Butterflies of Mexico and Central America

A Swift Guide to the

Butterfliesof Mexico and Central America

Second Edition

A Swift Guide to the Butterfliesof Mexico and Central America

Second Edition by Jeffrey Glassberg

Princeton University Press Princeton and Oxford Acknowledgements knowledge of foodplants and speciation. My wife, Jane Vicroy Scott, accompanied Thanks also to Roberto de la Maza, Jr., me on many trips, and made many sugges- Alida Enkerlin, Thomas and Domingo Ro- tions about the text and layout. Surpris- driguez, Jane Ruffin, Andres and Pilar Sada, ingly, my extended absences while in the Lucy Sada, Tino Sanchez, Jane V. Scott, field caused her some dismay. Her love and Jenn Sinasac, and Jim Springer for accom- big heart are what nurtured this project. See panying me on forays into the field. pg. 294. For the first edition, Barbara Ribble’s About 2800 of the approximately 3700 and Jane V. Scott’s careful eyes found photographs in this book were taken by the typographical errors while Jason Hall, Bob author, but other photographers provided Robbins, Jane V. Scott and Keith Wilmott, many wonderful photographs that increase looked over all or portions of this work the usefulness of this guide. Thanks to all and each made useful suggestions for its of them for allowing their photographs to improvement. My heartfelt thanks to all of be used in this work. I’d like to especially you. thank Will and Gill Carter, Kim Garwood, Dan Janzen and Winnie Hallwachs, John Kemner, Jean-Claude Petit and Pierre © 2017 by Jeffrey Glassberg Boyer, and Jane Ruffin, each of whom not only provided more than 50 photographs, Published by Princeton University Press, but also took the significant time and effort 41 William Street, Princeton, New Jersey to find the photos and to deliver them to me. 08540 Museum visits were crucial in prepar- ing the first edition of this work. Thanks to All rights reserved. No part of this publica- Dave Grimaldi, Jim Miller and Eric Quinter tion may be reproduced, stored in a retrieval at the American Museum of Natural His- system, or transmitted, in any form or by tory, New York, to John Burns, Don Harvey any means, electronic, mechanical, photo- and Bob Robbins at the United States copying, recording or otherwise, without National Museum, Smithsonian Institution the express and prior written permission of and to George Austin, Tom Emmel, Jackie the publisher. and Lee Miller, and Steve Steinhauser at the Allyn Museum/McGuire Center in Sara- sota/Gainesville, Florida for allowing me to ISBN 978-0-691-17648-2 work with specimens under their care. Library of Congress Control Number: A large number of people have contrib- 2017951508 uted to our understanding of Mexican and Printed on acid-free paper. ∞ Central American butterflies and so it would be impossible to thank them all here. For Printed in China Mexico, I need to single out the de la Maza 10 9 8 7 6 5 4 3 2 1 family, the UNAM-based group of Jorge Llorente, and John Kemner. Their distribution records contributed greatly to the maps in this book. For Central America, the Half-title page: A Wavy-lined Sunstreak (Arcas cypria) thorough work of Dan Janzen and Win- nie Hallwachs in the Guanacaste region of Title page: A Black-patched Bluemark (Lasaia Costa Rica has led to an explosion of our agesilas) Contents

Introduction Welcome to Mexico 1 1 Mexican Ecosystems 4 About the Plates 5 Abbreviations 6 About the Maps 6 Swallowtails Swallowtails 8 8

Whites and Whites 21 Yellows Yellows 29 21 Mimic-Whites 33 Gossamer- Hairstreaks 36 wings 36 Coppers and Blues 57 Metalmarks Metalmarks 58 58 Actinotes, True-Brushfoots 86 Brushfoots Admirals and Relatives 109 86 Leafwings and Emperors 136 Owls, Satyrs, Monarchs, Ticlears 150 Firetips 170 Eudamini 175 Skippers Spreadwing Skippers 175 Pyrgini 205 170 Skipperlings 234 Grass-Skippers 242 Conclusion Selected Bibliography 290 290 Selected Websites 292 About NABA 293 Dedication 294 Index 296 Photo Credits 299 Visual Index 302

