Owlet Caterpillars of Eastern North America [Course Book ed.] 9781400838295

Table of contents :
Contents
Preface
Acknowledgments
Photographic Credits
Introduction
Quadrifine Owlets (Family Erebidae)
Litter Moths (Subfamily Herminiinae)
Pangraptines (Subfamily Pangraptinae)
Snouts (Subfamily Hypeninae)
Straws (Subfamily Rivulinae)
Scalloped Moths (Subfamily Scoliopteryginae)
Fruitpiercing Moths (Subfamily Calpinae)
Necklace Moths (Subfamily Hypocalinae)
Scolecocampines (Subfamily Scolecocampinae)
Halflings (Subfamily Hypenodinae)
Fungus Moths (Subfamily Boletobiinae)
Phytometrines (Subfamily Phytometrinae)
Underwings, Zales, Witches, and Kin
Underwing Moths
Lesser Underwings
Grass Loopers
Chocolates and Kin
Wavy-lined Owlets
Witches
Eulepidotines
Elves (Family Euteliidae)
Nolas and Punkies (Family Nolidae)
Trifine Owlets (Family Noctuidae)
Loopers (Subfamily Plusiinae)
Tumblers (Subfamily Bagisarinae)
Resplendent Moths (Subfamily Cydosiinae)
Eustrotiines (Subfamily Eustrotiinae)
Bird-dropping Moths (Subfamily Acontiinae)
Pantheas and Yellowhorns (Subfamily Pantheinae)
Hieroglyphic Moths (Subfamily Diphtherinae)
Brothers (Subfamily Dilobinae)
Balsas (Subfamily Balsinae)
Daggers (Subfamily Acronictinae)
Foresters (Subfamily Agaristinae)
Paints or Hooded Owlets (Subfamily Cuculliinae)
Sallows, Brocades, and Kin (Subfamily Oncocnemidinae)
Amphipyrines (Subfamily Amphipyrinae)
Seedcoppers (Subfamily Stiriinae)
Sun Moths (Subfamily Heliothinae)
Groundlings and Kin (Subfamily Condicinae)
Fern Moths (Subfamily Eriopinae)
Cutworms, Lash-eyed Sallows, Sallows, and Kin (Subfamily Noctuinae)
Miranda Moth
Marvels
Midgets
Armyworms
Caradrines
Dipterygiines
St. Johnswort Moths
Angle Shades
Borers
Divers
Pinions, Sallows, and Kin
Pinions
Glaeas
Tentmakers
Antitypine Sallows
Griffins and Satyrs
Quakers, Woodgrains, and Kin
Bronzed Cutworms
Arches And Kin
Wainscots And Kin
Groundcats
Spanish Moths
Cutworms And Darts
Glossary
Cited Literature
Foodplant Index
Species And Subject Index

Citation preview

Owlet Caterpillars of

Eastern North America David L. Wagner, Dale F. Schweitzer, J. Bolling Sullivan, and Richard C. Reardon

With special acknowledgment for support received from the United States Department of Agriculture Forest Service FHTET and Discover Life in America

Princeton University Press Princeton and Oxford

This work is dedicated to our late friends and colleagues Douglas C. Ferguson and John G. Franclemont. Both shared a special interest in noctuoids, their immature stages, and life history studies.

Copyright 2011 © by David L. Wagner Requests for permission to reproduce material from this work should be sent to Permissions, Princeton University Press Published by Princeton University Press, 41 William Street, Princeton, New Jersey 08540 In the United Kingdom: Princeton University Press, 6 Oxford Street, Woodstock, Oxfordshire OX20 1TW press.princeton.edu All Rights Reserved Library of Congress Cataloging-in-Publication Data Owlet caterpillars of eastern North America / David L. Wagner ... [et al.]. — 1st ed.        p. cm.   Companion volume to: Geometroid caterpillars of northeastern and Appalachian forests.   Includes bibliographical references and index.   ISBN 978-0-691-15042-0 (pbk. : alk. paper)  1.  Noctuidae—East (U.S.)— Identification. 2.  Noctuidae—Canada, Eastern—Identification. 3.  Caterpillars— East (U.S.)—Identification. 4.  Caterpillars—Canada, Eastern—Identification.  I. Wagner, David L., 1956- II. Geometroid caterpillars of northeastern and Appalachian forests.   QL561.N7O95 2011   595.78’139097—dc23                                     2011016905 British Library Cataloging-in-Publication Data is available Publication of this book has been aided by support received from the United States Department of Agriculture Forest Service FHTET and Discover Life in America This book has been composed in Minion and Myriad Pro Printed on acid-free paper. Typeset and designed by D & N Publishing, Wiltshire, UK Printed in Singapore 10 9 8 7 6 5 4 3 2 1

Contents Preface4 Acknowledgments  5 Photographic Credits  6 Introduction  9 J Birds and Caterpillars (and

9 Importance of Owlet Caterpillars and Adults  10 J A Note on Nomenclature 11 Morphology  12 Larval Diets  15 Finding Caterpillars  15 Beating and Sweeping  17 Collecting Adults  18 Baiting  19 J Wine Ropes 20 J John Peacock’s Bait Recipe 21 Bait Trapping  21 Collecting, Vouchering, and Conservation 22 Feeding Adults  23 23 Obtaining Pairings  Obtaining and Handling Eggs  23 Rearing Caterpillars  25 Sleeving  26 Overwintering Larvae and Pupae  26 27 Natural Enemies  J Exotic Biological Control Agents 31 Climate Change)

J Guest Essay on Owlet Classification:  

Charles Mitter

Classification and Nomenclature  About This Book  Making Identifications  J Transformers

Voucher Data and Specimens  Diagnosing Owlet Caterpillars  Keys to Owlet Larvae Supplemental Digital Resources   Species Accounts Quadrifine Owlets (Family Erebidae) 

32 33 34 35 36 36 37 37 37

38

Litter Moths (Subfamily Herminiinae)  38 Pangraptines (Subfamily Pangraptinae)  61 Snouts (Subfamily Hypeninae)  63 Straws (Subfamily Rivulinae)  72 Scalloped Moths (Subfamily Scoliopteryginae)  73 J Caterpillar Mimicry 78 Fruitpiercing Moths (Subfamily Calpinae)  79 Necklace Moths (Subfamily Hypocalinae)  82 Scolecocampines (Subfamily Scolecocampinae)  84

Halflings (Subfamily Hypenodinae)  Fungus Moths (Subfamily Boletobiinae)  Phytometrines (Subfamily Phytometrinae)  Underwings, Zales, Witches, and Kin (Subfamily Erebinae)  J Dyar’s Rule

Eulepidotines (Subfamily Eulepidotinae) 

87 89 91 97 153 187

Elves (Family Euteliidae) 

192

Nolas and Punkies (Family Nolidae) 

198

Nolas and Punkies (Subfamily Nolinae)  198 Chloephorines (Subfamily Chloephorines)  204 Baileyas (Subfamily Risobinae)  207 Concanas and Kin (Subfamily Collomeninae) 210 Lichen Punkies (Subfamily Afridinae)  211 Trifine Owlets (Family Noctuidae) 

212

Loopers (Subfamily Plusiinae)  212 Tumblers (Subfamily Bagisarinae)  232 Resplendent Moths (Subfamily Cydosiinae)  235 Eustrotiines (Subfamily Eustrotiinae)  236 Bird-dropping Moths (Subfamily Acontiinae)  239 Pantheas and Yellowhorns (Subfamily Pantheinae)  249 Hieroglyphic Moths (Subfamily Diphtherinae) 254 Brothers (Subfamily Dilobinae)  255 256 Balsas (Subfamily Balsinae)  Daggers (Subfamily Acronictinae)  258 Foresters (Subfamily Agaristinae)  306 Paints or Hooded Owlets (Subfamily Cuculliinae)  311 Sallows, Brocades, and Kin (Subfamily Oncocnemidinae)  318 Amphipyrines (Subfamily Amphipyrinae)  324 Seedcoppers (Subfamily Stiriinae)  339 Sun Moths (Subfamily Heliothinae)  342 Groundlings and Kin (Subfamily Condicinae)  364 Fern Moths (Subfamily Eriopinae)  374 Cutworms, Lash-eyed Sallows, Sallows, and 378 Kin (Subfamily Noctuinae)  J Puddling 447 J Moths and Ears and Bats 476 J Moth Ear Mites 505 Glossary  Cited Literature  Foodplant Index  Species and Subject Index

552 555 563 569

4

Preface Owlet moths are one of our planet’s most successful lineages of organisms—more than 75,000 species are known worldwide, and thousands more remain unnamed. While some owlet caterpillars are important pests in woodlands and forests, agricultural landscapes, gardens, and orchards, the vast majority make a positive contribution to the health of our forests and other wildlands as forage for birds (especially during the nesting season), rodents, and other vertebrates. A few lineages serve as macrodecomposers, consuming fallen leaves and thereby accelerating the cycling of nutrients in wooded landscapes. Adult owlets pollinate many temperate plants and, like their larvae, are important to the diets of birds, bats, and other insectivorous vertebrates (see pages 9 and 10). Literature on North American owlet caterpillars and their life histories is widely scattered and, for many species, lacking. This book was written as a companion volume to Geometroid Caterpillars of Northeastern and Appalachian Forests (Wagner et al. 2002). Geographic coverage is expanded in this work to include the North American owlet fauna found east of the hundredth meridian. The ecological scope is broadened to include all community types where owlet caterpillars are found, including agricultural lands, gardens, and parks. While ostensibly it is an identification guide for caterpillars, we also added images of adults and other life stages, as well as relevant information about natural enemies, chemical ecology, insect behavior, invertebrate conservation, and myriad other aspects of lepidopterology. Collectively, the first three authors have been studying eastern noctuoids and related moths for more than 110 years; much unpublished data has been included. Introductory sections on finding caterpillars, rearing, baiting, and natural enemies are extensive, and include information not easily found elsewhere. Larvae of numerous species are illustrated and described here for the first time. We supply new foodplant records and observations on behavior, phenology, abundance, development, and overwintering stages. Many erroneous foodplant records, some of which have been repeated in the literature for upwards of 100 years, are corrected. Text for nearly 90 species (of the 815 treated in this work) is reproduced from Caterpillars of Eastern North America (Wagner 2005), although in many cases we modified the species account, e.g., by rewriting or adding text to the Remarks section. For nearly all of the shared species, we selected different larval images to illustrate this work so that the two books would be complementary. The text was written to be understandable and useful to educators and the general public, while also serving the needs of professional entomologists, forest managers, extension agents, and conservation biologists. From the outset, we were disposed to writing a book that could serve as a portal into entomology for students and those without training in the sciences. The expanded introductory sections and species remarks were written with these audiences in mind. We propose

many new common names for individual species and higherlevel taxa that more closely reflect current knowledge of evolutionary relationships, taxonomy, foodplant usage, or simply to distance this work from misapplied or ill-chosen names that have been carried forward for too long. An important goal was to assemble a work that would engender a greater appreciation for caterpillars and other aspects of biological diversity. Caterpillars can be attractive creatures and some noctuoids stunningly so. Dagger moth caterpillars are a study in diversity, some awash in reds, oranges, and yellows; cuculliine larvae are reminiscent of hand-painted and kiln-fired ceramic figurines; most foresters (agaristines) are exquisitely rendered with rings of black and orange. For nearly every species, we provide an image. In addition, we collaborated with a host of photographers to supply images of live adults in an attempt to foster an appreciation for living moths as they might be encountered in nature—this is the first book of New World moths to do so in a comprehensive manner. We have done our best to prepare a work that succinctly summarizes current knowledge. However, because many owlets are unstudied, we routinely had to wrestle with uncertainty regarding stated ranges, phenologies, foodplants, and which (if any) salient features would prove diagnostic. In some cases we extrapolated information about a species from admittedly limited collections or experience. The sheer scope of this effort and the preliminary state of our knowledge convince us that species accounts will need to be corrected or in other ways emended— DLW will maintain and make available an erratum and addendum on his website (see the link at http://press.princeton.edu/ titles/9420.html). The site will also include additional species accounts, supplementary images, and other on-line resources. For those willing to walk slowly, turn leaves, scan tree trunks, examine flowers, and venture out into yards and nearby woods, especially at night, there is much to be discovered about owlets and other caterpillars. Photographic studies capturing variation, head capsule patterns, developmental changes, and natural behaviors are needed; parasitoids and natural enemies are incompletely known for all but the most well-studied pest species; and detailed life history observations would add much to the existing literature. Even the casual observer, hobbyist, and student can expect to make worthwhile discoveries about these fabulously diverse and often handsome moths and their caterpillars. This work was originally intended to appear as a United States Forest Service Publication and as noted below much initial funding came from the U.S. Department of Agriculture’s Forest Service’s Forest Health Protection Program. Consequently, in the species accounts we made a special effort to treat all woodland and forest species likely to be of economic or ecological importance, and include accounts for owlet caterpillars of relevance in agricultural landscapes.

Acknowledgments More than 250 people directly contributed to this work—we feel extraordinarily fortunate to be able to coordinate and synthesize so much from so many.  And while this effort draws on collections made and images taken over the past two decades, much of the research and writing occurred over a seven-year period beginning in 2004, a good portion of which was made possible by several years of assistance to DLW by the United States Forest Service’s FHTET Program. Two historical works were quintessential in providing the foundation on which this effort is based: William T. Forbes’s Lepidoptera of New York and Neighboring States (1954, 1960) and Samuel E. Crumb’s The Larvae of the Phalaenidae (1956). Don Lafontaine was abundantly generous with his time over the course of this effort, and provided advice on matters of taxonomy and classification, draft copies of his checklist for North American noctuoids, DNA barcode data, editorial expertise, and common name suggestions. Considerable help with identifications, literature, and life history information was passed along by Dr. Tim McCabe (State Museum of New York), who has been a mentor and colleague to DLW, DFS, and JBS for more than two decades. In addition, Don and Tim graciously allowed us to share their previously unpublished images and life history information for dozens of species that otherwise would not have been represented in this volume. Steve Passoa helped with the Spodoptera text and was an enthusiastic champion of the effort (as he is of so many things involving the immature stages of Lepidoptera). Other colleagues who shared life history data and/or passed along taxonomic advice include Doug Ferguson, John Franclemont, Larry Gall, John Lill, Hugo Kons, Eric Quinter, and John Rawlins. Important distributional and phenological data was contributed James Adams, Vernon Brou, Larry Gall, Ed Knudson, Hugh McGuinness, Don Lafontaine, and Chris Schmidt. Others that sent along references and literature include Steve Passoa, Seth Johnson, Roy Parker, and John Calhoun. One hundred and ninety-one people contributed gravid females, eggs, or larvae for this work—we are especially appreciative of those who made multiple contributions or passed along livestock or images of taxa of special significance. Below we list contributors in decreasing order of numbers of collections that resulted in the 1450 larval images that illustrate this work: John Peacock, Eric Hossler, Tim McCabe, Monty Volovski, Henning von Schmeling, Valerie Giles, Lyle Buss, Laura Miller, Eric Quinter, Don Lafontaine, Jeff Slotten, Mike Thomas, Ryan Wagner, Ben Williams, Jeff Fengler, Fred Hohn, Darryl Willis, Tom Carr, James Wiker, Steve Johnson, Jean-Paul Laplante, Bryan Connolly, Mike Singer, George Godfrey, Mike Nelson, Nelson Bricker, Nancy Lowe, Mark Mello, Moria Robinson, Mary Jane Hatfield, Berry Nall, Scott Smedley, Henry Hensel, David Silsbee, Jessica Lowry, David Dussourd, Douglas Ferguson, Keith Hartan, Julia Joseph, Warren Kiel, John Lill, James Adams, John R. Byers, John DeBenedictis, Barbara Gray, Ron Panzer, Steve Passoa, Jorge Pena, and Jonathan Sylvestre. All others made fewer than three collections that resulted in illustrations in this work. A list of collectors, as well as the associated date and locality data, is posted on http://press. princeton.edu/titles/9420.html. Five colleagues—John Peacock, Eric Quinter, Jeff Slotten, Ben Williams, and Darryl Willis—repeatedly took it upon themselves to rear livestock, from the egg until the last instar, before passing along the caterpillars to DLW.

During the summer months and part time during the rest of the year, we had considerable help rearing caterpillars and maintaining records. Pre-project efforts (prior to 2004) were anchored by Valerie Giles, Eric Hossler, Jenny Jacobs, Julia Joseph, and Monty Volovski. Jess Watson ran DLW’s rearing facility through the summers of 2004–2006, Laura Ann McLoud in 2007, April Rodd in 2008, 2009, 2011, and Moria Robinson in 2009 and 2010. William Forrest reared many of the noctuines contributed by the Canadian National Collection, and most of these from eggs. Christine N. Weber (formerly of Valatie, NY) assisted with Tim McCabe’s caterpillar rearings from 1990 to 2002. Four colleagues, Don Lafontaine, Hugh McGuinness, Steve Roble, and Fred Stehr, our copyeditor, Sheila Dean, and April Rodd, read the text cover to cover and offered innumerable improvements. April’s efforts over the last two months were paramount to the quality of the final effort. Sheila’s attention to detail and knowledge of the Queen’s English were particularly helpful. Others who graciously read the draft or galley text, listed roughly in order of number of pages read, include Aurora Edington, Ben Gagliardi, Moria Robinson, Eric Hossler, Peri Mason, Angela Smilanich, Purbita Saha, Weston Henry, Judy Semroc, Christine Langdon, Bonnie Caruthers, Jim Wiker, Tony McBride, Ian Gauld (ichneumonids), Scott Shaw (braconids), Michael Singer, John Stireman (tachinids), and David Wall (ichneumonids). Ellen Foos and Robert Kirk, my editors at Princeton University Press, guided the effort and provided much expertise. April Rodd, Shawn Deford, Shawn Binns, Ben Gagliardi, Katie Rymsza, Brigette Zacharczenko, and Honey Suddapali checked the spelling and application of the scientific and common names. Namrita and David Price-Goodfellow, April Rodd, and Ben Gagliardi helped shoulder the weight of indexing. Much of the line art and image scanning was done by DLW’s daughter, Virginia. She and Virge Kask designed the cover; Virge carried out the work of image processing and layout for the cover. Additional line drawings were supplied by Valerie Giles and Amy Fernald. Preparing the color images that appear in this book was a colossal task. Shawn Kennedy processed, curated, and otherwise managed nearly 1900 larval and pinned adult images. Rene Twarkins and Amy Fernald were responsible for the same tasks for the live adult images. Rene spent many nights assembling the Catocala head plate. Namrita and David Price-Goodfellow, and Peter Davies at D & N Publishing, did an exceptional job with the design—we appreciate their efforts, patience, and skill, as well as their willingness to work with us through the entire publication process. Bryan Connolly collected and delivered foliage for a number of scarce foodplants. Ken King, Roger Hammer, Michael Heep, Les Mehrhoff, and Ken Metzler helped with plant identifications. They also provided directions to hosts of particular importance. Foliage for winter and early spring collections and foodplants that do not grow in Connecticut were supplied by the University of Connecticut greenhouses. Clint Morse, the greenhouse manager, could not have been more helpful. Ernie Cowan and Jim Duquesnel (Florida Keys), Keith Langdon and Sue Powell (Great Smoky Mountains National Park), Jimmy Paz (Audubon’s Sabal Palm Grove Audubon Center and Sanctuary), Donna Watkins (Florida State Parks), and staff at both the Connecticut Department of Environmental Protection and Massachusetts Division of Fisheries and Wildlife shepherded

5

6 Acknowledgments collecting permit requests. Their efforts significantly enhanced the coverage of the work. Principal financial support for this work came from two awards from the U.S. Department of Agriculture, Forest Services, Forest Health Technology Enterprise Team (Cooperative agreements 01-CA-11244225-215 and 10-CA-11420004-130). Additional support, lodging, and other assistance came from Discover Life in America’s “All Taxa Biodiversity Inventory” in Great Smoky Mountains National Park. Special thanks to Jeanie Hilten, Keith Langdon, and Becky Nichols who coordinated visits; Keith and Becky also accompanied us on outings, contributed samples, and poured on encouragement. Contracts from the Con-

necticut Department of Environmental Protection Long Island Sound Fund, Endangered Species/Wildlife Income Tax Check-off Fund, and State Wildlife Grants program, and from the Connecticut Chapter of The Nature Conservancy added dozens of species to this effort that otherwise would have been missed. An award from the College of Liberal Arts and Sciences at the University of Connecticut offset publication costs, and a second award from UConn’s Research Foundation funded the preparation of artwork. We thank our wives, Sylvia, Terry, Ashley, and Ann. Without their support this book, and especially much of the fieldwork and rearing efforts that are the foundation for this work, would not have been possible.

Photographic Credits The 2250 images that illustrate this work were taken over two decades by more than 100 photographers. Professional taxonomists, career entomologists, amateur collectors, students, foresters, extension agents, conservation biologists, and a small legion of nature photographers contributed. Many of those that sent images have become and will long remain colleagues and friends—we are fortunate indeed to have been able to work with so many. Credits are broken down by category below. Two-thirds of the larval images were taken by DLW and many of the remaining by students or technicians working in DLW’s lab (Hartan, Herrick, Hohn, Hossler, Jacobs, Joseph, McLoud, Robinson, Rota, Rymsza, Simmonds, Suddapalli, and Zacharczenko). Seventy-three people or institutes contributed larval images for this book. Cover credits appear elsewhere. Locality and other data associated with each image are posted on the senior author’s website (http://press.princeton.edu/titles/9420.html). We are much indebted to April Rodd, Tegan Wheeler, Sabina Perkins, and especially April for assembling, transcribing, and synthesizing the collection and image credit data for each of these images. Of special significance are the numerous contributions of Don Lafontaine at the Canadian National Collection and Tim McCabe at the State Museum of New York, who allowed us to use images of numerous species that have not been previously published, several of which represent unusual, rare, or otherwise poorly known taxa. Valerie Giles was an early catalyst for DLW’s interest in macrolepidopteran caterpillars—hundreds of her images taken between 1994 and 1997 anchor this effort. Lyle Buss was generous with his time and images, and substantially bolstered our representation of Spodoptera armyworms. Those contributing larval images to the body of this book are listed individually alphabetically below. DAVID ALMQUIST: Idia gopheri; LESLIE ANGEL: Elaphria grata; THÉRÈSE ARCAND (CANADIAN FORESTRY SERVICE): Achatodes zeae, Acronicta quadrata, A. tristis, Amphipyra tragopoginis, Anathix puta, Apamea devastator, Enargia decolor (2), E. fausta, Lacinipolia lorea, Polia purpurissata; STEVE BAKER: Stiria rugifrons (2); GIFF BEATON: Condica confederata; HERBERT BECK: Mesapamea fractilinea; THOMAS BENTLEY: Cucullia speyeri; CHARLES BORDELON: Androloma disparata; BOB BOWLES: Bellura obliqua; RICHARD BROWN: Acronicta near longa; VALERIE BUGH: Cerathosia tricolor,

Neogalea sunia; PAT BURKETT: Harrisimemna trisignata; LYLE BUSS: Acronicta clarescens, Anticarsia gemmatalis (2), Argyrogramma verruca, Catocala alabamae, Elaphria nucicolora, Litoprosopus futilis, Meganola new species, Morrisonia mucens, Noctua pronuba, Paectes nubifera, Spodoptera albula, S. dolichos (3), S. eridania (2), S. exigua, S. latifascia (3), S. ornithogalli; CANADIAN NATIONAL COLLECTION: Abagrotis bruneipennis, Actebia fennica, Agrotis venerabilis, Anaplectoides prasina, A. pressus, Anarta trifolii, Autographa mappa, A. rubidus, Brachionycta borealis, Caenurgina erechtea, Callopistria cordata, Catabena lineolata, Cerastis tenebrifera, Chytonix palliatricula, Coranarta luteola, Diarsia jucunda, Enargia infumata, Euclidia cuspidea, Eueretagrotis attenta, Eurois astricta, E. occulta, Euxoa detersa, E. messoria, E. pleuritica, E. velleripennis, Feltia geniculata, F. jaculifera complex (2), F. manifesta, F. tricosa, Graphiphora augur, Hemipachnobia monchromatea (2), Homoglaea hircina, Idia julia, Lithophane tepida, L. thaxteri, Lycophotia phyllophora (2), Macrochilo orciferalis, Macronoctua onusta, Mamestra configurata, Mythimna oxygala, Oligia chlorostigma, Orthosia revicta, Peridroma saucia (2), Plusia contexta, Protolampra brunneicollis, Pseudeva purpurigera, Pyrrhia exprimens (2), Sideridis rosea, Spaelotis clandestina (2), Sympistis badistriga, S. dentata, S. saudersiana, Syngrapha abstrusa (2), S. alias, S. epigaea, S. microgamma, S. montana (2), S. octoscripta (2), S. rectangula, S. selecta, S. u-aureum, Xestia badicollis (3), X. bicarnea, X. smithii, X. youngii, Xylena curvimacula (2), Xylotype arcadia, Zale minerea; JAMES CASTNER: Anticarsia gemmatalis, Litoprosopus futilis; PAUL CHOATE: Anticarsia gemmatalis; DALE CLARK: Ascalapha odorata; RAYMOND COREY: Androloma maccullochii; DAVID DUSSOURD: Amphipyra tragopoginis, Anagrapha falcifera; JEFF FENGLER: Acronicta lanceolaria, Nola pustulata, Orthosia rubescens, Panthea furcilla, Selenisa sueroides, Syngrapha viridisigma, Xestia elimata (2), Zale duplicata, Z. helata, Zanclognatha dentata; DOUGLAS FERGUSON: Acronicta rubricoma, Cutina albopunctella, Helicoverpa zea, Phoberia ingenua; ALEXANDRE FOURNIER: Alypia langtoni, Ufeus satyricus; BILL FRANK: Homophoberia cristata; LARRY GALL: Catocala desdemona; GRANT GENTRY: Callopistria floridensis, Selenisa sueroides; VALERIE GILES: Abagrotis alternata, Acronicta afflicta (2), A. hastulifera, A. impleta (2), A. noctivaga, Allagrapha aerea, Amphipyra glabella, A. pyramidoides, Anicla illapsa, Anterastria

