezuelan specimens typically have broader white postdiscal bands and a narrower orange DFW subapical marking, some of which approach A. iphicleola iphimedia and are marked with a f in the list of specimens examined below. Certain Trinidad specimens approach these Venezuelan specimens, but the postdiscal band is variable and usually narrower. Certain specimens have an orange DFW subapical marking similar to A. i. iphiclus, in addition to a darker VHW and

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288 straighter VHW postdiscal and submarginal series, and these may actually represent A. iphiclus. For the present, however, I treat them as A. iphicleola leucates, since typical A. iphiclus also seems to be present on Trinidad; one female specimen in the BMNH collected by M.E. Fountaine is very much like the syntype of exanima figured by Neild (1996), considered here a synonym of A. iphiclus iphiclus, with a narrow orange DFW subapical marking typically shaped as in A. i. iphiclus, relatively broad postdiscal bands, larger size, a dark VHW, and parallel VHW postdiscal and submarginal series. To date I have seen few specimens fi^om southern eastern Brazil, possibly because they are rare there, present and very similar to A. iphiclus ephesa, or simply absent. It is therefore unclear whether specimens fi^om Paraguay represent an isolated population, but they appear to differ little phenotypically from the holotype. Fruhstorfer (1915) described this subspecies based on a specimen from Bahia in the Tring Museum, and the holotype is now in the BMNH(R). I have been unable to locate the specimens he also refers to from Pemambuco in Staudinger's collection, there are neither A. iphiclus or A. iphicleola in the ZMHU from that locality. He also described daceleia on the same page, largely based on its smaller size, from an imspecified number of specimens from Trinidad. Due to the difficulties in assigning Trinidadian specimens to A. iphiclus or A. iphicleola, I designate the specimen in the BMNH(T) as the lectotype of Adelpha iphicla daceleia, with the following label data: "Fruhstorfer Coll. B.M. 1937285//iphicla daceleia Fruhst.//Trinidad//TYPE//Type//Syntype". This specimen has many of the ventral characters typical of A. iphicleola, except for rather sfraight VHW postdiscal series, and I therefore regard it as representing this species. For the reasons given above I also regard it as a synonym of A. iphicleola leucates. Hall (1938) synonymised both leucates and daceleia with phera, another Fruhstorfer taxon here regarded as synonymous with A. iphicleola iphimedia. Neild (1996) also used the name phera to refer to Venezuelan specimens treated here as A. iphicleola leucates, and referred to dacaleia as representing a distinct subspecies. I prefer to place daceleia as a

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289 synonym of leucates, rather than vice versa, since I am much more sure that the holotype of leucates is truly conspecific with A. iphicleola iphicleola, given the similar orange DFW subapical marking, shared ventral surface characters and apparent sympatry with typical A. iphiclus, whilst there is an element of uncertainty surrounding the identification of all Trinidadian specimens. The name daceleia is still available should the north Venezuelan and Trinidadian population prove to be distinct from typical leucates. Range: Coastal regions from northern Venezuela to eastern Brazil, Paraguay and Trinidad, and possibly southeastern Brazil. Habitat and adult ecology: This subspecies is common in parts of its range, such as Trinidad, but generally rare in continental South America. It has been recorded up to 1100m throughout the year, at least in Venezuela. Cock (pers. comm.) reports that it is common and widespread in or near primary or secondary forest in Trinidad, while Barcant (1970) states that males are encountered as solitary individuals in sunny areas, and also puddle at damp sand. Specimens examined (100m, 74f): t very wide white postdiscat bands, reduced orange DFW subapical marking, very similar to A. i. iphimedia but smaller; % large specimens, narrower postdiscal bands, orange DFW subapical marking shaped as in A. i. leucates; * DFW orange like A. i. iphimedia, resembles syntype phera mislabeled?; ? identification uncertain, may be A. iphiclus. VENEZUELA (21m, 25f): Aragua: El Limon 450m Feb. Jul. Im MUSM, Im AME; U Cesiva 700m Nov. Im JFL; Portochuelo Pass 1 100m Dec. If MUSM; Pozo Diablo, Maracay Oct. Im FSCA, 3m AME; Rancho Grande Apr. Jun. Im, If USNM; Valencia Sep. Im BMB; Bolivar. Bolivar IfJ BMNH; via El Dorado-Santa Elena km. 92 500m Jun. If? AFEN; Carabobo: Puerto Cabello Jul. 1ft BMNH; San Esteban Jun.-Sep. 2f? BMNH, Im, If BMB; Distrito Federal: Caracas 3000' Feb. Sep. Oct. Iff BMNH, Im AME, 2m, 2f BMB; Macuto 400' Mar. Iff BMNH; Lara: Sarare If MNHN; Monagas: Barrancas 3m, 6f AME; Tachira: Ciicuta If BMB; Not located: no specific locality 2m, 2f AMNH, Im, 2f(lt,lt) BMNH, Im, If MNHN, Im MCZ. TRINIDAD (44m, 330: Caroni: Brasso Im AMNH; Tabaquite, Narieva If BMNH; St. Andrew: Curuto Oct.-Dec. If BMB; Santa Cruz Mar. Im AMNH; St. George East: Arima Jan. Oct.-Dec. Im, If BMB, If AMNH; Arima valley Mar. Dec. 3f AMNH; Caparo Valley Im BMNH; Simla Dec. 2f AMNH; St. George West: Chanello Rd. Port of Spain Feb. 5m, 2f BMNH; Fondes Amandes Im AME; Hololo Mt. rd. Mar. Aug. 3m, 2f AMNH, If USNM; Maraval Oct.-Dec. Im BMB; St. Anns Sep. Oct. Dec. 2f AMNH, Im, If AME, 2m BMB; St. Patrick: Siparia Oct.-Dec. Im BMB; Victoria: Mt. Diable Jan. May 2f BMNH; Not located: Botanical gardens 2f AMNH; La Brea Sep. 2m FSCA; Moruga Im AME; Northern Mts. Dec. Jan. Im BMB; no specific locality Apr. May Oct. Dec. 16m, 5f BMNH, 2m, 2f AME, Im, If BMB, Im, 2f MNHN, 2f AMNH, 2m MCZ. GUYANA (If): Upper Takutu/Upper Essequibo: Aunai, Essequibo If? BMNH. FRENCH GUIANA (If): Laurent du Maroni: Maroni River If AME(may be mislabeled gortyna or iphicleola). "PERU" (5m): Not located: "PeruZ/Schaus" 5m USNM-error. BRAZIL (17m, 50: Bahia: Bahia Im BMNH; Itabuna Jan. Im AME; Ceard: Ceara 2m BMNH; Guanabara: Rio R. If BMNH; Mato Grosso: Corumba Apr. 4m BMNH; Urucum, 15 mi. S. Corumba Apr. 3m BMNH; Para: Para Im?, If BMNH; Pernambuco: Pemambuco 2m, 2f BMNH; Santa Catharina: no specific locality If (identical to A. i. iphicleola mislabeled?) BMB; Not located: no specific locality 2m BMNH, Im MNHN. PARAGUAY (Urn, 60: Bogueron: Estancia Cooper, Alto Paraguay Im, If BMNH; Central: Patiilo Cue 3m, 3f BMNH; Santisima Trinidad Jun. Aug. 2m AMNH; Guaird: Colonia Independencia Sep. Im AME; Paraguari: Sapucay lfi;W. Foster lO/XI/04) BMNH, Im BMB; San Pedro: Rio Jejuy Im BMNH; Not located: Central Im BMNH; no specific locality If BMNH, Im* USNM. ARGENTINA (Im)CorrientesItuzaingo Jul. Im AME. COUNTRY UNKNOWN (Im, 30: no specific locality Im, If BMNH, If AME, If MNHN Adelpha iphicleola iphimedia Fruhstorfer, 1915 Figs. 39e,f

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290 Adelpha iphicla iphimedia Fruhstorfer (1915: 523) TL: Cuba. Types: BIMNH(T): STm: "Fruhstorfer Coll. B.M. 1937-285//iphiniedia Fruhst.//Cuba//TYPE//Type//Syntype"; STf: "Fruhstorfer Coll. B.M. 1937-285//iphiniedia Fruhst./ZHelguin, Cuba H. Rolle Berlin SWI 1 /nTPE//Type" [both examined] =Adelpha iphicla phera Fruhstorfer (1915: 523) TL: locality stated as unknown. Types: MHNG: STm?: "Mexique/Ziphicla phera Fruhst." [photograph examined] Nymphalis basilea Cram., Poey (1847: 48); Adelpha basilea Cram., Gundlach (1881); Adelpha iphicla Linn., Holland (1916); Adelpha iphimedia Fnihst., Brown and Heineman (1972); Adelpha iphicleola iphimedia Fruhst., Willmott and Hall (1999) Identification, taxonomy and variation: This subspecies is distinguished from all others by the very broad postdiscal bands and narrow orange DFW subapical marking, which scarcely enters into cell M2 and is particularly narrow in cell R5. There is slight variation in the width of the postdiscal bands and orange DFW subapical marking. Fruhstorfer (1915) described this subspecies from an unspecified number of specimens from Cuba, mentioning both of the diagnostic characters, and two syntypes are in the BMNH(T). A fiirther three males in the BMNH(M) from Fruhstorfer's collection are also probable syntypes. The name phera was described by Fruhstorfer on the same page in comparison with his taxon exanima (figured by Neild, 1996), supposedly differing in wider postdiscal bands and relatively "small franscellular spots" on the ventral surface. He gave no indication of the number, sex or whereabouts of the type specimen(s), and stated that the locality was unknown. Hall (1938) interpreted phera to be what I have freated here as A. i. leucates, while Neild (1996) applied the name just to northern Venezuelan A. iphicleola (as A. iphiclus phera). Both these decisions were entirely logical given Fruhstorfer's almost worthless description, but a specimen discovered by Gerardo Lamas in the MHNG, which appears to be a syntype and bears a label stating "iphicla phera Fruhst." in Fruhstorfer's hand, is clearly not from northern continental South America. Although this specimen is labeled "Mexique", and Fruhstorfer (1915) claimed the locality was unknown, it may be that he made a mistake or the label was subsequently added. The specimen has the very reduced orange DFW subapical markings and wide white postdiscal bands typical of the Cuban subspecies iphimedia, but is also very similar to a specimen in the USNM labeled "Paraguay". The

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291 latter specimen may be mislabeled, or it may be that specimens from Paraguay have converged on the Cuban phenotype. I regard the name for the present as representing, and therefore being synonymous with, A. iphicleola iphimedia (see Willmott and Hall, 1999) This taxon has generally been treated as a subspecies of A. iphiclus or even a distinct species (Brown and Heineman, 1972), but the latter is untenable given the close similarity of certain northern Venezuelan specimens of A. iphicleola leucates. I place it as a subspecies of A. iphicleola for the same reasons as leucates, that it possesses all the characters typical of A. iphicleola iphicleola with the exception of the orange DFW subapical marking extending to vein M3. Range: This subspecies occurs only on the island of Cuba and the neighboring Isle of Pines (Riley, 1975). Habitat and adult ecology: Riley (1975) states that in Cuba this is a forest butterfly encountered typically as solitary individuals, and although it is widespread, it is not common. Alayo and Hernandez (1981) state also that it tends to fly high in the canopy, where it rests occasionally. Specimens examined (42m, 6 If): CUBA (40in, 571): la Habana: Havana 2f AMNH; U Havane If BMNH; Las Villas: Soledad, Santa Clara If MCZMalanzasMatanzas Apr. Sep. Im BMNH, 2m BMB; Orienle: Baracoa 2m USNM; Guantanamo If AMNH; Holguin lOm Uf BMNH SantiagoJul. Dec. 3fUSNM, ImAME, If BMB; Sierra Maestra 1000" 6f MCZ; Tanamo Jan. Mar. lm,4fUSNM, Im 2fAMNH If BMNH; Torqumo Mass 3500' Jul. 3ni, 8f MCZ; Torquino River 100' Im MCZ; Pinar del Rio: N. of Vifiales Sep. If AMNHSanta Cnstdbal If BMNH; Not Icoated: E. Cuba Im, 4f BMNH; Mocha Apr. 2m BMB; no specific locality 7m, If BMNH 2m If MCZ 3f BMB, 2m MNHN, Im, 2f USNM, 3m, 3f AMNH. "GUYANA" (Im): Cuyuni/Mazaruni: Kartabo Im AMNH-erri)r' COUNTRY UNKNOWN (Im, 40: no specific locality Im, 3fBMNH, If USNM. Adelpha iphicleola gortyna Fruhstorfer, 1915 Figs. 39g-j Adelpha iphicla gortyna Fruhstorfer (1915: 523) TL: Colombia, Cauca Valley. Types: BMNH(M): LTm: "TYPE//Bogota//Paratype//Fruhstorfer Coll. B.M. 1933-131"PLTmColumbien ex coll. H. Fnihstorfer//TYPE//Paratype" = iphiclus; BMNH(T): PLTm: "Fruhstorfer Coll. B.M. 1937-285//iphicla gortyna Fruhst.//Columbien ex. Coll. H. Fruhstorfer//Bogota//TYPE//Type//Syntype" = iphiclus [all examined] Adelpha iphicl^Lmr^ DeVries (1987) misid.; Adelpha iphiclus Linn., DeVries (1987: pi. 27, fig. 1) ?; Adelpha iphicleola ssp. nov.?Neild(1996:pI.3,flg. 130);^rfe/pA«,>A,c/eo/agorO'/iflFruhst.,WillmottandHall(1999) Identiflcation, taxonomy and variation: This subspecies is distinguished from the nominate by generally lacking the ventral characters which typically separate this species from A. iphiclus, and by having a

