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 distribu- tion 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 in- cluded. 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).

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