INTRODUCTION Welcome to Mexico and Central America and the World of Butterflies! Mexico and Central America, vast and varied, both old and new, are some of the greatest places on earth to see and enjoy butterflies. Mexico offers a hard to beat combination of great diversity and amazing abundance. The more than 2400 butterfly species found in Mexico and Central America (about 1750 in Mexico alone) represent more than 10% of the world’s total species. Many of these beautiful creatures can be found nowhere else. Just as importantly for your enjoyment, Mexico more consistently has greater concen- trations of butterflies than perhaps anywhere else. In northern Mexico, there are times during peak butterfly seasons when one has very limited visibility looking down trails, because the warm, late summer air is saturated with butterflies! In Chiapas, as I walked down a trail to an ancient Mayan ruin set deep in the Lacandonan rain forest, butterflies actually interfered with my search for other butterflies, as scores of Many-banded, Pale and Ruddy Daggerwings and other large brushfoots such as Orion Cecropians, landed all over me, including on my face and arms.

About This Book The first thing to consider when creating a book is “what is the purpose of the book?” The purpose of this book is to help people to identify the butterflies of Mexico and Central America. Very little information that is not directly related to this purpose is included. In addition, all text is embedded in the photographs, allowing swift access in the field. By separating biological information from field identification plates, one can create a more portable, affordable, field guide. Other key considerations for field guide effec- tiveness are image size and layout. At one end of the spectrum, some authors place one species per plate. Although this allows illustrations of butterflies at a large size, it doesn’t allow one to easily compare many similar species. At the other end of the spectrum, species are sometimes illustrated at sizes that are too small to see the features one needs to identify the butterflies. In this work, I have tried to steer a middle course. When possible, similar species are placed so that the reader can view and compare all of them at once. For example, all the blue heliconians appear on a two-page spread. In some cases, for example, “blue” leafwings, groups include too many similar species to place on a single two-page spread. In order to facilitate identification, images of butterflies are arranged so that similar species can be easily compared. This means rotating images to align the butterflies in similar orientations so that you don’t need to perform mental gymnastics to compare them.

Previous Page: A mudpuddle collection in Chiapas. Visible are Polydamus Swallowtails, Florida Whites, Great Southern Whites, Apricot Sulphurs, a Cloud- less Sulphur, Statira Sulphurs, Yellow Angled-Sulphurs, a Little Yellow, a Tailed Orange, Julia Heliconians, Many-banded Daggerwings, a Ruddy Daggerwing, and a Tailed Cecropian. 1 INTRODUCTION Whenever possible, the images are of live butterflies, taken in the field (photos of more than 1500 living species are shown). Living butterflies hold their wings in positions that differ from those of mounted museum specimens. For example, an obvious difference is that live grass-skippers spread their hindwings flat but open their forewings only partially, appearing very different from completely flat-spread museum specimens. A less often noted difference is that when landed with their wings closed, living skippers fold under the trailing edge of their hindwings, hiding about one-fifth of the wing. Thus, the shape of the hindwing, especially the length versus the width, may appear very different in the field than on museum specimens. In addition, relatively fresh individuals often have distinctive sheens that are useful for identification; these sheens are lost upon aging and after death, as are some markings on the butterfly’s body and especially its eye color. All photographs of live butterflies taken by the author are of wild, unrestrained butterflies, except for five photographs taken in butterfly zoos. In the great majority of cases, photographs are of butterflies in Mexico or Central America, because in some cases, populations of the same species flying outside of this area have a slightly (or sometimes greatly) different appearance. A minority of photographs were taken of butterflies that were flying in the United States or South America. When good quality photographs of live butterflies of a particular species were not available, then I photographed museum specimens. Museum specimens were normally photographed with a blue background, which was then removed in Photoshop and replaced with a green (usually) or other-colored background. The bluish tint around the fringes of some of the specimen images is an artifact of the original blue background. Please note that many of the museum specimens illustrated have pin holes and tears in their wings. These holes and tears can usually be distinguished from actual wing spots because they are usually not symmetrical on both wings (on topside views) and because the background color can be seen through the hole. Most users of this guide will already be familiar with butterflies and butterfly identification. Beginners should consultBut- terflies of North America (Barnes & Noble, 2002) or A Swift Guide to Butterflies of North America (Princeton University Press, 2016) for general information about butterflies. This work should allow you to identify to species most Mexican and Central America butterflies that you see (well) or photograph (clearly). However, there will be many groups that are difficult and some that will be close to impossible (see scintillant metalmarks and many of the brown-skippers). Prior to this guide, the only other modern work that attempted something like a complete treatment of a Central or South American country’s butterfly fauna was the two-volumeButterflies of Costa Rica by Phil DeVries. Understandably, that work did not include hairstreaks or skippers, because so little has been known about these groups. I have included hairstreaks and skippers, with the understanding that some of the illustrations will be mislabeled and that some of the suggested field marks, based upon limited sample size, will prove to be unreliable. In a fair number of cases you will have to settle for identifying the butterfly as one of a small number of possible species, because, so far as I am aware, present knowledge doesn’t allow us to separate these species in the field. In most of these cases photos of museum specimens identified as the species in question are shown without any other identification information, and the identification given should be considered to be tentative.