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292 reduced orange DFW subapical marking which usually has a straight basal edge and just touches vein M3, rather than broadly bordering this vein. It is thus very similar to A. iphiclus iphiclus, and its identification is discussed under that taxoa Adelpha iphicleola thessalita is distinguished by having a more extensive orange DFW subapical marking, typically broadly bordering vein M3, and the white DFW postdiscal band tapers from the anal margin to cell M3, with a less strongly concave distal edge, instead of being of even width. Adelpha iphicleola leucates has a wider white postdiscal band, and ventral characters typical of the nominate subspecies. There is slight individual variation in the width of the orange DFW subapical marking. The single specimen known from northwestern Ecuador differs from typical A. i. gortyna in having a narrower postdiscal band which is tinted greenish blue, and a narrower orange DFW subapical marking. Since these characters correspond to the wing pattern of the mimetic group of which this species is part in that area, it is probable that they occur stably in A. iphicleola throughout western Ecuador. However, given that I have only examined a single, slightly worn specimen, and the variability in this species, I do not describe it at present. Fruhstorfer (1915) described this taxon as a subspecies oi A. iphiclus, based on specimens from "Colombia, very common in the Cauca Valley". He compared it to A. iphicleola iphicleola, and said that it had a smaller orange DFW subapical marking and narrower postdiscal bands. I have located three syntypes in the BMNH, two which are typical Colombian A. iphiclus and a third which has a more extensive orange DFW subapical marking and white DFW postdiscal band which is broadest in cell M3, which I regard as A. iphicleola. Since I otherwise regard Colombian A. iphiclus as representing the nominate subspecies, and the name funalis is available should it prove to be distinct, Willmott and Hall (1999) designated the third of these syntypes as the lectotype A. iphicla gortyna and placed the name as a subspecies of A. iphicleola. Despite the lack of shared venfral characters between this taxon and nominate A. iphicleola, the similar shape of the orange DFW subapical marking, and its geographic

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293 position, between the nominate subspecies and A. iphicleola leucates, which also has ventral characters similar to the nominate, suggests it is conspecific with both of these taxa. Furthermore, certain specimens from Panama appear to be intermediate between this subspecies and the nominate, with the ventral surface as in the former, but the orange DFW subapical marking as in the latter. Range: Western Colombia to northwestern Ecuador, to northwestern Venezuela in the foothills of both slopes of the Merida range to northwestern Colombia (Meta province). Very probably also occurring in eastern Panama. Habitat and adult ecology: This subspecies is rare in collections. In western Ecuador I found a single male attracted to rotting fish in a river side trap at the edge of a road and primary forest, in bright sun, at the end of the wet season. This is the first record for the species from western Ecuador, where it appears to be considerably rarer than its close relative, A. iphiclus. Specimens examined (30m, 6f): t broader postdiscal bands; ? identification uncertain. VENEZUELA (2m): Tachira: Via Choiro del Indio San Cristobal Nov. Im AFEN; Hda. Panaga 800m Jul. Im AFEN. COLOMBIA (27m, 6f): Boyaca: Muzo Im? MNHN, Im AME; Caldas: Quebrada de Pidrias, Mpio. Anserma Apr. Im ESM; Quebrada Pidrias, cerca a la desembocadura al Cauca Mpio. Anserma Im MHNM; Cauca: Espejuelo, nr. Cali Cauca Mar. Apr. Im BMNH; Cundinamarca: Bogoti Im BMNH, If AMNH; Fusagasugi If AMNH; Mt. Redondo Dec. If AME; Tobia Feb. Im KWJH; Villeta Imf AMNH; El Cesar: Manaure Im, If BMNH; Magdalena: Rio Chiquinla, Atinquez, Santa Marta 800m Jul. If BMNH, Im? MNHN; Mela: Restrepo May Im JFL; "Villavicencio Meta Colombia 1800' July 13 1972 leg. R. Robbins//R.K. Robbins collection" 1ft USNM; "Villavicencio Col. Spring 1924 [collector name illegible]" Im AMNH; Tolima: Payande, Rio Frio 950m Feb. Mar. 3m AME; Rio Ata 1000m May 2m AME; Nol located: de Bogoti a Buenaventura Dec. -Feb. 4m BMNH; no specific locality 4m BMNH 2m AMNH. ssp. nov.? ECUADOR (Im): Esmeraldas: Rio Piguambi 800m Jul. Im KWJH. Adelpha iphicleola thessalita Willmott and Hall, 1999 Figs. 39k,l; 123a,b Adelpha iphicleola thessalita Willmott and Hall (1999: in press) TL: Finca San Carlo, Napo, Ecuador. Types: ECUADOR: KWJH: HTm: Finca San Carlo Sep. 600m; PTm: same data as HTPTm: El Capncho Napo 800m Oct.; 3PTm: Las Minas de Misahualli Napo 400m Jul.; 3PTm: Pimpilala Napo 600m Sep PTm' PTf: Yanna Rio Manduro nr. Coca 250m Sep.; 2PTm: Puyo-Canelos rd. km. 30 600m Oct.; BMNH(M): PTm: "Canelos' Ecuador, A. Simson//m//B.C.A. Lep. Rhop. Adelpha iphicla Linn. Godman-Salvin Coll. 1916.-4."BMNH(R)PTm"Napo Rio Napo, E. Ecuador July 1927 (Dr. Spillmann)"; AME: PTm: Puerto Misahualli Napo 6 Nov. 1983 D and J JenkinsMNHN2PTm: Rio Napo; DAT: PTm: "Talag Ecuador Alt. 800m D.A.Trembath 15.IX.95 No. 14179". PERUAMEPTf "Mo'vobamba! Pern, VI. 4. 1944//A.C.AllynAcc. 1965-5" Jun.[all examined] •

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294 Identification, taxonomy and variation: This subspecies is distinguished from A. i. gortyna and A. i. leucates under those subspecies, and from A. iphidus iphiclus under that taxon. The extent of the orange DFW subapical marking is sHghtly variable, sometimes onll bordering vein M3 for a short distance. Willmott and Hall (1999) described this taxon as a subspecies of ^4. iphicleola on the basis of the similar orange DFW subapical marking and hindwing shape, and apparent intermediate specimens in western Colombia. Range: To date known only from eastern Ecuador to northeastern Peru, but very probably extending into southeastern Colombia. Habitat and adult ecology: Although this subspecies is very rare in major museum collections, it is actually not uncommon in the field, in lowland forest habitats up to 850m. Males may be atfracted to rotting fish in ridgetop light gaps in primary forest, often several specimens to a single trap. They are also found perching in small groups in large clearings near rivers on tops of bushes 3-4m high, in a manner similar to A. serpa and A. thoasa. I have not observed this perching behavior in A. iphiclus. Specimens examined (18m, 2f): ECUADOR (18m, IQ: Napo. El Capricho 800m Oct. Im KWJH; Finca San Carlo 600m Sep. 2m KWJH; Las Minas de MisahualH 400m Jul. 3m KWJH; Rio Napo Im BMNH(R); Pimpilala 600m Sep. 3m KWJH; Puerto Misahualli Nov. Im AME; Rio Napo 2m MNHN; Talag 800m Sep. Im DAT; Yarina, Rio Manduro, nr. Coca 250m Sep. Im, If KWJH; Pastaza: Canelos Im BMNH(M)Puyo-Canelos rd., km. 30, 600m Oct. 2m KWJH. PERU (10: San Martin: Moyobamba Jun. If AME. Additional locality data: ECUADOR: Napo: Apuya Aug. Sep.; Chichicorrumi Jul.; Pimpilala Aug.; Rio Shandia Sep.; Tiguino Aug. (Willmott and Hall, unpubl. sight records). Adelpha abyla (Hewitson 1850) Figs. 40a,b; 124a,b; 206a,b Heterochroa abyla Hewitson (1850: 437, pi. IX, fig. 7) TL: Jamaica. Types: BMNH(T): STf: "Jamaica//B.M. TYPE No. Rh. 9852 Heterochroa abyla f Hew.//Type//Syntype"BIVINH(M): STf: "(Jamaica) Hewitson Coll. 79-69 Heterochroa abyla. 3.//Paratype//abyla//B.M. TYPE No Rh 9853 Heterochroa abyla f Hew."; STm: "(Jamaica) Hewitson Coll. 79-69 Heterochroa abyla. 1."; STf: ""(Jamaica) Hewitson Coll. 79-69 Heterochroa abyla. 2. [all examined]. Adelpha abyla Hew., Kirby (1 871); Adelpha iphicla abyla Hew., Hall (1 938); Adelpha abyla Hew., Riley (1975)

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295 Identification, taxonomy and variation: Adelpha abyla is easily distinguished from A. iphiclus and A. iphicleola by the pronounced hindwing margin at veins Cu2 and Cul, the whitish submarginal Hne on the DHW, and the broad and dark subapical orange marking on the DFW. There is Uttle variation in the specimens examined. Hewitson (1850) described the species from an unspecified number of specimens in the British Museum and his own collection, and I have located 1 male and 3 female syntypes in the BMNH. The figure of the dorsal surface of the right hand pair of wings in the original description and the type locality leave no doubt as to the identity of this taxon. The lack of sympatry of A. abyla and either of its two potential sister species, A. iphiclus and A. iphicleola, leave its specific status open to question. I have been unable to find any consistent genitalic differences between any of these three taxa, notwithstanding the claims of Schwartz (1989a) that the genitalia of A. abyla are indistiguishable from A. f. lapitha (and therefore would be very distinct from either A. iphiclus or A. iphicleola). I conclude that Schwartz's claim must have been based on misplaced specimens, since the genitalia that he describes as belonging to A. fessonia clearly belong to A. iphiclus, A. iphicleola ox A. abyla. While the majority of authors (Fruhstorfer, 1915; D'Abrera, 1987; Schwartz et al, 1989a,b; Smith et al., 1994) have regarded A. abyla as a good species, Hall (1938) placed it as a subspecies of A. iphiclus. I have been tempted to place it as a subspecies of A. iphicleola, but I retain it as a good species for three reasons: the modification in shape of the hindwing margin and the shape of the DFW orange subapical area, which extends over the inner submarginal series, are two very distinctive characters which occur in no other mainland subspecies of either A. iphiclus or A. iphicleola. These wing shape and pattern differences are greater than occur between either A. iphiclus and A. iphicleola, and between any of their constituent subspecies. The shape of the DFW orange subapical marking is more similar to A. iphiclus, only just extending into cell M2,

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296 while the hindwing shape and distally displaced white postdiscal spot in cell M3 on the DFW are more similar to A. iphicleola. I am therefore unable to say with certainty whether A. iphiclus or A. iphicleola is the sister taxon to A. abyla. There are also hints of morphological differences in the early stages in the two published descriptions (Swainson, 1901; Smith et al., 1994), discussed below. It is interesting to note that the dorsal wing pattern of this species more closely resembles that of Central Amercian A. iphicleola iphicleola and A. iphiclus iphiclus than the Cuban A. iphicleola iphimedia, geographically the closest and therefore the most plausible sister taxon if relatively recent dispersal were proposed to explain the distribution of the two taxa. Miller and Miller (1989) regarded the evolution of the Jamaican butterfly fauna to have been heavily influenced by the proximity of the island to Central America during the Eocene, and suggested that the ancestor of the endemic Jamaican swallowtail, Pterourus homerus (Fabricius), may have become isolated from the mainland population during this period. Since A. abyla appears to have a poor dispersal ability across open water, having failed to colonise or establish populations on any of the neighboring islands of the Greater Antilles, it seems possible that a similar vicariance explanation may also account for its origin and evolution on Jamaica. Range: The island of Jamaica. Immature stages: According to Ellwood and Harvey (1990), there is a watercolor painting of the larva and pupa by Lady Edith Blake, painted between 1889 and 1898, in the collection of the Entomology Library at the BMNH. A handwritten annotation on the painting notes that the adult eclosed in August, nine days after pupation. The depicted hostplant, Mussaenda treutleri (Rubiaceae), which shows larval feeding damage, is a cultivated ornamental, native to Asia. Swainson (1901) was the first to provide a description of the larva and pupa of this species, while Smith et al. (1994) summarised detailed notes on the early stages by Milner and Turner. The pale brown egg is laid singly beneath the tip of a young leaf on a sapling, and third to fifth instars are brown, greyish

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297 brown ventrally, with lateral black stripes, and pale grey dorsally on Al. The head capsule is dark brown with a paler median stripe, and in addition to the usual lateral chalazae it apparently has "numerous smaller, light brown spines". Since the front of the head capsule in A. iphiclus/iphicleola is notably smooth, this observation may be of taxonomic interest, or it may be simply referring to the usual, sparse, small setae which surround the chalazae and sides of the head capsule. There are several points of similarity between Mihier and Turner's notes and those of Swainson, which suggest that the morphology of the larval scoli and pupa may be rather different in this species compared to A. iphiclus/iphicleola (figured by Aiello, 1984). Curiously, both state that scoli begin on the first two thoracic segments, but since no dorsal or subdorsal scoli are known on Tl in any other Adelpha species, I assume both incorrectly numbered the segments and regarded T2 as the first segment. Making this assumption, both state that there are pairs of anteriorly curving subdorsal scoli on T2 and T3, with that on T3 the longer, a pair of short scoli on Al, and pairs of larger, posteriorly curving scoli on A2, A8 and AlO, the latter two described by Swainson as "branching stellate spines". Both descriptions also mention a series of subspiracular scoli, but any references to other subdorsal scoli are conspicuously absent. Since the subdorsal scoli of A. iphiclus/iphicleola are well developed on all abdominal segments from A2-A8, the implicit reduction of scoli on A3A7 in A. abyla is of potential taxonomic significance. Milner and Turner's notes describe the pupa as being brown with reddish coloration and golden dorsal spots, having the dorsal projection on segment A2 curving anteriorly to meet the projection of T2, while Swainson states "the shape is very curious, resembling the pictures of "Punch", long nose and all". Both of these descriptions imply that the projection on A2 is much more produced anteriorly in A. abyla than in A. iphiclus/iphicleola (see Aiello, 1984), while Milner and Turner state that the head horns are short, pointed projections, as in A. iphiclus/iphicleola.