Photographic Credits teratophora, Autographa precationis, Callopistria mollissima, Calophasia lunula (2), Calyptra canadensis, Catocala antinympha (2), C. blandula, C. epione, C. gracilis, C. insolabilis, C. luctuosa, C. marmorata, C. meskei (2), C. muliercula (2), Chytonix palliatricula (3), Cucullia intermedia, Eucirroedia pampina, Eudryas grata, E. unio, Euparthenos nubilis, Eupsilia tristigmata, Heliothis virescens, Hyppa xylinoides, Idia aemula, I. americalis, I. lubricalis, Lithophane laticinerea (2), Loscopia velata, Noctua pronuba (2), Nola pustulata, Ochropleura implecta (2), Orthosia alurina (3), O. hibisci (2), O. revicta, Palthis angulalis, P. asopialis, Parallelia bistriaris, Phalaenophana pyramusalis, Plusia putnami (2), Polia imbrifera, Protodeltote muscosula, Psaphida styracis, Scolecocampa liburna, Sericaglaea signata, Simyra insularis, Spodoptera ornithogalli, Sympistis chionanthi, Tarache aprica (2), Trichodestra legitima, Ulolonche culea, Xestia dolosa (2), X. smithii, Zale buchholzi, Zale intenta (2), Z. metata, Z. submediana, Zanclognatha cruralis (2); GEORGE GODFREY: Hydraecia immanis, Leucania commoides, L. insueta, L. multilinea, L. phragmitidicola, L. pseudargyria, L. scirpicola; PETER HAGGARD: Acronicta funeralis; KEITH HARTAN: Catocala relicta; MARY JANE HATFIELD: Helicoverpa zea (predation), Lithophane bethunei (eating scale insect), Schinia grandimedia, Tricholita signata; FRED HOHN: Chytolita morbidalis; ERIC HOSSLER: Abrostola urentis, Achatia distincta, Acronicta impleta, A. interrupta, A. ovata, A. retardata (2), Allotria elonympha, Catocala minuta, C. piatrix, Charadra deridens, Egira alternans, Galgula partita (2), Isogona tenuis, Lithophane georgii, L. innominata, L. viridipallens, Morrisonia confusa (2), Nephelodes minians, Orthosia hibisci, Panthea acronyctoides (2), Parallelia bistriaris, Phlogophora periculosa, Phosphila miseloides, P. turbulenta (2), Polygrammate hebraeicum, Pyrrhia cilisca, Spodoptera ornithogalli, Xestia dolosa, Zale helata (3), Z. metatoides, Z. undularis; INSTITUTO DE CIENCAS AGRARIAS: Spodoptera littoralis; JENNY JACOBS: Crocigrapha normani, Hypena humuli, Ponometia erastrioides, Lithophane unimoda, Xystopeplus rufago; SAMUEL JAFFE: Catocala badia; KELLY JONES: Zale perculta; JULIA JOSEPH: Cryptocala acadiensis, Idia diminuendis, Lacinipolia olivacea, L. renigera, Macrochilo litophora, Panthea furcilla, Renia factiosalis; KENPEI (WIKIPEDIA COMMONS): Spodoptera litura; KEN AND SELENA KING: Diphthera festiva; JOHN LILL: Acronicta funeralis, A. increta (beech);TIM MCCABE: Abablemma brimleyana, Amolita fessa, Arugisa lutea, Catabena lineolata, Cerastis tenebrifera, Chrysanympha formosa, Chytonix sensilis, Dargida diffusa (3), Deltote bellicula, Elaphria alapallida, E. festivoides, Eutelia pulcherrimus (2), Idia concisa, I. laurentii, Lacanobia subjuncta, Lesmone detrahens, Motya abseuzalis, Nigetia formosalis, Orthodes cynica, Pachypolia atricornis, Platypolia anceps, Pseudorthodes vecors, Sigela brauneata, Spirameter lutra; LAURA ANN MCLOUD: Catocala amica (2), C. muliercula, C. relicta (2), Lithophane querquera, Papaipema arctivorens, P. cataphracta; BERRY NALL: Amyna octo, Bagisara tristicta, Cropia templada (2), Diastema tigris, Melipotis acontioides; Ponometia semiflava; MIKE NELSON: Acronicta lanceolaria, Catocala pretiosa, Pyrrhia aurantiago, Zale lunifera; STEFAN NESER, ARC PLANT PROTECTION: Bellura densa; NORTH CAROLINA STATE UNIVERSITY EXTENSION: Alabama argillacea; OHIO AGRICULTURAL RESEARCH AND DEVELOPMENT: Abagrotis orbis; RON PANZER: Schinia gaurae; STEVE PASSOA: Alabama

argillacea, Spodoptera exigua; JEAN-CLAUDE PETIT: Cucullia postera (2); JOHN PICKERING: Lithophane querquera (eating Alsophila); ROBERT PILLA: Cydosia nobilitella; BENJAMIN POWELL: Bellura densa; NOBLE PROCTOR: Melanchra picta; MARLIN RICE: Spodoptera albula, Striacosta albicosta; MORIA ROBINSON: Acronicta albarufa, A. haesitata (2), A. lobeliae, A. radcliffei, A. vinnula, Bleptina inferior, Catocala ilia, C. serena, Crocigrapha normani, Elaphria georgii, Hadena ectypa (3), Hypenula cacuminalis, Hyperstrotia villificans, Maliattha synochitis, Metalectra albilinea, M. discalis, M. quadrisignata, M. tantillus, Niphonyx segregata, Orthosia revicta, Panopoda rufimargo, Phytometra rhodarialis, Spiloloma lunilinea (2); APRIL RODD: Idia scobialis; JADRANKA ROTA: Eupsilia new species; KATIE RYMZSA: Ponometia erastrioides; JUDY SEMROC: Acronicta funeralis, A. impleta; MERLE SHEPARD: Diphthera festiva (2); KIMMO SILVONEN: Agrotis volubilis, Lateroligia ophiogramma, Oligia strigilis, Rhizedra lutosa; DENISE SIMMONDS: Acronicta oblinita, Melanchra adjuncta, Schinia nubila; JEFF SLOTTEN: Catocala agrippina, C. atocala, Schinia mitis; SPANDANA SUDDAPALLI: Scolecocampa liburna; MIKE THOMAS: Acronicta radcliffei, A. rubricoma, Catocala illecta, Cucullia convexipennis (2), Psaphida resumens (eggs); JIM TUTTLE: Catocala parta; HENNING VON SCHMELING: Harrisimemna trisignata, Helicoverpa zea, Ponometia erastrioides, Spragueia apicalis, S. leo (2); REGINALD WEBSTER: Lithophane thujae; KIRBY WOLFE: Spodoptera androgea (3); BRIGETTE ZACHARCZENCKO: Mythimna unipuncta. Forty-three photographers contributed images of live adults. Hugh McGuinness and Lynn Scott sent the lion’s share: their efforts represent hundreds of hours at lights photographing adults and even more time curating their image holdings. Steve Walter and David Silsbee fully embraced our effort and generously offered use of their many images of living moths. We want to acknowledge Bob Patterson and his extraordinary efforts with the Moth Photographer’s Group (http://mothphotographersgroup.msstate.edu/MainMenu.shtml)—through this community site and BugGuide (http://bugguide.net/node/view/15740) we were able to track down photographers and images for all but two of the 372 species that were treated in a full species account. Individual credits follow (listed alphabetically). LISA AMES: Hypsoropha monilis, Spragueia dama; LESLIE ANGEL: Catocala vidua; BETSY BETROS: Stiria rugifrons; CHARLES BORDELON: Spiloloma lunilinea; BOB BORTH: Catocala amestris, C. serena; VALERIE BUGH: Acronicta vinnula, Alypia octomaculata, Melipotis acontioides, Spragueia guttata; PAT BURKETT: Harrisimemna trisignata; JONATHAN BURISHKIN: Zale metatoides; LYLE BUSS: Spodoptera eridania, S. latifascia; ILLIA-SHAUNDRA CHAVEZ: Melanchra assimilis; STEPHEN CRESSWELL: Phytometra rhodarialis; JOHN DAVIS: Lithophane georgii; MARK DREILING: Acronicta connecta, A. longa, A. rubricoma; MELINDA FAWVER: Pyrrhia cilisca; ERIC GOFREED: Thioptera nigrofimbria; LOUIS HANFIELD: Cucullia speyeri; BILL JOHNSON: Acronicta dactylina, A. haesitata, Allotria elonympha, Catocala coccinata, C. concumbens, Hypena baltimoralis, Litoprosopus futilis, Meganola minuscula, Noctua pronuba; MERRILL LYNCH: Baileya levitans, Pseudeustrotia carneola; STAN MALCOLM: Callopistria mollissima; JIM MCCLARIN: Tarache tetragona; HUGH MCGUINNESS: Abagrotis alternata, A. magnicupida, Acronicta afflicta, A.

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8 Photographic Credits americana, A. clarescens, A. dolli, A. exilis, A. fragilis, A. increta, A. innotata, A. interrupta, A. lithospila, A. modica, A.morula, A. ovata, A. tritona, Afrida ydatodes, Agnorisma badinodis, Agriopodes fallax, Agrotis ipsilon, Amphipyra pyramidoides, Anagrapha falcifera, Anticarsia gemmatalis, Anicla illapsa, A. infecta, Anomis privata, Argyrostrotis anilis, Autographa ampla, A. precationis, Bleptina caradrinalis, Callopistria cordata, Catocala amica, C. andromedae, C. ilia, C. palaeogama, Chaetaglaea sericea, C. tremula, Charadra deridens, Chytolita petrealis, Chytonix palliatricula, Cissusa spadix, Comachara cadburyi, Condica sutor, Cosmia calami, Cucullia asteroides, C. intermedia, Cutina albopunctella, Egira alternans, Elaphira versicolor, Euplexia benesimilis, Eupsilia morrisoni, E. sidus, E. tristigmata, E. vinulenta, Fagitana littera, Feltia subterranea, Helicoverpa zea, Himella intractata, Hypena eductalis, H. scabra, Idia lubricalis, Lacanobia grandis, Lacinipolia renigera, Lascoria ambigualis, Leucania adjuta, L. linita, Lithophane patefacta, L. unimoda, L. viridipallens, Loscopia velata, Lycophotia phyllophora, Macrochilo orciferalis, Marathyssa inficita, Meganola phylla, Melipotis indomita, Metaxaglaea violacea, Mocis texana, Morrisonia confusa, M. evicta, Nephelodes minians, Nola clethrae, Ochropleura implecta, Ogdoconta cinereola, Orthosia revicta, Palthis angulalis, P. asopialis, Pangrapta decoralis, Panthea furcilla, Panopoda rufimargo, Papaipema arctivorens, Parallelia bistriaris, Phlogophora periculosa, Phoberia atomaris, Phosphila turbulenta, Phyprosopus callitrichoides, Polia detracta, P. nimbosa, Polygrammate hebraeicum, Ponometia erastrioides, Psaphida resumens, P. styracis, Pyreferra hesperidago, Renia salusalis, Schinia rivulosa, Scolecocampa liburna, Sericaglaea signata, Spargaloma sexpunctata, Spodoptera exigua, S. ornithogalli, Sunira bicolorago, Sympistis perscripta, Trichordestra legitima, Ulolonche culea, Xestia dilucida, X. dolosa, X. elimata, Xylotype capax, Xystopeplus rufago, Zale aeruginosa, Z. helata, Z. lunata, Z. obliqua, Z. unilineata, Zanclognatha protumnusalis; ROBIN MCLEOD: Gonodonta nutrix, Hypena sordidula, Papaipema cataphracta, Sympistis saundersiana; TOM MURRAY: Condica videns; MIKE NELSON: Acronicta lanceolaria; RANDY NEWMAN: Argyrostrotis quadrifilaris, Melipotis jucunda, Mocis latipes, M. marcida, Panopoda repanda, Ponometia semiflava; DAMON NOE: Eupsilia cirripalea, Isogona tenuis, Lacinipolia explicata, Psaphida thaxterianus, Zale buchholzi, Z. obliqua; RON PANZER: Schinia gaurae; ROBERT PILLA: Cydosia nobilitella; NELSON POIRER: Autographa mappa; MATTHEW AND DELIS PRIEBE: Drasteria grandirena; MARTIN RICE: Striacosta albicosta; ROGER RITTMASTER: Basilodes pepita; MORIA ROBINSON: Acronicta lobeliae, Gondysia similis, Homophoberia apicosa, Hypsoropha hormos, Ponometia candefacta; HENNING VON SCHMELING: Ascalapha odorata, Spragueia leo; LYNN SCOTT: Acronicta grisea, A. hasta, A. impleta, A. impressa, A. laetifica, A. noctivaga, A. oblinita, A. retardata, A. spinigera, Achatodes zeae, Agrotis venerabilis, Amphipyra tragopoginis, Anarta trifolii, Apamea devastator, Baileya ophthalmica, Balsa labecula, B. malana, Calophasia lunula, Catabena lineolata, Calyptra canadensis, Catocala cara, C. mira, C. neogama, C. relicta, Cerastis tenebrifera, Colocasia flavicornis, C. propinquilinea, Condica vecors, Copivaleria grotei, Crambodes talidiformis, Cucullia convexipennis, Dargida diffusa, Diachrysia balluca, Egira dolosa, Euparthenos nubilis, Eupsilia devia, Euxoa messoria, Feltia jaculifera complex, Galgula partita, Homoglaea hircina, Hypena edictalis, Hyperstrotia pervertens,

Ipimorpha pleonectusa, Lacanobia subjuncta, Ledaea perditalis, Leucania pseudargyria, Leuconycta diphteroides, L. lepidula, Lithomoia germana, Lithophane grotei, L. innominata, L. pexata, Metalectra discalis, Morrisonia latex, Mythimna unipuncta, “Platypolia” mactata, Orthodes cynica, O. majuscula, Orthosia alurina, Panthea acronyctoides, Peridroma saucia, Phlogophora iris, Plusia putnami, Protodeltote muscosula, Pyreferra citrombra, P. pettiti, Pyrrhia exprimens, Raphia frater, Schinia arcigera, S. florida, Simyra insularis, Spaelotis clandestina, Spiramater lutra, Spodoptera frugiperda, Sympistis chionanthi, Xylena nupera, X. thoracica, Zale duplicata, Z. lunifera, Z. minerea, Zanclognatha cruralis; JUDY SEMROCK: Caenurgina crassiuscula, Eudryas grata, Idia aemula, Rivula propinqualis; MERLE SHEPARD: Diphthera festiva; DAVID SILSBEE: Abrostola urentis, Achatia distincta, Acronicta superans, Allagrapha aerea, Catocala crataegi, C. epione, Celiptera frustulum, Colobochyla interpuncta, Crocigrapha normani, Eudryas unio, Feralia jocosa, F. major, Hypena humuli, H. madefactalis, Hyppa xylinoides, Marathyssa basalis, Melanchra adjuncta, Nedra ramosula, Nola triquetrana, Orthosia hibisci, O. rubescens, Paectes oculatrix, Phosphila miselioides, Protorthodes oviduca, Psaphida electilis, P. rolandi, Renia discoloralis, Syngrapha abstrusa, Xestia badicollis, Zanclognatha jacchusalis; LEROY SIMON: Chrysodeixis includens; JANICE STEIFEL: Melanchra picta; MARK SZANTYR: Eutelia pulcherrimus, Lithophane querquera, Scoliopteryx libatrix; STEVE WALTER: Athetis tarda, Bellura densa, Catocala antinympha, C. badia, C. obscura, C. ultronia, Epiglaea decliva, Euxoa detersa, Heliothis virescens, Heliocheilus lupatus, Magusa divaricata, Metaxaglaea viatica, Megalographa biloba, Orthosia garmani, Paectes abrostoloides, P. pygmaea, Renia factiosalis, Schinia nundina, S. trifascia, Tarache aprica, Trichoplusia ni, Xanthopastis regnatrix, Xestia smithii; MACHELE WHITE: Cucullia alfarata; CAROL WOLF: Litoprosopus futilis. Images of species not listed above were taken by DLW. Pinned adult images were taken by Shawn Kennedy, Rene Twarkins, and DLW. Photographic credits for the introductory pages and essays: MAX ALLEN: eastern bluebird; VALERIE BUGH: Condica videns in Parasitoids Plate; RICK CECH: Spodoptera frugiperda with ants; LYNN FAUST: Photuris larvae; HAROLD GREENEY and LEE DYER: Voria in Tachind Flies Plate; STEVE JOHNSON: moths at bait; ALEXANDER BARRETT KLOTS: Mythimna unipuncta ear (moth ears); LEROY KOEHN: bait trap; HUGO KONS: Mythimna unipuncta ear (ear mite); JOHN PEACOCK: burlap banding; MIKE QUINN: beating; MORIA ROBINSON: bark spider, Podisus with Orthosia, assassin bug with erebine, Zelus with erebine; JAMES SCHROEDER: grizzly bear; SCOTT SMEDLEY AND TOM EISNER: puddling in Gluphisia septentrionis; MIKE THOMAS: Compsilura coccinata (adult), Ammophila with Panopoda caterpillar; Marja van der Straten: Spodoptera litura; HENNING VON SCHMELING: Podisus feeding on Coxina; JAMES WIKER: moths on wine rope. More than 75,000 images were taken and studied over the course of this effort that were not reproduced here. These supplementary images were important in our descriptions and understanding of a species and those related to it. We are grateful to all those that labored to take and share images that contributed directly and indirectly to this work.

Introduction Owlet moths represent one of the most successful branches on the tree of life, whether measured in terms of species numbers, mass, or ecological importance. Adults range in size from a little over 7 mm to what may be the New World’s largest insect, the White Witch (Thysania agrippina), which has a wingspan that sometimes exceeds 30 cm. Noctuoid taxonomy and classification is undergoing a renaissance. Current classifications differ markedly in the number of recognized families and subfamilies for this group of moths. The Noctuidae have been treated as a single family historically, but now are frequently divided into smaller and more natural assemblages, with the resulting classifications differing among workers. Moreover, both molecular and morphological studies indicate that the tiger moths (Arctiidae) and tussock moths (Lymantriidae) are evolutionary derivatives from within the Erebidae. Here, we follow Lafontaine and Schmidt’s (2010) recent reclassification and checklist for the North American fauna in which four families are recognized: Erebidae, Euteliidae, Nolidae, and Noctuidae. We refer to this group collectively as owlets or Noctuidae sensu lato (Noctuidae s. l.). We did not include the Arctiinae and Lymantriinae, now two erebid subfamilies, because the common eastern representatives of both groups were treated in Wagner (2005). This book illustrates 726 species of eastern noctuoids, of which 372 species are accorded full (page) accounts; an additional 89 species are diagnosed or in some other way made identifiable. Our western boundary is the hundredth meridian as it was for Klots (1951), a work that greatly influenced the early interests of the first three authors. Each treatment includes a diagnosis, brief notes regarding similar species, common habitats, an approximate range,

phenology, a statement about relative abundance, a brief accounting of larval foodplants, and a Remarks section that is a potpourri of life history, taxonomy, and other relevant notes. For most of these species, we include two adult images, one of a live individual in a representative resting posture and one of a spread specimen. An additional 354 species are treated in abbreviated accounts (without adult images). The set of species selected for inclusion emphasizes taxa likely to be encountered by the layperson or forest manager, attractive species apt to draw public attention, and those of economic significance. A number of scarce or otherwise obscure species are included simply to provide better taxonomic coverage, or because they have interesting life histories, are of conservation concern, or have caterpillars that we otherwise deem noteworthy. Treeand shrub-feeding taxa, such as the daggers, quakers, pinions, sallows, zales, and especially the underwings (a popular group with collectors), are richly represented. Introductory sections address the importance of owlets, morphology, larval diets, natural enemies, classification, and various topics related to finding and rearing owlet caterpillars. Within the body of the work, each of the subfamilies, most tribes, and several large genera are introduced separately. The book concludes with two indexes: one for foodplants and a combined taxonomic/subject index. Our target audience is intended to range from students or laypeople to Ph.D. entomologists with training in systematics. An overarching goal was to prepare a guide that could serve as a portal into the world of lepidopterology (the study of butterflies and moths) for those without training in biology. To make this possible we offer extensive introductory text,

Birds and caterpillars (and climate change) The arrival of birds to temperate forests each spring is timed to coincide with peak insect biomass. The numbers and weight of nestlings that can be reared, the rapidity of their growth, and the number of broods attempted are all influenced by the amount of food available to the nesting parents. In temperate forests, much of the insect forage is made up of caterpillars (Nagy and Holmes 2005, Rodenhouse and Holmes 1992), especially those of inchworms (Geometridae), leafrollers (Tortricidae), and owlets. Most of the owlet caterpillars available to foliage-gleaning birds come from just three of the 63 or so tribes represented in this work: the Orthosiini, Hadenini, and Xylenini (all Noctuinae). Daniel Janzen, who has studied caterpillars for more than two decades in the tropics, refers to caterpillars as the hamburger of forested ecosystems. While graphic and rich, the metaphor understates the importance of caterpillars to birds—we venture that caterpillars make up no less than 60% of the diet of many songbird nestlings. One of the dangers of climate change is that some plants and animals will find themselves mismatched with the seasonal or phenological cycles upon which their survival is dependent. When spring comes early, most insects can adjust, because much of their vernal activity is triggered by ambient temperatures. But, migratory birds wintering in the Caribbean or Latin America are timing their migrations, courtship, mating, and nesting activities by changing day lengths, oblivious to whether or not eastern North America is experiencing an earlier-than-average spring. Phenological mismatching and diminished nestling success tied to climate change has been documented in the great tit (Visser et al. 2006) and several other species in Europe. The extent to which this phenomenon is challenging birds and other wildlife elsewhere is in much need of study. J eastern bluebird on way to feed nestlings. J

9

10 Introduction emphasize common names, provide images of the live insects, infuse the species accounts with natural history and information of general interest, and offer an extensive glossary. We hope that we have not fallen short, as so much remains to be learned about these moths.

Importance of Owlet Caterpillars and Adults By virtue of their sheer diversity and abundance, owlet caterpillars and adults have become enmeshed in the ecological processes of forests, grasslands, and other terrestrial ecosystems. Owlet caterpillars are common on trees and most woody plants in the spring, when foliage is soft and rich with nutrients. Another peak of larval abundance occurs in late summer and fall, mostly on composites and grasses. Caterpillars are a staple for insectivorous vertebrates. Birds are particularly reliant on caterpillars, and many insectivorous species time their nesting activities to periods of peak larval abundance—nestling survival would be appreciably lower in forests lacking owlet larvae (see essay on page 9). Goatsuckers move with storm fronts to feed on the abundance of moths that push northward on the leading edge of tropical storm cells— we estimate that greater than 90% of the moth biomass in these migratory swarms is made up of armyworms (Mythimna and Spodoptera), loopers (various Plusiinae), flower moths (Helicoverpa and Heliothis), the Black Snout (Hypena scabra), cutworms (Agrotis ipsilon and Peridroma saucia), and other owlets. Insectivorous mammals, including mice, shrews, raccoons, skunks, foxes, and many others, consume cutworms and other owlet caterpillars (and their pupae) found near the ground; some mice ascend shrubs and trees during their nocturnal forays for insect prey. Some insectivorous bats are dependent on moths and, by default, noctuoids. The colony of Mexican free-tailed bats that roosts under the Congress Avenue Bridge in Austin, Texas, is said to harvest more than 35,000 pounds of insects during the course of some nights. And the colony of 20 million Mexican free-tails that pours forth at twilight from Bracken Cave, Texas, purportedly consumes more than 100 tons of insects each night before returning to its roost. No doubt a healthy fraction, and perhaps the largest, is made up of noctuoids. Even mammals as large as bears feed on cutworm adults. Of particular importance are aggregations of the Army Cutworm (Euxoa auxiliaris), which aestivates by the millions in talus slopes in the Rockies. The communal gatherings are a critical food store for the grizzly bears of Yellowstone National Park and presumably for bear populations elsewhere. A single bear is estimated to eat 20,000 to 30,000 moths a day during portions of the summer—as much as one-third of the calories

required for an entire year may derive from the consumption of owlet moths (Mattson et al. 1991; French et al. 1994; White et al. 1998a, b). The pollination services provided by noctuoids are underappreciated (Committee on the Status of Pollinators in North America 2007). Many cuculliines, hadenines, heliothines, plusiines, xylenines, and myriad others avidly seek nectar at flowers. Owlets are among the most common insects to visit flowers of apple, apricot, aster, basswood, buttonbush, campion, cherry, fireweed, goldenrod, jasmine, lobelia, milkweed, various orchids, phlox, pinks, red maple, wild plum, willow, white snakeroot and other eupatoriums, as well as other plants that offer nectar at night. Noctuidae may be the principal pollinators of Platanthera and other white-flowered orchids. Cucullia are able to take nectar while hovering, much like hawk moths (Sphingidae); similarly, plusiines scarcely alight, placing just their tarsi on the corolla while feeding. Other owlets, like xylenines, orthosiines, and hadenines, actively crawl over flowers while probing them with their relatively short tongues. Larvae of some owlets are macrodecomposers that consume and digest fallen leaves, wood, and other organic matter. Herminiines are especially important in this regard with more

J Grizzly bear flipping rocks to uncover aggregations of the Army Cutworm Euxoa auxiliaris. J

Introduction than 50 common, litter-feeding species in the East—the subfamily can be abundant in deciduous woodlands. One owlet serves as a biological control agent in our region: the Toadflax Brocade (Calophasia lunula) was introduced into Belleville, Ontario, to control butter-and-eggs, an introduced Linaria species. The moth has since spread south to at least Maryland. No less than two are used in biological control programs elsewhere. Because of their great diversity and relative ease of sampling and identification, noctuoids are used in biotic assessment and monitoring efforts. Across the East, and especially in the Northeast, noctuoids and other moths are taken into account by The Nature Conservancy, Natural Heritage Programs, state agencies, and like-minded organizations when considering land acquisition, management plans, and other conservation decisions. In Connecticut, 42 (54%) of the 78 lepidopterans listed as endangered, threatened, or of special concern are owlets (including tiger moths). The indirect effects of plant-animal interactions on the quality and culture of our day-to-day lives are too commonly overlooked. Some of our most pleasant fragrances, such as jasmine—a staple of the cosmetics industry—are volatiles produced and released to attract moths and other nocturnal pollinators. The sweet scents of gardenia, narcissus, and other flowers that we select for our bouquets and corsages are manufactured by plants to attract moths. The spices that flavor our food are secondary chemicals produced by plants to discourage herbivory by caterpillars and other enemies. Similarly, the turpentine, resins, and latexes manufactured by plants repel the legions of insects and pathogens that would otherwise despoil unprotected tissues and organs. Antiherbivory substances may have considerable pharmaceutical value: e.g., digitalis, salicylic acid (aspirin), and taxol. Hundreds of plant alkaloids such as nicotine, caffeine, and theophylline shape the culture of our daily lives. Among the most widespread defensive compounds

A Note on Nomenclature Suffixes for family-group and tribal names are standardized across all zoological classifications (see below). Common or vernacular names based on the scientific higher category names may be made by adding English suffixes (right-hand column in table). The first letter of higher category names must be capitalized (e.g., Acronictinae). Capitalization is optional for the common name based on this name (Acronictines or acronictines). Category

Latin suffix (example)

Latinized suffix common name (example) (English common name)

Superfamily

-oidea (Noctuoidea)

-oids (noctuoids)

  Family

-idae (Noctuidae)

-ids (noctuids) (owlets) (in part)

  Subfamily

-inae (Noctuinae)

-ines (noctuines) (cutworms, sallows, and kin)

   Tribe

-ini (Xylenini)

-ines (xylenines) (pinions, swallows, and kin)

   Subtribe

-ina (Antitypina)

-ines (antitypines) (no common name applied)

in woody plants are tannins; these phenolics are cross-linked to proteins, rendering ingested plant tissues indigestible. Humans have found other uses for tannins, as in the production of leather and red wines. In sum, such indirect effects of Lepidoptera have been great, and our collective human experiences much enriched. While the vast majority of owlets should be regarded as having a positive or inconsequential economic importance, the ranks of the Noctuoidea include more than two dozen significant agricultural and garden pests in the East. Armyworms (Mythimna and Spodoptera) occasionally reach densities so great that they lay bare acres of cereal and other field crops. Members of the genus Spodoptera become increasingly important southward and at times cause massive crop losses. Cutworms, so named because of their habit of severing young plants at or near ground level, can be particularly detrimental in the spring when a single larva may consume or otherwise damage many seedlings over the course of its development. Both garden and field crops are adversely affected, e.g., some loopers (Plusiinae) are serious pests of crucifers. Heliothine noctuids are flower and seed predators. The Corn Earworm (Helicoverpa zea) and Tobacco Budworm (Heliothis virescens) account for millions of dollars in annual crop losses across the United States. These two moths and most of the other owlets regarded as crop pests are migrants that move northward en masse from our southernmost states, Mexico, and the Caribbean on the winds of storm fronts that boil out of Mexico and the Gulf. Single storms carry millions of individuals—local and regional population eruptions sometimes follow, resulting in minor to complete defoliation of susceptible foodplants. Curiously, no native eastern noctuoid is a chronic forest pest and only rarely do any cause defoliation in forested ecosystems. In Canada, the Straw-eyed Tentmaker (Enargia decolor) occasionally strips aspen of all leaves (Wong and Melvin 1976). The Black-dotted Brown (Cissusa spadix) and Common Oak Moth (Phoberia atomaris) sometimes defoliate oaks over localized regions, and in the South and West, a number of melipotis sometimes strip their leguminous hosts of leaves. In all cases outbreaks are localized and ephemeral. The Gypsy Moth (Lymantria dispar) is the principal owlet known to defoliate tracts of forests across eastern North America, but it is an exotic lymantriine introduced from Europe and falls outside the coverage of this work. Caterpillars of a few acronictines, and especially those of the American Dagger (Acronicta americana), may tunnel into siding and other wooden structures to fashion pupal cells, thereby blemishing outdoor structures and furniture. Florida’s Cabbage Palm Caterpillar (Litoprosopus futilis) is a more significant nuisance. Wandering prepupal larvae release a reddish-brown fluid that deeply stains siding and other nearby objects; some enter structures, including homes, in search of suitable pupation sites. Cocooning Litoprosopus caterpillars

11

12 Introduction tear up rugs, upholstery, and other fabrics, and then weave the severed fragments into the walls of their cocoons. Adult owlets are rarely considered pests in the East, although in the Rockies, millers (mostly the Army Cutworm, Euxoa auxiliaris) sometimes aggregate by the thousands in garages, sheds, and other structures. In our region, aestivating individuals of the Copper Underwing (Amphipyra pyramidoides) will gather in large numbers under shutters and in other man-made structures. Their liquid excreta can be substantial, accumulating on shutter slats, siding, and adjacent outdoor furniture; fortunately the reddish staining is easily washed away.