2 INTRODUCTION For many difficult to identify butterflies, especially hairstreaks and skippers, it is useful to note the size of the butterfly, using the length of the forewing (measured from its base to its apex). Obviously, in the field you will not be able to measure the exact size of the wing, but with experience you should be able to estimate the approximate size or realize that it is the same size, or larger, or smaller, than other species with which you are familiar. It will also ultimately prove to be useful to note the elevation at which your unknown butterfly is seen, as many species have distinct elevational ranges, although there is little detailed information available yet for most Mexican and Central American species.

Names My intention in all cases, if a species has been given a name by the NABA Names Com- mittee, is to use both the English and scientific names used on theNABA Checklist and English Names of North American Butterflies(Cassie, et al. 2001)(available at www. naba.org). For species not on the NABA Checklist, I generally use the scientific names of the Atlas of Neotropical Butterflies (Lamas, G. [Ed.] 2004). Exceptions to the Atlas scientific names are for genus names where the NABA Checklist clearly intends another genus name. For example, Dark Kite-Swallowtail is Eurytides philolaus on the NABA Checklist, but placed in the split genus Protographium in the Atlas. Its relative, Short- lined Kite-Swallowtail, is not on the NABA list, but would clearly be placed in the genus Eurytides, not Protographium, and so I treat it as Eurytides agesilaus. For species not on the NABA Checklist, in most cases I have had to create an English name. Many of these names were created over the past twenty years for participants in Sunstreak Tours trips (www.sunstreaktours.com). When creating English names for butterflies I have tried, where possible, to coin names that would provide some identification help in the field. Some, such as Orange Kite-Swallowtail Eurytides( thyastes), are obvious choices. Others, such as Pointer Sister (Adelpha iphiclus), where the name refers to a small “pointer,” an extension on the forewing apical orange spot that is usually lacking on other sisters, are less obvious but very useful nonetheless. Because a mix of name types may be the easiest to learn, I also created names that refer to areas where the species is found (Oaxacan Swallowtail, Papilio esperanza), or refer to the person who described the species (Freeman’s Firetip, Pyrrhopyge hoffmanni). There are whimsical names, such as Phat Redhead, Esthemopsis pherephatte (Species in this group have red heads and phat is street patois for really good) or, in desperation, refer to the scientific name of the species (I say “in desperation” because some of these names are difficult to pronounce and because if and when the scientific name is changed — which happens quite often — then the English name is left referring to a void). In creating these names, I considered the butterfly fauna of the entire Neotropics, not just of Mexico and Central America, because butterfliers will eventually want to have English names for all these species. For example, Callicore pitheas is striped brilliant red and black above, unlike any other Mexican eighty-eight. But, there are quite a few other species of non-Mexican eighty-eights that have similar patterns above. There- fore, I named this species, Two-eyed Eighty-eight, referring to a unique pattern on its underside. Also, in considering the entire Neotropics, new group names for butterflies, 3 INTRODUCTION

Mexican Ecosystems After De la Maza, R. Mariposas Mexicanas. 1987. Fondo de Cultura Economica S.A. de C.V. p.g 52.