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298 Habitat and adult ecology: Riley (1975) reports that this species is uncommon but widespread in foothills and mountains in open, sunny forest glades. Brown and Heineman (1972) quote Walker saying that the species, presumably males from the description of the behavior, usually perches 4.5-6m above the ground, confirmed by Smith et al. (1994), who state that favored perches are the tips of twigs where individuals typically rest with the wings open. Specimens examined: 61 (33Tn, 28f) JAMAICA (33m, 27f): Baron Hill 1200' Jun.-Aug. 3m, 3f AME, Im BMB; Bath Jul. Im AMNH; Blue Mts. 1200' Jul. Im BMNH; Coleville If AMNH; Crownland Jan. Im, If AMNH; Cuna Cuna Jul. If USNM; from a box of Jamaican things Im MCZ; Halfway Tree Jul. If AMNH; Holly Mount Aug. Im AME; Jacks Hill Im AMNH; Jackson Town? Jul. If AME; Manchester Jan. Feb. 2m, If MUSM; Montego Bay Im AMNH; Moore Town 3m, 3f BMNH, Im BMB; Mt. Diablo Apr. Aug. Im, If AMNH, Im, If AME; Oracabessa Oct. Im AME; Orangefield Feb. Im AMNH; Rio Bueno Feb. Im AMNH; Rochmore Mar. Im AMNH; St. Thomas Jan. If AMNH; Stony Hill Oct. Nov. 2m BMNH, Im, If BMB; Wilson Run Feb. If AMNH; no specific locality 5m, 6f BMNH, Im AMNH, If ZMHU, Im, If BMB, 2f USNM. COUNTRY UNKNOWN (If): no specific locality If BMNH. Additional locality data: JAMAICA: Pantrepant, Spring, Warsop, Islington, Rockmore, Ecclesdown, Com Puss Gap, John Crow Hill, Hermitage, Kingston, Long Mountain, Pollyground, Rock Hall, Christiana, Cave River, Accompong (Brown and Heineman, 1972). Adelpha melona (Hewitson, 1847) Figs. 41; 125; 207 Identification, taxonomy and variation: This species is recognised on the DFW by the broad orange postdiscal band and lack of any subapical spots representing the postdiscal series, by the orange filling the area in the VHW discal cell between the second and third cell bars which continues as a band to the vein Sc+Rl, and, in most subspecies, by having orange along the anal margin just basal of the DHW tomus. A unique character visible in all subspecies except some specimens of the nominate and A. m. pseudarete is a curving, thin red-brown line distal of the pale postdiscal band in the VFW subapical area. This red-brown line seems to be composed of the inner submarginal series in cell M2, the space between the outer postdiscal and inner submarginal series in cell Ml, and possibly the outer postdiscal series in cell R5. The male genitalia are unique within the genus in having the dorsal tip of the gnathos extended to form a long anterior arm. There is variation in the presence or absence of a white DHW postdiscal band, the amount of orange or white in the DFW

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299 postdiscal band and the extent to which this band is constricted at vein M3, and the expression of the postdiscal and submarginal series on the ventral surface, and five subspecies are recognised. Adelpha melona, as treated here, embraces three taxa typically considered as distinct species, the nominate subspecies, A. melona leucocoma and A. melona deborah. All however share the diagnostic characters discussed above, particularly the unique male genitalia. The wing pattern of A. melona is highly modified and its relationships within the genus are difficult to ascertain. However, several characteres point to it being most closely related to A. ethelda and A. epione. The valvae of the male genitalia, which have dorsally pointing spines near the posterior tip and a relatively long clunicula, are similar to those of both these species. All three of these species also share, on the VFW, the fusion of the postdiscal series and the postdiscal band and the loss of the submarginal series in cell M3, the entire loss or reduction of the submarginal series in cells Ml and R5, the second and fourth discal cell bars converging to touch posteriorly, and on the VHW, the replacement in at least some subspecies of the inner and outer postdiscal series with a red-brown line, the merging in the discal cell of the of the first and second cell bars, and the third and postcellular bar, and orange scaling filling the area between the second and third discal cell bars forming a band extending to vein Sc+Rl. This consensus of characters suggests the placement adopted here, although the relationship oiA. melona, A. ethelda and A. epione with respect to members of the A. iphiclus group is not clear. Range and status: Central Panama to northwestern Ecuador and northwestern Venezuela, to Bolivia, Brazil, Paraguay, northeastern Argentina, the Guianas and Trinidad. Not uncommon to very rare, in lowland forest up to 1200m. Specimens examined: 271 (182m, 89f)

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300 Adelpha melona melona (Hewitson, 1847) Figs. 41a,b Heterochroa melona Hewitson (1847: 258, pi. XX, fig. 2) TL: Colombia. Types: BMNH(M): STm; [not identified]; BMNH(T): false STf: "Heterochroa melona Hewiteon f Syntype det. R.I. Vane-Wright 1983//Illustrated in The Butterflies of Venezuela A. Neild, 1996//162//B.M. TYPE No. Rh. 9843 Heterochroa melona f Hew.//melona Hew7/Para//Type//Syntype" = melona leucocoma [examined] =Heterochroa arete Menetries (1857: 118) syn. nov. TL: Bahia. Types: ZISP: ST [not examined] =cibyra Fruhstorfer (1915: 529) stat. nov. TL: locality stated to be unknown. Types: MHNG: HTf: [not examined] Adelpha melona Hew. , Kirby (\il\) = A. melona leucocoma; Adelpha arete M£n., Kiiby (1871) Identification, taxonomy and variation: This subspecies is distinguished from A. m. pseudarete by having the DFW postdiscal band orange to the anal margin. Males are distinguished from A. m. leucocoma by the reduction of the ventral postdiscal and submarginal series and the loss or reduction of the pale postdiscal spot in cell M3 distal of the postdiscal band, in addition to several characters discussed below. Females of this subspecies have a broad white submarginal series on the VHW, but still lack the whitish submarginal series in the VFW tomus and the reddish brovra postdiscal line in cells Cul and M2, in addition to having reduced postdiscal series on the VHW. Hewitson (1847) described this species based on an unspecified number of specimens which he stated were in the British Museum, from Colombia, and figured the dorsal surface of the left hand wing pair on plate 20, fig. 2. hi the BMNH(T) there is a putative female syntype, labeled as from Para (Brazil), and all subsequent authors have regarded the name as applying to the Amazonian subspecies of A. melona. However, a careful reading of Hewitson's description, coupled with an examination of his original black and white figure, leads me to believe that this is incorrect for a number of reasons. Hewitson's descriptions of other species in the same paper (Hewitson, 1847) are typically very detailed and accurate, as are the figures, since they generally correspond very closely with the syntype specimens in the BMNH. Even were the BMNH syntype labeled with the same locality as that given in the original description, I would still not believe it to be

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301 the specimen illustrated by Hewitson, since the illustrated specimen is clearly a male from the shape of the hindwing, which is much more triangular, has a narrow, elongate tomus and a relatively straight distal margin, and has a narrower and more evenly edged DFW orange band, which has a smoothly curved distal edge in cells M2-R5, is only slightly indented at the basal edge at vein M3, and has no white at the anal margin. The only Colombian specimen of A. melona in the BMNH is from Florida, on the Rio Putumayo, and would not have been in the collection at the time of the description. Since A. melona is otherwise very rare in Colombia, it seems highly unlikely that Hewitson's specimen originated there, and much more plausible that it was either mislabeled, or Hewitson gave an erroneous locality. In fact, the shape and color of the DFW band and the shape of the DHW band in Hewitson's figure correspond exactly to specimens from southeastern Brazil, the arete of authors. Hewitson's description of the ventral surface also fits much more closely the southeast Brazilian population, in the following respects: 1. The underside is described as "vinous brown", a perfect description, while Amazonian specimens have only a dark brown ground color which is largely obscured by paler markings; 2. the postdiscal band on the VFW is said to have the "widest portion ... clouded with brown", and this fits the VFW postdiscal band which widens continuously from anal margin to costa and has brown shading in cells M2 and Ml, whereas in Amazonian specimens the band is of varying widths throughout and less noticeably obscured by brown scaling; 3. there are said to be four to five white dots in the VFW apex, and three narrow, white lunules in the VHW tomus, as in southeast Brazilian specimens, but there is no mention of the orange-brown submarginal line on the VFW or the prominent white submarginal markings in the tomus, or the broad white submarginal series and the two orange-brown postdiscal lines on the VHW, such as occurs in Amazonian specimens; 4. the discal cell of the VFW is said to have a "lilac" spot, corresponding to the area between cell bars two and four, whereas this area is pure white in Amazonian specimens. The only part of the description that does not fit typical

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302 specimens of the southeastern Brazil population is the mention of three blue spots distal of the VFW discocellulars; these are typically present in Amazonian melona, but usually absent or reduced in southeast Brazilian melona, though they are present in a single specimen in the BMNH. In conclusion, the weight of evidence points to Hewitson's specimen having come from the southeast Brazilian population. In the BMNH(M) there are four male specimens of this subspecies which lack locality data, several of which appear to be old specimens, and it seems very plausible that at least one of these is a syntype oi melona and the label has either been lost or Hewitson guessed what he thought was a probable locality. Notwithstanding the above discussion, the situation is fiirther complicated by Hewitson (1867: fig. 2) again figuring melona, but this time the ventral surface, and this is clearly a specimen of the Amazonian population. He figured a specimen of the southeast Brazilian poplation under the name arete beside it, and stated that melona was distinguished by having the orange band "more sinuated on its inner border, more so usually than is represented in my former figure". I still believe that Hewitson's original description was based on a specimen of the southeast Brazilian population, which was after all in the British Museum rather than his own collection, and perhaps later obtained Amazonian specimens which he identified as melona, causing him to make an error when he refigured the taxon 20 years after the original description. Menetries (1857) described Heterochroa arete based on an unspecified number of specimens from Bahia, and although I have not seen any syntype specimens, which should be in the ZISP, the description is sufficiently detailed to recognise the taxon described. Menetries mentions the orange DFW band being somewhat as in Adelpha nea nea, the venfral surface being of a violaceous color, accurately described the postdiscal band and more basal white band, states that the VFW cell pattern is as in A. melona, and makes no mention of ventral white submarginal markings. He distinguished it from A. melona by lacking the blue spots distal to the VFW discocellulars, which I regard as individual variation, and lacking the white "lunules" in the VHW tomus, although he

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303 appears to have just misunderstood Hewitson's use of the word. Fruhstorfer (1920: pi. 1 lOAb) correctly figured typical arete as having almost no white in the DFW postdiscal band, since Menetries stated that the white band "sur les superieures se perd immediatement en se fondant dans le bande fauve". The three male specimens that I have examined of this subspecies from Bahia are identical to Hewitson's figure of melona on the dorsal surface, and so I place arete as a synonym of melona (syn. nov.). Fruhstorfer (1915) described a subspecies of arete, cibyra, based on a single female in the Geneva museum, which lacked any trace of white on the DFW. Since, however, this description corresponds exactly to arete. Hall (1938) placed it as a synonym of that name, and I further place it as a synonym of melona (stat. nov.). Range: This subspecies is only known fi-om three localities, Bahia, Itaparica and Rio, with all remaining specimens lacking any acciu-ate data. The latter in particular is a general locality for specimens fi-om all over southeastern Brazil. It is therefore difficult to say what the true range of this phenotype is, and whether or not it is sympatric with A. m. pseudarete. Habitat and adult ecology: This subspecies appears to be very rare and no field observations have been published. Specimens examined (1 3m, 5f): t trans, to A. m. pseudarete. BRAZIL (7m, 4f): Bahia: Bahia 3m BMNH; Itaparica Dec. If BMNH; Guanabara: Rio 2m, Bfljlf) BMNH; Not located: no specific locality 2m BMB. COUNTRY UNKNOWN (6m, If): no specific locality 4m, If BMNH, Im USNM, Im MNHN. Adelpha melona pseudarete Fruhstorfer, 1915 stat. nov. Figs. 41c,d Adelpha arete pseudarete Fruhstorfer (1915: 529) TL: locality stated to be unknown. Types: BMNH(T): STm: "Fruhstorfer Coll. B.M. 1 937-285//arete pseudarete Fruhst.//Fruhstorfer//TYPE//Type//Syntype" [examined] =Adelpha arete Men., Hall (1938) form Identification, taxonomy and variation: This subspecies differs from the nominate in having the postdiscal band on the DFW white in cell Cu2, and often in cell Cul. In the case of the latter, the orange in cell

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304 Cul may extend, gradually tapering, almost to the anal margin, otherwise it terminates at vein Cu2. Specimens from further west, especially Paraguay, usually have the orange terminating at vein Cu2. Fruhstorfer (1915) described this subspecies as differing from arete in having the white on the DFW extend to vein Cul, and this closely matches the single Fruhstorfer specimen in the BMNH which is a syntype. Hall (1938) regarded this as representing no more than a form of A. m. melona, and there is much evidence that this is the case. I have seen one female from Bahia which is intermediate to A. m. melona, and another male, from Morro do Chapeu, which is more typical A. m. pseudarete. The ranges of both phenotypes, as currently known, seem to widely overlap, clearly inconsistent with the subspecies concept. I retain pseudarete as a distinct subspecies (stat. nov.) since I have examined so few specimens in collections with accurate data that is is impossible to say whether or not it is sympatric with A. melona melona. It is also probable that the two phenotypes are maintained through mimicry, and it is therefore of potential interest that A. melona melona differs from the typical southeast Brazilian mimicry complex, perhaps indicating that it does not occur with typical members. Finally, since the few specimens I have examined from further west, such as Sao Paulo and Paraguay, seem to be phenotypically relatively stable, it seems possible that the two forms really do represent valid subspecies. I hope that by retaining them here it will provide more information and encourage workers in the field to seek to determine accurately their ranges and so test the validity of this taxonomic arrangement. Range: The range of this subspecies is difficult to determine, but it has been reported from inland areas in Bahia throughout the Atlantic coastal area to Sao Paulo, in Minas Gerais, and across to Paraguay and northeastern Argentina. More information is needed on its distribution with respect to A. melona melona. Habitat and adult ecology: Nothing has been reported on the biology of this rare subspecies, which has been captured up to 1200m.