Morphology Like all insects, caterpillars have three body sections: head, thorax (segments T1, T2, and T3), and abdomen (segments A1–A10) (Fig. 1). Because the thorax and abdomen are little differentiated in caterpillars, they are sometimes collectively referred to as the trunk or body. The head (Figs. 2–3, 6–8) is comparatively simple and unmodified; it is usually rounded, and either smooth and shiny or granulated in texture. An important landmark is the triangle or frons (Fig. 2), which is bounded

1

2

4

3

5

J Larval morphology. Fig. 1: Lateral view of whole body. Fig. 2: Frontal view of head. Fig. 3: Lateral view of head. Fig. 4: Hypopharyngeal complex. Fig. 5: Mesal or inner surfaces of mandibles. J

Introduction on its sides by the adfrontal sutures. The height of the triangle relative to the length of the middorsal suture is constant within a species and is sometimes used to diagnose members of a genus. Many owlet caterpillars have an elongate, darkly pigmented patch, the coronal bar, to either side of the middorsal line. Caterpillars have six stemmata or lateral eyes on each side of the head, more or less arranged in a semicircle, with the fifth displaced toward the base of the antenna (Figs. 3, 6)—the relative size and spacing of the stemmata differs among owlet tribes and subfamilies. The relative length of the antenna also varies among groups. The mouthparts, unsurpassed in their importance to larval taxonomists, are not given their due in this work, because they often must be dissected and examined under a compound microscope.1 The mandibles, and in particular their inner surfaces, provide many useful species-level characters; details of the hypopharyngeal complex (Figs. 4, 8), e.g., number and development of spines over the upper surface, are routinely used by specialists to verify species-level identifications. Features of the spinneret and labial palpi (Fig. 8) also 1 Godfrey (1987: 4) provides a simple protocol for preparing dissections of the mandibles and hypopharyngeal complex. Sometimes it will be helpful to clear and stain preparations, using potassium hydroxide and stains in the same way that they are used for studying genitalia.

6

8

may be of value in making determinations. The various forms of the spinneret (e.g., the position of the pore, whether or not it bears flanges, to what degree its apex is fringed, or the presence of a dorsal groove) have much taxonomic value at the subfamily level. The spinneret’s length, relative to that of the labial palpus, is also diagnostic for many species, genera, and tribes. The thorax is composed of the three leg-bearing segments. The degree to which the prothoracic shield is differentiated from adjacent areas of T1 (prothorax) varies across and within subfamilies. In owlets that feed on foliage, the shield may be scarcely differentiated. The opposite extreme occurs in many borers where the shield is sclerotized (blackened) and conspicuous in relation to the rest of the thorax. The dorsum of the last abdominal segment bears a variously modified anal plate or shield. (Both the prothoracic and anal plates provide a rich source of taxonomic characters for some groups, especially among those that are internal borers.) Spiracles, the ventilation openings into an insect’s respiratory system, are found on segments T1 and A1–A8, with the first and last spiracles enlarged. The spiracular color, though variable, is often used in identification. Many workers use the height (longest diameter) of the spiracles as a standard against which the lengths of the body setae (hairs) are measured. As in other caterpillars,  Scanning

7

electron micrographs of the Androgea Armyworm (Spodoptera androgea). Fig. 6: Lateral

9

view of head. Fig. 7: Ventral view of head and t1. Fig. 8: Dorsal view of mouthparts; note dorsal groove in spinneret (characteristic of cutworms and other higher Noctuidae). Fig. 9: Proleg with circa 26 crochets. 

13

14 Introduction

10

11

12

 Larval chaetotaxy and patterning. Fig. 10: Schematic diagram of first two thoracic and six abdominal segments showing names for primary setae. Fig. 11: Trunk regions. Fig. 12: Lateral view of A10 with primary setae indicated. 

prolegs are usually present on A3–A6 (collectively called the midabdominal prolegs) and on A10 (anal prolegs), although the anterior pairs are frequently reduced in owlets (e.g., many erebines, Condicinae, and others) or are absent entirely (e.g., Acontiinae, Bagisarinae, many Hypenodinae, most Plusiinae, etc.). (Early instars of many trifids have reduced prolegs on A3 and A4.) The apex of the proleg bears a series of hooks called crochets (Fig. 9). The individual crochets being either one length (uniordinal), or two alternating lengths (biordinal), help assign a caterpillar to a tribe or subfamily. While the number of crochets on a given proleg may differ between closely related species, we seldom refer in this work to characters of the crochets, again because larval preservation and microscopic examination are needed to assess crochet number and character, and because numbers often vary within a species.

We use the term stripe for markings that run parallel to the body axis and extend along a number of segments. An approximate terminology has been adopted to convey where the stripes are located: moving from the top of the body (dorsum) to the bottom (venter) are the middorsal, addorsal, subdorsal, supraspiracular, spiracular, subspiracular, subventral, adventral, and midventral stripes (Fig. 11).2 Markings that extend around the caterpillar’s body are called rings or, when thicker, bands. Line is used for shorter markings, regardless of their orientation. Chaetotaxy refers to the presence or absence, relative lengths, and positions of the body setae. Formal keys for the identification of caterpillars are heavily dependent on 2 We use these terms to define relative position on a hypothetical cylinder and, for the purposes of this text, ignore arguments of homology based on setal insertions or studies of larval development.

Introduction chaetotaxy, particularly the primary setae, i.e., those that are broadly homologous across lepidopteran families. A few lineages also possess additional secondary setae. The primary setae have specific names based on their position (Figs. 10, 12) (Hinton 1946, Stehr 1987). The basic number of primary setae on a body segment is eleven: two dorsal (D1 and D2), two subdorsal (SD1 and SD2), three lateral (L1–L3), three subventral (SV1–SV3), and one ventral (V1) (Fig. 10); this number varies between segments (e.g., the prothorax bears two additional setae, the XD1 and XD2, and many of the 11 primary setae are lost rearward of A8). Variation in number and position occurs among species, genera, and families; accordingly, setal arrangements are often used in taxonomic diagnoses. For example, the SD1 seta on A9 is undifferentiated from other dorsal or lateral setae in most erebids (it may be greatly thinned in some euclidiines) and basal noctuids, but it tends to be appreciably finer in some plusiines, hadenines, noctuines, and their kin. Serious students will find chaetotaxy invaluable, and will want to consult Stehr (1987). Thoracic and abdominal setae are borne from pinacula, the rounded, hardened bases that are often more darkly pigmented than adjacent cuticle. A glossary with biological, morphological, and other specialized terms is provided at the end of this book.

Larval Diets Where there are terrestrial plants there are owlet caterpillars: nearly all woody plants, grasses, most forbs, and even ferns host one or more species. In addition, algae, fungi, lichens, leaf litter, and various other organic substrates are exploited. Although none of our eastern owlets is truly aquatic, a number feed on aquatic plants, and our divers (Bellura) are reported to be capable swimmers. Numerous lineages, especially among the Noctuidae, are denizens of early successional habitats: grasslands, meadows, weed lots, and coastal strand communities. Included in this same set of species are several agricultural pests and the unwelcome owlet caterpillars that frequent our vegetable gardens. Fewer noctuoids than might be expected feed on shrubs, trees, and other woody plants, although these include several diverse groups with many species: e.g., daggers (Acronicta and kin), underwings (Catocala), zales, pinions (Lithophane), and other xylenines. Many owlets that feed on woody plants are univoltine species that are dependent on young leaf tissues in the spring. Members of two tribes, the pinions and sallows (Xylenini), and the arches, quakers, and kin (Orthosiini), are among the most abundant spring-feeding forest macrolepidopterans and no doubt play an important role in the spring diets of insectivorous birds and mammals. Exceptions are many: daggers (Acronictinae), yellowhorns and kin (Pantheinae), and some others feed on mature, hardened foliage of shrubs and trees—

as might be expected, their caterpillars tend to have large heads and mandibles. Leaf litter of varying degrees of dampness and decomposition is consumed by various erebids and noctuids. Some noctuines that overwinter as caterpillars consume fallen leaves during the winter and spring. Many Herminiinae feed on downed plant matter and senescent, aboveground organic matter. Included among the litter moths are two of our most specialized owlets: Idia majoralis is apparently an inquiline in pack rat (Neotoma) nests, and Idia gopheri is a denizen of gopher tortoise and armadillo burrows where its caterpillars feed on droppings. Algae and lichen feeders include lichen punkies (Afridinae), some halflings (Hypenodinae), litter moths (Herminiinae), midgets and kin (Elaphriini), Metalectra, and others (see Wagner et al. 2008b). Several genera feed on fungi, e.g., Chytonix, Metalectra, and presumably a number of halflings. Of the 815 eastern owlets treated here, only a few genera have carnivorous members; these include Cosmia, Elaphria, Enargia, and Lithophane, all of which are facultative predators (they can and often do mature on leaf tissue). Predaceous owlet caterpillars feed principally on other lepidopteran larvae and pupae, especially those that are appreciably smaller or otherwise vulnerable (e.g., in or near a molt). Scale insects and other sessile Hemiptera are also consumed. In addition to the above are those species that are facultatively cannibalistic. Such behavior is frequent in flower and seed feeders, stem and root borers, and among taxa where the food supply may be limited and/or the risk of movement to a new feeding site is high.

Finding Caterpillars The most sporting and rewarding means of finding caterpillars is to discover them individually by examining leaves, flowers, stems, and bark; by opening shelters and galleries; and by conducting other direct visual searches. In so doing, one learns about the animal’s feeding and resting behaviors, its alarm responses, and other basic aspects of a caterpillar’s biology. (Some of our colleagues whimsically refer to such searching efforts as “zenning” for caterpillars.) Watch for signs of feeding, especially damaged leaves and feculae (droppings). New growth or actively growing plants are often favored. Additional tips on locating caterpillars are given in Wagner (2005) and references therein. Sizeable evolutionary radiations of grass feeders are found within the Erebinae, Eustrotiinae, and Noctuinae (Apameini, Hadenini, Leucaniini); dozens of genera scattered across other owlet subfamilies can be added to these. Grasses are important winter foods for some Noctuinae that later switch to feeding on forbs or even woody plants as they become available in the spring. Most grass feeders feed on the blades, but some specialize on the flowers and seeds. Numerous apameines bore into

15

16 Introduction stems or roots or feed externally on these tissues, belowground. Grass feeders are cryptic in both coloration and habit and may shelter in concealed sites by day. Nighttime searches will be more productive. Taller grasses can be sampled with a beating sheet or sweep net. Species that stay within a hummock or feed close to the ground will have to be rooted out individually. A large guild of species (especially among the apameines) either bore into aboveground tissues or tunnel belowground in rhizomes and roots. The most familiar of these is the large genus Papaipema, with more than 50 eastern species. Their presence is usually revealed by the frass that is pushed from the larval tunnel and accumulates on the ground below the tunnel entrance. Related genera with species that tunnel in roots are far less conspicuous in habit and poorly known. A great many apameines are specialist feeders on grasses, sedges, and related plants—wetlands have a rich fauna. Locating larvae requires special effort, but the potential to make new and worthwhile biological discoveries is great. As a group, owlet caterpillars tend to be cryptic in both coloration and habit, and many are nocturnal. They can be exceedingly difficult to locate by day, hiding in shelters, in bark fissures, and beneath leaf litter or, in the case of many cutworms, tunneling belowground. Expect nocturnal searches to yield different sets of species than those recovered from leaves during the day. Beating or sweeping is usually the most efficient method for sampling noctuoid caterpillars, as explained below. Given that all Lepidoptera are more abundant as earlier instars and early instars are more apt to rest on foliage, it pays to keep a watchful eye for young caterpillars. Learn to distinguish small and early instar noctuoids from their more numerous microlepidopteran cousins. Many microleps possess a clearly demarcated prothoracic shield, and most have the ability to wriggle or scoot backwards, often with great rapidity (see Wagner 2005: 11 for additional tips for distinguishing between macrolepidopterans and microlepidopterans). Members of several lineages perch on bark during the day, and coloration is often a clue to their identity. In general, green caterpillars will be found resting on foliage and other green tissues. Bark-resting caterpillars tend to be gray or brown, and are often hairy or have rootlet setae along the subventer; many are either somewhat flattened or have the ability to flatten their bodies. Beating is an effective technique for acquiring bark resters (especially those that position themselves along twigs and branches). Burlap banding also works well. At eye level, wrap a six-inch-wide band of burlap around a stem/trunk twice. Cut slits up from the bottom to within an inch of the upper edge of the burlap “skirt” or bands so that flaps of the material can be lifted and the bark beneath examined (right). The method works best on smooth-barked trees, and is all but useless on shagbark hickory and other trees that have an abundance of refugia. Of course, different tree species host different caterpillars. Don’t expect to find more than a caterpillar or

two for every ten skirts on a given day. Late spring, before foliage has hardened, is the most productive season. Dale Habeck had moderate success sweeping tree trunks with a soft-bristled hand broom or wide paintbrush over a beating sheet. More so than for other Lepidoptera, searching on the ground, belowground, or on objects near the soil surface can be a fruitful means of locating erebid and noctuid caterpillars. By day, cutworms secret themselves on the ground (some even beneath it), and then emerge from their hiding places after nightfall to feed. A number seek shelter under objects, especially the grass and forb feeders. To find these, flip boards, logs, rocks, fallen vegetation, trash, and leaves in contact with the ground. We have not deployed small squares of wood or other sheltering objects, but these have been used by others to acquire cutworms and other caterpillars. In agricultural fields, Crumb (1929) collected larvae under bundles of cut vegetation that he had strapped together and placed out as “bait.” When looking for caterpillars in low vegetation, we avoid areas where mouse runways are plentiful. Members of Agrotis, Euxoa, Striacosta, and a few Schinia hide belowground during the day. The Sandhill Cutworm (Euxoa detersa) illustrated in this work was found while screening sand. Friable and/or sandy soils are more likely to produce caterpillars than clays and other dense soils. Larval coloration and form are a clue to habit—stocky, pale to brown caterpillars with a smooth integument, inconspicuous setae, and short prolegs are often diggers that tunnel underground. As noted above, leaf litter can be searched. During the growing season, check around the bases of trees, especially below those with smooth bark that lack an abundance of hiding sites along the bole. Wintertime searches for herminiines, hadenines, noctuines, and others that overwinter as halfgrown caterpillars will be less productive, so set your expectations accordingly. Watch for feculae. The feeding damage of

J Burlap bands are a good means to sample late instars of bark-resting owlets, especially on smooth-barked trees that offer few comparable refugia. J

Introduction herminiines is characteristic. Begin looking in the transition zone between the dry upper surface litter and the wet, darkened leaves that are decomposing. We do not know if Berlese funnels and Winkler traps are efficient at extracting caterpillars from soil and litter, but our impression is that neither works well, perhaps because moth caterpillars are adapted to periods of dryness and thus are not easily driven from their feeding sites. Because herminiines are quick to release their grip and feign death, it might be possible to use a set of coarse screens to sift through leaf litter. Another option would be to sift leaf litter with quarter-inch mesh screening over a bed sheet. Two additional methods merit comment for grass- and sedge-feeding species as well as other ground dwellers. Rains occasionally flood these habitats, and at such times caterpillars will crawl onto emergent plants. Searching such areas while water levels are high can be rewarding (and fascinating). A final option is to cover low-growing plants overnight with a sheet anchored at the corners, and then after daybreak, inspect the underside of the sheet for adhering caterpillars. For some foodplant specialists, we have had success collecting random shoots of appropriately aged foliage, placing them in a large, clear, plastic bag, and then checking daily for the production of feculae. We obtained eggs and early instars of the Sharpblotched Nola (Nola pustulata) and two Paectes in this way. We recommend this method only for small species with rapid development, at locations where adults are known to occur in great numbers, and where random foliage collections will not have an impact on the hostplant population. Sleeving is another option: simply place a net bag over plants likely to be hosting eggs or early instars. Many of the foodplant records reported in this work were obtained serendipitously, i.e., where caterpillars turned up as “contaminants” in sleeves that DFS was using to rear other species. We have had much success checking plants nearing complete defoliation due to the outbreak of a pest species. Co-occurring caterpillars on denuded plants become concentrated on the remaining leaves, making their presence evident. If you are searching for caterpillars and encounter large numbers of foliage-gleaning ants (especially Formica or Solenopsis), be aware that they will have removed most of the larvae before your arrival. High numbers of spiders, lady beetles, and other predators, coincident with seemingly low numbers of caterpillars, is also telling…push on. When a single species is targeted, study its range, habitat, common foodplants, and phenology before starting your search. Consult literature on related species to direct your effort—congeners will be similar in appearance and many aspects of their life histories will be shared. Make a point to check what is known about related Old World owlets, since the early stages of many European and Japanese owlets are well studied. Keep in mind that while larger and last instars are more easily observed and photographed and are more likely to

garner one’s attention, the smaller, early, and middle instars are always more numerous and have the added benefit of sometimes being free of parasitoids. Climbing cutworms and many tree-feeding underwings (Catocala) will turn up in beating samples as early instars, but once grown these same caterpillars retreat by day into litter or bark crevices, respectively, where they can be exceedingly difficult to sample.

Beating and Sweeping The most efficient way to locate owlet caterpillars is to beat; use a stick, pipe, or baseball bat to rap vegetation over a light-colored drop cloth. The handheld beating sheets sold by entomological supply houses are a good investment. An inverted umbrella can be used in a pinch. To beat larger limbs or trees, scale up and use one or more bed sheets as drop cloths. Take along students, friends, or colleagues so that corners of the sheet can be positioned more quickly, moved into awkward positions, and held above patches of poison ivy. If there is appreciable debris on the sheet, after a short time, tip the sheet at an angle steep enough to roll or slide the loose material down to one side of the sheet. Many caterpillars will attach themselves to the fabric and adhere rather than tumble away. Rake your fingers through the debris and watch for any movement (especially of smaller, early instars). Return unwanted caterpillars to the plant from which they were dislodged. Ecotones such as forest edges and isolated plants usually yield more caterpillars than accessible portions of plants

J Use of a beating sheet. Smooth, white fabrics are best—these allow one to quickly locate even small, early instar caterpillars. Senior author beating juniper, May 2011. J

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18 Introduction growing in forest interiors or larger stands. Moreover, edges usually have overhanging limbs that lend themselves to beating. Sample one plant species at a time so that you know what to feed your collected larvae, and make sure that your sheet is clean before moving to a second plant species. If possible, avoid windy days because caterpillars hold on to foliage with greater tenacity during such weather, and as a consequence beating will produce fewer numbers of caterpillars (and working with a sheet in wind is difficult). Sweeping can be employed to collect caterpillars from shrubs and low-growing plants. Take a net with a stiff rim, pass it rapidly through vegetation, and then examine the contents of the net bag; we find it helpful to dump the contents onto a white (beating) sheet. This method is effective for collecting caterpillars from grasses, sedges, forbs, and low heaths. In both beating and sweeping samples, a fair number of the discovered caterpillars will be in the process of molting and therefore not able to attach to their foodplant securely. Treat molting caterpillars with care—the less they are handled the better. They will need to reattach to something to complete their molt. While most will attach adequately to paper toweling, a leaf, twig, or bark, those that have difficulty can be given silk—DFS keeps pieces of the inner cocoons of Cecropia or Promethea Moths on hand to offer molting caterpillars. Many caterpillars are best beat or swept after nightfall, such as those that hide in duff (many noctuines), on or under bark (some pinions and many late instar underwings), or off plant on nearby vegetation (some legume-feeding underwings). Lowbush blueberries and other low plants, as a rule, can be expected to yield more caterpillars at night. While flashlights are fine for nocturnal searches, headlamps free up one’s hands, and are enormously helpful when sweeping and using a beating sheet at night.

Collecting Adults The standard for collecting moths is to set up a light trap or sheet. Blacklight, mercury vapor, sunlamp, and other bulbs that emit ultraviolet wavelengths yield the greatest catches, although incandescent and fluorescent lights can also be effective. The small portable blacklights that are sold by biological supply houses are adequate to very good for herminiines, noctuines, and most other owlets, but larger blacklights or mercury vapor lamps are more effective for attracting erebines, xylenines, and others. At sites without electricity, use a generator to power a 175- or 250-watt mercury vapor light. Set up both blacklights and mercury vapor lamps when circumstances allow. Some collectors add sunlamps to their setups. In addition to high ultraviolet output, mercury vapor lights and sunlamps have the added advantage of providing considerable illumination for one to work about the sheet. But this added light is

also a disadvantage in that many individuals will settle in the darkness of nearby vegetation. A trick employed by beetle collectors that is seeing increased use by lepidopterists, is to run both blacklights and a mercury vapor light on the same sheet and to disconnect the mercury vapor light once every couple hours for a half hour or more. Moths (and beetles) whose activity was suppressed by bright illumination will frequently reactivate and move to the sheet. Traps can be run without a killing agent to secure gravid females for egg stock. Add inverted egg cartons so that trapped moths will have places to hide and settle. In the morning, gently “unpack” the catch by slowly turning over the egg cartons and inspecting each cell. We also find it worthwhile to run traps in light rains and on cool nights; enigmatically, gravid females sometimes make up a significant proportion of the catch on such nights. The standard means for obtaining gravid females is to “run a sheet.” A white bed sheet is hung vertically adjacent to a light—the sheet reflects the light and provides a landing platform for attracted insects. Place a second sheet on the ground as a drop cloth to prevent attracted moths from working their way into underlying vegetation. While most collectors use white bed sheets, another option is to use sheer curtain material or screen fabrics so that any light will be visible from both sides of the sheet. Leave a few folds or wrinkles in the sheet and the drop cloth to give incoming moths a place to shelter— some will be reluctant to come to rest in the full illumination of a mercury vapor light; e.g., Garman’s Quaker (Orthosia garmani) is more likely to secret itself in folds or under the drop cloth than to perch beside a bright bulb. Moths have particular times of the night when they are active; e.g., some daggers (Acronicta) will not appear in number until after midnight and many stalk borers (Papaipema) have specific flight times that last about 45 to 90 minutes on a given night. Moths will accumulate at a light through the night; thus predawn visits to the sheet will usually be rewarded with high numbers of both species and individuals. Turn off the light bulbs well before dawn to give the moths time to disperse, before birds arrive to feed on them. It is a good idea to periodically tap nearby vegetation to dislodge moths that have settled away from the sheet. On nights of modest moth activity, we will spend the better part of the evening away from the sheet searching vegetation for caterpillars, returning every half-hour or so to see what new moths have arrived. Full moons when accompanied by clear skies are unfavorable for light trapping. Many collectors plan their field trips around the new moon. Warm stormy nights, regardless of the moon phase, can be exceptionally good, especially if the storms remain distant or rains are light. Winter (2000) and Covell (2005) supply much useful advice on light trapping. A more challenging and engaging way to locate female moths is to actively search for them by day or night. The same

Introduction flowers that host butterflies by day may be frequented by dozens of other owlet species have been recorded feeding owlets after nightfall. Apamea, paints (Cuculliinae), loopers at the tree (Winn 2008, unpubl. checklist). Over a six-week (Plusiinae), and flower moths (Heliothinae) are avid flower period beginning in late July, John Peacock (1981) observed visitors. A patch of buttonbush, fireweed, or milkweed in full hundreds of adult Catocala representing 18 species visiting a bloom may yield dozens of moths over the course of an eve- dying American elm that was oozing sap through entry tunning. Apricot and chickasaw plum blossoms can be outstand- nels bored by scores of elm bark beetles. Black birch and sugar ing in the spring. Flower visitation often peaks at twilight or maple sap are attractive to a range of early spring owlets; any very soon after dark, but of course there are moths that wait broken branch, recent cut, or other wound is bound to draw until midnight or even dawn to take to the wing. Expect some the attentions of moths and other insects. very good nights, many slow nights, and some with no activity—this is the way with moths. Walking suitable habitat and inspecting vegetation with a flashlight is a good way to locate moths, especially some of the Apameini and litter moths (Herminiinae). When wearing Sugary plant secretions can be mimicked by concocting fera headlamp, one can watch for orange eye shine. Many spe- menting mixtures of sugar, fruit, beer, wine, molasses, etc. cies of underwings (Catocala) and grass-feeding hadenines The recipes of some of our colleagues border on alchemy. (Dargida and Leucania) can be found while they are oviposit- After preparation, which may include a period of aging to ing on trunks or stems of the foodplants. While light trapping allow fermentation, the mixtures can be used to bait owlat a sheet, search nearby vegetation; invariably you will locate ets. Baiting or “sugaring” is a time-tested technique for sammoths that either ignored your lights completely or settled pling owlets, especially erebines, herminiines, xylenines, and well away from the sheet. Keep in mind that the moths that a variety of fall migrants (including hadenines, noctuines, arrive at lights tend to be males, whereas those located by condicines, and as nectar starts to wane in late fall, even avid searching vegetation will yield a more balanced proportion of females. Many moths can be found during the day. We have reasonable success flushing moths from open habitats such as beaches, sand plains, and grassy fields. Marshes and other wetlands are also good, especially for renias, zanclognathas, and other litter moths (herminiines). As might be guessed from their bright hindwing coloration, underwings (Catocala) can be flushed during the day. Tapping, the preferred method for obtaining female underwings, is discussed on page 99. Males and females of most owlets take nourishment from the sugary solutions provided by flowers, broken tree limbs, oozing plant wounds, rotting fruits, and accumulations of honeydew excreted by aphids and other homopterans. Tree wounds are attractive to  Moths at beer and brown sugar a diversity of moths—any moist bark bait. Above: A good October patch that has an abundance of flies, baiting night in north central wasps, and especially butterflies during Pennsylvania. Most of the moths the day is sure to be a flurry of activity are the Bicolored Sallow (Sunira by night. A well-studied tree wound in bicolorago) and five species of an aged English oak in New York City’s pinions (Lithophane). Above right Central Park makes the case: as of this and right: Fall-migrating Armyworm writing some 23 underwing species and (Mythimna unipuncta) fueling up. 

Baiting

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20 Introduction Wine Ropes Ropes dipped in wine (or bait) will draw an array of owlets and other moths. Tim McCabe uses cheap red or white wine to which he adds sugar until no more will go into solution. In a five-gallon bucket with a secure lid, Jim Wiker pours a pound or two of brown sugar and a dash of amyl acetate (banana oil) into two to three gallons of sangria. Ropes made of cotton, hemp, or jute are then immersed in the mixture and hung from limbs or strung between two sticks in treeless landscapes. Wiker’s wine bucket contains more than a hundred 1.5 meter rope sections that he hangs just before dark. You may want to wear rubber or plastic gloves when deploying and retrieving the ropes. Wine ropes work well in the fall and winter for myriad xylenines, many migratory owlets, and various others.

flower visitors such as plusiines). A primary advantage of baiting is the quantity of moths that come on the best nights. Few other methods have the potential to produce hundreds of individuals over the span of an hour’s time, and no other method rivals the technique in late fall, winter, and early spring. Moth diversity studies based solely on light samples are often conspicuously deficient in abundant and diverse groups that are easily sampled at bait. Baiting is an effective method for obtaining females because bait catches are much less male biased than light-trap catches. The main disadvantage of baiting is its unpredictability; many nights the results will fall short of expectations and the resources invested, as discussed below. Bait concoctions vary from simple to complex (and bizarre) (Holland 1968, Sargent 1976, Winter 2000). Many experienced baiters swear by one ingredient or another like blackstrap molasses or their own, sometimes secret, additives. DFS uses a fairly simple recipe: one can (12 fl. oz.) of cheap beer (or apple cider), 1½ cups sugar, ¼–½ cup molasses, about four overripe bananas (or the equivalent in apples, peaches, pears, or quinces), a brewer’s yeast tablet (optional), and a spoonful of cornmeal (optional). The mix is then run through a blender, although hand mixing will suffice. The bait can be used immediately but will work better if allowed to ferment for a few days. Do not store bait in a sealed container—there is a risk it could explode from gas pressure. Good bait should smell strongly of alcohol and yeast. Do not clean the container between batches; just add fresh ingredients to whatever remains. Not surprisingly, fermenting fruits, such as overripe watermelon, also draw moths and sometimes in great numbers (Himmelman 2002: 21). Wine mixtures, used extensively in Europe, are seeing broader use by owlet collectors (see text box above).