Pliego, P. E., Siguenza, A.G.N and Peterson, A.T. 1993. A Geographic, Ecological, and Historical Analysis of land bird diversity in Mexico. pg 286. In Biological Diversity in Mexico. Origins and Distribution. Ramamoorthy et al. (Eds.). Oxford University Press

Picachos

Chipinque Dr. Arroyo & Galeana Zacualtipan

Canon del Novillo Orizaba Gomez Farias Xilitla La Esperanza Mimbres Teocelo Patla Uxpanapa Chapala desert/scrub Catemaco Chamala seasonal forest Uruapan Palenque Bonampak pine-oak forest Chilpancingo El Faisanal Chajul San Agustin Etla cloud forest Santa Rosa Tuxpan Santa Maria tropical semi-evergreen forest Oaxaca Chimalapa Mapastepec Arriaga tropical evergreen forest Matias Romero

especially for hairstreaks, metalmarks and skippers, needed to be coined to deal with the thousands of species in these groups. For quite a few hairstreak and metalmark group names, the words “streak” (as in greenstreak or dreamstreak) or “mark” (as in bluemark or jewelmark) were chosen as part of the group name to quickly convey the fact that species in these groups were hairstreaks or metalmarks. For example, Baeotis zonata is a small metalmark with a pattern of rather pale yellow and faded black and no other Mexican metalmark has a very similar pattern. Thus, a name based on this pattern might have been chosen, but, as it turns out, there are around 20 additional species in the genus Baeotis with similar patterns found throughout the Neotropics. Accordingly, the group “yellowmarks” was chosen and Baeotis zonata was named the Square-spotted Yellow- mark. For the most part, as with the yellowmarks, group names were given to related species, usually a genus, that a knowledgeable observer would recognize as belonging to that group, even if the observer was unfamiliar with the species. For some groups, the group name helps in identification. Beamers (genusPhocides ) are named for radiating “beams” of turquoise on the base of the forewings. Very similar species of blue firetips lack these “beams.” Complete checklist of Mexican and Central American butterflies can be seen and downloaded from www.sunstreaktours.com.

4 INTRODUCTION About the Plates

FIELD MARKS A red line draws your attention to a feature that may be helpful in identifying this species

SIZE The length of the blue/white bar that is placed directly above the name of the species, is the actual length of the FW, measured from its base to its apex.

CATERPILLAR Acacia If a caterpillar foodplant or foodplants have been reported for a spe- FOODPLANT cies, the name of the foodplant species, genus, or family is given in a green color rectangle. If all species on a page use the same group of foodplants, that information is given at the top right of the page.

HABITAT pine-oak If a species seems to be restricted to special habitats, that is indicated by a gray color rectangle.

FLIGHT April-May Unlike most United States species, whose flight period is restricted to PERIOD a short period of the year, the great majority of Mexican and Central American species can be found flying throughout much of the year. In cases where a species’ flight time seems to be restricted to a small por- tion of the year, that is indicated by a black color rectangle.

RANGE Colima Swallowtail A green color-wash over the species name indicates that the species Battus eracon is, so far as known, endemic to Mexico, although the species, e.g., Sonoran Marble, may stray across a border. Quite a few of the species that are shown ranging to the Guatemalan border will eventually be found in Guatemala.

Desert Orangetip A blue color-wash over the species name indicates that the species Anthocharis cethura is found only in Mexico and the United States and, in some cases, Canada.

Pearl Crescent A blue to yellow color-wash over the species name indicates that, in Phyciodes tharos addition to occurring in Mexico and the United States (and, in some cases, Canada), the species ranges south of Mexico, but not as far south as Costa Rica.

Beautiful Beamer A yellow color-wash over the species name indicates that, in addition Phocides belus to occurring in Mexico, the species ranges south of Mexico, but not as far south as Costa Rica.

Victorine Swallowtail An orange color-wash over the species name indicates that, in addition Papilio victorinus to occurring in Mexico, the species ranges south to Costa Rica, but not to Panama.

Blazing Skipper No color-wash over the species name indicates that in addition to oc- Vinpeius tinga curring in Mexico, the species’ ranges south to at least Panama (many species range south throughout tropical America).