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305 Specimens examined (14m, 60: t trans, to ,4. m. melona. BRAZIL (12m, 40: Bahia: Bahra 1ft BMNH; Morro do Chap6u 1200m Apr. Im USNM; Espiriio Santo: Linhares Aug. 2m, If AME; no specific locality 2m BMNH, Im MCZ; Guanabara: Rio Janeiro If MNHN; Minas Cerais: Leopoldina Im ZMHU; Para: Etat de Para Im BMNH-error; Sao Paulo: Borhumi If BMNH; Sao Paulo Im BMNH; Not healed: no specific locality Im BMNH, Im ZMHU, Im MNHN. PARAGUAY (Im, 20: Paraguari: Sapucay If BMNH; Not located: Central Paraguay Im BMNH; no specific locality If BMNH. COUNTRY UNKNOWN (2m, 20: no specific locality 2m, 2f BMNH. Additional locality daU: ARGENTINA: Misiones (Hayward, 1951). Adelpha melona leucocoma Fruhstorfer, 1915 stat. rest. Figs. 41e,f; 125a,b; 207a-c Adelpha melona leucocoma Fruhstorfer (1915: 529) TL: Upper Amazon. Types: BMNH(T): STm: "Fruhstorfer Coll. B.M. 1937-285//melona leucocoma Fruhst.//Fruhstorfer//TYPE//Type//Syntype//Published as leucocoma Fruhst. (S.T.)"; STf: "Fruhstorfer Coll. B.M. 1937285//leucocoma Fruhst./ZAmaz. S. Fruhstorfer//TYPE//Type"; BMNH(M): STm: "TYPE//Peru H. Fruhstorfer//no indication of what this is a type of?//apparently a paratype of leucocoma (G.T.)" [all examined] =Adelpha melona meridionalis Fruhstorfer (1915: 529) syn. nov. TL: Santa Catharina. Types: BMNH(R): HTf: "melona meridionalis Fruhst./ZSanta Catharina//TYPE//Syntype" (examined] =Adelpha biedermanni Fruhstorfer (1915: 530) syn. nov. TL: not stated. Types: BMNH(M): STm: "Joicey Bequest Brit. Mus. 1934-120//Syntype//61. 28/^iedermanni m Fruhst. Amazonas" [examined] =Adelpha melona nonsecta Kaye (1925: 413) syn. nov. TL: Trinidad. Types: AME: STf: "Type H.T.//Trinidad R. Dick//Adelpha melona Hew. 1847 nonsecta Kaye 1925"; ST?f: 'Trinidad, 1898, May, N.J. Kaye// Adelpha melona Hew." [both examined] Adelpha melona Hew., Kirby (1871), D'Abrera (1987); Adelpha melona melona Hew., Fruhstorfer (1915), Hall (1933) misid.; Adelpha melona ihesprolia Fldr., Fruhstorfer (1915) misid.; Limenitis melona meridionalis Fruhst., Brown and Mielke (1967) Identification, taxonomy and variation: This subspecies is distinguished from the nominate and A. melona neildi under those taxa. It differs from A. m. pseudarete by the same venfral characters as it differs from the nominate. This subspecies is very similar on the dorsal surface to A. nea, but easily distinguished by the VFW discal cell pattern, which is has the white area between the second and fouth cell bars divided into three by a black line in A. nea. There is variation in this subspecies in the width of the orange DFW postdiscal band, the extent to which it is constricted at vein M3, in some specimens being almost completely broken, the amount of white in the postdiscal band at the anal margin, and whether or not the pale spot distal to the postdiscal band in cell Cul on the VFW is isolated from the band or not. This taxon has historically been regarded as the nominate subspecies, but for the reasons outlined above under A. melona melona, I do not believe this to be the case. Fruhstorfer (1915) described A. melona leucocoma, and later figured it (Fruhstorfer, 1920; pi. llOAa, missp. as leucosoma), based on specimens from the Upper Amazon,

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306 which had white at the DFW anal margin, broader white DHW postdiscal bands and a paler ventral surface. The figure shows a probably slightly melanic specimen with the orange DFW band completely broken. Hall (1933) was right to regard there as being no distinct populations of A. melona within the Amazon, but since his synonymy of leucocoma with melona was based on a misidentification of melona, I reinstate leucocoma as a subspecies of melona (stat. rest.). Fruhstorfer (1915) described a second subspecies a few lines later, as A. melona meridionalis, and later figured it (Fruhstorfer, 1920: pi. llOAa), based on a single female labeled "Santa Catharina" in the Tring Museum. The holotype is in the BMNH(R) and does not differ fi-om typical females of this subspecies, and the locality, as Fruhstorfer suspected, is clearly erroneous. Given the latter fact, I prefer to use the name leucocoma to refer to the Amazonian and Guianan population of ^. melona, and so synomyise meridonalis with leucocoma (syn. nov.). Without giving any indication of the number or sex of specimens, where they were collected or in which collection they were deposited, Fruhstorfer (1915) described a species, A. biedermanni, which closely matches a single specimen in the BMNH labeled with Fruhstorfer' s handwritten label "biedermanni m Fruhst. Amazonas". This specimen was figured by D'Abrera (1987), and it is clearly a melanic aberration, and therefore a synonym of, A. melona leucocoma (syn. nov.). Notable characters which identify it as A. melona include very close discal cell bars on the VHW with orange filling the space between cell bars three and four, the shape and placement of the VFW discal cell bars, with a broad, gradually tapering area between cell bars two and four, and the orange shading at the DHW anal margin near the tomus. Kaye (1925) described A. melona nonsecta based on an imspecified number of Trinidadian specimens, which are now in the AME, on the basis that the DFW orange band was less constricted at vein M3. This however is a variable character which typically occurs also in lower Amazon specimens, and I therefore regard it as not sufficiently

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307 distinct or consistent to warrant subspecific recognition, and synonymise nonsecta with leucocoma (syn. nov.). This subspecies has typically been considered to represent a distinct species from the nominate, but I treat them as conspecific since females of the nominate show several of the ventral characters of this subspecies, and the genitalia are identical. Range: This subspecies occurs east of the Andes from Venezuela to Bolivia, throughout Amazonian Brazil as fas southeast as the Brazilian plateau, and in the Guianas and Trinidad. Immature stages: Moss (1933) reared this species in Brazil (Para), and figured both the last instar larva (pi. I, fig. 6, 7) and the pupa (pi. H, fig. 7). Early instars are typical of the genus in appearance and habits, and later become greenish with very dense scoli, very closely resembling a fragment of moss. The scoli are relatively long with lateral spines on all segments except Al, though they are longest on T2, T3, A2, A7 and A8. The pupa is dull maroon and very distinctive in the dorsal projection on segment T2 terminating in a sharp, triangular point, being extended to touch the projection from segment A2, which is more typical in size. The head horns are leaf-shaped, elongated at the tip and sharply curved posteriorly. The larval foodplant is a creeper on tree trunks, and the larva feed within 2m of the ground. Habitat and adult ecology: This subspecies is not uncommon in relatively intact lowland rain forest habitats up to 1200m, where it flies throughout the year. I have frequently found males in light gaps caused by logging in primary forest, almost always attracted to rotting fish in traps, though they also may be found feeding on sap from freshly cut trees, and on rotting banana. Females are rarer, and generally are encountered in the afternoon in large light gaps with secondary growth. I have never observed the species perching, suggesting it may be a canopy species. Specimens examined (127m, 74f): VENEZUELA (2m, 2f): Amazonas: Gavilan, 35 km. S.E. Puerto Ayacucho 100m Mar. 2m AFEN; Bolivar. El Dorado-Santa Elena, km. 107, 520m Aug. If MUSM; Not located: San Ignacio de Yuruani Jun. 950m AFEN; no specific locality If BMNh! COLOMBIA (4m): Amazonas: Florida Im BMNH; Uticia Oct. Im LMC; Puerto Nariilo Dec. Im LMC; Not located: no specific locality Im BMB. ECUADOR (11m, 2f): Napo: Apuya 600m Aug. Im KWJH; Chichicorrumi 450m Sep. Im KWJH; Finca San

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308 Carlo 600m Feb. Oct. 2m DAT, If KWJH; Pimpilala 600m Feb. Im KWJH; Rio Napo Im BMNH; Santa Rosa Apr. If MJP; Sinde Oct. Im MJP; Pastaza: Puyo 2m JFL; Sucumbios: Garzacocha, La Selva, Rio Napo 250m If Nov. PJD; Limoncocha Feb. Im USNM; Not located: no specific locality Im USNM. PERU (45m, 60: Cuzco. Cosilipata Valley Im BMNH; Huanuco: Cord, del Sira 800m Aug. Sep. Im MUSM; Junin: Satipo 2m AMNH; Loreto: Balsapuerto 220m May Im MUSM; Cocha Shinguito, Rio Samiria 130m Jun. Im MUSM; Rio Sucusari 140m Sep. Im MUSM; Iquitos Mar. Apr. 2m, If BMNH, Im, If ZMHU, 2m, If AME, Im USNM, Im AMNH; Iquitos-Nauta, km. 15, 120m Sep. 2m MUSM; Naranjal, km. 82 Tarapoto-Yurimaguas Im, If MUSM; Pebas Nov. 2m BMNH, Im ZMHU; Quebrada Shihua, Rio Momon 120m Sep. Im MUSM; San Roque Im AME; Yurimaguas Im ZMHU, Im BMB; Madre de Dios: Boca Rio La Torre 300m Oct. Im MUSM; Puna. Chaquimayo 2500-3500' Aug.-Oct. Im BMNH; Yahuarmayo 1200' Feb. Mar. Im BMNH; San Martin: Achinamiza Dec. Im AMNH; Jepelacio 7m, If AMNH; Rio Negro, nr. Rioja Sep. Im MUSM; Not located: Rio Huallaga Im USNM, 4m AMNH; Rio Ucayali Im BMNH; no specific locality 2m BMNH, Im USNM, If BMB. BOLIVIA (8m): Cochabamba: Cristal, Chapari May 600m Im MUSM; La Paz: Caranavi 1200m Feb. Im MUSM; U Paz 1000m Im BMNH; Rio Songo 2m ZMHU; Santa Cruz: Prov. del Sara Im BMNH; Rio Juntas 300m Im ZMHU; Not located: no specific locality Im MCZ. BRAZIL (29m, 32f): Amazonas: Manaus Jan. -Mar. 3m BMB; Manicori Im ZMHU; Sao Paulo de 01iven(;a Im BMNH; Maranhao: "Maranham" If BMNH; Para: Itaituba, 15 km. S., Im USNM; Obidos Apr. Im, If BMNH, Im BMB; Para 1 6m, 25f BMNH, Im MNHN; Santarem Oct. Im, If BMNH, If ZMHU, Im AME; "Santa Catharina: Santa Catharina" If BMNH-error; Not located: S. Brazil If BMB; no specific locality 2m ZMHU, If BMB. GUYANA (Im, 7f): Cuyuni/Mazaruni: Bartica Dec. Im BMNH, If BMB; Carimang R. If BMNH; Potaro/Siparuni: Potaro River Oct. Nov. If AME; Not located: no specific locality 4f BMNH. FRENCH GUIANA (8m, 1 10: Cayenne: Camp St. Elie, pk. 15-5 on D21 Nov. If USNM; Cayenne Im BMNH, Im ZMHU, Im, 2f USNM; Matoury Dec. If MUSM; Laurent du Maroni: La Mana Im MNHN; Maroni River 2f AME; St. Jean du Maroni Im, If BMNH; Not located: Rochambeau Mar. Im FSCA; no specific locality 2m, 2f BMNH, If USNM, If BMB. SURINAM (6m, 30: no specific locality 6m, 3f BMNH. TRINIDAD (3m, 30: St. George West: Hololo Jan. Im AME; St. Anns Oct. -Dec. 2f BMB; Not located: Northern Mts. Dec. Jan. Im BMB; no specific locality 1000' Feb. Im, If AME. COUNTRY UNKNOWN (1 Im, 70: Amazonas Im BMNH; Amazon 3f BMNH, 2m USNM, 2m BMB, Im MCZ; Amaz. S. Im BMNH; lower Amazon Im MCZ; Tarinas 2f BMNH; U. Amazon Im BMB; no specific locality If BMNH, If ZMHU, Im USNM ImBMB. Additional locality daU: ECUADOR: Morona-Santiago: Santiago Sep.; Napo: Apuya Oct Dec.; Chichicorrumi Feb.; El Capricho Oct.; Finca San Carlo Aug. Sep. Dec.; Pimpilala Aug.; Rio Tiputini Jun.; Satzayacu Sep.; Yasuni Jul.; Sucumbios: Pafiacocha Oct. (Willmott and Hall, unpubl. sight records). BRAZIL: Goias: Campinas Jan. (Brown and Mielke, 1967). TRINIDAD: Not located: Las Lappas Trace 2000' Sep.; Mome Catherina Mar. (Cock, pers. comm.). Adelpha melona neildi Willmott, ssp. nov. Figs. 41g,h Description and diagnosis: FW length of HT: 30 mm. This subspecies is distinguished from A. melona leucocoma by having a broader orange DFW postdiscal band that is typically less indented at vein M3 at the basal edge, by typically having a narrower white DHW postdiscal band, increased orange scaling separating the pale postdiscal spot in cell Cul on the VFW from the postdiscal band, and by having the inner postdiscal series on the VHW marked as a red-brovra line, instead of silvery gray. All of these characters are variable to some extent. It is distinguished from A. melona deborah in having a white DHW postdiscal band. Types: Holotype male: PANAMA: Canal Zone: Colon (Sta. Rita), 1500', 12 Jan. 1972 (S.S. Nicolay); in the USNM. Allotype female: Distrito de Panama, Cerro Jefe, 900m, 1517-m-78 (Gordon B. Small); in the USNM.

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309 Paratypes: PANAMA: Canal Zone: 1 female: nr. Colon, 1000', March 1969, G.B. Small; in the USNM; 1 male: Summit, Panama, C.Z. V-2-64; in the USNM; 1 male: Summit, Panama, C.Z. III-22-64; in the USNM; Panama: 1 female: same data as AT; in the USNM; 1 female: same data as AT except IV-20-78; in the USNM. Etymology: I name this subspecies for my good friend Andrew Neild, who has willingly provided an enormous amount of help in photographing specimens and recording data from Venezuelan collections, been always eager to enter into long discussions on taxonomic problems, and whose work on Adelpha (Neild, 1996) represents the most careful and insightful taxonomic work on the genus to date. Taxonomy and variation: This subspecies is somwhat variable in the width of the white DHW postdiscal band, which may be ahnost obsolete, and the extent to which the basal edge of the orange DFW band is incised at vein M3. It is superficially very similar to A. m. leucocoma, but the shape of the orange DFW band and the generally darker ventral surface, in addition to its closer geographic proximity to A. m. deborah, which isolates it from A. m. leucocoma, show it to be most closely related to A. m. deborah. The variation in the white DHW postdiscal band may well indicate some intergradation to A. m. deborah, which perhaps occurs in the Darien. Range: All the specimens of this subspecies that have been collected to date are from cenfral Panama, and it presumably occurs further west, quite possibly into Costa Rica, though it was unrecorded by DeVries (1987). Habitat and adult ecology: This subspecies is clearly very rare and all records are from lowland rain forest below 500m, from January to August. Orellana (pers. comm.) reports to me that he encountered two specimens feeding on fermenting fruits hanging on a Bunchosia cornifolia (Malpighiaceae) tree in August. Specimens examined (3m, 4f): PANAMA (3m, 40: Canal Zone: Colon (Sta. Rita) 1500' Jan. Im USNM; nr. Colon 1000' Mar. If USNM; Summit CX Mar May 2m USNM; Panama: Cerro Jefe 900m Mar. Apr. 3f USNM.