Volatile substances that occur naturally in fermenting sugar baits can be used to attract moths (or augment a bait mixture). Foremost among these are isobutanol (2-methyl-1-propanol) and acetic acid (Meagher and Mislevy 2005). Landolt (2000) reported another short-chain alcohol, 3-methyl-1-butanol, as attractive to some erebids and noctuids. In summer you can apply your bait to tree trunks before sunset. In cooler seasons, if at all possible, apply by noon or even the day before and retouch around the edges about 30 to 60 minutes before dusk. Do not rebait the entire patch. Such prebaiting can make a significant, and sometimes a more than tenfold difference, especially in fall and winter; we have even experienced occasions when trees baited the evening before attracted more moths than our set of newly “painted” trees. In summer the difference between seasoned and fresh bait trails is less, but applying the bait early and retouching along the edges is still a good idea. If trees are in short supply, use wine ropes (see left) strung between two poles or sponges, soaked in bait, and then hung from branches or set in shallow dishes, on plastic lids, or even placed directly on the ground. While both butterflies and moths that have been imbibing bait for a time will become intoxicated, others are quick to fly off. Approach the bait slowly and keep your flashlight beam off the bait patch (and moths) at first. Once light has been directed on a moth, do not interrupt the beam. Some collectors carry a net in one hand for especially active moths like underwings (Catocala) and the Cabbage Palm Caterpillar (Litoprosopus futilis). Startled xylenines tend to drop to the ground (into leaf litter) and feign death, so if you are carrying a net place it beneath the bait patch before drawing your headlight beam across the bait patch. Some collectors clear leaves from the base of the tree at the time they apply the bait. To “paint” tree trunks use a broad, clean brush (8–10 cm) and apply bait in a patch roughly 20–30 cm long at head height. Don’t worry about bait dripping down the trunk. Chunks of banana and other fruit in the bait can be mashed against the bark with the brush. To ensure that you are able to locate painted trees, follow a trail, edge, or roadside, or bait areas with relatively little understory. Flagging tape can be used to advantage. Select trees with smooth bark or plates of smooth bark: pines, hickories, cherries, and birches are all good. Corky or absorbent bark (like white oak or cedar) is best avoided. Vary the direction and wind exposure. In cool seasons, be sure to bait sunny portions of the trunk. Most moths will arrive within the first hour or two—this is especially true of winter moths but also for many summer-active taxa, including species that normally do not appear at a light until after midnight. Baiting is unpredictable, and often there will be no obvious explanation for poor or spectacular results. In our experience, it is most effective in temperate woodlands and forests

Introduction where bait has analogs in sap flows, tree wounds, and senescing fruits that moths would be visiting regularly. Dry pine and oak woods can be exceptional. In northeastern forests and barrens, one sometimes sees dozens of moths on a single bait patch. On rare occasions you will attract thousands of pinions and sallows (xylenines), or hundreds of underwings and zales. A 200-meter bait trail with 20–40 baited trees in southern New Jersey will yield as many as 40–50 species in a single June evening. By contrast, June can be the worst month for baiting with yields of less than one moth per tree. The efficacy of baiting sometimes drops off in mesic woodlands, deserts, and elsewhere where the natural analogs of bait are either too abundant or too scarce—we often hear colleagues grouse that where they live, the technique is neither worth the time or effort it takes to prepare the bait, nor the waste of a good can of beer. Abundant honeydew, overripe fruit, and nectar availability will depress the effectiveness of bait. In this regard it is important to note that visitations over the winter to spring bait season will drop off abruptly when the first red maples burst into flower. In general, moth numbers at bait during the growing season will not approach those seen during the fall and winter months, with notable exceptions: zales are often exceptionally common in pinelands, as are underwings (Catocala) in oak-hickory woodlands. Few entomological experiences rival a good underwing baiting night (see Holland 1968: 146 for a splendid essay about one such evening, presumably in western Pennsylvania). In late fall, winter, and early spring, warm days over 18° C (64° F) with a mild evening are best; high humidity, light rain, or drizzle can help, especially in xeric habitats that have been dry for some time. Moths will begin arriving before dark, peak during the early evening, and continue to arrive in decreasing numbers until temperatures drop. (On warm and stormy nights moths will remain active all night.) In general, the minimum sunset temperature for good cool-season baiting varies from about 7° C (45° F) in New England to about 10° C (50° F) in southern New Jersey. In summer, temperature is less important and baiting can be good on cool evenings. Baiting is only slightly affected by moon phase. On bright nights trees in the open may yield fewer moths than those in the shadows. In suburban areas it is best to avoid baiting trees that are near lit porches or street lights. Sugar baits attract more than moths and butterflies. Bears and skunks will lick at a bait patch. White-footed mice may add to the enjoyment of checking a bait line; in the pinelands of New Jersey and southward, one will sometimes encounter a gray fox wrapped around a tree trunk, up on its haunches, licking at bait. Baiting is also a good way to observe flying squirrels; some individuals eat mainly bait, while others will feast on the moths. Finding wings at the base of a bait patch usually indicates a flying squirrel. In winter, eastern screechowls are occasionally seen hunting a bait trail for moths and

John Peacock’s Bait Recipe The bait has two basic components: overripe, fermented bananas and a solution of beer, brown sugar, apple juice, grape juice, and molasses. For the banana “base,” take 10 or more pounds of peeled, overripe bananas and mash them in a large Tupperware® container, and then let the mush ferment for two or three days in the closed container at warm temperatures. Occasionally stir the mixture during the first two days. Decant and discard any liquid that accumulates. Prepare the solution that will be added to the fermented banana base as follows. In a 5-gallon pail, mix 48 oz. of red grape juice, 32 oz. of apple juice, 10 lbs. of dark brown sugar, 5 bottles of beer, and 16 oz. of “robust” molasses. (Avoid ingredients with preservatives.) Keep the pail loosely covered and at warm temperatures for three weeks or more prior to use, stirring every few days to be sure all of the sugar is dissolved. Prior to use, stir in a few cups of the above liquid mixture to the banana base, and then spoon the resulting banana/liquid mixture into a round, flat plastic bait pan, and place in the trap. The bait will improve after a few days in the field. When humidity is low, add some of the liquid mixture to the bait every other day or so to keep the bait adequately moist. Also, check to be sure visiting moths haven’t imbibed all the free liquids; this sometimes happen on nights with elevated activity. It is best if the bait is a little “tacky” so that there is a substrate on which the moths can rest while feeding. You may use the same starting bait over a three- to five-week period, adding a spoonful or so of the fermented bananas and a little of the liquid mixture from time to time. After using the bait repeatedly, it may appear “caramelized,” but will still be effective.

mice. Caterpillars, especially climbing cutworms and tiger moths (Arctiinae), also may be found imbibing bait. Floral scents are attractive to many flower-feeding adults, including armyworms (Spodoptera), the Corn Earworm (Helicoverpa zea), darts (Agrotis), the Velvetbean Caterpillar (Anticarsia gemmatalis), Mocis, and a range of loopers (Plusiines) (Meagher 2002). Like sugar baits, floral scents tend to attract both sexes. Use caution when working with any such chemicals, as some may prove to be health risks.

Bait Trapping An alternative to painting bait on tree trunks is to employ bait traps. Traps require less maintenance and attention and often yield splendid catches of owlets, as well as nymphalid butterflies, hawk moths, and other insect life. Bait traps have other advantages: they run all night and they can be tied to a cord and pulled high into (canopy) trees, or placed in localities that would be inconvenient or inaccessible for running a bait line. Additionally, they can be placed out at any time and picked up well after sunrise or even a day or two later (although with standard trap designs many moths will manage to find an egress from the trap after daybreak). Initially, you may want to run a trap in your yard over the course of a few weeks or months to get a handle on your local fauna and to develop a bait and routine that works well for you. We venture that

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22 Introduction you will find the experience engaging and rewarding, and at the very least will see owlets and other insects that you would rarely encounter otherwise. While the same bait concoctions noted above can be poured into a pan in your bait trap, we recommend using a more solid bait, e.g., a mash of a half-dozen or more overripe bananas over which liquid bait is poured to encourage further fermentation. (We share John Peacock’s proven recipe— fine-tuned and field-tested for underwings (Catocala)—on the previous page). Leroy Koehn makes an apple-peach bait by combining a cup of sliced apples, an equal measure of sliced peaches, and a cup of sugar in a plastic bag; he then keeps his “ambrosia” until it starts to ferment. In traps, semisolid baits prevent drowning of insect visitors, remain active over longer periods, and are easier to service. Over warm and dry periods, mash baits will need to be rehydrated, often in late afternoon or early evening, with more liquid bait, water, or as is sometimes used by Weston Henry, a mixture of ethanol and water. The most common bait trap design has a narrow entry area near the bottom where bait is positioned and a large upper screened area that collects moths that have finished

J Catches in bait traps can be prolific (and interesting) and, just as often, disappointing, even over the span of just a few days—see text. J

feeding. Traps range from less than 25 cm in diameter to large, 75 cm diameter constructions that can easily retain 100 or more underwings. A band of dark fabric around the top will encourage settling, and consequently ensure better catches. Some collectors add a few leaves, or halved or quartered egg cartons at the bottom of the trap to provide sheltering nooks to visiting moths. It is best to service traps an hour or so before daybreak. Such predawn checks work for underwings, litter moths (Herminiinae), and most other owlets. Xylenines, and other winter moths are more adept at finding their way out of traps after they have fed; check your traps more frequently, especially in early evening when activity is at its peak. To discourage visits from flies and hornets, cover the bait pan during the day (leaving it in the trap), and then uncover the bait an hour or so before dusk. See Winter (2000) for additional information on bait trapping.

Collecting, Vouchering, and Conservation Much remains to be learned about the life histories of eastern owlets, so much so that even the weekend biologist or student can expect to make worthwhile contributions to our knowledge of this extraordinarily diverse group of moths. As you read through this book, taxonomic problems and uncertainties about the foodplants and life cycles will surface by the dozens. Pick a group and start rearing; take notes, images, and videos; save appropriate voucher specimens. Larval, pupal, and adult vouchers that are well preserved and labeled will become a scientific legacy. They can be examined and reexamined, redetermined, dissected, or their DNA can be sequenced. An obvious case for the importance of vouchers is seen when a species proves to be more than one entity. Many eastern litter moths (Herminiinae) are proving to be complexes of two or more biological species. Long into the future, biological specimens will be “mined” for their DNA, each having the importance of a well-preserved fossil. The possibilities are almost beyond our comprehension, e.g., trace amounts of plant material in the gut could be sequenced to determine a caterpillar’s diet at the time it was preserved. Toxicological residues in a specimen could be used to study the extent of environmental contamination at some fixed point in the past. Many future possibilities will be limited only by the lack of preserved specimens. Our collections will be our longest legacy, attesting to times, locales, and climates of the recent past. We are advocates of collecting and life history studies. These activities are rewarding and scientifically justifiable, but just as important, for many children, students, and amateurs, they provide an entry point into the world

Introduction of entomology, systematics, and invertebrate conservation. Many of today’s most active insect conservationists and researchers were first collectors (Wagner 2007e). The National Wildlife Refuge System in this country was established by funds generated by duck stamps purchased by bird hunters. Collecting efforts must be carried out in an ethical and responsible fashion. The Lepidopterists’ Society statement on collecting (http://www.lepsoc.org/statement_on_collecting. php) should be read and followed, regardless of whether the intent is to release or voucher the reared adults (and reared adults should be released only near their place of origin and during their natural flight season when they are more likely to reproduce successfully). Never remove more than a fraction (about 5–10%) of the number of individuals you believe to be present in a population. Species with conspicuous caterpillars or those that are otherwise easily collected—for example, many stalk borers (Papaipema) and flower-feeding flower moths (Heliothinae)—could easily be affected as a result of a single overzealous collecting effort. If one were to collect the visible host flowers in a small Schinia colony, a population could be seriously diminished. Be especially careful if the foodplant is highly localized or scarce. When appropriate obtain and carry (necessary) permissions and permits. Paradoxically, undercollecting is a greater peril to most Lepidoptera than overcollecting. This is because we still need to know a great deal about most of the species in this guide before we can recognize which taxa might be declining, understand a species’ biological requirements, or develop management practices for an imperiled colony. Outside of the Northeast and upper Midwest comparatively few moths receive legal protection, are considered in management practices, or are used in conservation planning, simply because we know too little about them. Numerous aspects of the life history may be learned by collecting, rearing, and observing caterpillars: e.g., the number of annual broods, seasonal and daily activity patterns, and cocooning and overwintering behaviors. Hostplant-use patterns, and how these vary geographically, are remarkably incomplete for most of the species in this guide. Such knowledge is critical in the design of management plans for species of conservation interest. We desperately need collections documenting the impacts of introduced biological control agents on native caterpillars. Should you engage in appreciable rearing, an exercise which we find intellectually rewarding, as did Douglas Ferguson and John Franclemont to whom this book is dedicated, remember that an important part of this process is good record keeping. Save and label any parasitoids, and above all, share your findings. A useful reference for handling and preparing specimen vouchers is Winter’s (2000) Basic Techniques for Observing and Studying Moths and Butterflies.

Feeding Adults To be kept alive, adult owlets should be fed daily or every other day with sugary solutions. The exact proportions are not important. A two- to five-part water to honey mixture or maple syrup solution will work fine, but table sugar or almost any other sweet may be used. More dilute mixtures are acceptable—adults will simply imbibe more and secrete the excess water. We offer a solution in a saturated cotton ball, small squares of sponge, or a tiny wad of toilet paper. Fruit juices and small slices of fruit, e.g., moistened apples and grapes, also can be offered. Change fruit pieces after a day or two, because they soon spoil. We travel with raisins; small sections are cut, soaked, and placed where the females will have easy access. Sport drinks are eagerly accepted by a gamut of Lepidoptera—we sometimes add Gatorade® or Powerade® to the food of long-lived underwings (Catocala) and zales. Wine will also serve. If egg number is important, try adding a trace of egg white to the solution to enrich the diet.

Obtaining Pairings With few exceptions, field-collected females already will have mated. Pinions and sallows that overwinter as adults do not mate until late winter or spring. To obtain pairings, place one female and two or three males in a spacious container with a screen or mesh cover that will allow good air flow (for circulation of courtship pheromones). All moths should be well fed. Although mating may occur soon after the sexes are placed together, expect the majority to delay pairing—many moths are reproductively active for specific periods during the night. Generally, mating pairs will stay together for at least an hour, although some species remain coupled until nightfall of the following day.

Obtaining and Handling Eggs Most owlets oviposit readily in captivity. Female owlets can be expected to produce between about 300 and 1500 eggs if kept well fed, although conifer feeders and owlets that feed on hardened summer foliage produce fewer, larger eggs. Some moths (e.g., sallows in the genus Eupsilia) require multiple matings if they are to continue producing viable eggs. Place gravid females in small vials and jars. Hang a strip of paper towel vertically, usually with one end draped over the lip of the jar and with the other end bent across the bottom. A minute drop of water can be added to the strip, especially

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24 Introduction if no food is provided initially. (No free water should be visible.) Most species will lay their eggs on the paper strip, others will lay on the container walls and lid, and some (including many noctuines) will drop their eggs about the bottom of the container. If you believe the species oviposits in cracks or crevices in bark or about buds, add a tightly rolled piece of paper towel or strip of corrugated cardboard into which the female can insert her eggs. A long-pointed abdomen usually indicates that a species positions its eggs in soil or deep crevices. For example, female darts in the genus Euxoa oviposit into sand or soil. (If the sand placed in a container has already been screened with fine mesh, you can rescreen the sand to recover any eggs.) Underwings (Catocala) and others oviposit in bark crevices—offer these a strip of bark of a preferred host (see also page 100). Lash-eyed sallows (psaphidines) and a few other owlets active in fall, winter, and spring should be held with a twig of their foodplant, with terminal buds, and a paper towel strip. In addition, we sometimes add a piece of dead leaf of a favored host. Grass feeders such as wainscots (Leucania) and related genera prefer to oviposit in the leaf sheaths around grass stems—include dried, dead stems in the vial. Daggers (Acronicta) prefer to oviposit on smooth surfaces, so hold gravid females in clean, smooth plastic containers or glass jars. Most owlets require food (see Feeding Adults, above). Sugar solutions can be supplied on a small bit of cotton (less than 5 mm in diameter) placed at the bottom or top of the jar— but not in contact with the paper strip. Punch two or three small air holes into the lid and place the container outdoors, on the north side of your house or on a shady porch. Most species will oviposit one to three nights after their first meal. Some Lepidoptera require elevated temperatures and light to

J Noctuid eggs range from hemispherical to flattened with intricate surface ornamentation. The clear chorions of the Reddish Speckled Dart (Cerastis tenebrifera) eggs are all that remain after hatching. Dark eggs in the image contain first instars that failed to eclose. J

stimulate egg-laying behaviors. Such seems to be the case for many diurnal moths (e.g., many chocolates, Argyrostrotis). Females held in vials and jars without foliage, for example those in transport between field and lab, will often lay if they are exposed to elevated temperatures (32–34° C or 90–95° F) for an hour or two. A reliable means of securing eggs is to sleeve females on an appropriate host. Females should be fed before they are introduced into the netting. Alternatively, add a saucer of fruit (such as crushed grapes), or place a plug of cotton saturated with honey or sugar water inside the sleeve or on top of the sleeve, positioned so that the female can feed through the bag. DFS places conifer feeders in a clean ice cream carton, plastic box, or widemouthed glass jar of at least a half-gallon capacity. He adds a sprig of the foodplant, with new growth if available, so that the upper needles touch the cover. The container is then covered with nylon netting or other airy cloth. Finally, he adds a feeding plug, placing it inside the container or simply setting it upon the netting cover. Each evening the containers are slightly moistened (misted). Feeding plugs are changed every two or three nights. Conifer foliage held in a vase or florist water pick will last ten or more days, and sometimes long enough for the eggs to hatch and the first instars to establish. Newly laid eggs are often cream in color but may turn yellow, orange, pink, red, or gray within a day. Infertile eggs often fail to change color and/or collapse. Typically noctuoid ova are hemispherical with lacelike ribbing. Some are handsomely banded. A day or so before hatch, the eggs may darken, in large measure because the developing caterpillar becomes increasingly visible. The chorion, or egg shell, is translucent in nearly all noctuoids; hatched eggs of the Reddish Speckled Dart, Cerastis tenebrifera, are shown in the photograph to left. Hatching rates are generally better if the eggs experience natural humidity in well-ventilated containers. Eggs laid on flowers or leaves quickly mold if left in sealed containers. Adding leaves to vials, even when the ova are on the verge of eclosion, may prevent hatch in sallows, pinions, and others. The volatiles from cut pine foliage also are purported to depress hatching rates—here again, ventilated containers are recommended. If eggs normally hatch within four to ten days, no watering is needed, although a light misting may be helpful under dry conditions. Be sure that the ova then have time to air-dry if they are to be housed in an airtight container. For eggs laid in the fall (e.g., most Xylenini) minimal care is needed. Keep the eggs in a shady outdoor place in a small jar with a tight, but not completely sealed, lid. Egg jars need protection from direct sun and from heavy precipitation. Air holes are optional—they provide ventilation and regulate moisture, but can be a liability in that they make a container subject to flooding and provide a means of escape for newly hatched caterpillars. While a garage or enclosed porch

Introduction prevents flooding, discourages predation, and moderates cold winter temperatures, be forewarned that eggs held there may hatch before food is available in spring. Eggs that overwinter on trees and shrubs are adapted to swings of heat and cold, and unless the stock comes from a much warmer climate than where you live, the ova will not need protection from extreme temperatures. Periodic mistings will prevent desiccation, but make sure the eggs are air-dried afterwards. The greater danger will be the growth of mold because the tendency will be to supply considerably more moisture than is needed. John Peacock overwinters eggs in his garage but, come March, he transfers the eggs to an airtight plastic container with a slight bit of moisture; he places this in his refrigerator to prevent early hatching. As appropriate foliage becomes available, he removes the eggs. This strategy has two other advantages: it allows one to stagger cohorts of larvae, starting some early and some late, and more importantly, egg hatch can be timed around one’s travel plans. Many lepidopterists sleeve their eggs for ease of handling and to avoid the uncertainties surrounding moisture management. If females normally oviposit directly on leaves, position the eggs so that they are adjacent to suitable food: e.g., eggs laid on paper strips can be cut away and stapled to the underside of host leaves. While sunny twigs are generally best, branches that experience long periods of full midday sun, with temperatures exceeding 32° C (90° F), are best avoided. If the caterpillars are active and normally have to disperse to find food, e.g., underwings (Catocala), and many pinions and sallows (both Xylenini), one can simply place the eggs in the sleeve. However, for most owlets, early instar survival will be enhanced if neonates are started in vials, and later introduced into sleeves as late first to third instars.

Rearing Caterpillars Owlet caterpillars are easily reared if provisioned with fresh foliage, their containers are kept clean, and they are offered suitable pupation substrates. Monitor both growth and feculae production daily; most species will show visible growth over any one- to two-day window. Only during a molt, and prior to pupation, should feeding slow down. If growth rate or feculae production falls off (and the caterpillar is not approaching or coming out of a molt) consider trying foliage of a different age or species. There is no single best method—or stated differently, no two lepidopterists use the same rearing methods. Much advice on rearing caterpillars appears in Wagner et al. (2002) and Wagner (2005), and Todd Stout’s website on raising butterflies is loaded with terrific information about caterpillar husbandry (http://www.raisingbutterflies.org). Rather than repeat information found in these places or other widely available

works (e.g., Winter 2000 and Covell 2005), we address a few points and offer suggestions geared toward the rearing of owlet caterpillars. Most spring and summer species will complete their development quickly, within four to five weeks. Conversely, those that overwinter as larvae may spend eight to ten months in this stage. Because owlets are largely nocturnal, it is often possible to speed development by rearing them in darkness or at least by minimizing the number of hours that they are exposed to light. As a group, owlets are reasonably hardy and can be reared at moderate densities. Some ventilation will enhance survival, especially if multiple caterpillars are being reared in a single container, but it is more crucial that foliage remain fresh. Rare, desirable, or cannibalistic caterpillars are best reared individually to aid study, diminish the spread of disease, and circumvent cannibalism. Also, if data on parasitoids are desired, it is best to raise each caterpillar in its own container. Neonates will imbibe water. Access to water is especially important if the caterpillars have hatched and wandered without encountering food. As a matter of routine, we lightly mist newborn larvae either before or immediately after their transfer to host foliage. Lethargic caterpillars, e.g., Paectes, are best picked up individually with a needle or fine brush (e.g., a #1 camel hair, pointed artist brush), and placed onto appropriate foliage. More mobile caterpillars, e.g., underwings (Catocala) and various other erebines, easily find their way onto suitable host tissues. Many spring-active noctuids can be reared on young cherry or walnut foliage. Other hosts that are accepted by a large fraction of temperate caterpillars include alder, apple, basswood, birch, blueberry, and oak. Plants that continue to produce young leaves late into the spring or early summer such as alder, birch, cherry, walnut, and willow will allow one to supply new foliage long after oaks and hickories have hardened for the summer. Noctuines include many generalists on nonwoody plants and will accept a range of forbs and grasses. Most will find something suitable in a salad of fleabane, clover, dandelion, and grasses. Organically grown lettuce or the inner leaves on non-organic heads are options, even in the winter months; romaine and other leafy varieties are presumably more nutritious, and less likely to cause diarrhea. Many owlets can be reared on artificial diets such as those sold by biological supply houses. BioServe® makes dozens of diets for a wide range of insects. Bergomaz and Boppré (1986) describe a diet made from beans that is acceptable to a variety of noctuoids. Tim McCabe has successfully reared a variety of erebids and noctuids on BioServe’s pinto bean diet; he occasionally makes customized batches for foodplant specialists by drying leaves of the hostplant, and adding ground up leaves to the diet before it is poured into diet cups. Many borers will accept

25

26 Introduction carrot, potato, and sweet potato: core out a hole about the same diameter as the caterpillar and then introduce the larva head first; sometimes it helps to then seal the hole with a plug or to position the entrance such that the caterpillar cannot easily back out. Vases and floral water picks can be used to great advantage in open-air containers and other situations in which the foliage would wilt within a few hours. The Oasis® foam sold by flower shops can be purchased in bricks and cut to size to keep foliage fresh for days. Grasses and many forbs often do not hold up well once they have been cut—consider potting these and then covering the foodplant (and larvae) with netting. Many owlets require a layer of soil or peat in which to pupate. Failure to provide an appropriately moistened pupation substrate is probably the most common cause of mortality in reared owlet prepupae and pupae. A deep layer (5 cm or more) (2 in.) of lightly moistened peat, potting soil, or dirt should be offered to any caterpillar that wanders for extended periods, especially if the caterpillar is showing no interest in eating or otherwise appears to be prepupal. We use peat for all univoltine, spring-active caterpillars. Many balsas (balsines), brothers (dilobines), dagger moths (acronictines), and foresters (agaristines) fashion their pupal chambers in wood. Offer them chunks of soft pulpy wood, e.g., fallen branches and trunks with wood that is light and soft.

Sleeving Fine mesh bags or sleeves can be used to enclose females for oviposition or for rearing cohorts of larvae outdoors. In addition to being the most time- and labor-efficient means of rearing caterpillars, sleeving has other advantages: the larvae are less susceptible to stunting, caterpillars can be left unattended for a week or more, and, because sleeves are well ventilated, diseases are rare. Finally, the sleeved caterpillars are exposed to normal photoperiods and temperatures, and consequently will be synchronized with native populations. Make sure to vigorously shake or rap any branches that are to be sleeved in order to dislodge ants, spiders, lacewing larvae, assassin bugs, ladybugs, and other natural enemies. Predaceous stink bugs often accumulate on the outside of sleeves and feed on the enclosed caterpillars through the netting. Upon occasion a bird will peck through a sleeve to feed on the occupants; hornets or wasps may also chew through and raid the sleeve. Valuable livestock can be protected from both by double sleeving or using hardware cloth. Pillowcases, because they are made of denser fabric and are nearly opaque, offer more protection. Small sleeves must be moved frequently; larger sleeves have the advantage of being low maintenance.

J Sleeves of fine netting or fabric, such as the fivegallon paint filter bag used here, have many advantages over container rearing—see text. J

Prepupal larvae of most owlets, especially those that pupate in soil or wood, should be removed from the sleeve at least every other day and placed over peat or other suitable substrate; left untended, some will chew their way through the sleeve in an effort to locate a suitable pupation site (or die of dehydration and exhaustion). Other species will cut bits of the sleeve for use in the walls of their cocoon. If you know the species pupates in wood, this can be added to the sleeve. A safe option is to periodically harvest nearly mature larvae from the sleeve and rear them indoors for the last few days. Sleeves may be purchased from biological houses or made by hand. We have made large sleeves for enclosing entire branches from synthetic tent screening fabrics. The one- and five-gallon paint filter bags sold at home improvement stores are economical and handy for rearing small larval cohorts. Pillow cases can be purchased used from secondhand stores or at flea markets. When necessary, or in case of disease problems, launder sleeves with ordinary detergent and some added chlorine bleach.

Overwintering Larvae and Pupae Our general suggestions for overwintering stages undergoing diapause are given in Wagner (2005). We recommend that

Introduction rearing lots be held outdoors in garages, sheds, or porches, where they will be exposed to normal temperatures, humidities, and photoperiods. Those who live in the North must take measures to protect their livestock from subfreezing temperatures—prolonged exposure to temperatures below about −3° F (26° C) can be fatal, especially for larvae or pupae that are usually underground or sheltered near the ground under leaves. DLW overwinters collections in a garage, periodically misting the lots through the winter. DFS, who lives in coastal New Jersey, simply places his pupal containers in foam coolers packed with dry leaves, and places the coolers in a shady, sheltered spot outdoors, e.g., an outside porch or crawl space that stays above −3° C (26° F). Another option is to cover the rearing vials with fabric or a lid with several holes, place them on a tray or in a cage, and set them into a shallow pit so that they are elevated just above the ground. Then cover the vials with a deep layer of leaves. Either invert the vials so they do not fill with water or lay a sheet of plastic or wood over the top before covering them with leaves. We have lost collections to mice—take measures to safeguard your lots. In most years, we bring our collections indoors in February or March to accelerate emergence so that we have more time to work with emerging adults, in advance of the field season. Owlets that overwinter as caterpillars may feed intermittently during periods of mild weather: included are many darts (Noctuini), litter moths (Herminiinae), and some arches (Hadenini). Winter-active caterpillars eat fallen leaves or are general feeders on forbs and/or grasses. A method with which we have had much success is to place the livestock in a clay flowerpot partially filled with soil and a few layers of food leaves. The pot is then covered with screening or muslin and partially buried outdoors, exposed to normal rainfall, snow cover, and temperatures. When especially heavy rains are expected, we temporarily cover exposed pots under a sheet of wood or plastic.