PHOTOGRAPHS Photographs are of both live butterflies and of museum specimens. Almost all photographs of live butterflies (about 1500 species) were taken in the field, about seven were taken in butterfly zoos. All photographs of live butterflies taken by the author are of unrestrained, unmanipulated individuals. Museum specimens are shown with solid backgrounds (mostly pale green, sometimes yellow or gray). 5 INTRODUCTION

Abbreviations and Glossary (see page 7 for Mexican state abbreviations) costa the leading edge of the forewing FW forewing ground ground color (the main background color) Grstrk greatstreak Hairstrk hairstreak HW hindwing lead the leading edge of the hindwing MF multiple familes (used as foodplants) MM metalmark pg page pm postmedian (past the middle of the wing, away from the body) pop. population scrophs Scrophularaceae sm submarginal (not actually at the wing margin, but a short way inward) sk. skipper sp. species ssp. subspecies SW southwest trail the trailing edge of the hindwing w. west w/ with w/o without

About the Maps Despite the considerable efforts of many people, the distribution of Mexican butterflies is still not well-known. The range maps in this book are, in most cases, very general representations of where particular species are likely to be found. I have constructed them based upon locality labels on museum specimens, upon information in published books and articles and on the web, and upon my personal experience. They are best viewed as providing information about whether a species is mainly restricted to northern, western, eastern or southern Mexico, rather than allowing one to pinpoint exactly where a species is found. Even so, it is almost certain that some species shown as occurring only in eastern or western Mexico will eventually be found to occur more widely. A red question mark on a map indicates that the species has been reported from this area but that given the other information about the species’ range, it is possible that the report is in error. Reported occurrences are sometimes in error because of misidentification of the butterfly, because of mislabeling the location where a museum specimen was collected, or because the location label was ambiguous (for example, a village name found in more than one state). A blue question mark on a map indicates that although the species has not been reported from this area, it possibly occurs there. Some species are illustrated that have not been reported, as yet, from Mexico. Instead of a range map, the name of the northernmost country from which the species has been reported is given. Many of these species probably occur in southeastern Mexico.

6 INTRODUCTION

BCN

Common Map Views SON and Locations of CHH BCS COA Mexican States DUR NLE SIN ZAC TAM NAY SLP

QUE HID YUC QRO

MEX DFE MOR

AGU TLA COL CAM GUA PUE JAL VER TAB

GUE COL OAX CHP MIC

Abbreviations AGU: Aguascalientes MOR: Morelos A green area indicates that the species is known to oc- BCN: Baja Cal. Norte NAY: Nayarit cur here or that I believe that it is likely to occur here. BCS: Baja Cal. Sur NLE: Nuevo Leon CAM: Campeche OAX: Oaxaca COA: Coahuila PUE: Puebla A red area indicates that the species is known to occur COL: Colima QRO: Quinana Roo here but that it appears to be genuinely rare. CHH: Chihuahua QUE: Queretaro CHP: Chiapas SIN: Sinaloa indicates that this species has been reported DFE: Distrito Federal SLP: San Luis Potosi ? DUR: Durango SON: Sonora from this area but that it is possible that the report GUA: Guanajuato TAB: Tabasco is in error. GUE: Guerrero TAM: Tamaulipas HID: Hidalgo TLA: Tlaxcala indicates that although this species has not been JAL: Jalisco VER: Veracruz ? MEX: Mexico YUC: Yucatan reported from this area, it may occur here. MIC: Michoacan ZAC: Zacatecas 7 SWALLOWTAILS Archaic and Battus Swallowtails

Acacia elongated pale spot along costal margin

May- July

acacia forest

Archaic Swallowtail Baronia brevicornis Aristo- lochia

one row of large orange spots

iridescent iridescent blue blue Pipevine Swallowtail Battus philenor Aristo- lochia

cream

red spots

no marginal pale spots Polydamas Swallowtail Battus polydamas Aristo- lochia mainly July- Oct

marginal pale spots blue spots spots yellow green- ish spots males w/ marginal yellow pale spots abdomen Colima Swallowtail Battus eracon 8