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310 Adelpha melona deborah Weeks, 1901 stat. rest. Figs. 41 i J Adelpha deborah Weeks (1901: 353) TL: Colombia. Types; MCZ: STm: "Strecker can't May '99//Syntypus//Adelpha deborah//767 Slide no. M.C.Z7/Type//ln British Museum from Bogota unnamed/AThis specimen photographed for G. Lamas by A. Aiello Jan. 1981"; BMNH(T): STm: "Illustrated in The Butterflies of Venezuela A. Neild, 1996//10//[Bogota] 98-234 ftxim H. Ellis Bolivia/ZType" [examined] Adelpha melona deborah Weeks, Lamas and Small (1992); Adelpha deborah Weeks, Neild (1996) Identification, taxonomy and variation: This subspecies is distinguished from all others by the lack of a white postdiscal band on the DHW. There is slight individual variation in the width of the orange DFW band. Weeks (1901) described this taxon in a paper entitled "Description of nine new Bolivian butterflies", but the habitat is listed as "Colombia", and the only information on the specimen(s) on which the description was based is a comment that a single specimen "of this has been found in the British Museum collection without a name". This is the BMNH syntype, though the MCZ specimen, from Weeks collection, is presumably that on which the description was founded, since Weeks (1911: pi. 34) later figured a specimen which corresponds exactly in wing pattern to this specimen. Most subsequent authors have regarded the taxon as a distinct species (Fruhstorfer, 1915; Hall, 1938; D'Abrera, 1987), but Lamas and Small (1992) placed it as a subspecies of A. melona, since the variation in the DHW postdiscal band in A. melona neildi, from Panama, shows the presence or absence of this band to be a relatively weak character. Furthermore, the genitalia do not differ from typical A. melona and the taxon is clearly parapatric with respect to A. melona neildi and A. melona leucocoma. Neild (1996), probably unaware of the note by Lamas and Small (1992), accorded it specific status, with reservations, and I place it once more as a subspecies of A. melona (stat. rest). Range: This subspecies occurs west of the Cordillera de Merida in Venezuela to northwestern Ecuador, west of the Andes. The variation in the white DHW postdiscal band in A. m. neildi suggests that A. m. deborah may also occur in eastern Panama, and

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311 intergrades to A. m. neildi in central Panama. The single record from "Palo Alto, Rio Tigre" in Venezuela is probably an error, since this locality appears to be in northeastern Venezuela with the range of A. melona leucocoma. Habitat and adult ecology: This subspecies is rare throughout its range, and occurs in wet lowland rain forest up to 800m. I have only encountered a single male, which was attracted to a trap 5m above the ground baited with rotting fish in bright sun in a deep river canyon lined with forest, at around 11am. After circling the trap several times the specimen landed on my hand and began to feed on sweat. This record was at the end of an unusally severe wet season, and since it was accompanied by the discovery of several other rare Adelpha species, it suggests that the subspecies may very well be seasonal. Specimens examined (25m): VENEZUELA (2m): Trujillo: La Gira Betijoque 570m Dec. Im AFEN; Not located: Palo Alto, Rio Tigre Oct. Im USNM-error. COLOMBIA (20m): Boyaca: Muzo Im BMNH; Otanche Feb. Apr. Aug. 4m JFL, Im ESM; Caldas: Rio de la Miel Mar. If ESM; Cundinamarca: Bogota 2m BMNH; Bogota-Bolivia Im BMNH; env. de Bogota 3m BMNH; Santander: El Centro If AMNH; La Soledad Dec. Im AMNH; Tolima: Rio Chili Apr.-Jun. 2m BMB; Valle del Cauca: Zacarias, Rio Dagua 70m Mar. Im, If LMC; Not located: no specific locality 2m BMNH, Im ZMHU, Im AMNH. ECUADOR (Im): Esmeraldas: Rio Piguambi 8OO111 Jul. Im KWJH. "PERU" (Im): no specific locality Im BMNH-error. COUNTRY UNKNOWN (Im): no specific locality Im MCZ. Adelpha ethelda (Hewitson, 1867) Figs. 42; 126; 208 Identification, taxonomy and variation: This species can be distinguished from A. epione by the less rounded hindwing which has a more angled apex, a broader tomus and a more dentate distal margin. The majority of subspecies also have, on the VHW, the distal margin of the postdiscal band slightly concave, a complete row of silvery gray submarginal spots, a more complete redbrown outer postdiscal series, and an orange DFW band. There is variation in the color of the DFW band, its width and orientation, the presence on the DHW of an orange band, and the expression of the VHW postdiscal band and distal series, and six subspecies are recognised. The systematic status of the taxon treated here as A. ethelda is open to debate, since it appears to be paraphyletic with respect to A. epione. Both A. epione and A.

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312 ethelda share many distinctive local and universal synapomorphies, including: the complete loss of the postdiscal series anterior of vein Cul on the forewing (the postdiscal series and postdiscal band could also be fused, but an examination of A. melona, probably closely related to these species, suggests a loss is more likely), the entire loss of the outer submarginal series on the ventral surface on both wings and its replacement with a redbrown border, the replacement of the inner and outer postdiscal series on the VHW by red-brown bands, and the VHW postdiscal band being displaced basally to border the third discal cell bar, while the first and second appear to be fused to form a single dark line. The fusion of many of the principal character elements on the VFW and otherwise highly modified wing pattern makes it difficult to place these two species within the genus. However, the male genitalia, which are indistinguishable, have dorsally pointing spines at the distal tip of the valvae and a relatively elongate clunicula, strongly resembling those of A. melona, A. iphiclus and close relatives. Sparse orange-brown scaling at the base of the DFW, and the expression of the outer postdiscal series on the DHW, are two characters usually found in primitive Adelpha, where I have preliminarily placed these taxa for the present. DeVries (1987) states that the immature stages are similar to A. cytherea, also consistent with a relatively primitive placement in the genus. Since I am imable to isolate with certainty the sister species of A. ethelda/epione, although it may well be A. melona (see under that species), an analysis of the relationships of their constituent taxa is more complicated. The reduced white VHW postdiscal band occurs only in galbao, epione and agilla, as does the extensive orange scaling in the cell posterior to vein 3A on the VHW. Since these characters do not occur in A. melona or any A. iphiclus group members, or other relatively primitive Adelpha, I regard them as synapomorphies. The taxa epione and agilla, treated here as A. epione, both share a rounded hindwing shape and white DFW band, both of which appear to be local synapomorphies uniting them as sister taxa. There are thus several possible ways of classifying the taxa of A. ethelda and A. epione, depending on taxonomic philosophy.

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313 Adopting an evolutionary species concept, since A. epione and A. ethelda form a monophyletic group, and excluding A. epione would make the resulting group paraphyletic, the taxa should be regarded as forming a single species. My reasons for not following this approach concern the close proximity of A. epione agilla and A. ethelda zalmona in Colombia and the complete lack of evidence that these taxa intergrade, or that there is, or has been for a substantial period of time, any gene flow between them. Since areas of the Cordillera Oriental extend below 1500m, these taxa, which both occur above 1500m elsewhere in their range, might be expected to cross from one slope to the next. This seems to happen in A. lycorias, which has a similar altitudinal range, and intermediate specimens between the east Andean A. lycorias lara and A. lycorias melanippe are known to occur. To date there are unsufficient locality data from Colombia to determine whether these A. epione agilla and A. ethelda zalmona are sympatric, but a consensus suggests that they are not, A. epione agilla occurring on the eastern slopes and A. ethelda zalmona on the west. Adelpha epione agilla is more closely related to A. epione epione from southeastern Brazil, and then to A. ethelda galbao, than to A. ethelda zalmona, from which it is separated by only a few tens of kilomefres. Adopting a biological species concept, 1 regard it as highly unlikely that A. epione agilla and A. ethelda zalmona would be able to, or do, freely and successfiilly interbreed under natural conditions, and therefore I prefer to regard them as distinct species for the present. The systematic position of galbao is purely arbitrary, but since there is no evidence that A. ethelda is monophyletic with its exclusion, and given its closer overall similarity to A. ethelda in the more visually obvious characters of wing shape and pattern, I prefer to consider it as a subspecies of A. ethelda for the present. Range and status: Mexico (Chiapas) to western Ecuador, throughout Colombia west of the Cordillera Oriental to to northwestern Venezuela on the western slopes of the Cordillera de Merida, with a single disjunct subspecies in the Guianan highlands from

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314 Venezuela to French Guiana. Males not uncommon in lower montane rain forest and cloud forest habitats from 100-2000m, females very rare. Specimens examined: 261 (254m, 7f) Adelpha ethelda ethelda (Hewitson, 1867) Figs. 42a,b Heterochroa ethelda Hewitson (1867a: pi. Heterochroa, fig. 3, 4) TL: Quito [Ecuador]. Types: BMNH(T): STm: "B.M. TYPE No. Rh. 9816 Heterochroa ethelda m Hew.//Quito Hewitson Coll. 79-69 Heterochroa ethelda. 1 .//Type//Syntype" [examined] Adelpha ethelda Hew., Kirby (1871) Identification, taxonomy and variation: This subspecies is distinguished from all others by the orange band on the DHW. There is little variation in the specimens examined. Hewitson (1867a) described and figured both surfaces of this species from an unspecified number of male specimens, and there has never been any confiision as to its identity. A syntype is in the BMNH. Although this subspecies is superficially very distinct from the remaining subspecies, it is parapatric, shares all important wing pattern and genitalic characters with these subspecies and occurs at similar altitudes in similar microhabitats. Furthermore, I have examined two specimens, one of which is in the AME and was figured by Steinhauser and Miller (1977) as an aberration of Adelpha eponina (fig. 19, 20), which are phenotypically intermediate between ethelda and typical A. e. eponina. These specimens have a white DFW band as in A. e. eponina, but the band is more vertical in cells Cu2 to M3 and tinged orange in cell Cu2, while the DHW has a thin orange band along the inner postdiscal series, with sparse orange scaling in the space basal of the outer postdiscal series. Despite the similarity of the dorsal surface with A. leuceria, the species are not closely related and the pattern is achieved in each species through different modifications of the wing pattern ground plan. In A. ethelda, the orange DHW band represents the fiision of the inner and outer postdiscal series, while the

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315 postdiscal band is absent, whereas in A. leuceria the orange DHW band is the postdiscal band and the postdiscal series are absent. The same may also be true of the DFW band, although is is difficult to say how this is derived in A. ethelda (see species introduction). Range: Western slopes of the Andes fi-om Colombia (Tambito) to northern central Ecuador. Habitat and adult ecology: This subspecies remained something of a rarity for many years following its description, the type being the only specimen known to Fruhstorfer (1915). Samson (1978) summarised all the data then known on this subspecies, concluding that it was restricted to cloud forests of the western Ecuadorian Andes. In our experience this is the case, and we have found it to be common during the late wet season and early dry season in both heavily disturbed and primary forest from 7001900m. Males are typically encountered along rivers and forest edges, where they puddle at damp sand, and may be attracted to traps baited with rotting fish or banana. I have never seen the female in the field, and only a single specimen in collections. It is tempting to speculate that the marked difference in the dorsal pattern of this subspecies with respect to others is the result of mimicry, although the only species in western Ecuador with a similar pattern is A. leuceria. This species is much more rarely encountered, at least by the lepidopterist, although it does occupy a similar altitudinal range. Specimens examined (23m, 1 f): COLOMBIA (3m): Cauca: Tambito 1200m 22/3/96 T. Pyrcz Im JFL; Nariiio: El Palmar, Aug. '45 1300m K. von Sneidem Im AMNH; Santander: La Belleza Aug. Im ESM-error. ECUADOR (19m, If): Carchi: Las Juntas, Rio Golondrinas 1400m Nov. Im KWJH; Chimborazo: Huigra 25-3000' Feb. 2m BMB; Imbabura: Paramba Im BMNH, Im BMB; "Napo": Misahualli Im MUSMerror; Pichincha: Alluriquin Aug. Im USNM; Quito Im BMNH, If AMNH; Rio Las Palmeras 1900m Aug. Im KWJH; Rio Tanti 750m Jul. Aug. 3m KWJH; Sto. Domingo 4m AMNH; Not located: Bailos/Sto. Domingo area 2m BMNH; W. Ecuador Feb. Im BMB. COUNTRY UNKNOWN (Im): no specific locality Im BMB. Additional locality data: ECUADOR: Esmeraldas. El Encanto Jul.; Rio Piguambi Jul.; Carchi. Lita, ridge east of Rio Baboso Jul.; Pichincha: Rio Tanti May Jun. (Willmott and Hall, unpubl. sight records). Adelpha ethelda eponina Staudinger, 1886 stat. nov. Figs. 42c-f Adelpha epione eponina Staudinger (1886: 143) TL: Provinz Cauca, Columbien. Types: ZMHU: HTm: "Eponina Stgr.Z/Cauca [illegible]//Origin//Syntype" [examined]

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316 =Adelpha eponina volupis Fruhstorfer (1915: 512) TL: Rio Aquaca Valley, Colombia. Types: BMNH(T): STm: "Fruhstorfer Coll. B.M. 1933-131//eponina volupis Fruhst.//Rio Aguacatal Colomb. W. Cord. 2000m Coll. Fassl//TYPE//Type// Syntype" [examined] Adelpha zalmona eponina Stdgr., Hall (1938); =Adelpha zalmona Hew., D'Abrera (1987) form Identification, taxonomy and variation: This subspecies is easily distinguished from all others by the white DFW band. It also differs from A. e. zalmona and A. e. sophax in having a ftiUy developed silvery gray VHW postdiscal band and a well developed orange-brown outer postdiscal series. It is distinguished from A. epione by the concave instead of convex distal edge of the redbrown band distal of the silvery VHW postdiscal band, by the silvery inner submarginal series of spots on the VHW, by the less rounded hindwing and by the white instead of orange cell posterior of vein 3A. The thickness of the DFW band is subject to individual variation and it may or may not reach tomus. Staudinger (1886) described this taxon as a subspecies of A. epione based on a single specimen from Cauca Province in Colombia, and the holotype is in the ZMHU. Hall (1938) placed it as a subspecies of A. zalmona, Fruhstorfer (1915) and Steinhauser and Miller (1977) regarded it as a frill species, while D'Abrera (1987) considered it to be an infrasubspecific form of ^. zalmona. Since it is closely parapatric and has all the VHW characters typical of A. ethelda (see general identification section), I place it as a subspecies of that species (stat. nov.). Fruhstorfer (1915) described the taxon volupis from an unspecified number of specimens from the Rio Aquaca valley at 2000m, and stated that it differed from A. e. eponina in having a narrower and less extensive white DFW band. Any differences apparent in the syntype in the BMNH merely represent individual variation and the name was synonymised with .4. e. eponina by Hall (1938). Range: Adelpha e. eponina occurs along the western slope of Cordillera Occidental in Colombia, as far south as Rio Calima, south of which it is replaced hy A. e. ethelda. Other fragmentary data indicate that it probably occurs along the western slopes of the Cordillera Central at least as far north as Antioquia and along the eastern slope of the