Natural Enemies Caterpillars are attacked by a menagerie of pathogens, parasites, parasitoids, and predators. Soil-dwelling species are susceptible to attack by Beauveria, Cordyceps, Metarrhizium, and other entomophagous fungi. Owlet larvae also fall victim to protozoan and nematode parasites. In the southern Appalachians, our spring caterpillar collections often yield a mermithid nematode (Parasitoids Plate, row 1, left); quakers (Orthosia), pinions (Lithophane), and other spring xylenines are among the most common casualties. The early stages of owlet moths host a variety of hymenopteran (wasp) and dipteran (fly) parasitoids. These attacks are usually fatal; and hence, from an ecological viewpoint these enemies function as predators. For this reason,

entomologists refer to them collectively as parasitoids (as opposed to parasites, which often do not kill their hosts). Parasitoids that immediately arrest the development of their host are idiobionts; many egg and pupal parasites are in this category. Koinobiont parasitoids allow their host to feed and develop. Koinobionts may begin feeding as soon as they hatch from their egg, or delay development until their host reaches a certain instar, stage, or size. In univoltine species, koinobionts may diapause for months before commencing their own growth. Most parasitoids grow slowly at first, consuming only blood (hemolymph), their own nutritive cells released into the host, and/or nonvital host cells and tissues, then finish off the host in a burst of feeding and growth, often triggered by the host larva’s size or hormonal state. A common strategy is to wait until the host larva has spun its cocoon or constructed a pupal cell (and thus fashioned a safe haven for the parasitoid), before consuming the prepupa. While the vast majority of the parasitoids attacking owlets are endoparasitoids, developing within the body of their host, a few such as Euplectrus (Eulophidae), are ectoparasitoids that live affixed to the outside of the host’s body (Parasitoids Plate, row 2). Owlet larvae fall victim to a diversity of braconids, ichneumonids, encyrtids, eulophids, and numerous other families of parasitic wasps. The third volume of Krombein et al. (1979) includes a host index with dozens of records for hymenopteran parasitoids. All early stages are attacked. Owlet eggs are parasitized by minute mymarid, scelionid, and trichogrammatid wasps. Minute microhymenopteran wasps—eulophids, encyrtids, among others—attack first or second instars. The importance of such early mortality factors is underappreciated, and we confess that much of our treatment below pertains to the parasitoids one is likely to rear out from collections of late instar owlets. Two microhymenopteran lineages that issue from late instars are worthy of special mention because of their widespread prevalence. Euplectrus and related eulophids can be very common, especially in the South: once fully fed, the larvae crawl beneath the vanquished host and spin a series of dirty white to brown cocoons. The blackened cadaver with its array of cocoons appears to be enveloped by fungus—no doubt, a biological ruse that affords the wasp pupae protection. Polyembryonic encyrtids parasitize many loopers (Plusiinae)—a single egg laid by the female wasp will divide mitotically to yield more than two thousand genetically identical sisters (Strand 1989, Ode and Strand 1995). The whole of the looper’s body will become packed with minute encyrtid larvae and ultimately with the pupae (Parasitoids Plate, row 1, center). Three braconid subfamilies are reared from owlet larvae with regularity: Microgastrinae, Meteorinae, and Rogadinae. At maturity, microgastrines bore out through the body wall and spin small, yellow to white, oval to barrel-shaped cocoons, usually with loose silk over the exterior, on or near

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28 Introduction

Introduction

J Tachinid Flies. Row 1: Three tachinid (macrotype) eggs on Lithophane petulca (left); white arrow points to breathing tube of tachinid inside body of Eucirroedia pampina (center); tachinid maggot just issuing from cadaver of Orthosia hibisci (right).

Row 2: Larva and newly forming puparia of Voria ruralis in cadaver of Autographa metallica (left); tachinid puparium adjacent to (euclidiine) caterpillar host (note necrotic hole from where the larva issued) (center); nearly a dozen tachinid puparia alongside the cadaver of their vanquished Acronicta increta host (right). J

the victimized larva (Parasitoids Plate, row 5). Gregarious microgastrines may issue from the larva in groups of a dozen or more (Parasitoids Plate, rows 4 and 5). More commonly, and especially in the spring, you will encounter solitary microgastrines attacking early to middle instar daggers (Acronictinae), hairy-eyed owlets (Hadenini and Orthosiini), and pinions, sallows, and kin (Xylenini). Upon emerging from the host caterpillar, prepupal Meteorus larvae drop down on a strand of silk and then spin a hanging, often structurally diagnostic cocoon (Parasitoids Plate, row 6, left). Some Meteorus have the ability to snap their cocoons about in a fashion analogous to a jumping bean. Rogadine braconids mummify their victims, pupating within their cadavers (Parasitoids Plate, row 3). Most are solitary wasps that issue from middle instars, although one common eastern Aleiodes is gregarious and emerges from final instar hosts in number (Parasitoids Plate,

row 3, right). Shaw (2006) illustrates the common eastern rogadines that attack owlets. Many Ichneumonidae parasitize Lepidoptera.3 Some, such as the Pimplinae, Cryptinae, and Ichneumoninae, seek out, sting and paralyze, and oviposit onto or into the pupae or the cocooned prepupae of moths. The wasp larva develops by consuming its immotile food resource (i.e., as idiobionts). Some ichneumonine oviposit into the final instar while it is seeking a pupation retreat, and delay their own development until the host caterpillar has constructed its cocoon and/or molted into the pupal stage (i.e., as koinobionts). The habit of attacking a caterpillar and then allowing it to complete its larval development (koinobiosis) is common in a number of ichneumonid subfamilies, including the very diverse and

Opposite J Parasitoids. Row 1: Nematode from Lithophane joannis (left); Trichoplusia ni attacked by Copidosoma, a polyembryonic encyrtid (center); eulophid larvae on Oligocentria lignicolor (Family Notodontidae) (right). Row 2: Eulophid pupae on same Oligocentria lignicolor (Family Notodontidae) (left); Euplectrus larvae on Spodoptera ornithogalli (center); Euplectrus cocoons on same Spodoptera ornithogalli caterpillar-cadaver (right). Row 3: All Aleiodes mummies (Family Braconidae): A. nolophanae on Hypena (left); A. quebecensis on Acronicta

sp. (center); A. stigmator on Acronicta americana (right). Row 4: All microgastrines (Braconidae) on Condica: Larvae exiting C. albigera (left); cocoon bundles of same braconid on C. albigera (center); cocoon bundle and the host, C. videns (right) (image from Valerie Bugh). Row 5: All microgastrines (Braconidae): Cocoon on Orthosia hibisci (left); cocoons on Catocala badia (center); cocoon bundle on Mythimna unipuncta (right). Row 6: Meteorus cocoon from Spragueia jaguaralis (left); campoplegine (Ichneumonidae) cocoon on unidentified noctuoid (center); campoplegine (Ichneumonidae) cocoon on Phyprosopus callitrichoides (right). J

3

Text for the Ichneumonidae was contributed by Ian Gauld in May 2006.

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30 Introduction

J Predators: Row 1: Bark spider (Thomisidae) (left); three Photuris (Lampyridae) larvae consuming pyralid caterpillar (center); Podisus stinkbug (Pentatomidae) attacking larva of Orthosia hibisci (right). Row 2: Assassin bug (Reduviidae) feeding on erebine (left); Zelus (Reduviidae) feeding on erebine (center);

Podisus stinkbug (Pentatomidae) feeding on Coxina cinctipalpis (right). Row 3: Ants (Formicidae) consuming caterpillar of Spodoptera frugiperda (left); Ammophila (Sphecidae) carrying away paralyzed larva of Panopoda rufimargo to be entombed in an underground brood cell, where it will be slowly consumed by the Ammophila wasp’s larva (center). J

frequently encountered Campopleginae, Banchinae, Cremastinae, Anomaloninae, and Ophioninae. In almost all cases, the parasitoid larva begins feeding in the host caterpillar and then suspends its own development until the host has sought out a pupation retreat; the parasitoid’s development is then reactivated, the host caterpillar is consumed, and the wasp spins its own cocoon. Species of Anomaloninae are unusual in delaying development until after the host has pupated. Only among some of the Campopleginae is the host caterpillar killed before it has found a pupation retreat; many spin a black and white mottled cocoon that resembles a bird dropping (Parasitoids Plate, row 6, right) suspended from vegetation by a thread, concealed beneath the cadaver of the host caterpillar, or secreted within the mummified remains of the host. The majority of endoparasitic braconid and ichneumonid wasps attack exposed, externally feeding caterpillars, and consequently tend to have short ovipositors. While most are

diurnal, some, such as Ophion and Enicospilus (Ophioninae), search for hosts at night. A few Banchinae, Campopleginae, and Cremastinae have long ovipositors that they use to probe for concealed hosts, such as those under bracts, in shelters, or boring inside shoots and stems. Although the overwhelming majority of koinobiontic Ichneumonidae are endoparasitic, one common group of Tryphoninae are ectoparasitic koinobionts. These sting the host caterpillar and then attach a black egg externally by a short stalk that is usually placed just behind the host’s head—positioned so that the host caterpillar cannot chew through and destroy the egg. The solitary wasp larva develops externally. Summaries of ichneumonid biologies can be found by visiting the Genera Ichneumonorum Nearcticae pages posted on the American Entomological Institute’s website (www.amentinst.org). Tachinid flies, primarily in the subfamilies Tachininae and Exoristinae and to a lesser extent in the Dexiinae, are among

Introduction Exotic Biological Control Agents It is common practice to release parasitoids and predators from other countries to control introduced pests that have established in our forests and croplands without their normal complement of natural enemies. More than 300 exotic parasitoids have been introduced into North America in the past hundred years to control pest species (Hawkins and Marino 1997). In their compilation, Hawkins and Marino noted more than 50 instances where such exotic parasitoids have been reared from native, nontarget hosts. One example receiving much attention in the East involves the tachinid fly (Compsilura concinnata) (right), which was introduced from Europe to control two lymantriine tussock moths: the Gypsy Moth (Lymantria dispar) and the Browntail Moth (Euproctis chrysorrhoea). The tachinid fly is credited with virtually eradicating the latter (Elkington et al. 2006). In addition to these two introduced pests, the fly parasitoid is known to attack more than 180 native species of Lepidoptera from more than a dozen families, including many noctuoids (Webber and Schaffner 1926, Schaffner and Griswold 1934, Schaffner 1959, Arnaud 1978, Clausen 1978). During Gypsy Moth outbreaks, Compsilura densities sometimes reach 10,000 adult flies per hectare (William et al. 1992). Because Gypsy Moths are only present for the fly’s spring generation, the fly’s second, third, and fourth generations must seek out and parasitize native caterpillars. Boettner et al. (2000) demonstrated staggeringly high mortality rates from Compsilura in field trials of two native giant silk moths in Massachusetts. There is growing concern that Compsilura and other introduced parasitoids are responsible for the widespread decline of silk moth (Saturniidae) populations across many areas of the northeastern United States (Howarth 1991, Boettner et al. 2000, Kellogg et al. 2003, Elkinton and Boettner 2004). In addition to several species of saturniids, it is our impression that some hawk moths (Sphingidae) and datanas (Notodontidae)

the most commonly reared parasitoids of owlets in eastern North America. All are endoparasitic koinobionts. Most tachinids, especially the exoristines, lay their eggs on the surface of the caterpillar; upon hatching, the first instar maggot burrows through the integument, and into the body of the caterpillar. The smooth, white, oval (macrotype) eggs, commonly seen on caterpillars, belong to tachinids (Tachinid Flies Plate, row 1, left). Ovipositing flies tend to place their eggs near the host larva’s head so that the caterpillar will be unable to chew into and destroy the eggs. Hard-shelled macrotype eggs take a few days to develop and may be shed if the caterpillar molts before the fly larvae hatch. More commonly, exoristine eggs are thin-walled; these tend to have shorter incubation periods and hatch soon after they are laid. Other tachinids lay (microtype) eggs by the thousands on leaf surfaces, where they are consumed by feeding caterpillars. These minute eggs hatch within the gut of the host and the neonate fly larvae then tunnel into the host’s body. A few tachinids are unusual in possessing a piercing ovipositor that allows the female to insert eggs or larvae directly into the body of the host, e.g., Eucelatoria females oviposit into loopers (Plusiinae). About half of our tachinids are solitary (a single maggot develops with a host), and the other half are gregarious, with as many as a dozen individuals issuing from a single host larva (Tachinid Flies Plate, row 2, right). At maturity, the maggots of most tachinids crawl from the cadaver, drop to the ground, work their

are not as common as they were four decades ago in the Northeast (see discussion in Schweitzer et al. 2011). The recent declines of some butterflies, e.g., Chlosyne harrisii and C. nycteis, also remain unexplained (Wagner 2007d). Among the taxa treated in this volume, the mysterious declines of three Acronicta from large parts of the Northeast—Barrens Dagger, Paddle Dagger, Witch Hazel Dagger (A. albarufa, A. funeralis, and A. hamamelis, respectively), and the Zebra Arches (Melanchra picta)—may relate to changes in their natural enemy complexes. In sum, the impacts of introduced biological control agents on native species are in critical need of study and assessment.

way into leaf litter, and then pupate within the skin of the last instar maggot, forming a hardened, smooth-walled puparium (Tachinid Flies Plate, row 2 center). Others pupate within the host cadaver (Tachinid Flies Plate, row 2, left). Other lineages delay their development until the host caterpillar has pupated: depending on the species, pupation may occur within the victim’s pupal casing or exterior to it, within the host’s cocoon or pupal cell. Arnaud’s (1978) catalog contains dozens of host records of tachinids reared from moths and other insects. While most tachinids are host specialists, others are generalists—the known hosts of Compsilura concinnata include over 200 species of Lepidoptera (many of which are owlets) as well as a few Hymenoptera (Schaffner and Griswold 1934, Arnaud 1978). Stireman et al. (2006) recently reviewed the biology and evolutionary history of this important group of parasitoids. Parasitism rates generally range between 10% and 25% from collections of wild, late instars, but at times can climb much higher. Attack rates sometimes exceed 75% in collections of stalk borers (Papaipema) and paints (Cucullia). Note that such rates (from single collection events) represent but a slice of the insect’s life cycle and that natural enemies are culling larvae over the entirety of a species’ development. Reared parasitoids should be saved, pinned, and fully labeled, with the identity of the host larva recorded when known. It is also good practice to save associated cocoons, puparia, and mummified hosts. Unhatched puparia and wasp

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32 Introduction Guest Essay on Owlet Classification: Charles Mitter (University of Maryland) Noctuid classification was long dominated by the monumental work of Hampson (1898–1913), despite early recognition of the nonphylogenetic nature of Hampson’s arrangement. Over the past 15 years, however, noctuid phylogeny and classification have been under very active study, resulting in continual change as new evidence emerges. The snapshot of current understanding offered here is based on DNA sequence data, from which especially rapid progress has been made. Its degree of agreement with more traditional evidence from anatomical features is still being evaluated (Lafontaine and Fibiger 2006). Morphological and molecular evidence agree that about 30% of the species of Noctuidae as here defined, including a majority of the major pests, belong to a clade (= monophyletic group) known as the “trifines.” (The name refers to a feature of adult hindwing venation pointed out by Hampson.) There is also strong evidence that about half the trifines, in turn, belong to a clade, sometimes called the “true cutworms” (Lafontaine 1993), that subsumes nearly all of the subfamilies Xyleninae, Hadeninae, and Noctuinae of previous authors. These are

provisionally placed with the Heliothinae and several smaller groups, also of significance as pests, in what has been termed the “Pest Clade” (Mitchell et al. 2006). Relationships among the remaining subfamilies, here informally termed the “lower trifines,” are as yet poorly resolved. Molecular data strongly support the inclusion of most of the remaining Noctuidae, some 23,000 species, in what has been termed the “L.A.Q.” clade (Mitchell et al. 2006). As the acronym is meant to signify, this group consists of the traditional families Lymantriidae and Arctiidae plus most of the so-called “quadrifine” (as opposed to “trifine”) Noctuidae in traditional classifications. Relationships are uncertain for two “quadrifine” groups not included in the “L.A.Q.” clade, though the most recent results suggest that the Nolidae are related to the “trifines.” Relationships within the vast “L.A.Q.” clade, heretofore poorly understood, are now undergoing intensive molecular and morphological study (R. Zahiri, J. Zaspel, pers. comm.). In addition to its importance for classification, a phylogeny is of cardinal utility in helping us understand how, and in what Earth-history context, the

Introduction Guest Essay on Owlet Classification: Charles Mitter (University of Maryland) continued enormous life history diversity of noctuids has evolved. Full exploration of this subject, barely begun, will require much new information on life history and phylogeny. A speculative example of the kind of insights that may emerge is provided by several broad differences in use of hostplants and geographic distribution between noctuid clades, visible in the phylogeny provided here (see detailed discussion in Mitchell et al. 2006). Although exceptions abound, the trifines appear to be predominantly herb feeders, whereas the remaining noctuids show a greater proportion of lineages feeding on trees, probably the ancestral habit for Noctuidae and most other so-called macrolepidopteran families. Trifines also show on average much greater association than other noctuids with open habitats, and with both high latitudes and high elevations (Holloway and Nielsen 1998). Accumulating evidence links these trends to the dramatic global climate changes of the Cenozoic era. Noctuids may have

cocoons can sometimes be identified to genus and thus have value in a collection. There is a special need for data on the host ranges of introduced (nonnative) parasitoids employed as biological control agents (see text box page 31). Owlet caterpillars are attacked by hundreds of invertebrate predators (Predators Plate): spiders, assassin bugs, predaceous stink bugs, lacewing larvae, ladybugs, ground beetles, sand wasps, vespid and related wasps, and ants. Yellow jackets, paper wasps, and ants are chronic threats to owlet larvae. Caterpillars that feed or take refuge on the ground are exposed to other enemies including centipedes and ground-dwelling spiders, ants, beetles, etc. The importance of mites that attack eggs and early instars is undoubtedly considerable but not well understood. Caterpillars are a staple of songbirds and comprise the main, or in some cases, the sole food for both nestlings and adults (Holmes 1990; text box on page 9). As such, birds have been a major selective force in caterpillar evolution, influencing what caterpillars look like and how they behave. Birds have been implicated not only in what some caterpillars eat but also when and how they eat (Heinrich 1979, 1993). Rodents, especially mice, shrews, and voles, are caterpillar and pupa hunters, and replace birds in importance as predators of grounddwelling caterpillars.

Classification and Nomenclature The Noctuoidea is one of two hyperdiverse superfamilies of Lepidoptera, with world species diversity likely to exceed 75,000 species. It is rivaled only by the Pyraloidea whose tropical diversity, in places, seems almost unbounded. Given the extraordinary evolutionary success of the Noctuoidea, perhaps it is to be expected that the higher classification of the superfamily remains in a state of flux, even after more

originated as tree feeders in the mesic to wet tropical forests that covered much of the earth starting early in the Cretaceous period (145–65 million years ago). Beginning about 50 million years ago, there has been an overall trend toward cooler and drier climates, latitudinal zonation of vegetation, and, particularly at higher latitudes, expansion of strongly seasonal and open habitats. The latter, in turn, prompted diversification of herbaceous angiosperm groups. While other moth lineages probably became increasingly restricted to the tropics, trifines and a few other insect groups (e.g., Winkler et al. 2009) seem to have been predisposed for adapting to this new ecological opportunity, analogous to the diversification of grazing mammals that accompanied the spread of open habitats including grasslands. These same habitats, of course, fostered the origins of human agriculture, pitting us against the trifines in a continuing global smackdown.

than a century of study and taxonomic shuffling. Commonly adopted classifications for the North American owlet fauna have drawn heavily from the framework established in Franclemont and Todd’s (1983) checklist, which in turn was based on Hampson’s (1898–1913) many works. A number of rearrangements to the Franclemont and Todd system were proposed by Poole (1995) in his phylogenetic treatment of the higher classification of the trifid noctuids. Recent studies on the classification of the superfamily have been fueled by studies of internal (especially male genitalia) and external morphology, immature stages, and molecular characters (e.g., Poole 1995; Speidel et al. 1996; Mitchell et al. 1997, 2000, 2006; Kitching and Rawlins 1998; Beck 1999, 2000; Fibiger and Lafontaine 2005; Lafontaine and Fibiger 2006; and Zahiri et al. 2011). Molecular data strongly suggest that two large lineages formerly regarded as full families, the tiger moths (Arctiinae) and tussock moths (Lymantriinae), derive from within the Erebidae. With but a handful of exceptions, we follow Lafontaine and Schmidt’s (2010) North American checklist to arrange the species treated in this work. Their spellings for species names are also adopted. A summary classification appears below for those taxa represented here by one or more species. We selectively include tribes when these help to subdivide large subfamilies and/or heterogeneous subfamilies, e.g., Erebinae and Noctuinae. For a complete listing of the higher taxa within the North American Noctuoidea consult Lafontaine and Schmidt. Other authors treat subfamilies recognized in this work as full families: e.g., Mitchell et al. (2006) suggested that the Herminiinae, Hypeninae, and others could be given family-level status. The phylogenetic positions of many groups (e.g., Nolidae) may change, given the disagreement among existing studies. Due to the sheer enormity of the superfamily, and because many tropical, diminutive, and taxonomically isolated taxa have yet to be included in molecular studies, any treatmentmust be regarded as provisional—expect much taxonomic shuffling in the Noctuoidea over the coming decades.

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34 Introduction Higher Classification for the Owlets Treated in this Work4 Family Erebidae Leach, [1815] Subfamily Lymantriinae Hampson, [1893] [omitted] Subfamily Arctiinae Leach, [1815] [omitted] Subfamily Herminiinae Leach, [1815] Subfamily Pangraptinae Grote, 1882 Subfamily Hypeninae Herrich-Schäffer, [1851] Subfamily Rivulinae Grote, 1895 Subfamily Scoliopteryginae Herrich-Schäffer, [1852] Subfamily Calpinae Boisduval, 1840 Subfamily Hypocalinae Guenée, 1852 Subfamily Scolecocampinae Grote, 1883 Subfamily Hypenodinae Forbes, 1954 Subfamily Boletobiinae Guenée, [1858] Subfamily Phytometrinae Hampson, 1913 Subfamily Erebinae Leach, [1815] Tribe Catocalini Boisduval, [1828] Tribe Melipotini Grote, 1895 Tribe Euclidiini Guenée, 1852 Tribe Poaphilini Guenée, 1852 Tribe Ophiusini Guenée, 1837 Tribe Thermesiini Guenée, 1852 Subfamily Eulepidotinae Grote, 1895 Family Euteliidae Grote, 1882 Family Nolidae Bruand, 1846 Subfamily Nolinae Bruand, 1846 Subfamily Chloephorinae Stainton, 1859 Subfamily Risobinae Mell, 1943 Subfamily Collomeninae Kitching & Rawlins, [1998] Subfamily Afridinae Kitching & Rawlins, [1998] Family Noctuidae Latreille, 1809 Subfamily Plusiinae Boisduval, [1828] Tribe Abrostolini Eichlin & Cunningham, 1978 Tribe Argyrogrammatini Eich. & Cunning, 1978 Tribe Plusiini Boisduval, [1828] Subfamily Bagisarinae Crumb, 1956 Subfamily Cydosiinae Kitching & Rawlins, [1998] Subfamily Eustrotiinae Grote, 1882 Subfamily Acontiinae Guenée, 1841 Subfamily Pantheinae Smith, 1898 4 The backbone of the classification used in this work was reproduced from Lafontaine and Schmidt (2010). However, our concept of the Amphipyrinae differs. We treat the Stiriinae as a separate subfamily (not as a subordinate tribe within the Amphipyrinae), and we regard the Phosphilini to be part of the Amphipyrinae, not a tribe within the Noctuinae. Additionally, we believe the larval characters and biology of Ufeus are so divergent as to be worthy of tribal recognition. (Lafontaine and Schmidt (2010) treat the Ufeina as a subtribe within the Xylenini.) As noted previously, two recent additions to the Erebidae, the Lymantriinae (tussock moths) and Arctiinae (tiger moths), were treated in Wagner (2005).

Subfamily Diphtherinae Fibiger & Lafontaine, 2005 Subfamily Dilobinae Aurivillius, 1889 Subfamily Balsinae Grote, 1896 Subfamily Acronictinae Heinemann, 1859 Subfamily Agaristinae Herrich-Schäffer, [1858] Subfamily Cuculliinae Herrich-Schäffer, [1850] Subfamily Oncocnemidinae Forbes & Franc., 1954 Subfamily Amphipyrinae Guenée, 1837 Tribe Amphipyrini Guenée, 1837 Tribe Phosphilini Poole, 1995 Tribe Psaphidini Grote, 1896 Subfamily Stiriinae Grote, 1882 Subfamily Heliothinae Boisduval, [1828] Subfamily Condicinae Poole, 1995 Subfamily Eriopinae Herrich-Schäffer, [1851] Subfamily Noctuinae Latreille, 1809 Tribe Pseudeustrotiini Beck, 1996 Tribe Elaphriini Beck, 1996 Tribe Prodeniini Forbes, 1954 Tribe Caradrinini Boisduval, 1840 Tribe Dypterygiini Forbes, 1954 Tribe Actinotiini Beck, 1996 Tribe Phlogophorini Hampson, 1918 Tribe Apameini Guenée, 1841 Tribe Arzamini Grote, 1883 Tribe Xylenini Guenée, 1837 Tribe Ufeini Crumb, 1956 Tribe Orthosiini Guenée, 1837 Tribe Tholerini Beck, 1996 Tribe Hadenini Guenée, 1837 Tribe Leucaniini Guenée, 1837 Tribe Eriopygini Fibiger & Lafontaine, 2005 Tribe Glottulini Guenée, 1852 Tribe Noctuini Latreille, 1809

About This Book Each family, subfamily, and less frequently tribes and larger genera, are introduced with information about diversity and distribution and general interest tidbits, followed by an abbreviated diagnosis and life history notes. Tips for collecting and rearing are sometimes added at the end of an introductory section. Additional information is salted into the Remarks sections for the species within a given group. Members of a genus are usually listed in alphabetical order, except in large genera (e.g., Catocala, Lithophane, and Zale), where related species and/or those similar in appearance or habit are grouped. Each species account begins with a common name followed by the scientific name. Many of the former are our own, and are often based on an English rendering of the scientific name or a derivation of a related British species. Recent synonyms

Introduction (enclosed by parentheses and preceded by an equals sign) sometimes follow the scientific name. For those species where different common names have been applied to the caterpillar and adult, we often use the name for the larva and place the adult name in brackets after the scientific name. Additional common names are sometimes added to the brackets or mentioned in the Remarks section, although we make no effort to provide a full reporting of common names, which can be numerous for economically important species. Our use of common and scientific names is idiosyncratic. We generally supply both, but not always; for example, we often skip mention of the Latin name when it is an obvious derivative the common name (e.g., Bethune’s Zale and Zale bethunei). We favor common names for familiar species but switch to Latin names when dealing with more difficult taxonomic matters, issues, or species. When mentioning a second species in an account other than the subject species, we apply boldface to draw attention to the fact that some included information applies to a second entity. We generally adopt the common name accepted by the Entomological Society of America and BugGuide, but propose replacement names where we regard the accepted names to be factually misleading or inappropriate based on current taxonomy, hostplant data, etc. Additionally, we propose new common names for a number of unfamiliar moths for which no common name has so far been applied. For several subfamilies, we abandoned nonsensical or poorly considered names and applied a uniform naming system to bring the common names in line with existing classifications, e.g., Hypeninae are dubbed snouts and Plusiinae are called loopers. Common names of moth species are capitalized. Full species accounts include four sections: Recognition, Occurrence, Common Foodplants, and Remarks. In the Recognition section we provide a description of the last instar; especially useful characters are italicized. (For taxa with one or more close relatives in our fauna and when we are uncertain that our description is diagnostic, no features have been italicized.) Each description ends with a generous estimate of the caterpillar’s length. If known, and where coloration differs from that of the final instar, we frequently note the appearance of the middle and/or penultimate instars. The Recognition section often includes mention of congeners or other species likely to be confused with the focal species. The Occurrence section begins with a listing of habitats frequented by a given species followed by a synopsis of the range, usually beginning with the northwestern corner of the distribution and cycling clockwise. We have endeavored to summarize the North American range east of the hundredth meridian; occasionally, and especially for widespread species, we mention the western and southern range limits. By convention, the range is followed by a synopsis of the number of annual generations and phenology (seasonal activity). For most Lepidoptera, the number of generations and the overlap

between generations increase as one moves southward where species begin activity earlier in the year and fly later into the fall. Unless noted otherwise, we emphasize the life cycle and phenology that would be expected from northern Georgia to southern Canada. This section concludes with an abbreviated assessment of a species’ abundance. Common Foodplants are listed for every species. We have not attempted to provide a complete listing of known host plants, especially for those species with catholic diets. Robinson et al. (2002, 2011) are the best single-source compendia for foodplant data. Other important sources include Forbes (1954, 1960), Crumb (1956), Prentice (1963), Tietz (1972), Ferguson (1975), Rings et al. (1992), Covell (2005), and Heppner (2007). Taxon-specialized references are listed in the boxes that introduce the subfamilies. For general feeders, hosts are listed in alphabetical order. When foodplant preferences are apparent, we make an effort to mention them. Common names for the plant hosts closely follow those provided by the USDA website PLANTS Database (http://plants. usda.gov/index.html), although we do adopt a few considered exceptions. For example, we continue to use the traditional sense of the genera Aster and Solidago because we do not find that the proliferation of newly proposed generic names helpful for understanding the owlet diets in this guide. When the common name of the foodplant is likely to be unfamiliar or ambiguous, we provide the host’s scientific name parenthetically. Many of the host records are our own; only occasionally do we insert our initials (e.g., in instances where we deem our record to be significant). The Remarks section is a potpourri of information on a species’ life history, behavior, taxonomy, economic importance, and other observations likely to generate interest in or appreciation for the insect and its caterpillar. Notes on husbandry and tips on how to locate larvae are given here. Extensive unpublished data appear in this section. Information relevant to members of a given genus or group has sometimes been divided among the Remarks sections of related species. For example, if the overwintering stage is constant across a genus, mention of this may be made only once, either in the introductory box or in the Remarks section of the first member of the genus treated. Less obviously, when there was available space on a page, we frequently filled the Remarks section with information pertinent to other members of the focal species’ genus, tribe, or even subfamily.