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317 Cordillera Oriental. More accurate data are needed to assess the range of this subspecies with respect to A. e. zalmona and A. e. ethelda. Habitat and adult ecology: Little has been reported on the habitat or behavior of this subspecies, but it appears to be locally common in cloud forest habitats from 10002000m, where it appears to fly throughout the year. Its geographic and altitudinal ranges closely match those of the white-banded A. lycorias melanippe, and the two taxa are probably involved in mimicry. Specimens examined (68m): t trans, to A. e. ethelda. COLOMBIA (68m): Antioquia: Antioquia Im ZMHU; Caldas: Manizales Im BMNH; "Boyaca": Muzo Apr. Im USNM-error; Cauca: no specific locality 2m BMNH, Im ZMHU; Choco: San Jose del Palmar Jan. 3m JFL; "Cundinamarca": Bogota 3m BMNHerror; Risaralda: Distrito de Pereira 2m BMNH, Im BMB; Quebrada Rio Negro, area a Pblo. Rico 1550m Aug. 2m MHNM; Rio TamanS 2500' Aug. Im BMNH; Valle del Cauca. Cali-Buenaventura 1000m Mar. 2(lt)m ESM; Farallones, km. 55 1200m Jul. Im KWJH; Juntas 2m BMNH, Im Dec. ZMHU; Ugo Calima Oct. Im ESM; Queremal 1200m Jul. Im LMC; Rio Aguacatal 2000m 2m BMNH, Im BMB, Im MCZ; Rio Anchicaya 1000-1 1 50m Oct. Dec. Jan. Feb. 12(lt)m AME; Rio Bravo, Calima May Im LMC; Rfo Calima 1300m Nov. Jan. 3m AME; Rio Dagua 600-1600m Sep. 6m BMNH, 5m ZMHU, Im LMC; Valle Im JFL; Yatacue, Alto Anchicaya Aug. 2m LMC; Not located: bet. Queremal and Buenavista 3500-4000' Feb. 2m AMNH; Quibdo-Bolivar 1000m Feb. Im ESM; Villa Eloira Im BMNH; no specific locality Im BMNH, Im BMB, Im AMNH. Adelpha ethelda zalmona (Hewitson, 1871) stat. nov. Figs. 42g,h Heterochroa zalmona Hewitson (1871 : 165) TL: New Granada. Types: BMNH(T): STm: "Illustrated in The Butterflies of Venezuela A. Neild, 1996//I4// N. Granada Hewitson Coll. 79-69 Heterochroa zalmona 1.//B.M. TYPE No. Rh. 9796 Heterochroa zalmona m Hew.//zalmona//Type//Syntype"; BMNH(M): ST?m: "N. Granada Hewitson Coll. 79-69 Heterochroa zalmona 2."; ST?m; "N. Granada Hewitson Coll. 79-69 Heterochroa zalmona 3" [all examined] Adelpha zalmona Hew., Kirby (1871); Adelpha zalmona zalmona Hew., Neild (1996) Identification, taxonomy and variation: This subspecies is distinguished from A. e. sophax by the more irregular distal edge of the orange DFW band in cells Cul and M3, which is straight in A. e. sophax, and by the duller black-brovra postdiscal area on the VHW, with the outer postdiscal series represented by a pale grayish line instead of a red brown line. Although these differences are slight, they are consistent and also suggest that A. e. sophax may be more closely related to A. e. eponina, which shares both of these characters, than to A. e. zalmona. Adelpha ethelda ssp. nov. is distinguished under that taxon. There is little variation in this subspecies, except in two specimens, listed below, which are transitional to A. e. sophax and A. e. eponina.

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318 Hewitson (1871) described this taxon based on an unspecified number of specimens in the collection of Saunders and Hewitson, and I have examined a syntype and two further possible syntypes in the BMNH which correspond well with the original description. This taxon and remaining subspecies have usually been considered as a species distinct from A. ethelda ethelda, but for the reasons discussed under that subspecies I treat them all as conspecific and therefore place zalmona as a subspecies of ethelda (stat. nov.). Range: This subspecies occurs from Colombia, where it is found from the northern tip to the eastern slopes of the Cordillera Cenfral south to southern Tolima, and on the western slope of the Cordillera Oriental from Bogota to Santander, northwards to the western slopes of the Cordillera de Merida in Venezuela, and although it has yet to be recorded in the Sierras de Perija and Nevada de Santa Marta in Colombia, it probably occurs there. Habitat and adult ecology: Judging from the number of specimens in collections this subspecies is locally common throughout its range. Neild (1996) reports that males are typically encountered as solitary individuals along forest edges, often puddling on fracks. It has been recorded from 4001000m throughout the year. Specimens examined (124m): f trans, to A. e. sophax; % trans, to A. e. eponina, whitish in middle of DFW band. COLOMBIA (1 13m): Antioquia: Frontino 2m(I}) BMNH-error?; Medellin 3m BMB; Mesopotamia Im AMNH; Valdivia 3m BMB; Boyaca: Muzo 4-800m Jun. Jul. Im BMNH, 3m AME, Im USNM, 20m BMB; Otanche Jan. May Aug. 5m JFL; Caldas: Victoria Im ESM; Cundinamarca: Bogota 17m BMNH, 4m USNM, 17m BMB, Im MNHN; env. de Bogota I Im BMNH; head Rio Carare 25000' 2m USNM; Santander: La Lechera 850m Feb, 2m AMNH; La Soledad Dec. Im AMNH; Rio Opon Dec. Im AMNH; Rio Quirata 5-700m Nov. Dec. Im AMNH; Tolima: Rio Chili Apr. -Jun. 3m BMNH, Im BMB; Valle del Cauca: Cali 1000m Oct. Nov. 2m MUSM-error?; Nol located: Pipiral Im MNHN; no specific locality 20m AMNH, 4m BMNH, 2m ZMHU, 3m BMB. VENEZUELA (2m): Trujillo: La Gira Betijoque 520m Dec. 2m AFEN. "PERU" (Im): no specific locality Im BMB-error. COUNTRY UNKNOWN (8m): no specific locality 6m MCZ, 2m USNM. Adelpha ethelda sophax Godman and Salvin, 1 878 stat. nov. Figs. 42ij; 126a,b; 208a,b Adelpha sophax Godman and Salvin (1878b: 265) TL: Cache, Costa Rica. OTL: Costa Rica; New Granada. Types: BMNH(T): LTm: "B.M. TYPE No. Rh. 9797 Adelpha sophax m G. and S.//Type. sp. figured/ZB.C.A. Lep. Rhop. Adelpha sophax G. and S. Godman-Salvin Coll. 1916.-4.//Cache Costa Rica H. Rogers/ZType H.T.//Syntype//m"; BMNH(M): 2PLT?m: "B.C.A. Lep. Rhop. Adelpha sophax G. and S. Godman-Salvin Coll. 1916.-4.//Cache Costa Rica H. Rogers"; PLT?m: "B.C.A. Lep. Rhop. Adelpha sophax G. and S. Godman-Salvin Coll. 1916.4.//Cache Costa Rica H. Rogers//m//16 71"; PLT?m: "Bogota, Colombia C. Felder/ZB.C.A. Lep. Rhop. Adelpha sophax G. and S. Godman-Salvin Coll. \9\6.-4." =ethelda zalmona [all examined] Adelpha zalmona sophax G. and S., Fruhstorfer (1915), Lamas and Small (1992)

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319 Identification, taxonomy and variation: This subspecies is distinguished from A. e. zalmona and A. ethelda ssp. nov. under each of those subspecies. Costa Rican specimens differ from Panamanian specimens by the orange on the DFW extending over the outer submarginal series in the tomus. This is a minor and slightly variable difference, and there is otherwise little variation in the series examined. Godman and Salvin (1878b) appear to have been unaware of the Hewitson's (1871) description of zalmona when they described this taxon, since they only mentioned A. epione and A. boreas in the original description and described it as a full species. The description was based on an unspecified number of male specimens from Costa Rica, collected by H. Rogers, and Colombia, all in their own collection, and I have examined a number of valid and possible syntypes in the BMNH. The single Colombian specimen is referable to the older name A. e. zalmona, so to preserve the name sophax for the Costa Rican subspecies, as it has been used by subsequent authors (DeVries, 1987; Lamas and Small, 1992), I designate the specimen figured as this taxon by Godman and Salvin (1884) as the lectotype of Adelpha sophax, with the following label data: "B.M. TYPE No. Rh. 9797 Adelpha sophax m G. and S.//Type. sp. figured/ZB.C.A. Lep. Rhop. Adelpha sophax G. and S. Godman-Salvin Coll. 1916.-4.//Cache Costa Rica H. Rogers//Type H.T.//Syntype//m". For the reasons discussed under the nominate subspecies I consider sophax to be a subspecies of A. ethelda (stat nov.). Range: To date this subspecies is known only from central Panama to Costa Rica, where it occurs on both slopes of the Cordillera Central and Talamanca. It may well also occur in Nicaragua. Immature stages: The egg is pale green and deposited singly, up to several times per plant (DeVries, 1986, 1987: Costa Rica). The mature larva is mottled dark and pale brown, with scoli similar to A. cytherea, and the head capsule is dark brown with short lateral spines (DeVries, 1987). The pupa is apparently similar to A. cytherea. Both these

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320 brief descriptions suggest a rather different early stage morphology from A. melona, but unfortunately I have been unable to examine any early stage material or adult voucher specimens, and the relationships of this species within the genus remain uncertain. Larvae are solitary and feed on new, then old leaves of plants along roadsides and riparian edges, in bright sun (DeVries, 1986). They also make frass chains and rest in later instars on damaged portions of the leaf (DeVries, 1987). Habitat and adult ecology: DeVries (1987) reports this subspecies to be locally common from 5001200m on the Atlantic slope in cloud forest habitats, particularly in the February-March dry season. Males may be encountered perching as solitary individuals along forest edges and river sides during sunny periods in the morning, up to around 10:00 a.m. Hall and Willmott (1993) reported the species from the Pacific slope, where males were observed perching around 10m above the Rio Angel, in small groups. In my experience, the general behavior of this subspecies is similar to the nominate. Specimens examined (2Sin, 3f): PANAMA (13m, If): Chiriqui: Bugaba 2m BMNH; Chiriquf 3m, If ZMHU, 2m USNM, Imf BMB, Im MCZ; Colon. Pifia Jul. Im AME; Panama: Cerro Campana 2500' Jun. Sep. Im FSCA, 2m USNM. COSTA RICA (12m, 2f): Alajuela: Rio Sucio, nr. Cariblanco 750m Sep. 2m, If KWJH; Cartago: Cache 4m, If BMNH; Juan Viflas Im USNM; Tuis Im USNM; Not located: no specific locality Im BMNH, Im ZMHU, 2m BMB. Adelpha ethelda ssp. nov. Figs. 42k,l Adelpha sophax G. and S., de la Maza and de la Maza (1985); Adelpha zalmona sophax G. and S., de la Maza and de la Maza (1993). Identification, taxonomy and variation: This subspecies is distinguished from A. e. sophax by the much more diagonal DFW orange band, which is thinner, of more even width, and does not cover the outer submarginal series in the DFW tomus. There is little variation in the specimens examined. This subspecies is represented in several collections, including in addition to those listed below, the collection of Mast de Maeght in Brussels, where there is at least one male from Chiapas, according to a photograph taken by Yves Hiemaux in the drawer

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321 containing A. e. zalmona at the BMNH. I do not describe it here since Lee and Jacqueline Miller (pers. comm.) have been intending to describe it for some time and will do so shortly. Range: To date this subspecies is known only from Mexico, from the Atlantic slopes in the states of Oaxaca, Veracruz and Chiapas, though the majority of specimens are from the latter state. It almost certainly occurs in Guatemala, and possibly also to Honduras. Habitat and adult ecology: De la Maza and de la Maza (1993) report this subspecies, as A. zalmona sophax, from humid lowland rain forest in Chiapas from 100-500m. Judging from the number of specimens in collections and the range of sites recorded by de la Maza and de la Maza (1993), it is not uncommon in the field. It has been collected from August to December and once in May. Specimens examined (12m, 3f): MEXICO (12m, 3f): Chiapas: "Chiapas, Mex. coll. W. Schaus" Im USNM; Comitan Nov. If AME; Rio Chixoy Aug. 6m AME; Oaxaca: Chimalapa Dec. If AME; Rio Sarabia Sep. Oct. Im, If AME; Zuzla [??] May Im AME; Veracruz: "Franca Vieja, V.C., Mexico VIII '44. M. Guerra, coll. Frank Johnson Donor" Im AMNH; "Pajaritas, V.C., Mex. VIII 1944 M. Guerra, coll. Frank Johnson Donor" Im AMNH; San Manuel Oct. Im AME; Not located: photo in drawer of specimen from Mexico Im BMNH. Additional locality data: MEXICO: Chiapas: Chajul 150m (de la Maza and de la Maza, 1985); Cuenca de Grijalva; Cuenca de Tuliji; Macizo Central; Selva Lacandona (de la Maza and de la Maza, 1993). Adelpha ethelda galbao Brevignon, 1 995 stat. nov. Figs. 42m,n Adelpha epione galbao Brevignon (1995: 17, fig. 25, 26) TL: SaOI, French Guiana. Types: LCB: HTm: SaUl, 3 Sep. 1993 [original illustration examined]; JYG: 2PTm: same locality data as HT except Sep. 1982, 5 Sep. 1987; Musie Departmental de Cayenne: PTm: same locality data as HT, Nov. 1970, coll. R. P. Barbotin [not examined] Identification, taxonomy and variation: This subspecies is distinguished from all others by the broader, more vertically orientated orange DFW band, which is slightly paler in the basal half, by the silvery gray VHW postdiscal band being reduced to a spot near the costa, and by the orange shading over the cell posterior to vein 3A. The two specimens I have examined are very similar in wing pattern. Brevignon (1995) described this taxon as a subspecies of A. epione, with which it shares the reduced VHW postdiscal band, and the cell posterior of vein 3 A on the VHW

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322 being orange. These two characters appear to be local synapomorphies which group A. epione epione. A, epione agilla and galbao. The systematic placement of galbao depends, however, on one's point of view regarding what constitutes a species, and I place it as a subspecies of A. ethelda (stat. nov.) for the reasons given in the discussion of this species. Range: To date this recently discovered subspecies is known from two sites, one in southem Venezuela and one in French Guiana, but clearly it is more widespread, presumably occurring throughout the Guiana shield area in montane areas. Even so, it is clearly disjimct in range from remaining A. ethelda subspecies. Habitat and adult ecology: This subspecies is very rare in collections, and I have seen no specimens in any that I have examined. In addition to the original description, I have also examined a photograph, courtesy of Andrew Neild, of a specimen captured by Roberto de la Fuente in southem Venezuela. According to Brevignon (1995), males may be encountered in areas of secondary growth during the hottest hours of the day. Specimens examined (2m): VENEZUELA (Im): Amazonas: Ashidowa Ten Alto Siapa 1 m(photograph) DLF. FRENCH GUIANA (Im): Laurent du Maroni: Saul Sep. lm(photograph) LCB. Adelpha epione (Godart, [1824]) Figs. 43; 127; 209 Identification, taxonomy and variation: This species is most closely related to A. ethelda, from which it is distinguished by the rounded hindwing shape, with the hindwing apex less angular and the distal margin less dentate, and the oblique white band on the DFW, which also occurs in A. ethelda eponina. Adelpha ethelda eponina is further distinguished on the VHW by having a complete silvery gray postdiscal band, a complete orange-brown outer postdiscal series, and a complete submarginal series of silvery gray spots. I discuss other distinctive characters of this species under /i. ethelda. There is geographic variation in the shape of

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323 the white DFW band and the expression of the VHW postdiscal series, and two subspecies are recognised. This species appears to be monophyletic, but its recognition as distinct from A. ethelda probably makes that species paraphyletic, and my reasons for retaining both species are discussed under A. ethelda. Range and status: Eastern slopes of the Andes from Colombia to Bolivia, to southeastern Brazil. Not uncommon in lower montane cloud forest habitats up to 1600m, but also occurring in lowland rain forest as low as 200m up to 200 km. away from the Andes. Specimens examined: 342 (332m, lOf) Adelpha epione epione (Godart, [1824]) Figs. 43a,b Nymphalis epione Godart [1824]: 405) TL: Bresil. Types: MNHN?: STm: [not located] Adelpha epione Godt., Kirby (1871) Identiflcation, taxonomy and variation: This subspecies is distinguished from A. epione agilla by the narrower white DFW band, which is of more even width and has more even distal and basal margins, by the VHW having fraces of the silvery gray spots of the inner submarginal series, by the postdiscal band being narrower with a straighter distal edge, and by the red-brown outer postdiscal series being uniformly faint throughout instead of more pronounced in its anterior half There is little variation in the specimens examined. Godart ([1824]) described this species based on males from Brazil, and since the description refers to the oblique white DFW band and five reddish brown bands on the VHW, corresponding to the basal band, the discal cell band, and the inner and outer postdiscal and outer submarginal series, there can be no doubt as to its identity. I have

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324 been unable to locate any syntype specimens in the MNHN. The name maina, introduced by Martin et al. ([1923]) as a subspecies of A. epione, is a nomen nudum. Range: The nominate subspecies is known from southeastern Brazil, from Espirito Santo to Sao Paulo and Minas Gerais. Habitat and adult ecology: This subspecies is uncommon in collections and there are no published observations of it in nature. Specimens examined (27m, 5V): BRAZIL (27m, 30: Espirito Santo: no specific locality Im, 2f BMNH; Guanabara: Rio de Janeiro Im ZMHU; Minas Gerais: no specific locality Im BMNH; Rio de Janeiro: Petrdpolis 3m BMNH; Sao Paulo: Arafatuba 450m 3m BMNH; Bahuru 3m BMNH; Casa Branca 2m BMNH, Im ZMHU; Rio Batalha Im BMNH; S5o Paulo 6m BMNH, 2m BMB; Not located: no specific locality Im, If BMNH, Im BMB, Im MNHN. COUNTRY UNKNOWN (20: no specific locality 2f BMNH. Adelpha epione agilla Fruhstorfer, 1907 Figs. 43c,d; 127a,b; 209a,b Adelpha epione agilla Fruhstorfer (1907: 172) TL: Colombia. Types: BMNH(T): HTm: "epione agilla Fruhst.//Fruhstorfer Coll. B.M. 1933-1 3 l/ZCoIumbien ex. coll. Fnjhstorfer//TYPE//Holotype//Type" [examined] Adelpha epione Godt., Staudinger (1 886) in part Identification, taxonomy and variation: This subspecies is distinguished from the nominate under that account. There is little variation except in a single specimen from Upper Paraguay River, which appears intermediate in wing pattern to the nominate subspecies, having a broad but evenly edged white DFW band. Fruhstorfer (1907) described this subspecies based on a single male collected by Stichel in 1896 in Colombia, and the putative holotype is in the BMNH. He described several differences such as the broader white DFW band and dark silver gray VHW postdiscal band, and cited the latter character as distinguishing it from A. ethelda eponina. Range: This subspecies occurs from Colombia to Bolivia, across to either Paraguay or Brazil on the "Upper Paraguay River". In Colombia it seems to occur only on the eastern slopes of the Cordillera Oriental. There are several specimens labeled as from localities west of here, such as Antioquia and Muzo, but I regard these as probable mislabellings and believe that its presence outside the eastern slope has yet to be demonstrated.

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325 Habitat and adult ecology: This species is rather local but can be common in relatively undisturbed habitats, and typically occurs in cloud forest from 6001600m, although individuals may also be found as far as 200 km. from the base of the Andes in flat lowland rain forest at around 200m. Males are usually encountered along mountain rivers puddling at damp sand, and are also attracted to rotting fish in traps along forest edges or in light gaps. They may also be seen perching 5-7m above the ground on the tops of bushes in large forest light gaps along streams. Females are much rarer than males, the only individual I have seen was flying low along a wide ridgetop path with abimdant secondary growth in the early afternoon. The subspecies seems to fly throughout the year. Specimens examined (305m, 5f): t trans, to A. epione epione. COLOMBIA (58m, IQ: "Antioquia": Rio Cocoma 800-lOOOm Aug. Im AMNH-error; "Boyaca": Muzo 2m AME-error; Santa Maria Dec. Im ESM; Cundinamarca: Bogota 14m BMNH, 2m, If USNM, Im BMB; env. de Bogota 9m BMNH; Meta: Villavicencio Jan. Im ESM; Not located: no specific locality 21m AMNH, 2m BMNH, Im ZMHU, Im AME, 2m USNM. ECUADOR (65m, If): Chimborazo: Rio Bamba Im AME; Morona-Santiago: Macas Oct. 2m AMNH, Im BMNH; Mindez 850m Oct. Im MJP; Mendez-Santiago rd., km. 40, Oct. Im MJP; Rio Abanico 1600m Nov. If KWJH; 2°-4''S, 78°W 875m Nov. Dec. Im AMNH; Napo: Talag Sep. Im DAT; Paslaza: Hda. Moravia, nr. Shell Feb. 1200m Im KWJH; Lorocachi, Rio Cururay 200m Jul. Im KWJH; Mera Oct. Im AME; Puyo-Tena rd. Jun., km. 17, Im DAT; Rio Llandia Nov. Im DAT; Sarayacu 200m Mar. Im AME; Tungurahua: Rio Chinchin Grande 1400m Oct. Im AME; Rio Negro Jul. 2m FSCA; Rio Tigre 1200m Aug. Im AME; Rio Topo 4500' Apr. Jun. Jul. Oct. 2m DAT, Im JFL, Im BMNH, Im FSCA, Im USNM; Rio Zuiiac 1300m Jul. Im KWJH; Santa Inez Im MCZ; Yunguilla, nr. Bailos Apr. 2m USNM; Zamora-Chinchipe: Quebrada Chorillos 1250m Apr. Im KWJH; Zamora 3-4000' 16m BMNH; Not located: Oriente 3m AMNH; Pacific slope 2m USNM-error; no specific locality 3m BMNH, Im MCZ, Im JFL. PERU (105m, If): Amazonas: Falso Paquisha 800m Oct. Im MUSM; Huambo 3700' Im BMB; Huayabamba 3500' 2m BMNH, Im ZMHU; Cuzco: Callanga 1500m Im ZMHU; Cosilipata Valley Im BMNH; Hda. Cadena lOOOm Im MUSM; Marcapata 4500" 2m MUSM, Im BMNH; Quincemil Nov. Im MUSM; Huanuco: Divisoria Jun. Im AME; Pozuzo 800-lOOOm lOm BMNH, Im MUSM; Puente Cayumba, Rio Huallaga 800m Apr. 7m AMNH, Im MUSM; Tingo Maria 800m Mar. Jun. Jul. Nov. 2m FSCA, 3m AME, Im MUSM; U. Marailon Im BMNH; no specific locality 1400m Apr. Im USNM; Junin: Alto Yurinaqui 1000m May Im MUSM; Chanchamayo 1200-1400m Sep. 2m BMNH, Im ZMHU, Im, If AME, Im USNM, 2m BMB; La Merced Im BMNH, Im MUSM, Im AMNH; Rio Colorado 2500' Mar. Apr. Aug. Oct. 2m BMNH, Im AME, 3m BMB; Rio Ipoki May Im MUSM; San Ramon 3000' Oct. Im BMB; Satipo May Dec. 2m AME; Loreto: Iquitos Im BMNH, Im AMNH; Marotta Jul. Im AME; Nauta Im BMNH; Pebas 2m BMNH, Im ZMHU; Rio Cachiyacu Im BMNH; Rio Morona Jan. Im AMNH; Yurimaguas Im ZMHU; Madre de Dios: Albergue Amazonia 600m Oct. Im MUSM; Boca Rio la Torre 300m Oct. Nov. 2m MUSM; Puerto Maldonado 250m Aug. Im AME; Pasco: Pichis trail 600-I200m Apr. Im ZMHU, Im BMB; Puno: La Oroya-Agualani Oct. Im BMNH; La Union, Rio Huacamayo 2000' Dec. 4m BMNH; Rio Tavara 450m Aug. Im MUSM; San Gaban 2500' March, April Im BMNH; Sto. Domingo Im BMNH; Yahuarmayo 1200' Feb. Mar. Oct. Nov. 3m BMNH; San Martin: Jepelacio Im AMNH; Moyobamba Im, If BMNH, Im BMB; Rio Huambo Feb. Im MUSM; Not located: E. Peru Im BMB; middle Rio Ucayali Jun. Jul. 5m AMNH; Ucayali Im ZMHU; S. Peru Im BMB; no specific locality 2m AMNH, Im BMNH, Im USNM, Im MCZ. BOLIVIA (64m, \{): Cochabamba: Alto Palmar 1 1 00m Im MUSM; Charapaya 1300m Jan. Im BMNH; Cochabamba 4m BMNH; Cristal Rayo, Chapare Mar. Im JFL; El Palmar May 2m AME; 5 days N. Cochabamba 4m MCZ; San Mateo Im BMNH; Yunga del Esp. Santo Im BMB; El Bent: Cachuela Esperanza Sep. 2m AME; Muschay, Rio Beni Aug. 3m BMNH; Reyes Jul. 2m BMNH; Rio Quiquibby 600m Sep. 2m KWJH; La Paz: Coroico 1200-2000m May Im BMNH, Im ZMHU, Im AME; Cusilluni May Im BMNH, Im MCZ; Guanay 1500' Aug. Oct. Im JFL, Im BMNH; Rio Songo 1200m Im ZMHU; San Ernesto 1000m Sep. Im BMNH; Yungas 1200m Nov. 2m BMNH; no specific locality 3m BMNH, Im ZMHU, Im AME; Santa Cruz: Buenavista 400m Oct. Im AMNH, 3m, If AME, Im MUSM; Prov. Sara Mar. Apr. 10m BMNH; Santa Cruz 2m USNM; Not located: no specific locality 3m AMNH, 3m BMNH, Im BMB, Im MNHN. COUNTRY UNKNOWN (13m, If): no specific locality 4m BMNH, 3m, If AME, 2m USNM, 3m MCZ; Santa Cruz, Up. Paraguay R Sep Imt BMNH. Additional locality data: ECUADOR: Napo: Pimpilala 600m Sep.; Pastaza: Shell 1000m Apr.; Zamora-Chinchipe: Quebrada Chorillos 1250m Nov. (Willmott and Hall, unpubl. sight records). Adelpha syma (Godart, [1824]) Figs. 44a,b; 128a,b; 210a,b

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326 Nymphalis syma Godart ([1824]: 374) TL: Bresil. Types: MNHN?: ST: [not located] =Adelpha pravitas Fruhstorfer (1913: pi. 107f; 1915: 522) syn. nov. TL: [no locality stated]. Types: BMNH(R): STf: "syma forma pravitas Fruhst.//f. pravitas Fruhst.//R//107 f3//Syntype//TYPE" [examined] Adelpha syma Godt., HObner ([1827]: 37, fig. 571, 572); Helerochroa syma Godt., Westwood (1850); Adelpha syma Godt., Fruhstorfer (1915) Identification, taxonomy and variation: The distinctly banded ventral surface and the very broad orange DFW marking distinguish this species from all others. There is little variation. Godart ([1824]) described this species based on specimens from Brazil, and mentions the very large orange DFW marking and the venfral surface being entirely redbrown with white bands, lined with black, of which there were six on the forewing and five on the hindwing. There is therefore little doubt as to the identity of the species, despite the fact that no type specimens are known to exist. Fruhstorfer (1913) figured a specimen, as pravitas, which had the DFW postdiscal and DHW tomal orange markings entirely white, and later stated that he believed it to be an aberrant form of A. syma (Fruhstorfer, 1915). Since the female specimen in the BMNH(R), which is clearly that figured in the original description, does indeed represent an aberration, I synonymise pravitas with syma (syn, nov.). Due to the highly modified wing pattern it is difficult to determine this species' relationships within the genus, but the male genitalia, the straight, continuous white postdiscal band on the VFW and the straight postdiscal series, suggest it may be related to A. cytherea. Range: This species is known from the eastern Brazilian coast at Espirito Santo, south as far as Uruguay, and west across to northeastern Argentina and Paraguay. There are single records from Bahia and Natal, in Rio Grand do Norte, which require confirmation, and three records from the Amazon which are certainly erroneous. Immature stages: Hoffinan (1937) reared and described in detail the immature stages and behavior of this species in Santa Catharina, and except where noted, the description