Making Identifications Descriptions in this treatment apply to the last instar. Unfortunately, there is no easy way to know when an unfamiliar caterpillar is fully grown. We still occasionally fail to recognize last instars of very small species collected in the wild before

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36 Introduction transformers Caterpillar guides are given to treatments of the last instar, in large measure because this is the most conspicuous; in the case of pests, the last instar is the most destructive and therefore receives a lot of attention. For greater than half the species in this guide, our last-instar image will allow one to identify a penultimate or even a middle instar of that same species. But in others, remarkable changes in form and habit transpire between the nascent and mature larva. Those that we record for many daggers are especially noteworthy (pages 258–99)— differences between the last two instars may exceed those that occur between the genera and subfamilies of other moths. More remarkable still is that a few of these “masters of metamorphosis” even change their phenotype and modus operandi within a single instar. Take the case of the Greater Oak Dagger (Acronicta lobeliae) (below). The early and middle instars resemble leaf blemishes and rest near their feeding site. The pale last instar invariably rests off of foliage, on twigs, where its coloration is matched to oak bark. But late in the last instar the caterpillar changes yet again, transitioning to a black-headed, dark caterpillar with pale striping that resembles some Datana (Notodontidae). Our guess is that this is an adaptive form, suited to the ground and the business of tunneling into downed wood, the preferred pupation site. Even more remarkable changes occur in its distant kin, Cadbury’s Mystique (Comachara cadburyi) (page 303) and the Hebrew (Polygrammate hebraeicum) (page 302).

1st instar

3rd instar

5th instar

they pupate. While one might generalize that many noctuoid caterpillars exceed 3 cm in length at maturity, the number of exceptions is legion; e.g., nolines are under 1.5 cm when mature, and hypenodines can be much smaller. For many species, we provide an ancillary image of the penultimate instar when its coloration differs significantly from that of the mature larva. Familiarize yourself with the scope of this guide prior to using the work for identification purposes: flip through the species accounts, read the subfamilial and tribal boxes, and look over the images to develop a “gestalt or feel” for the major groupings. Conversely, when closing in on an identification, be sure to back up and read tribal and subfamilial boxes to make sure that your caterpillar possesses the features common to the group. Variation in coloration and patterning is the rule, especially in the darts (Noctuini). Don’t expect your caterpillar to exactly match any image; conversely, even when a caterpillar seems to be a match for a species figured here, make sure your unknown possesses the characters given in the diagnosis. Features such as head capsule shape, setal lengths and patterns, proleg development, and warting are less variable—these should mirror the features of your unknown. Keep in mind that some shape features, such as the form of the rump, are dependent on whether the caterpillar is ambulatory or at full rest. Characters that we feel are especially reliable and helpful are italicized. In many subfamilies, knowledge of the foodplant will help reveal a caterpillar’s identity. With unusual hosts, it is possible to zero in on an identification by working backward from the foodplant index. Do not assume that the plant on which you have found a caterpillar is the host; caterpillars wander. Make a point to check for recent feeding damage when you discover a caterpillar or, in the case where a caterpillar has been brought home, watch for signs of feeding and/or the production of feculae.

last instar (feeding)

prepupal last instar

If you cannot find your caterpillar in this book, you might also consult works on the noctuoids of Europe and eastern Asia, e.g., Sugi (1987), Porter (1997), Beck (1999, 2000), and Ahola and Silvonen (2005, 2008, 2010). In many cases you will be able to get an approximate identification by checking these richly illustrated volumes. Bug Guide (http://bugguide.net/node/view/15740), Moth Photographer’s Group (http://mothphotographersgroup.msstate.edu/), and list serves can be powerful and immediate sources of help with troublesome larval (and adult) identifications. Cast around and do some homework before making posts—others are more likely to lend assistance when it is evident that a person has consulted easily accessible sources. Include pertinent data that can help in identification: host, date, and locality. Posts that contain such data and other observations serve a greater good.

Voucher Data and Specimens For the majority of the species in this work, larval and adult specimen vouchers, original photographic images, rearing notes, and associated databases are those of the senior author and have been deposited at the University of Connecticut. Lesser numbers of larval and adult specimen vouchers are housed in the collections of DFS, Eric Quinter, and Tim McCabe. With the exception of 32 species loaned by Eric Quinter, all vouchers for the pinned adult images are at the University of Connecticut. The adult specimens imaged for this work bear the label “USDA Forest Service Noctuid Cater. Guide.” Data for the adult images (state only) and figured larvae (locality, date, foodplant, collector, and photographer) are available from the senior author. While we endeavored to use images of larvae

Introduction reared from eggs deposited by gravid females or wild-collected caterpillars that yielded identifiable adults, we have included images of caterpillars that were not successfully reared to the adult stage (for this reason and others, some images may be wrongly associated with a given species).

Diagnosing Owlet Caterpillars Owlet caterpillars are fantastically rich in species and heterogeneous in form. No single character will ensure immediate recognition.5 They range from slender to chunky, nearly glabrous to densely hairy, brown and nondescript to boldly aposematic, adorned in reds, yellows, or oranges. Setae may be straight or barbed. All have the crochets arranged in a series more or less parallel to the body axis (a mesoseries) (Fig. 9), never in a circle or incomplete circle. In most, the crochets are of a single length. Many have a midventral prothoracic gland, the adenosma (often visible as an everted tubular gland in preserved material). The head is usually smooth and rounded; a coronal bar to either side of the dorsal midline, and reticulate mottling over each lobe, are common. There are only two prespiracular (L group) setae on T1 (Fig. 10). In nearly all, there is only one SV seta on T2 and T3, although Arctiinae, some Plusiinae, and others have two (Beck 2009). Setae L1 and L2 on A3–A6 are usually below the level of the spiracle. The classic cutworm is smooth and stocky, with relatively short prolegs and reduced, inconspicuous setae; these are usually colored in earth tones. The SD1 seta on A9 is fine and hairlike (narrower in diameter than the width of the dorsal setae) in the cutworms and kin. Middorsal, subdorsal, and spiracular stripes are widespread. Conspicuously striped or ringed macrolepidopterans have a high probability of being owlets. Most have a full complement of prolegs (on A3, A4, A5, A6, and A10), although in some taxa the first, or first and second, pairs are either reduced or entirely absent. It is easier to exclude taxa that are not owlets: none have branched spines (scoli) on the body or head. With the exception of the Green Marvel (Agriopodes fallax), they are not sluglike with minute secondary setae and the head is never fully retracted within the thorax (as in Lycaenidae and Limacodidae). If secondary setae are present, there is no anal point (between the anal prolegs, extending up toward the anus). Thyatirine drepanids, with two extra L group setae on A1–A8, come closest in 5 In Stehr’s (1987) key to the families of lepidopteran larvae, owlets appear in nearly a dozen places, all in the second half of his 225-couplet/ triplet key.

appearance (see Wagner 2005: 141). The anal prolegs are not reduced, greatly lengthened, or otherwise modified, as is commonly the case with prominents (Notodontidae). The labral notch, which engages the leaf when a larva is feeding, is V-shaped in notodontids and usually U-shaped in erebids and noctuids. In sum, if the crochets are arranged in a mesoseries, prolegs are present on A5, the anal prolegs are normal, and you have eliminated the common macrolepidopteran families as a possibility, you are probably looking at an owlet.

Keys to Owlet Larvae Below we list works that include larval keys for large numbers of eastern North American species.6 Full citations for each publication are given in the “Cited Literature” section at the end of this book. Works treating the larvae of subfamilies, tribes, or smaller numbers of species are mentioned in the introductory boxes for those taxa. Crumb, S. E. 1956. Comprehensive treatment of North American owlet larvae with keys to most North American subfamilies, tribes, genera, as well as many species. Forbes, W. T. M. 1954. Species keys for northeastern Acronicta, Catocala, Zale, and a few others; no familial or subfamilial key. Godfrey, G. L. 1972. Keys and descriptions for approximately 45 genera and 150 species of North American Hadenini. Godfrey, G. L. 1987. Key to about 70 species of commonly encountered North American owlets, including all major pest species. Whelan, D. B. 1935. Key to 25 common pest species attacking corn in Nebraska; most are darts (Noctuini).

Supplemental Digital Resources DLW’s website, linked at http://press.princeton.edu/titles/9420. html, includes an erratum and other ancillary resources to this book. Several owlets not treated in this work have accounts on the website. Others that appear here as abbreviated accounts have full accounts on the website. A draft key on the website will enable users to identify larvae of many eastern owlet caterpillars to subfamily and, in some cases, genus. 6 Beck’s (2000) key to European owlet caterpillars will work for many North American taxa, at least to the level of subfamily and tribe, and occasionally to genus.

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Quadrifine Owlets Family Erebidae

E

rebids are a splendidly heterogeneous lot, varying substantially in form and life history. The family is fantastically diverse in the tropics, where thousands of new species await description. We omit mention of two erebid subfamilies, the Lymantriinae and Arctiinae, which have been treated elsewhere (e.g., Ferguson 1978 and Wagner 2005). Phylogenetic relationships within the erebids were the focus of a recent, much-needed molecular study by Zahiri et al. (2011). In the adult, the hindwing appears to have four veins leaving the lower reach of the discal cell (and hence the common name quadrifine or quadrifid owlets). We know of no larval characters that unite the family’s members: the trunk is often slender and the anal prolegs directed backward (at rest). Most have three SV setae on A1 and A2, and SD1 on A9 is unmodified (except in some Euclidiini where it is fine and hairlike). In many the anterior prolegs are missing or reduced. The majority feed on trees and shrubs, although diets vary widely across the Erebidae: among their ranks are lineages that feed on algae, fungi, detritus, rotting wood, grasses, and sedges. Given the great diversity in kind and habit across the family, our strategy is to abbreviate what would be an unwieldy and likely flawed attempt to characterize the family; instead we emphasize treatment of the various erebid subfamilies.

Litter Moths

Family Erebidae: Subfamily Herminiinae Litter moths are among the most numerous owlets in many woodlands, forests, and marshes, where they sometimes outnumber all other owlet subfamilies in midsummer blacklight trap samples. Herminiines are also diverse (and woefully understudied) in the Neotropics. Adults display a marvelous array of scent-dispersing organs that are used during the courtship and mating of these otherwise mostly unheralded owlets. Forbes (1954) includes mention of 54 northeastern species of herminiines. Additionally, the eastern fauna includes several undescribed taxa, including a few that are widespread and well known to collectors. The brown and rather undistinguished caterpillars are slow moving with short, inconspicuous, and often peglike setae. Many bear a dark pseudospiracle on T2 and T3. Renia, Zanclognatha, and related genera bear a curious network of red reticulations over the body. In many, the trunk appears corrugated (narrowed between segments). The head tends to be small and rounded (Sugi 1987). Typically, the integument is roughened with minute creases or granules, warts, teeth, or dense, short setae. SD2 arises below the lower margin of the prothoracic shield (Crumb 1956). T2 bears only one SD seta in Renia; several genera possess just a single L seta on T2. In most, the SV group is trisetose on A1. The first pair of prolegs is reduced (or absent) in Idia. Herminiine caterpillars are surprisingly varied in form and coloration given their secretive habits, i.e., they

are essentially concealed feeders. Idia larvae are especially diverse in character. Differences in form (and one might guess microecology as well) may occur between instars—see, for example, the account for the Undulate Renia (R. factiosalis). Herminiines appear to have been a favorite with Crumb—in his 1956 monograph he provided larval descriptions for 30 of the eastern species. Owada (1987) is a rich source of information for the Asian fauna. Herminiines consume a wide range of organic matter including living foliage, algae and lichens and other epiphytic growth, as well as fallen leaves, flowers, and fruits in various states of decomposition; a few consume dead insects, fungi, vertebrate dung (Rawlins 1984, Owada 1987), and organic matter in and about animal nests. The degree to which they are seen in beating samples varies markedly among genera.

! Forest floor-dwelling herminiines graze on dead leaves, removing all tissues but the veins. This leaf from a lab colony was skeletonized by many larvae over the course of a week !

Litter Moths Subfamily Herminiinae 39 Lascoria, Palthis, and some Zanclognatha feed above the ground and thus are taken by beating; in contrast, few Idia and most Renia do not feed on vegetation, preferring fallen leaf litter. Four Idia have been reared from animal nests. The Common Idia (I. aemula) occasionally breeds in squirrel nests. American Idia (I. americalis) caterpillars have been found in the nests of ants (Formica) (Crumb 1956), squirrels (Pete Kovarik, unpubl. data), and birds (Eric Quinter, unpubl. data). The Major Idia (I. majoralis) was once reared from pupae found in a nest of the eastern woodrat (Rings et al. 1992). I. gopheri, so far as is known, lives in the burrows of the Florida gopher tortoise and nine-banded armadillo (evidently the larvae feed on organic matter within the tunnels) (see Schweitzer et al. 2011). Several feed on living plant tissues; in Asia some of these are green and strikingly different in appearance and habit from our earth-colored North American species (see Sugi 1987). Sigal (1984) reared Zanclognatha theralis from old man’s beard (Usnea). Litter moth caterpillars are lethargic and play dead when molested. Most shun light. The majority overwinter as half-grown caterpillars in leaf litter; at least a few pass the winter as pupae. As conditions allow, larvae may feed throughout the winter months. Prepupae, which sometimes take on a characteristic red-orange cast in Zanclognatha and others, spin a sparse cocoon into which soil and frass may be woven.

Long-horned Owlet Hypenula cacuminalis

At least South Carolina to Florida, west to Texas. Multiple generations in Deep South with mature caterpillars essentially year-round. Common. Presumably mostly dead organic matter and especially dead leaves, but ours avidly consumed recently picked pokeweed.

Angulate Fan-foot

Macrochilo litophora [Brown-lined Owlet] Wisconsin east to Quebec, and south to Carolinas, Mississippi, and Texas. One generation in North, two generations in southern Ohio and Missouri with mature caterpillars in early spring, then again in midsummer. Generally infrequent. Live and dead grasses, clover, and presumably other organic matter.

Adults come readily to light and bait. Net collecting during the day can be an effective means of securing females: e.g., many renias can be flushed in number during the day. Searching vegetation at night with a headlamp or flashlight may also produce good numbers of females relative to adjacent sheets or light traps. Herminiines lay eggs readily in captivity, depositing their ova singly or, more commonly, in clusters. While most will lay in an empty vial, it sometimes helps to add a strip of slightly moistened paper, corrugated cardboard, or oak leaf litter as an oviposition substrate. We rear most on dead oak leaves placed over a 2–4 cm bed of lightly moistened peat; dead birch or black cherry leaves also work well. We occasionally augment the litter with dead grass or sedge blades, and in the springtime with fallen oak or hickory catkins. Many wetland species can be reared on withered sedge or grass leaves. If conditions allow, use a container in which you can set up a moisture gradient in the litter: from damp (or even decaying) leaves at the bottom to dry leaves on top. Some ventilation is recommended. Watch out for mite infestations, especially if you are tending multiple lots—if you find pale, unpigmented mites on your larvae, remove them, and then transfer surviving larvae to new containers with freshly collected litter. If you have access to a dissecting microscope, it is easy to remove unwanted mites with a camel brush or needle.

40 Litter Moths Subfamily Herminiinae

Common Idia Idia aemula ! recognition Smoky brown, dark gray, or occasionally with pinkish hue, and dark, blurred middorsal and subdorsal stripes. Proleg on A3 reduced with crochets shortened. Peglike setae (which may be orange in color) borne from elevated black pinacula. Setae over dorsum about three to four times length of pinaculum diameter. D1 setae directed forward and D2 setae rearward. Integument granulose. Head with brown reticulate patterning; length of median line (from top of triangle to rear of head) twice height of triangle. Brown to black spiracles broad with narrow rims. Larva less than 2 cm. Larva perhaps indistinguishable from that of the closely related, but rarer, Pale-winged Idia (Idia new species = I. concisa of Forbes) (page 42), with which it co-occurs over much of the East. Also resembling American Idia (I. americalis), but that species lacks prolegs on A3 (see page 42). ! occurrence Woodlands and forests from Canada, south to Florida and Texas. Three generations over much of East with mature caterpillars occurring nearly year-round. Abundant. ! common foodplants Various forms of organic matter. Crumb (1956) reported larvae to be an occasional pest of corn fodder. Our ex ova caterpillars were reared on dead leaves; Warren Steiner reared adults from larvae collected from dry oak leaves of a squirrel nest; Mary Jane Hatfield reared an adult from a larva found tunneling in an earthball (Scleroderma cepa); and an adult in the United States Museum of Natural History was bred from an (evidently abandoned) bald-faced hornet’s nest. ! remarks The Common Idia is one of the most abundant moths of eastern woodlands, yet we have not found its larva in the wild. Presumably most caterpillars feed on fallen leaves and other forest floor litter. The network of veins left by grazing larvae is lacelike—skeletonized white oak leaves are reminiscent of lace doilies (page 38). Given the numerical dominance of herminiines in the East, it seems likely that they play a modest role in macrodecomposition of plant litter, nutrient cycling, and soil formation in some forest types—their numbers can be especially high in oak woods and forests. Underwing (Catocala) hunters who use sugar baits are well acquainted with the genus. In most woodlands and summers, Idia will outnumber and outlast Catocala, flying into November.

Litter Moths Subfamily Herminiinae 41

Glossy Black Idia Idia lubricalis ! recognition Smoky with roughened integument and short, amber peglike setae borne from raised or hornlike pinacula. Black (D1) setal bases alternate with tan, yellow, or orange (D2) setal bases over abdomen. Projections smaller and in linear band across thoracic segments. Lateral setae arising from black bases perched on low, scaly warts that may be somewhat orange in color. Proleg on A3 little more than a knob; crochets absent. Head black, granulose with conspicuous peglike setae. Larva less than 2.5 cm. Middle instars with dorsal setae borne from more dramatically raised “horns” (inset). Raised dorsal protuberances of Glossy Black Idia distinguish it from other eastern Idia with the exceptions of I. denticulalis (page 42) and I. laurentii (page 44). ! occurrence Woodlands and forests from Canada to Florida and Texas. One principal generation with at least partial second generation over much of East: mature caterpillars from May to August. Multiple broods southward. Common. ! common foodplants Fungi and lichens; perhaps generalized on organic matter of forest floor, and at least occasionally on low living foliage. Ours reared on leaf litter. ! remarks We wonder if the thickened setae of various herminiines are secretory in nature. Interestingly, similar peglike setae also occur in litter-feeding geometrids (e.g., Idaea). Size and pattern variation among adults is extensive. Likewise, larvae differ dramatically in appearance; some have proportionately enormous dorsal warts while in others these warts are reduced. Barcodes of eastern collections segregate into at least four distinct groupings (JBS)—it seems certain that more than one species is going under the name lubricalis in collections. Herein lies an easy target for someone with a penchant for rearing: put females of different “forms” up for eggs and photodocument the offspring and the adults that you rear out. In the wild, caterpillars of the Glossy Black Idia are often found on or close to fallen wood (Crumb 1956).

42 Litter Moths Subfamily Herminiinae

American Idia Idia americalis

Southern Canada to Florida and Texas. Multiple generations with mature caterpillars year-round. Very common. Dead leaves and other detritus; lichens. Close to I. aemula but proleg on A3 absent, crochets longer, and spiracles more narrowed.

Pale-winged Idia

Idia n. sp. = I. concisa of Forbes 1954 Range unclarified, but widespread across much of eastern North America. One generation in Northeast with mature caterpillars from spring into June. Locally common. Several records from conifers (see Prentice 1962 and Rings et al. 1992); dead cherry leaves (Tim McCabe, unpubl. data). Presumably close to I. aemula, but diagnostic features unstudied.

Toothed Idia Idia denticulalis

Wisconsin to New England, south to Alabama and Texas. Apparently one generation in Connecticut with mature caterpillars into early summer; two or more generations southward. Locally common. Dead leaves and other detritus; lichens. A1–A6 with black dorsal triangles toward rear of segment, D1 wart encircled by pale ring over abdominal segments; often with D2 setae on pale, elevated warts on A7–A9.

Litter Moths Subfamily Herminiinae 43 Orange-spotted Idia Idia diminuendis

Wisconsin to Nova Scotia, southward to Florida and Texas. One principal generation in Connecticut with mature caterpillars from spring into late June; more broods southward. Locally common. Dead leaves and other detritus. Dorsal setae short, apically broadened, and prothoracic shield dingy orange; compare with I. rotundalis.

Forbes’s Idia Idia forbesii

Wisconsin to Quebec, south to Florida and Texas. One principal generation in Connecticut with mature caterpillars from spring into June and July, and smaller brood in late summer (Eric Quinter, unpubl. data); additional broods southward. Locally common in dry woods. Dead leaves and other detritus. Integument minutely spined; dorsal setae thin, curved, pale. Head red orange, unpatterned.

Gopher Tortoise Idia Idia gopheri

Georgia to central Florida, west to Louisiana. Multiple generations through growing season. Local and rare. Presumably tortoise scat and detritus; also associated with armadillo burrows and nest. Comparatively large, body unpigmented (translucent) with prominent dorsal peglike setae.

Julia’s Idia Idia julia

Southern Canada to Georgia and Texas. One principal generation in Connecticut with mature caterpillars from spring into late June; two generations on Long Island and multiple broods southward. Common. Dead leaves and other detritus. Nearly black, integument fuzzy with abundant pale setae; some dorsal setae swollen apically.

44 Litter Moths Subfamily Herminiinae

Laurentine Idia Idia laurentii

Central New York to North Carolina (mountains) (Forbes 1954). One generation with mature caterpillars from spring into early summer. Uncommon to rare northward, locally common in southern Appalachians. Reared on dead cherry leaves by Tim McCabe (unpubl. data). Our larval images and bar codes suggest I. laurentii is two species. We do not know if our figured individual is nominal I. laurentii.

Rotund Idia Idia rotundalis

Southern Canada to Florida and Texas. One principal generation in Connecticut with mature caterpillars from spring into summer; more broods southward. Very common. Dead leaves, fungi, and other forest floor organic matter. Elongate, mostly unpigmented; reduced A3 proleg without crochets; dorsal setae thickened.

Smoky Idia Idia scobialis

Michigan, southern Quebec, and Maine to Florida and at least Kentucky. Much of East with one principal and partial second summer generation; mature caterpillars from spring into early summer, then again in July and August. Locally common. Dead leaves and other detritus. Ground color dark with smoky middorsal stripe; peglike setae as long as distance between D1 and D2 pinacula.

Dark-banded Owlet Phalaenophana pyramusalis

Saskatchewan to Nova Scotia, south to Florida and Texas. Two generations over much of East with mature caterpillars throughout year; additional broods southward. Common. Dead leaves. Godfrey (1981a) provides additional details in his larval description of P. extremalis.

Litter Moths Subfamily Herminiinae 45

Early Zanclognatha Zanclognatha cruralis ! recognition Brown, spindle shaped with yellow spotting between reddish reticulations. Dark middorsal stripe usually absent. Diameters of D1 pinacula larger than those of D2 pinacula over abdominal segments. Larva to 2.5 cm. There are more than a dozen Zanclognatha in our region. Larvae of Dark Zanclognatha (Z. obscuripennis), Wavy-lined Zanclognatha (Z. jacchusalis) (formerly known as Z. ochreipennis—see Lafontaine and Honey 2009), and others are similar in appearance. Larvae of the Variable Zanclognatha (Z. laevigata) and Grayish Zanclognatha (Z. pedipilalis) bear a smoky middorsal stripe. See Wagner and McCabe (2011) for a preliminary key to larvae of ten eastern Zanclognatha species, and additional images. ! occurrence Woodlands and forests from Wisconsin, east through southern Canada to Florida and Texas. Two generations from southern Quebec to at least North Carolina with mature caterpillars nearly year-round. Common. ! common foodplants Diet incompletely known; caterpillars have been found on beech, hazel, hemlock, maple, nettle, and red spruce. In captivity larvae will mature on dead oak leaves. Wilted leaves of herbs are also readily consumed. ! remarks John Lill (unpubl. data) has found egg clusters of the Early Zanclognatha, containing 2 to 3 dozen eggs, on the underside of living beech leaves (lower inset). Presumably the early instars start feeding on foliage, bark, and detritus. Eventually the caterpillars drop to the ground where they mature. In the fall, late instars of the Early Zanclognatha can be taken in beating samples from an array of forbs and woody plants, including conifers. After Palthis, Zanclognatha larvae are our most frequently encountered herminiines in beating samples—most of the other genera treated here tend to be associated with dead leaves, fungi, and other organic matter of the forest floor.

46 Litter Moths Subfamily Herminiinae

Wavy-lined Zanclognatha Zanclognatha jacchusalis (= Z. ochreipennis of previous authors)

! recognition Ground color yellow or beige to orange-brown with

conspicuous yellow and red mottling; without dark middorsal stripe. Dorsal abdominal pinacula with diameter of D1 pinacula about one-third distance that separates them across midline; diameter of D1 pinacula more than twice that of D2 over abdominal segments. Oblique, yellow subdorsal patches nearly join over dorsum towards rear of A1 to A8. Larva under 2 cm. Closely similar to Early Zanclognatha (Z. cruralis), Dark Zanclognatha (Z. obscuripennis), and perhaps others. ! occurrence Barrens, oak woodlands, and forests from Wisconsin to Maine, south to Georgia and Louisiana. One principal generation with mature caterpillars from early to midsummer; two generations in Missouri. Abundant northward and in mountains southward. ! common foodplants Presumably detritus; figured individual reared on dead oak leaves. ! remarks Zanclognathas are among the most common moths in blacklight traps in the Northeast, and at least in Connecticut, the Wavy-lined Zanclognatha is the most abundant of these in July and August. Dry oak woodlands are especially likely to yield herminiines in number—in some light traps they constitute more than 10% of the trap catch (e.g., Hohn and Wagner 2000). Larvae shun light and seek the shelter of darkness when exposed to bright light. When touched the larva presses the rear of the body down against the substrate. The effect of the flattened caudal end is headlike: the anal prolegs serve as the antennae, and the spiracles on A8 serve as the darkened eyes. At rest the body is somewhat laterally compressed and the larva appears “taller” than other caterpillars of the same length. While we have beat caterpillars of the Early Zanclognatha (Z. cruralis) from vegetation on numerous occasions in the fall, we have never knowingly taken larvae of the Wavy-lined Zanclognatha in our samples. Perhaps the latter species is more closely associated with leaf litter. Like other Herminiinae, the larvae are incredibly slow, more apt to feign death than crawl from danger, and even then their movements are excruciatingly slow. When disturbed, early instars may spin down on a belay line; older larvae simply drop from their perches and curl up. The genus is in great need of taxonomic revision—what we recognize as the Wavy-lined Zanclognatha may well prove to be a complex of species (e.g., see Handfield et al. 1999). The Zanclognatha theralis-minoralis-gypsalis-inconspicualis complex is especially problematic.

Litter Moths Subfamily Herminiinae 47

Conifer Zanclognatha Zanclognatha protumnusalis ! recognition Comparatively smooth, mottled in dull red and yellow with creamy subdorsal stripe, edged below with black. Reddish dorsum with dark middorsal stripe that widens and becomes diffuse over prothorax. Dorsal pinacula small, becoming smaller rearward. D1 pinacula twice diameter of D2 pinacula. Lateral areas more yellow than red. Larva under 2 cm. This is one of three widespread Zanclognatha in the East associated with conifers. Caterpillars of the Pine Barrens Zanclognatha (Z. martha)—a species of pitch pine woodlands—are separable by their dull charcoal ground color, corrugated profile, and the absence of red mottling (page 49). Our captive larvae matured on new stem tissue of pitch pine (some young needles also were consumed). Caterpillars of the recently described Toothed Zanclognatha (Z. dentata) can be recognized by the creamy supraspiracular spot on A7. Also, the dorsal abdominal pinacula are wartlike, and conspicuously wider and higher than those borne over the thoracic segments. Its larvae are found routinely in beating samples from hemlock, fir, as well as a variety of deciduous woody trees and shrubs (page 48). ! occurrence Woodlands and forests from southern Canada to Florida and Texas. One principal generation northward with mature caterpillars in early summer; partial second generation in New Jersey, two or more broods southward. Locally common. ! common foodplants Specimens in Canadian National Collection reared from fir, pine, and spruce (Don Lafontaine, pers. comm.). We have reared ex ova cohorts on dead oak and black cherry leaves. ! remarks While literature indicates that larvae of the Conifer Zanclognatha eat a variety of conifers (Atlantic white cedar, balsam fir, black spruce, jack pine, and white spruce), we suspect that many of these records apply to the Toothed Zanclognatha (Z. dentata). For example, most of the caterpillars beaten from various conifers by Chris Maier and his colleagues over the course of their eightyear conifer survey (Maier et al. 2004) are referable to Z. dentata.

48 Litter Moths Subfamily Herminiinae

Toothed Zanclognatha Zanclognatha dentata

Wisconsin to Nova Scotia, south to South Carolina and northern Georgia. One principal generation over most of range, with mature caterpillars in spring. Locally common. Chris Maier et al. (unpubl. data) have collected or reared larvae from balsam fir, Douglas-fir, hemlock, and pitch pine; ours were beaten from witch hazel and Morrow’s honeysuckle; we reared an ex ova cohort on dead oak leaves (Wagner and McCabe 2011). Pale supraspiracular spot on A7, and dorsal pinacula wartlike; compare with Lettered Zanclognatha below.

Variable Zanclognatha Zanclognatha laevigata

Manitoba to Nova Scotia to northern Florida and Missouri. One generation with mature caterpillars from late spring through midsummer. Common, but absent from much of Atlantic Coastal Plain. Dead leaves and other detritus. Dark middorsal stripe; diameter of dorsal pinacula on abdomen less than height of spiracle on A8.

Lettered Zanclognatha Zanclognatha lituralis

Wisconsin to Nova Scotia, south to Florida and Texas. One generation northward, with at least partial second generation from Ohio and Connecticut southward; mature caterpillars late spring through midsummer. Locally common. Dead leaves and other detritus. Pale supraspiracular spot on A7; dorsal pinacula not wartlike; compare with Toothed Zanclognatha above.

Litter Moths Subfamily Herminiinae 49 Pine Barrens Zanclognatha Zanclognatha martha

Wisconsin to Maine, south to Georgia (one record from Florida Panhandle; Hugo Kons, pers. comm.) and Texas, although absent from many areas within range. One generation with mature caterpillars in early summer. Locally common. Ours from pitch pine. Smoky to charcoal; body corrugated, with red reticulations.

Dark Zanclognatha

Zanclognatha obscuripennis Missouri to Quebec (rare), south to Florida and Texas. Two generations over much of East with mature caterpillars throughout year; continuously brooded in Florida. Local and rare north of New Jersey; common southward along coastal plain. Dead leaves and other detritus; one reared from hazel. D1 and D2 pinacula on abdominal segments nearly equal in diameter.

Grayish Zanclognatha Zanclognatha pedipilalis

Wisconsin to Nova Scotia to southern Florida (rare) and Gulf states. One principal generation with partial second brood in Connecticut, two broods in Missouri, continuously brooded in Florida; mature caterpillars nearly year-round. Common. Dead leaves and other detritus. Dark (not red) reticulations; no spot on A7; dark middorsal stripe.

Morbid Owlet

Chytolita morbidalis Manitoba to Maine, south to North Carolina and Texas. One generation with mature caterpillars in spring. Dead leaves; Forbes (1954) lists grass, dandelion, hazel, and lettuce. Very common. Ours reared on dead oak leaves. Pinacula smaller than those of Zanclognatha, and embedded in yellow spot.

50 Litter Moths Subfamily Herminiinae

Stone-winged Owlet Chytolita petrealis ! recognition Yellowish to reddish brown with dark pinacula and reticulate red patterning. Integument granulated. Reddish-brown to smoky middorsal stripe weak over T1 and rearward of A8. D1 and D2 pinacula small, round, diameter about half height of black spiracles; D1 pinaculum slightly larger than that of D2, and both seated in yellowish spot. Larva less than 2 cm. We are not aware of characters that distinguish its caterpillar from those of the Morbid Owlet (C. morbidalis) (page 49). Also closely resembling many zanclognatha caterpillars, but spinneret more elongate, four to five times width (length only three times width in zanclognathas). ! occurrence Swamps, wetlands, mesic woodlands, and forests from Alberta to Nova Scotia, south to Florida and Texas. One principal generation with mature caterpillars in spring; at least partial second brood from Missouri to New Jersey southward. Common. ! common foodplants Ours reared on dead, dried, white oak leaves that were periodically moistened. See Remarks. ! remarks Chytolita are among the most common herminiines: adults readily come to both lights and baits. In culture the caterpillars were often seen feeding on the upper surface of leaves and did not seem “light shy.” Older instars chewed through entire leaves, consuming more vascular tissues than idias, which fastidiously skeletonize oak leaves (see page 38). Caterpillars of Chytolita and Zanclognatha share the same general form, texture, and coloration, including a characteristic reddish reticulate patterning. Both genera flatten the rear of the body in a manner that suggests a false head. Presumably, the Stone-winged Owlet is a generalist on low-growing vegetation and fallen plant matter. Its congener, the Morbid Owlet (C. morbidalis) (see previous page), has been reared on grass, lettuce, dandelion, and hazel leaves (Forbes 1954). Summer-generation adults of the Stone-winged Owlet are appreciably smaller than those of first brood, a trait shared with Zanclognatha, other herminiines, and double-brooded geometrids. The cocoon is a wispy affair spun in a leaf fold, reminiscent of a spider egg case; filmy silk may be spun as an amorphous sheet distant from the pupa. The red markings are accentuated in the prepupa (right). As with other herminiines, the larva overwinters.

Litter Moths Subfamily Herminiinae 51

Streaked Fan-foot Macrochilo orciferalis [Bronzy Macrochilo] ! recognition Somewhat trapezoidal in cross section, reddish tan-brown with subdorsal stripe that extends from T2 to A10. Abdominal segments with two to three shallow creases rearward on each of first eight segments. Dorsum with numerous fine, vague, wavy reddish lines. Setae minute; pinacula small, black or brown. Legs short, held under body. Prothoracic plate pale brown with black pinacula. Head pale brown with faint reticulations. Sides of A9 with slight bulge. Larva less than 2 cm. ! occurrence Wetlands, mesic grasslands, heathlands, dunes and other coastal strand communities, and woodlands from Wisconsin to Nova Scotia, south to Florida and Texas. At least two generations in Northeast with mature caterpillars in late spring, then again in midsummer. Common. ! common foodplants Figured individual was collected by beating, and reared to maturity on mixture of blueberry and dead grass. Donald Lafontaine (pers. comm.) swept a larva from a meadow dominated by timothy (Phleum pratense), and reared the individual to maturity on grass. ! remarks When at rest, the caterpillar’s short prolegs are scarcely visible beneath the body. Its coloration suggests that it rests along leaves and stems of dead grasses and sedges by day. We have reared other Macrochilo species solely on diets of dead sedge leaves, and suspect that graminoid leaves constitute a dietary staple for the genus. A revision of the genus was published by Ferguson (1982).

52 Litter Moths Subfamily Herminiinae

Slant-lined Fan-foot

Macrochilo absorptalis [Slant-lined Owlet] Manitoba to Nova Scotia, south to Missouri, New Jersey, and the Smokies. (A related and as yet undescribed species occurs along Atlantic Coastal Plain from New Jersey to at least North Carolina.) One generation over most of range with mature caterpillars mostly in early summer; at least a partial second brood in Virginia and Missouri. Locally common in wetlands, e.g., those with abundance of upright sedge (Carex stricta). Ours reared on yellowed and long-withered sedge leaves.

Shadowy Phalaenostola Phalaenostola eumelusalis [Dark Phalaenostola]

Dakotas to New Brunswick, south to Georgia and Iowa. Evidently two generations over most of East with mature caterpillars in spring, then again in midsummer. Generally infrequent. Natural hosts unknown; ex ova individual figured here reared on dead grass. Dark middorsal stripe that narrows anteriorly distinguishes members of genus.

Black-banded Phalaenostola Phalaenostola larentioides

Wisconsin to New Brunswick, south to Florida and Texas. Evidently two or more generations with mature caterpillars nearly year-round. Very common. Dead and living grasses; also leaf litter and (living) clover leaves. Some individuals bear more reddish markings (Crumb 1956).

Florida Tetanolita Tetanolita floridana

Wisconsin to Long Island, south to Florida and Texas. Two or more generations with mature caterpillars nearly year-round. Common. Presumably on fallen organic matter; ours reared on moistened oak leaves from previous season. Dorsal warts increasing in size rearward.

Litter Moths Subfamily Herminiinae 53

Bent-winged Owlet Bleptina caradrinalis [Variable Snout] ! recognition Stout, foreshortened, brown to brown-black larva with short

mushroomlike setae (appearing glandular); coarsely granulose integument. Segments A9 and A10 reduced. Tannish bars, forming broken stripe, just above spiracles, on A1 to A7 (A8). Often with pale to orange-red addorsal spot at rear of each abdominal segment. Anal plate dark brown, contrasting with paler ground color; prothoracic shield with pale middorsal and lateral spots. Anterior abdominal spiracles enlarged, twice as high as broad (Crumb 1956). Larva to 1.5 cm. Bleptina sangamonia flies with the Bent-winged Owlet from Ohio southward (but absent from Pennsylvania, New Jersey, and Delaware). In the coastal plain of North Carolina three Bleptina species occur: B. caradrinalis, B. near sangamonia, and B. inferior (with the last, shown in the lower inset, ranging up the Atlantic coast to New Jersey). In the Carolinas, both B. inferior and B. near sangamonia fly mostly in Carolina bays, xeric sand hills, and seaside communities, whereas B. caradrinalis occurs statewide in many community types. Crumb’s (1956) key to larvae treats three eastern Bleptina. ! occurrence Woodlands and forests from Canada, south to Florida and Texas. One principal generation with mature caterpillars in late spring northward, with partial second brood most years from at least Ohio and Connecticut southward. More generations in Deep South. Abundant. ! common foodplants Covell (2005) lists barberry, clover, hickory, as well as dead leaves; we collected one caterpillar on orache (Atriplex), and reared an ex ova cohort on dead oak and cherry leaves. ! remarks Across much of its eastern range, the Bent-winged Owlet is our most abundant and ecologically generalized herminiine, but we have not encountered its larva in the wild. This suggests that the caterpillars live on or near the ground and feed on dead leaves and other detritus. Adults can be abundant in light trap samples from grasslands, power lines, and open, grassy woodlands. The caterpillar’s height relative to its short body length and peglike setae suggest an evolutionary relationship with Renia.

54 Litter Moths Subfamily Herminiinae

Discolored Renia Renia discoloralis ! recognition Reddish brown, mottled, with coarsely roughened integument. Extensive red network dominates coloration. Smoky to reddish, diffuse middorsal stripe; vague, pale, mostly broken subdorsal stripe (most evident over thoracic segments). Dorsal setae very short; that on D1 shorter than diameter of blackened pinacula from which it arises. D1 pinaculum distinctly darker and larger than D2 pinaculum on abdominal segments. Most setae pointed at apex (many Renia have blunt setae). Setae below spiracles and those on A10 long, whitish, bladelike, directed downward. Spiracles tan with dark brown-black outer rim. Head, prothoracic shield, and anal plate coarsely roughened. Often with smoky coronal bar to either side of triangle. Larva less than 2 cm. Caterpillars superficially resemble many zanclognathas, but renias can usually be recognized by their modified (blunt-tipped or, in the case of R. discoloralis, bladelike) setae; those from D2 pinacula are usually directed backward (as on some idias). The head and body of renias tend to be coarsely roughened. ! occurrence Mesic to dry forests from Missouri to extreme southern Canada to Florida and Texas (but see below). One generation with mature caterpillars in late spring and early summer, although see below. Very common. ! common foodplants Dead leaves and other detritus. ! remarks The creamy eggs, with sparse, reddish pigment spots, are laid in rafts of 20–50 or more. A crude cocoon, sparse in silk, is fashioned in a leaf fold or other secluded niche. Renias are diverse and abundant in dry oak woodlands. Adults come to light, although there are nights when searching vegetation by flashlight can be much more productive, especially for females. The most efficient way to encounter the genus is to bait in July and August. Forbes (1954) correctly suspected that the double-brooded southern populations of the Discolored Renia, which are on average smaller in both broods, represent a distinct species. The moths are sympatric from eastern North Carolina to northern Delaware, often visiting the same baited trees, but there is no overlap of flight seasons or adult size in New Jersey.

Litter Moths Subfamily Herminiinae 55

Undulate Renia Renia factiosalis [Dark-banded Renia] ! recognition Smooth, straw colored to brown, with reticulate network of reddish markings; red or brown middorsal stripe running length of body sometimes absent (left inset). Dorsal pinacula brown, not raised; D1 twice diameter of D2, bearing minute seta. Remaining pinacula small and little differentiated. Setae very short except over head and anal shield; ventral and subventral setae in contact with substrate also longer. Head with dark pigmentation about setal insertions. Larva approximately 1.5 cm. Early instars differ in appearance with rugose integument and longer, thick setae that are flared at their apices (right inset). Dorsal setae from raised, black pinacula; D1 directed forward and longer D2 seta directed backward. Lateral setae borne from raised fleshly protuberances. Our description, based on a series of ex ova larvae, differs markedly and inexplicably from that of Crumb (1956). ! occurrence Woodlands from southern Canada to Florida and Texas. One principal brood in Northeast with mature caterpillars from late May into July; two broods in Missouri, southern Ohio, and North Carolina. Second generation evidently partial over much of the South. Common. ! common foodplants Dead leaves and other organic matter. ! remarks The lethargic larva perches on the underside of dead leaves. At rest the body of the last instar is somewhat flattened with the sides of abdomen forming a shelf that wholly obscures the midabdominal prolegs. From above, the rear of the body resembles a head of sorts (with spiracles on A8 serving as the eyes and the anal prolegs as the antennae). The differences in body form between middle and late instars borders on hypermetamorphosis—at first we thought our lots had contaminants and contained caterpillars of two separate species. We wonder if the two larval forms have different microecologies, with each morph adapted to a different lifestyle or niche. In the Northeast, the Undulate Renia is among the most common Renia in both dry and mesic woodlands.

56 Litter Moths Subfamily Herminiinae

Dotted Renia Renia salusalis ! recognition Smoky brown, roughened, nondescript with pale peglike dorsal setae arising from black pinacula; body broader rearward. Pattern subdued with vague reddish reticulations (most evident along venter). Dusky middorsal stripe poorly defined. Prothoracic shield often with pale middorsal line and/or pale patch along anterior margin, fused with prespiracular pinaculum (that bearing L setae). Subdorsum of A9 with two black pinacula (D1 and SV1) fused. Head small, dark, roughened or tuberculate. Larva under 2 cm. There are eight widespread Renia in the East as well as additional undescribed ones; e.g., there are two species going under this name in the Southeast. Crumb (1956) provides a larval key to six common species. ! occurrence Mesic to dry forests from Colorado, Ohio, and Massachusetts, south to Florida and Texas. Single-brooded in southeastern Massachusetts; two broods in New Jersey with mature caterpillars from spring into early summer, then again in midsummer. Probably three generations in southern Missouri. Common. ! common foodplants Dead leaves and other detritus. Feeding preferences of renias in nature essentially unknown. ! remarks The caterpillars shun light and crawl deeper into litter when exposed. Like other herminiines, Renia caterpillars feign death when handled. The individual figured here was reared from a cohort fed on dead oak leaves. The caterpillars generally ate through the entirety of fallen leaves, i.e., they did not carefully eat about vascular tissues as do many leaf-feeding Idia (see page 38). Eggs obtained in mid-June from a female collected at bait yielded three-quarter-grown larvae by mid-August, which then more or less stalled until the following spring. Our images, taken in November, are thought to represent a penultimate instar. The lethargic larvae remain active though the winter months, feeding sporadically during warm spells.

Litter Moths Subfamily Herminiinae 57 Speckled Renia Renia adspergillus

Wisconsin and Southern Canada to Florida and Louisiana; multiple broods in Southeast. Very common in New Jersey; absent or rare in Connecticut. Larvae feed beneath wispy web of silk on dead leaves and other organic matter.

Even-lined Renia

Renia flavipunctalis [Yellow-spotted Renia] Southern Canada to Florida and Texas. One brood in Northeast with mature caterpillars from late May into July. Common northward; uncommon on coastal plain of New Jersey and North Carolina. Dead leaves and other organic matter. (Image of middle instar taken in October.)

Chocolate Renia Renia nemoralis

Illinois to southeastern Massachusetts, south to Florida and Texas. One brood with mature caterpillars in July and August in New Jersey, later southward. Common southward. Dead leaves and other organic matter.

White-spotted Redectis Redectis vitrea

Illinois to southeastern Massachusetts, south to Florida and Texas. Evidently two generations in Virginia and New Jersey with mature caterpillars in spring, then again in midsummer. Moderately common southward. Autumn olive, New Jersey tea, crabgrass, and presumably many other plants. Larva constructs and feeds within a crude silken web-shelter.

58 Litter Moths Subfamily Herminiinae

Ambiguous Litter Moth Lascoria ambigualis ! recognition Elongate, dark brown herminiine with subtle deep red reticulations. Body tapered and elongated rearward; A8 humped in profile. A6–A9 with white middorsal stripe. White subdorsal spots form broken line over thoracic segments and A1; vague, shallowly oblique, subdorsal white lines above spiracles on A2–A6 (or A7). A7 with protruding bulge above spiracle. Inconspicuous setae borne from blackened pinacula. Spiracles narrow. Thoracic legs comparatively short. Larva to 2 cm. ! occurrence Shrubby fields, woodlands, and forests from Wisconsin and southern Canada to Florida and Texas. Two generations in Connecticut with mature caterpillars from October through May, and then again in midsummer; multiple broods southward. Common. ! common foodplants Polyphagous on live and dead plant tissues. We have found caterpillars on aster, blackberry, dead grasses, bindweed (Ipomaea), crownbeard or wingstem (Verbesina), winterberry (Ilex verticillata); we reared ex ova caterpillars on dead oak leaves. Crumb (1956) also lists chrysanthemum. ! remarks The figured individual was beaten from blackberry in early September. It fed on living leaves for several weeks before it overwintered. Lascoria larvae share their climbing habit with members of the genera Palthis and Redectis. The lethargic caterpillar rests fully extended: the thoracic segments are somewhat flattened and the anal prolegs trail behind the body. Adults appear in early spring, leading us to suspect that at least some individuals overwinter as late instars or pupae.

Litter Moths Subfamily Herminiinae 59

Dark-spotted Palthis Palthis angulalis ! recognition Small, mottled, brown caterpillar with low hump on A1 and large angulate hump on A8. A7 with oblique white line that changes to black on A8 and extends up side of hump. Additional faint, oblique, subdorsal white lines on anterior abdominal segments; oblique stripe on A1 usually best developed. Integument with abundant minute spinules. Pinacula black. Larva under 2 cm. Larva of Faint-spotted Palthis (P. asopialis) with larger, more toothlike integumental spinules—these especially evident over white areas on A8. ! occurrence Woodlands and forests from Canada to Florida and Texas. Two generations with mature caterpillars from June until late fall over much of East; three or more generations from Missouri southward. Very common. ! common foodplants Extremely diverse in diet. Forbs, woody plants, including conifers: e.g., alder, aster, basswood, birch, chestnut, fir, gale, goldenrod, ninebark, rhododendron, scrub oak, and spruce. ! remarks The caterpillar eats both live and dead organic matter, be it leaves, flowers, or fruits. It has even been reared from an old larval nest of the pyralid moth, Pococera robustella, where it was feeding presumably on accumulated feculae. The caterpillars are commonly taken in beating samples. They can also be found by searching low vegetation at night, especially during the fall. While Forbes (1954) reports that the larva overwinters, and this may be the most frequent overwintering stage in nature, we have had captive individuals overwinter as pupae in cocoons fashioned in leaf litter.

60 Litter Moths Subfamily Herminiinae

Faint-spotted Palthis Palthis asopialis ! recognition Small, coarse, brown caterpillar with vague white and black markings—exhibits main features noted above for Dark-spotted Palthis (P. angulalis). Eighth abdominal segment strongly humped; white oblique line from hump to anterior edge of A7, touching upper end of spiracle on A7. A1 slightly humped and swollen. Dorsum of A1–A7 with dark rearward pointing triangles edged with pale, oblique subdorsal lines; pale edging more prominent anteriorly. Integument roughened, covered with mixture of short, coarse, toothlike spines and hairlike spinules. Larva to 1.5 cm. ! occurrence Swamps, barrens, and woodlands from Wisconsin to southern Quebec, to Florida and Texas. At least two generations in Northeast with mature caterpillars nearly year-round; more generations southward. Common, but less so than Dark-spotted Palthis. ! common foodplants Live, senescing, and dead tissues of many plants; recorded hosts include aster, bean, beggarticks, corn, and wild indigo (Baptisia). ! remarks We find the caterpillars on low-growing plants, especially during the fall. Caterpillars that were offered aster flowers cut away the ray flowers and fed on the central disk flowers (and seeds therein). The caterpillars are slow moving. When disturbed they drop from their perch and feign death. Adult herminiines are among the most mundane owlets and often dismissed by collectors: too small, too easily abraded, and too difficult to identify—not one of the more than 50 eastern species is brightly colored. While superficially undistinguished, and always some shade of brown, their scent-releasing structures (used in close range courtship behaviors after nightfall) are marvelously diverse. Specialized male courtship scent scales are the rule among herminiines and there seems to be few limits to where these are found or the extent of their development; the antennae, legs, and mouthparts may bear scent scales (androconia). Palthis males possess an enormous androconial brush on their labial palpus (right), and a second brush along the foretibia.

Pangraptines Subfamily Pangraptinae 61

Pangraptines

Family Erebidae: Subfamily Pangraptinae Recent molecular studies by Zahiri et al. (2011) indicate that the Pangraptinae fall outside of any known erebid subfamily, clustering with Herminiinae, Arctiinae, and Aganainae (an Old World subfamily). The prolegs are reduced on A3 and A4.

Decorated Owlet Pangrapta decoralis complex ! recognition Small, green, smooth, spindle shaped. Body essentially unmarked except for dark green (middorsal) heart stripe. Head pale green, shiny, often with thin black line over each lobe. Prolegs on A3 and A4 reduced, about half size of those on A5 and A6. Body setae fine and inconspicuous, only a quarter to a third length of segment bearing them. Anal prolegs long, extended out from end of body; sometimes with pinkish cast. Spiracles small, pale orange. Larva to 2 cm. ! occurrence Barrens, woodlands, bogs, and heathlands from Manitoba to Nova Scotia, south to Florida and Texas. Two to three generations over much of East with mature caterpillars from June onward; flying throughout growing season in Florida. Very common. ! common foodplants Blueberry, huckleberry, sourwood, staggerbush, and likely other heaths. ! remarks Early instars graze patches of tissue from lower leaf surfaces, leaving the upper epidermis intact. This widespread moth appears to be a complex of at least three sibling species—the degree to which the members sort themselves out by foodplant, habitat, and range is in much need of study. Our main image is of a caterpillar beaten from huckleberry in Connecticut; the prepupal caterpillar (inset) came from a sourwood in eastern North Carolina. Decorated Owlet moths, easily flushed during the day, can be common in heathlands and woodlands with an ericaceous understory. Adults come to both light and bait. The pupa overwinters.

62 Pangraptines Subfamily Pangraptinae

Buttonbush Owlet Ledaea perditalis [Lost Owlet] ! recognition Bright green, somewhat spindle shaped with reduced prolegs on A3 and A4. Usually with white to yellow subdorsal stripe running length of body. Middorsal stripe either pale and narrow, or broad and reddish (inset). Supraspiracular stripe represented by mostly broken line of small yellowish spots; an even more rudimentary line of spots above prolegs. Head with maroon coronal line that extends forward and drops down side of triangle. Dorsal setae very short, only one-tenth length of segment that bears them. Spiracles small, pale orange. Larva to 2 cm. ! occurrence Wetlands from southern Canada to Florida and Texas. Two principal generations in Connecticut with mature caterpillars in early summer, then again in August and September; at least three generations in coastal Carolinas with caterpillars from May onward. Common in wetlands. ! common foodplants Buttonbush. ! remarks The broad maroon stripe occurs only in the last instar. We are uncertain if the form with a narrow middorsal stripe (main) represents a penultimate instar or simply an alternative form of the mature caterpillar. Larvae are often common in beating samples taken from buttonbush in August. Prepupae turn red. Overwinters as a pupa.

Snouts Subfamily Hypeninae 63

Snouts

Family Erebidae: Subfamily Hypeninae This small subfamily is represented by about 16 species in our area, all but two of which are placed in the nominate Hypena. Classifications before Poole (1989) had eastern Hypena species assigned to four genera: Bomolocha,1 Lomanaltes, Ophiuche, and Plathypena. Hypenine caterpillars are bright green, elongate, with a tapering abdomen, and anal prolegs that splay outward; setae are long especially rearward; the head is as wide or wider than the prothorax, and often bears a few large and numerous small black spots. The proleg on A3 is reduced in size or absent; that on A4 also may be reduced. Setae are sometimes borne from raised bases (pinacula) that may be reddish or black (with the degree of melanization or colorization varying among individuals). The SV group is trisetose on A1 and A2 but appears bisetose because the SV3 seta is shifted

upward, proximate to L3 (Beck 1999, 2000). In Hypena, the tacky fecal pellets are discharged with a flick of the caudal body segments. Hypenine caterpillars snap and contort violently when handled. Diets are specialized and varied such that knowledge of the foodplant will often lead to a correct identification. No less than five Eastern species feed on the nettle family (Urticaceae) (this total includes Hypena atomaria, which was recently reared from stinging nettle by Jim Sogaard (pers. comm.). The life history of H. appalachiensis is unknown, but it too may be an Urticaceae feeder. Caterpillars are readily sampled with a beating sheet—August and September are especially good months over much of the East to search for Hypena larvae. Most members of the genus (traditionally treated as Bomolocha) overwinter as pupae in slight cocoons spun among leaf litter; a few pass the winter as an adult (where the winters are mild), and possibly one as a middle instar.

Mallow Snout

Hypena minualis (= Ophiuche minualis) Gulf states but migrating north to Vermont in fall. Multiple generations with mature caterpillars throughout the growing season. Common. Globemallow (Sphaeralcea), false mallow (Malvastrum), and sida.

Variegated Snout

Hypena palparia (= Bomolocha palparia) Minnesota to Nova Scotia, south to northern Florida and eastern Texas. At least two generations over much of East with mature caterpillars from June onward. Locally common. American hornbeam, hophornbeam, and hazel.

Species traditionally treated in Bomolocha are not migratory, commonly feed on shrubs or trees, overwinter as pupae, and almost never come to bait. By contrast, many Hypena overwinter as adults or larvae, at least three migrate, and adults feed at bait. To our minds, former classifications that recognized multiple genera, within what is now treated as a single genus (Hypena), had much biological utility. 1

64 Snouts Subfamily Hypeninae

Baltimore Snout Hypena baltimoralis (= Bomolocha baltimoralis)

! recognition Elongate, emerald green with somewhat yellow intersegments;

proleg on A3 about half size of that on A4–A6. Translucent, tracheal trunk visible within body. Setae long, dark, arising from somewhat orange-red or blackened setal bases. In some forms, anterior dorsal seta (D1) and that above spiracle (SD1) borne from blackened spots (lower right); SD1 pinaculum on A8 largest. Head green and unmarked except for black lateral eyes or with black around setal bases. Larva to 3 cm. Several other snouts have similar caterpillars: foodplant associations will help establish identities for all. ! occurrence Edges of watercourses, swamps, woodlands, and forests from Wisconsin to Nova Scotia, south to Florida and Texas. At least two generations in New England with mature caterpillars from June to October; additional generations southward. Very common. ! common foodplants Red and silver maple. Prentice (1962) also reports sugar maple; we have not yet encountered it on this host in our surveys. ! remarks After the Black Snout (H. scabra), this is the most widespread and abundant Hypena over much of the East, especially in swamps, floodplains, and mesic woodlands. By day caterpillars rest on the undersides of leaves. DLW once collected the Baltimore Snout (while bass fishing) by rapping low red maple branches with an oar—larva floated on the water surface and were easily recovered. When disturbed, Hypena either launch themselves from their purchase or drop on a belay line. Godfrey (1987) reared adults from trap nests deployed to survey for cavity-nesting bees; evidently prepupal caterpillars sometime enter holes in trunks or logs, before spinning their cocoons.

Snouts Subfamily Hypeninae 65 White-lined Snout

Hypena abalienalis (= Bomolocha abalienalis) Southern Canada to northern Florida and Texas. At least two generations with mature caterpillars from June to October over much of East. Common. Elm, especially American, slippery, and rock.

Dimorphic Snout

Hypena bijugalis (= Bomolocha bijugalis) Southern Canada to Florida and Texas. At least two generations with mature caterpillars from June to October over much of East; additional generations southward. Locally common. Dogwood, especially shrubby species such as red osier.

Deceptive Snout

Hypena deceptalis (= Bomolocha deceptalis) Manitoba to Quebec, south to Florida and Texas, but absent from much of coastal plain. Two generations with mature caterpillars from June to October over much of East. Locally common. Basswood. Larvae commonly with black spotting (about pinacula) and head spots.

Flowing-line Snout

Hypena manalis (= Bomolocha manalis) Minnesota to Nova Scotia, south to Florida and Texas. Two to three generations with mature caterpillars from June to October over much of East. Common in wetlands and mesic environs. False nettle.