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327 below is compiled from his notes. The egg is deposited singly at the point of a lateral "tooth" of a hostplant leaf, and up to four eggs may be laid on a single leaf The egg is typical of the genus, and the first instar is brownish gray, later brown, wth a brown head. The second instar is matt brown with paler developing scoli, and remains similar in form up to and throughout the fourth instar. Miiller (1886) also reared this species in Santa Catharina, but reported that the early instars were green. The last instar is greenish with a brown dorsal mark near the centre of the body (Miiller (1886) reports black dorsal coloring on segments T1-A2, and A8 and sometimes A7, similar to A. plesaure and A. viola viola), and has pairs of brownish, fleshy, spined tubercles on segments T2, T3 and A2, the first pair inclined anteriorly, the second vertical, and the third inclined posteriorly. There are similar but smaller pairs of scoli on A7 and A8, while segments A3-A6 only have very small scoli. The prepupa is pale brownish. The pupa is light brown with a gold gloss, most pronounced on the thoracic segments, and differing in form from A. serpa. The abdominal segments are laterally compressed, with the wing edges protruding. There is a flat, dorsal keel which is composed of the projections on T2 and A2, and these appear to almost meet in the middle, similar to A. melona. The remaining abdominal segments each have a pronounced dorsal keel, ending sharply at the end of each segment. The head ends in two flat horns, which are dorso-ventrally compressed. The spiracles are darkbrown. The "first abdominal segments" have small projections in the same position as the last instar scoli. Hoffinann also stated that the pupa "rests" 31-35 days during the winter, possibly indicating diapause, and that the adults eclosed in July. Hayward (1931) also reared this species in Argentina and states that the larva is dark green and covered with protuberances with terminal spines, changing to a yellowish brown, presumably prior to pupation, in the last instar. The development time was Hsted by Hoffinann (1937) as follows: egg, 5 days; l" instar, 10 days; 2"'*-4* instar, 10-13 days (oviposition-moult of last instar, 48 days); S"' instar, 10 days; pupa, 31-35 days. Hoffinan speculated that this protracted development

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328 time, 89-94 days, was due to the low temperatures on June and July, when his rearing records took place. Certainly this time is much greater than other reported Adelpha, the next longest being yi. bredowii californica, at 65 days (Comstock and Dammers, 1932). The early instar larval behavior of extending the leaf midrib with frass fastened with silk is also typical of the genus, while the fifth instar abandons these perches and rests on top of the leaf The larva appears to rest in the FrontArched-Rear-Up Position of Aiello (1984), figured by Harry (1994: fig. 5), with the head pressed to the leaf Hoffinann also reported parasitisation of larvae by an unidentified Hymenoptera ("wasp"). Habitat and adult ecology: This is one of the most common species throughout its range, and its phenotypic stability suggests it may well be a model species in mimicry rings fi-om this region. It flies throughout the year, and although there is a single record of 2100m, this requires confirmation since all other records are below 1400m. Ebert (1969) found it to be common in Minas Gerais in a variety of habitats, fi-om primary to secondary forest, along rivers and in hilly areas. Brown (1992) reports it to be common in disturbed forest areas in the Serra do Japi, Sao Paulo, Brazil. Hoffinann (1936) reported that the species could often be found feeding at flowers of Mikania in Santa Catharina, Brazil. Specimens examined: 367 (289m, 78f) BRAZIL (269m, 740: "Amazonas": Canuma Apr. Im AMNH-error; Bahia. Bahia Im BMNH; Guanabara: Rio 5m, 2f BMNH; Distrito Federal: Brasilia Im ZMHU, 2m USNM; Espirilo Santo: Baixo Guandii Dec. 2m, If AME; Santa Teresa Oct. Im MUSM; no specific locality Oct. If BMB, Im AMNH; Minas Gerais: Campo Bello Im BMNH; Itamonte, 15 km. S.E., Feb. If USNM; Marianna 14m AME; Nova Lima Apr. Aug. Im USNM, Im AME; Ouro Preto Apr. Im BMNH; Passa Quatro 1000m 2f BMNH, Im MNHN; Rio-Belo, km. 290, May Im USNM; Sena do Espinha90 Apr. 3m AME; no specific locality 2100m Feb.-Apr. Aug. 17m BMNH, If MNHN; "Para": Obidos Im, If AME-error; Parana: Castro 2900' Jan. Mar. Apr. 3m, 2f BMNH, Im AME, Im, If USNM; Caviuna 2f AMNH; Curitiba Mar. Im FSCA, 2f MNHN; Femandes Pinheiro 2600' Apr. 2m AME; Gurrahaia Apr. Im AME; Igua^u Oct. 2m BMNH; Ponta Grossa 3500' May Dec. 3m AME, 3m BMB; Sao Luiz do Puruna 900m Mar. Im FSCA; Tibagy 2400' Mar. Im AME; Tijucas do Sul 850m Apr. If MUSM; Uni§o da Vitoria 2000' Dec. Jan. Mar. 6m, 2f BMB, Im MUSM; Rio de Janeiro: Itabapoana Im BMNH; Laguna de Sacuaresma 2m BMNH; Nova Fnborgo 3000' Jan. If BMNH, Im AMNH; 2m AME, 8m USNM, 6m BMB, Im MNHN; Petropolis Mar. Nov. 10m, 4f BMNH; Teresopolis 3m, If BMNH; Rio Grande do Norte: Natal If MNHN; Rio Grande do Sul: Northeast If BMNH; Pelotas Jan.-Apr. Jun. Dec. 47m, 8f AMNH, 6m, 6f MCZ, If BMNH, 2m MUSM, If AME, 8m, 4f USNM; Rio Grande do Sul May Jun. 2m, 4f BMNH, Im, If ZMHU, 2m, If AME, 3m MNHN; Santa Cruz Im AME; Santa Catharina: Rio Natal, nr. Rio Vermelho 900-1450' Mar. Im FSCA; Blumenau 24m, 4f MCZ, If BMNH, 2m ZMHU; Cauna Mar. Apr. 2m, If AMNH; Joinville If AMNH; Pinhal Apr. Im USNM; Rio Negro Im MNHN; Rio Vermelho 33503550' Mar. Im FSCA; SSo Bento do Sul 850m Feb. Mar. 6m, 1 f AME; Trombudo Alto Jan. 2m AME; Villa de Lages 2m BMNH; no specific locality 5m, 3f AMNH, 4m MCZ, If ZMHU; Sao Paulo: Alto da Serra Santos 2600' Feb. Apr. Im BMNH, 2m AME, If MCZ; Bahuru 7m BMNH; Itaquaquetetuba Jun. Im BMB; Jacarei Jaguari May 2m AME; S3o Paulo 2500' Nov Dec Feb Apr 3m 2f BMNH, Im, If ZMHU, Im USNM, 3m, 2f BMB, Im MCZ; Villa Mathias, Santos Nov. Im MNHN; Not located: Rio River If BMNH; S. Brazil If BMNH; no specific locality 4m BMNH, Im BMB, 3m MNHN, 3m, If AMNH. ARGENTINA (Im): Misiones San Ignacio May Im USNM. PARAGUAY (9m): Caaguazu: Yhii Sep.-Dec. 2m BMNH; Central: Trinidad Jul.-Sep. Im AMNH; Paraguari: Sapucay Jan. Sep. Nov. Dec. 3m BMNH, Im BMB; Not located: S. Paraguay 2m BMNH. URUGUAY (Im 2o' Montevideo: Montevideo Nov. Im, 2f AMNH. COUNTRY UNKNOWN (9m, 20: Im, If BMNH, Im AME, 5m If MNHN 2m

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329 Additional locality data: BRAZIL: Minas Gerais. Po?os de Caldas 1000-1400m Jan.-May Nov. Dec. (Ebert, 1969); Rio Grande do Sul: Cascata; Monte Benito Mar. Apr. May (Biezanko, 1949); Santa Catharina: Rio Negrucho 850m (Hoffmann, 1936). Adelpha cytherea (Linnaeus, 1758) Figs. 45; 129; 211 Identification, taxonomy and variation: This species is distinguished from all similar species by having the orange postdiscal series on the DFW fused with the white postdiscal band anterior of vein M3 at least, usually to vein Cu2, forming an even band to the costa. The presence of the third discal cell bar on the VFW helps to distinguish it from species which may be superficially similar, such as A. cocala, A. mythra and A. plesaure. The male genitalia are distinctive in having a high dorsal medial lobe on the valva, positioned very close to the dorsal basal lobe, and a short clunicula. There is considerable variation in the width of the white DHW postdiscal band, the presence or absence of orange along the postdiscal series on the DHW, and the extent of orange on the DFW in cell Cu2, particularly along the inner postdiscal series which immediately borders the distal edge of the white postdiscal band. I recognise seven subspecies, several of which may prove to simply represent clinal variation. The relationships of this species within the genus are somewhat obscure, but there is some evidence to suggest that it may be relatively closely related to A. salmoneus and A. viola. All of these species share similar male and female genitalia and the presence of a third discal cell bar on the VFW. The latter character is found in the most primitive members of the genus, but its presence in these three species may represent a reversal and therefore a synapomorphy. In addition, the larval hostplants and early stages of all three species are similar.

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330 Range and status: Widespread from Mexico to western Ecuador and northwestern Venezuela, to Bolivia, Brazil, northeastern Argentina, the Guianas and Trinidad. In secondary growth habitats from sea level to 1500m, very common. Specimens examined: 1689 (1256m, 433f) Adelpha cytherea cytherea (Linnaeus, 1758) Figs. 45a,b Papilio cytherea Linnaeus (1758: 481, n. 139) TL: Indiis. Types: Linnaean coll.: STm: [photograph examined] =Papilio eleus Linnaeus (1758: 486, n. 176) stat. rest. TL: Indiis. Types: Coll. unknown: ST: [not located] =Adelphalanilla•mhsX0T^Qx{\9\^: 108b; 1915: 521) TL: [Peru; Bolivia; Mato Grosso]. Types: BMNH(T): STm: "Fruhstorfer Coll. B.M. 1 937-285//cytherea lanilla Fruhst.Z/Yungas Bolivia//TYPE//Type//Syntype"; STf: "Peru H. Fruhstorfer//TYPE//Type"; BMNH(M): STm: Peni H. Fruhstorfer/Zcytherea lanilla Fruhst.//Fruhstorfer Coll. B.M. 1937-285"; 2ST?m: Mato Grosso/ZFruhstorfer Coll. B.M. 1937-285"; BMNH(R): STm: "cytherea lanilla Fruhst.//Pozuzo Huanuco 800-1 000m (W. Hoffinannsy/R" [all examined] Papilio cytherea Linn., Linnaeus (1764: 305, n. 123), Clerck (1764: tab. 39 [fig. 3, numbered in register at back]); Papilio eleus Linn., Linnaeus (1764: 312, n. 130), Herbst (1793: 133, pi. 147, fig. 3, 4); Papilio elaea [sic] Linn., Muller (1774: 613, pi. 19, fig. 10); Papilio elea [sic] Cramer (1779: 83, pi. 242, fig. D, E); Najas cytherea Linn., Hubner (1808: pi. 62, fig. 1, 2, 3, 4); Adelpha elea [sic] Linn., HQbner ([1819]); Adelpha cytherea Linn., HObner ([I8I9]); Nymphalis cytherea Linn., Godart ([1824: 373]); Heterochroa cytherea Linn., Westwood (1850); Adelpha cytherea Linn., Kirby (I87I); Adelpha cytherea Linn., Aurivinius(1882) Misapplication of names: Papilio cythereus [sic] Linn., Herbst (1793: 137, pi. 148, fig. I, 2) = Adelpha iphiclus iphiclus; Papilio cytherea Linn., Muller (1774: 619, pi. 19, fig. 6) = Speyeria adalia; Papilio cytherea Linn., Cramer (I78I: 170, pi. 376, fig. C, D) = Adelpha thoasa caltiphictea Identification, taxonomy and variation: The nominate subspecies is distinguished from other subspecies by the outer postdiscal series on the DFW extending to the anal margin as an orange line, but the inner series terminating at vein Cu2, with the result that the white postdiscal band is separated from the orange dash of the outer postdiscal series by a dark brown area in cell Cu2. Adelpha c. marcia also has a similar arrangement in cell Cu2 on the DFW, and although sometimes indistinguishable, it typically has a narrower orange DFW band. I retain the two as distinct since they are clearly disjunct in range. Adelpha c. aea typically has no orange in cell Cu2 on the DFW, and the white extends to vein Cul. Both A. c. despoliata and A. c. insularis have the inner postdiscal series on the DFW orange in cell Cu2, while the latter also has orange scaling lining veins Cu2-M2 in the postdiscal area of the DHW.

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331 This subspecies is rather variable both individually and geographically, but I do not regard any of the observed differences as sufficiently stable to warrant separate subspecific recognition. Specimens from the lower Amazon and the Guianas often have a scalloped distal edge to the DFW orange band, and some have bright orange band in the DFW discal cell between the second and fourth discal cell bars (the latter also occurs in specimens from the Orinoco delta (Neild, 1996)). The white hindwing band tends to thin clinally from west to east across the Amazon basin, especially in females, with specimens from Santarem and Maranhao having very thin white bands. Specimens from Pemambuco have the orange on the DFW extending to the white band in cell Cu2, while a single male from Quipapa (Pemambuco) and one from Tombador Falls (Mato Grosso), have the white band extended along the inside edge of the orange as far as vein M3. I have seen insufficient specimens from Pemambuco to determine whether any of the variation there represents a distinct subspecies and freat those specimens here as the nominate subspecies. This is another species which has suffered a varied taxonomic history, again due to a poor original description by Linnaeus. Linnaeus' (1758: 481) description of Papilio cytherea is as follows (words in square parentheses added by me): "P. [Papilio] N. [Nymphales] alis crenatis flavis fascia communi argentea: subtus fascia lanceolata argentea. Alae supra cinereo nebulosae: primores disco flavo.". Clerck (1764: tab. 39) then figured both surfaces of a specimen under this name which clearly belongs to A. cytherea as treated here (from now on referred to as A. cytherea), except that he omitted the orange tomal spot on the DHW. Several pages later, and shortly after his description of Papilio iphiclus, Linnaeus (1758: 486) described Papilio eleus as follows: "P. [Papilio] B. [Barbarus] alis dentatis fiisco-nebulosis concoloribus: fascia primoribus flava, positicis alba." This species was not, however, figured by Clerck, but Linnaeus (1764) embellished his somewhat curt original description as follows: "Alae primores supra basi margineque exteriore fusco nebulosae. Fascia disci lata, flava. Posticae supra

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332 flava-nebulosae. Fascia alba, transversa, angusta, quae etiam tangit anteriorum. Subtus omnes flavescentes margine albo undulato. Fascia alba fenestrata f. perspicua". This second description corresponds very well with A. cytherea as treated here, and it is th