66 Snouts Subfamily Hypeninae

Large Snout Hypena edictalis (= Bomolocha edictalis) ! recognition Bright lime green with faint, smoky middorsal heart stripe and broken white spiracular stripe. Spiracular stripe best developed from A8 rearward; occasionally reappearing as broken line on anterior abdominal segments. Prolegs absent on A3. Segment A8 longer than A7 and twice length of A9. Head mostly unmarked with peppering of small reddish-black spots over each lobe. Setae rusty. Larva to 3 cm. Middle instar shown to lower right. The Sordid Snout (H. sordidula) and Hop Vine Snout (H. humuli) also occur on woodnettle with the Large Snout. The former is distinguished by the presence of a rudimentary proleg on A3 and shiny black spots on the head. The Hop Vine Snout bears thin, white subdorsal and spiracular stripes and is appreciably smaller. ! occurrence Mesic woodlands and floodplain forests from Manitoba to Maine south to North Carolina (mountains), although absent from significant portions of this range (and some literature records may be based on misidentified individuals). One principal generation with mature caterpillars in late spring, but perhaps a partial second brood from Ohio south through Appalachians (Rings et al. 1992). Local but sometimes common where found. ! common foodplants Woodnettle (Laportea) and perhaps some Urtica. ! remarks Caterpillars can be common on woodnettle in middle to high elevation forests of the Appalachians. Look for the larvae on upper leaf surface and leaf petioles of newly produced foliage. Most throw a loop into the body, holding A1–A3 above the others. Caterpillars are capable crawlers, sometimes moving to different plants or resting on nearby vegetation, especially in the last instar. Only search plants with damaged leaves: the larvae are relatively neat feeders, removing leaf sections, often back to the midrib. If pinched or otherwise molested, the caterpillar wriggles violently, launching itself from its perch—some individuals continue to wriggle spasmodically for seconds. We have found larvae in the early spring at middle elevations in the Smokies (May and early June), weeks before adults appear. We suspect that middle instars overwinter, as in The Snout (H. proboscidalis), an Old World member of the genus.

Snouts Subfamily Hypeninae 67

Alder Snout Hypena eductalis (= Lomanaltes eductalis) ! recognition Elongate, bright yellow to lime green with dark red to black spots about setal bases: pinacula above spiracle often blackened and those below more apt to be reddened. Dorsum sometimes vaguely marked with subtle, mostly broken middorsal, addorsal, and supraspiracular reddish to smoky-green stripes; creamy subdorsal stripe sometimes present. Pinacula above spiracle (SD1) on A1–A8 may be enlarged and more conspicuous than others. Thoracic legs and prolegs frequently reddened. Setae dark, generally less than length of segment that bears them except over A8–A10. Proleg on A3 a third size of that on A4, crochets withdrawn. Anal proleg often with black or red line on outer face. Head usually with dark spots about setal insertions and sometimes with additional black spots forming vague arc over each lobe. Larva to 2.5 cm. ! occurrence Swamps and edges of watercourses, but more generalized northward. Saskatchewan to Nova Scotia, south to Florida and Texas. Two generations over much of East with mature caterpillars from June onward. Widespread and common in Canada, becoming increasingly localized southward. ! common foodplants Alder and river birch. ! remarks The caterpillar feeds and rests on underside of leaves, feeding either from a leaf edge or chewing holes out of the middle of the lamina. The alarm response is greatly reduced in duration and intensity relative to that of other snouts. The caterpillar weaves leaf fragments and feculae into the walls of its cocoon. The pupa overwinters.

68 Snouts Subfamily Hypeninae

Hop Vine Snout Hypena humuli ! recognition Many individuals more pale green than most Hypena; thin,

white subdorsal and spiracular stripes usually present. Obscure addorsal stripes to either side of dark heart stripe. Proleg on A3 completely absent; that on A4 about one-quarter smaller than those on A5 and A6. Setae fine, pale golden orange, nearly same length as segment that bears them. A7 and A8 about same length, each about twice length of A9. Spiracles pale, small. Integument bearing minute, upright, spinelike granules (Crumb 1956). Head shiny green and unmarked or with scattered, minute black spots—no large, shiny black patches. Larva to 3 cm. Nettlefeeding snouts tend to be darker green and lack the subdorsal and spiracular stripes of the Hop Vine Snout: see accounts for Large Snout (H. edictalis) and Sordid Snout (H. sordidula). Larva also similar to that of Black Snout (H. scabra) (page 70)—which we have not seen on nettles. ! occurrence Wet meadows, mesic woodlands, and riparian areas. Transcontinental in Canada, south in East to Florida and Arkansas. Two generations over most of East with mature caterpillars in late spring and early summer and then again in late summer; possibly additional generations southward. Common (although not easily sampled as an adult). ! common foodplants Hops, nettle, and woodnettle (Laportea). ! remarks During the day look for caterpillars extended along the midrib on the underside of leaves. The feeding damage is conspicuous with large sections of a blade removed. If pinched, the larva snaps about wildly; others draw themselves into a tight ball. In the Smokies we have found the caterpillar feeding alongside those of the Large Snout (H. edictalis). While generally regarded as uncommon (e.g., Rings et al. 1992), our multiple encounters with the larva would suggest that the moth is common and widespread, but that the adults are only weakly attracted to light. Baiting is often a better option for securing adults. The winter is passed as an adult. While nettles are scarce in Port Norris, New Jersey, especially in drought years, this moth is sometimes common in DFS’s yard through the winter months, and can be seen flying on warm spring evenings (but it is virtually absent from April to September), which suggests that the species is migratory.

Snouts Subfamily Hypeninae 69

Gray-edged Snout Hypena madefactalis (= Bomolocha madefactalis) ! recognition Pale green elongate body with well-developed, cream to yellow subdorsal stripe; blackened setae borne from minute, reddened or blackened pinacula; occasionally with pinacula embedded in larger black and/or red spot (inset). Body with relatively long, darkened dorsal setae, longer than segment that bears them. Head pale green and unmarked, or with two black spots on each lobe. Larva to 3 cm. ! occurrence Orchards, floodplains, woodlands, and forests from extreme southern Canada to Georgia and Texas. At least two generations with mature caterpillars from June to October over much of range. Common. ! common foodplants Mostly walnut; one reared from shagbark hickory (DFS). ! remarks Hypena typically rest fully extended with the legs splayed outward and backward on the underside of leaves. When moving forward they loop along, but not to the extent as would inchworms or true loopers; hence they and other owlets are often referred to as semiloopers. The alarm response is impressive: snouts contract violently for a period of two to four seconds, spasmodically hurling their bodies to and fro. In the wild, this response would drive the caterpillar deep into leaf litter or other protected places. Larvae toss their feculae by snapping the rear of the body to one side at the same time the pellets are released from the body, presumably to distance themselves from the volatiles in their droppings that are used by parasitic and predatory wasps to locate prey (see Weiss 2003 and 2006 for interesting discussions on “feculae flicking” by caterpillars).

70 Snouts Subfamily Hypeninae

Black Snout Hypena scabra (= Plathypena scabra) ! recognition Lime green with faint subdorsal and strong spiracular stripes. Weak, frosty addorsal stripe sometimes evident. Proleg on A3 absent. Setae often rusty to black, long, exceeding length of segments that bear them rearward. Dorsal setae on both T2 and T3 from common, raised wart. Prolegs on A4–A6 frequently with pinkish cast. Head shiny green, unmarked, wider than prothorax (i.e., larva with “neck”). Penultimate instar more strongly striped. Larva to 3 cm. The caterpillars are easily confused with those of Ommatochila mundula, a species that we have reared from blackbrush acacia (Acacia rigidula) in Texas. ! occurrence Waste areas, fields, croplands, and woodlands from Canada, south to Florida and Texas. Multiple generations with mature caterpillars throughout growing season. Abundant. ! common foodplants Partial to low-growing legumes (alfalfa, bean, beggarticks, bush clover, clover, pea, soybean, vetch, etc.), but also woody legumes such as false indigo (Amorpha) and locust. Other hosts include birch, cherry, corn, elm, hackberry, New Jersey tea, poplar, ragweed, sida, strawberry, and willow. ! remarks The Black Snout, more widely known as the Green Cloverworm, is an occasional pest of legumes. The young caterpillars “window” the leaves of sida, removing patches of tissue from the leaf underside while leaving the upper leaf surface intact. Like other hypenines, the caterpillars hurl themselves from their perch when disturbed by rapidly contracting and twisting their bodies in a fashion reminiscent of a tightly wound rubber band, and stay balled up until danger is perceived to have passed. The Black Snout is a migrant that recolonizes the northern portion of its range each summer. Adults arrive in Ohio and New England in early spring, sometimes as early as February and March. Over much of the East, the moth becomes increasingly common through summer and autumn. In southern New Jersey, adults can be abundant at light and bait by November, and often remain among the most common moths at bait through the winter. Adults overwinter where the January mean temperature is above freezing, roughly from southern Missouri eastward. As in most overwintering moths, adults are in a state of reproductive diapause through the winter months (see Johnson 1995). In frostfree areas larvae occur year-round.

Snouts Subfamily Hypeninae 71

Sordid Snout Hypena sordidula (= Bomolocha sordidula) ! recognition Bright lime green with faint middorsal, subdorsal, and supraspiracular smoky green stripes; these most apparent over thorax. A3 proleg small, a quarter size of others. Setae rusty to smoky depending on viewing angle. Tracheal trunk, running between spiracles, visible through body wall. Head with comma- or C-shaped shiny black patch over each lobe, with additional black spots at base of other setae and/or about lateral eyes. Larva to 3 cm. Readily distinguished from Large Snout (H. edictalis) and Hop Vine Snout (H. humuli), which occur with it on woodnettle, by its diminutive prolegs on A3 and the prominent, black spots on the head. ! occurrence Mesic woodlands and floodplain forests from Manitoba to Maine, south to northern Florida (Kimball 1965) and Texas, although absent over much of this range. One principal generation with mature caterpillars in midsummer (see below). Uncommon and local. ! common foodplants Woodnettle (Laportea). ! remarks Two reported hosts, alder and butternut, are in error. In the Smokies, where both the Sordid Snout and Large Snout (H. edictalis) feed on woodnettle, the activity of the two is offset: larvae collected during July will produce adults of H. sordidula, whereas those secured in May and the first half of June will yield H. edictalis. The phenology charts in Rings et al. (1992) are suggestive of two generations for both insects, but we have yet to encounter caterpillars of the second (possibly partial) generation of either. The fecal pellets of the Sordid and Large Snouts, both woodnettle feeders, often adhere to various surfaces about the larval perching sites (and to other caterpillars when reared together). Like the Large Snout, the caterpillars are capable crawlers and may wander off the host between feeding bouts. The Sordid Snout, too, has a highly animated and prolonged wriggling response (see Remarks for Large Snout).

72 Straws Subfamily Rivulinae

Straws

Family Erebidae: Subfamily Rivulinae This subfamily is represented in North America by nine species in three genera: Rivula, Oxycilla, and Zelicodes. At least in the nominate genus, L3 and SV3 share a common pinaculum on the first two abdominal segments, and the long primary setae are minutely barbed (Beck 1999, 2000). (We have not examined Oxycilla for this feature.) All four sets of midabdominal prolegs are well developed. Larvae feed on low plants.

Spotted Grass Moth Rivula propinqualis ! recognition Small, green caterpillar with long, minutely barbed setae. Setae

increasing in length rearward, where their length may exceed that of two segments. Prominent white subdorsal stripe, thickened over A3–A7 and narrowed over T1. Head somewhat yellowish, rounded, at least as wide as prothorax. Body wall translucent: tracheal trunk visible as thin spiracular stripe and yellowish testes evident in abdominal segment five of males. Larva to about 1.5 cm. The largely Caribbean and Central American R. pusilla occurs in southern Florida northward at least to Gainesville and in south Texas. R. stepheni occurs on the coastal plain in North Carolina and Louisiana; it is associated with small streams in mesic woodlands (Sullivan 2009). We have not seen the larva of either of these southern taxa. ! occurrence Meadows and fields, woodlands, and other grassy habitats from southern Canada to Florida and Texas. Two generations in Northeast with mature caterpillars in spring, then again from late July into September; multiple generations in Deep South. Common. ! common foodplants Grasses. ! remarks Adults are easily flushed during the day. Porter (1997) provides a brief account for the Straw Dot (R. sericealis), a British congener, whose larva can be obtained by sweeping at night, especially during the spring. Caterpillars of the Straw Dot fashion a cocoon between two blades of grass. The larvae share two characters with the tiger moths (Arctiinae): they lack a midventral prothoracic gland (present in most owlet larvae) and the setae are minutely barbed. Old World species overwinter as partially grown larvae (Porter 1997, Ahola and Silvonen 2005).

Scalloped Moths Subfamily Scoliopteryginae 73 Arizona Straw

Oxycilla tripla (Subfamily Rivulinae) At least southern Arizona. Mature caterpillars mostly during the summer rainy season in August and September. Very common. Ticktrefoil (Desmodium). While O. tripla is extralimital, two members of the genus occur in our area—we are not aware that either has been reared. Presumably our eastern species are at least roughly similar to the Arizona Straw in appearance.

Scalloped Moths

Family Erebidae: Subfamily Scoliopteryginae Members of this subfamily were previously classified in Catocalinae (now Catocalini), or most recently, in Calpinae. The 11 North American species fall into three genera: Scoliopteryx (Tribe Scoliopterygini) and Anomis and Alabama (Tribe Anomini). The SV group includes three setae on both A1 and A2. Caterpillars “possess an extra setigerous tubercle” below the second dorsal seta (D2) on A1–A6 (or A7) (Crumb 1956). The first, and sometimes the first two, pairs of prolegs are frequently reduced or absent. Several members of the Anomini have the dorsal pinacula surrounded by a faint halo or ring. The crochets at

Gulf Scalloped Moth Anomis editrix

Mostly southern Florida and Texas. Mature caterpillars throughout growing season. Locally common. Slippery burr (Corchorus siliquosus) in south Florida (DLW), Malvastrum in Belize (Tim McCabe), and Apeiba tibourbou in Costa Rica (Janzen and Hallwachs 2011) (all Malvaceae). (Given the range of foodplants and taxonomic uncertainties of species identities across the genus, we suspect that more than one species is represented by this name.)

the ends of each series are reduced in size in the Scoliopterygini (a character shared with Arctiinae). Inducible forms (i.e., color variants that result from crowding and perhaps other environmental conditions during development) occur in Anomini, and most notably in the Cotton Leafworm (Alabama argillacea), a cotton pest (Johnson et al. 1985). Its larvae become heavily marked with black during outbreaks. Other Anomini become reddish when reared at high density, and most take on red tints as prepupae. Many Anomini are animated wrigglers that thrash when alarmed. So far as known, Anomini feed on members of the mallow family (Malvaceae). Our single species of Scoliopteryx feeds on willow. Adults are strongly attracted to bait.

74 Scalloped Moths Subfamily Scoliopteryginae

The Herald Scoliopteryx libatrix ! recognition Elongate, bright green larva with yellow subdorsal stripe. Body wall transparent with tracheae visible within. Subdorsal stripe, often edged with black along lower margin, running length of body but weakening over T1 and A10. Head broadly rounded, usually unmarked; sometimes with fine black middorsal line. Prolegs with pinkish cast; those on A3 and A4 somewhat reduced in size. Crochets at either end of mesoseries much reduced. Spiracles pale tan, orange, or red-orange. Larva to 4.5 cm. ! occurrence Wetlands, riparian areas, woodlands, and forests. Transcontinental in Canada, south in East to Georgia (mountains) and Texas. Seasonality uncertain; most of our caterpillars from collections made in July and August in Northeast. European literature notes that a partial second generation occurs in some areas (Porter 1997). Common northward. ! common foodplants Primarily willow, but also poplar (Salicaceae); birch also reported (Prentice 1962, McCabe 1990). ! remarks The Herald occurs in Europe and North America. Mitochondrial sequence data indicate that North American populations are distinct from those of Europe (Lafontaine, unpubl. data), but it remains to be seen if the New World entity is distinct enough to warrant full species status. We have taken caterpillars while beating in midsummer. The larva commonly pupates in leaves on the host and emerges about three weeks later. Like other moths that overwinter, adults feed at sugar baits. The Herald winters in caves and man-made structures, including sheds and garages, and tolerates temperatures well below freezing.

Scalloped Moths Subfamily Scoliopteryginae 75

Hibiscus Scalloped Moth Anomis privata (= A. commoda) ! recognition Smoky tan, pinkish gray, or brown to gray or blue-gray. Conspicuous, raised, black, dorsal pinacula ringed with white. Subdorsal stripe yellow, about twice width of whitish middorsal stripe, somewhat less developed rearward. Prothoracic shield with spots in two prominent rows. A3 prolegs half size of those on A4. Top of head yellow-orange, although this portion may be pulled under prothoracic shield. Dorsal setae black, about half length of segment that bears them. Spiracles black. Larva to 4 cm. Early instars green like other Anomis, with yellow subdorsal stripe (lower right). ! occurrence Gardens, yards, parks, and nurseries. An Old World species, now established along Eastern Seaboard from at least Massachusetts to Virginia. Three broods in New Jersey and Virginia with mature caterpillars from late May or June into November. Common. ! common foodplants Hibiscus, often rose of Sharon (Hibiscus syriacus). ! remarks A. privata is introduced from Asia, likely from nursery stock. The color and behavior of larvae is a testament to natural selection. Last instars, mottled in grays, rest on bark by day, frequently with head down; whereas earlier instars of the Hibiscus Scalloped Moth are green and rest on leaves. In contrast to many other members of the Anomini, which are known to be migratory, the Hibiscus Scalloped Moth appears to be comparatively sedentary. From its introduction near Moorestown, New Jersey about 70 years ago, the moth has spread only to eastern Massachusetts and southeastern Virginia, a distance the following Anomis species might cover in two nights. During summer and fall, until the first frosts, caterpillars are easily collected by beating. At night first instars frequently dangle on a silk thread and are easily located by flashlight. Two such caterpillars found by DFS on 16 September and reared outdoors, spun their cocoons on 9 and 14 October. These and other pupae held outdoors in his yard produced moths in April.

76 Scalloped Moths Subfamily Scoliopteryginae

Yellow Scalloped Moth Anomis erosa [Abutilon Moth] ! recognition Pale green with thin, often white to yellow subdorsal stripe and large, pale, smooth, unmarked head. Proleg absent on A3; that on A4 also reduced. Middorsal, supraspiracular, and subspiracular stripes variably expressed (sometimes wanting). Dorsum sometimes with minute white circlets (visible with lens); these diminishing in number below subdorsal stripe and absent below subspiracular stripe. Spiracles yellow-orange to beige. Setae long, brownish, especially rearward; head with longest seta nearly equal to width of head. Larva to 3 cm. Similar to White-pupiled Scalloped Moth (A. flava) as a larva and adult; according to Crumb (1956), its setal insertions on the prothoracic shield are bounded by a fine black ring and the caterpillar is sometimes marked with dark addorsal and subdorsal stripes. It occurs along the coastal plain from Virginia to Texas. We have never encountered the caterpillar and know of few recent collections, leaving us to wonder if the White-pupiled Scalloped Moth is declining possibly due to decreases in the acreage of cultivated cotton and okra in the South. ! occurrence Gardens, fields, marshes, hammocks, scrublands, and other mostly open habitats. Mostly southern, but migrants have been recorded from Manitoba, Quebec, and Maine in the fall; it rarely reaches New Jersey in time to breed successfully. Mature caterpillars year-round in southern Florida and Texas. Very common. ! common foodplants Abutilon, bladdermallow (Herissantia), cotton, hibiscus or rose of Sharon, false mallow (Malvastrum), hollyhock, marshmallow, okra, rosemallow, sida, velvet leaf (Abutilon theophrasti), and other Malvaceae. ! remarks Migrants arrive in New Jersey on average every other year, as early as late August, but mostly in October and November. Regardless of when they arrive, all disappear (e.g., from bait trails) after the first hard freeze. Our caterpillars readily accepted fully yellowed leaves, a useful adaptation to latesummer and year-round breeding, but when available, preferentially consumed new tissue. In addition to leaves our larvae ate buds, sepals, and petals. The tacky feculae are flicked away from the body with a snap of the abdomen. In nature feculae often reveal a larva’s whereabouts, especially on those nights and mornings when dew has accumulated on the leaves and trapped the droppings. Scallop Moth caterpillars bred in crowded containers tend to be more darkly patterned than individuals encountered in the wild. A sparse cocoon is spun either on the underside of a living leaf or in leaf litter.

Scalloped Moths Subfamily Scoliopteryginae 77 Windowed Scalloped Moth Anomis exacta

Texas. Mature caterpillars throughout growing season. Locally common. Turk’s cap or wax mallow (Malvaviscus), malva de caballo (Malachra capitata), and probably other Malvaceae.

Straight-edged Scalloped Moth Anomis luridula

Florida and Texas. Mature caterpillars throughout growing season. Locally common. Portia tree (Thespesia populnea) in Florida. (The species name we apply may not be correct—the genus is in need of taxonomic revision.)

Gray-eyed Scalloped Moth Anomis impasta

Gulf states. Mature caterpillars throughout growing season, at least in southern Florida and Texas. Common. Ours reared from Turk’s cap (Malvaviscus) in Texas. Penultimate instar (upper); last instar (lower).

78 Scalloped Moths Subfamily Scoliopteryginae

Cotton Leafworm Alabama argillacea

[Cotton Moth]

! recognition Yellow-green to nearly black with strong

white to yellow middorsal and subdorsal stripes; latter twice width of former. Green-form larvae with little to no black; some larvae with dark addorsal edging to either side of middorsal stripe. One form with dorsum blackened between subdorsal stripes; more melanic forms blackened above subventral stripe. Black pinacula often surrounded by pale ring; dorsal pinacula raised. Proleg on A3 small, less than half size of those of A5 and A6. According to Godfrey (1987) the black spotting on the head is diagnostic (above center). Larva to 4 cm. ! occurrence Agricultural fields. Central and South America, and Caribbean; formerly breeding through cotton belt with late-summer migrants reaching north to Canada. Extirpated, or nearly so, from the United States (see Wagner 2009b). ! common foodplants Cotton; reports from Hampea (Parencia 1978) may be due to misidentification. ! remarks Black-form larvae occur during outbreaks— the adaptive significance of the dark phenotypes, assuming there is one, is unknown. Through much of the 1800s the Cotton Leafworm was the most significant cotton pest in the United States (Riley 1885). Migrants arrived from Mexico in the spring and populations built up over the summer months. By fall, the moth frequently made its way north as far as southern Canada. Holland (1903) wrote of immense swarms that reached Pittsburgh in the fall. Forbes’s (1954) account is equally telling: “Moth basically tropical, but breeding each summer in cotton belt and often flying north in millions in October; also a very active migrant in the tropics, where large populations may appear at a place and shortly move on again.” Like the passenger pigeon, this once abundant animal has had a dramatic reversal in fortune. The insect was last seen in the Northeast in the late 1970s. Johnson (1995) suspected the moth’s decline was due to the elimination of cotton growing in former source regions of northern Mexico. No doubt, widespread insecticide use, cultural practices, and more recently, the use of transgenic varieties resistant to caterpillars, also have played roles in the insect’s demise (Wagner 2009b). In parts of South America, the Cotton Leafworm is still considered an important defoliator of cotton. The adult feeds on fruits such as peaches and grapes (Parencia 1978), and presumably comes to bait.

CATERPILLAR MIMICRY Mimicry of one animal by another was first explained in 1862 by Sir Walter Henry Bates upon his return from 11 years as an insect collector in the Amazon Basin (the first four of which were spent with his friend Alfred Russel Wallace). Writing about butterfly wing patterns, Bates noted, “…on these expanded membranes Nature writes, as on a tablet, the story of the modifications of species…” Bates proposed that longwing butterflies and other Lepidoptera gained protection by mimicking distasteful species—and that the resemblances among such unrelated insects lent support to Darwin’s and Wallace’s newly proposed theory of natural selection. While various forms of mimicry have been identified from across the tree of life, reports of mimicry by caterpillars of other distasteful larvae are curiously rare (e.g., DeVries 1994 and Berenbaum 1995). It is our impression, after having reared and photographed hundreds of owlet species for this work, that caterpillar mimicry is in fact common and widespread in the Noctuoidea (several putative cases are referenced in the index of this volume). Some of the most compelling cases of larval mimicry obtain from the Acronictinae (pages 258–305), whose North American members include mimics of datanas (Notodontidae), lappets (Lasiocampidae), woolly bears (Arctiinae), and tussock caterpillars (Lymantriinae). The Yellow-haired Dagger (Acronicta impleta) (below) is frequently mistaken for a tussock of the genus Dasychira (bottom). Whether this and other acronictine examples represent Batesian mimicry (where the mimic is palatable) or Mullerian mimicry (where both model and mimic are protected) is an evolutionary conundrum waiting to be solved.

Fruitpiercing Moths Subfamily Calpinae 79

Fruitpiercing Moths Family Erebidae: Subfamily Calpinae

Calpines are especially diverse in tropical regions: only two of the 15 North American species are resident in Connecticut. By contrast, Knudson and Bordelon (2000) list seven for Texas. Adults use their armored tongues to pierce fruit (or, in one Asian lineage, mammalian skin) to feed. The body tends to be elongate and the first and sometimes the first two pairs of prolegs are reduced or absent. On A1 (and A2) the

Moonseed Moth

Plusiodonta compressipalpis Minnesota, extreme southern Canada, south to northern Florida and Texas. Two generations over much of East with mature caterpillars from June onward; year-round in southern Florida and Texas. Common southward. Moonseed and snailseed vines. Middle instar (upper) and final instar (lower).

Moonseed Fruitpiercer Gonodonta sinaldus

South Texas. Presumably with continuous generations and mature caterpillars year-round, at least in extreme southern Texas. Common. Moonseed vine. Coloration variable.

distance between SV1 and SV2 is only slightly longer than that between SV1 and SV3 (Crumb 1956). A8 is often swollen or humped. Many are brightly colored and feed in exposed locations on leaves, shoot apices, or stems, suggesting that the larvae are chemically protected or mimetic or both. For example, caterpillars of the Meadow Rue Owlet (Calyptra canadensis) resemble sawfly larvae. Coloration sometimes changes dramatically across instars, and, in Gonodonta, varies confoundingly among individuals in a population.

80 Fruitpiercing Moths Subfamily Calpinae

Meadow Rue Owlet Calyptra canadensis [Canadian Owlet] ! recognition Unmistakable as last instar: dorsum waxy white with scattered middorsal black spots. Subventer and venter normally black (the upper individual figured above with a green venter had recently molted). Head yellow with three black spots on each side. Proleg on A3 reduced. Larva to 3.5 cm. Early and middle instars waxy yellow-green with or without dark subdorsal spots (lower right). ! occurrence Fields, roadsides, wet meadows, edges of rivers, woodland edges, and other mesic open habitats from Canada to Georgia (mountains) and Texas. Two generations in Connecticut mature caterpillars in May and July. Locally common. ! common foodplants Meadow-rue (Thalictrum). ! remarks The color and behavior of late instars are reminiscent of sawflies. The large black lateral spot on the head, which includes the lateral eyes, adds to the “sawfly” ruse. It is not known if the Meadow Rue Owlet is chemically defended, but the caterpillar’s bold coloration suggests so. Alternatively, the caterpillar’s coloration may be cryptic in the same way that many bicolored (dark above and pale below) fish and damselflies are. In aquatic environments pale colors of the venter blend in to the sky to any predators lurking below, and darker colors of the dorsum match those of a pond, and thus challenge the senses of birds and other aerial predators. An analogous phenomenon may occur in Calyptra given that it often feeds and perches ventral (dark) side up. Thus from above, the caterpillar is far less conspicuous than one might imagine, especially in the spring-generation when plants are low to the ground and soils wet and dark. Early instars make smooth-edged, round to elongate holes in the center of leaf blades—the damage is diagnostic and easily located once learned. Often the same patch of meadowrue will be denuded, year after year, with only stems and petioles left, and in some years not even those. In Connecticut, the presence of adults well into September with mature larvae by the 20th of May, suggests that the species overwinters as an egg or partially grown larva. Adults can be taken more reliably at bait than light.

Fruitpiercing Moths Subfamily Calpinae 81

Citrus Fruitpiercer Gonodonta nutrix ! recognition Body black, sometimes with white rings, and prominent orange to salmon spots. Coloration highly variable. A8 humped with A9–A10 reduced and directed downward (from hump); proleg on A3 absent, while that on A4 reduced and less than half size of prolegs on A5 and A6. A1–A8 with prominent, elliptical subdorsal orange spots. Roughly triangular subventral orange spots less developed rearward. A1–A3 bearing an orange midventral spot. Prothoracic shield with three pairs of white spots along leading edge. Anteclypeus and basal antennal segment waxy white. Crochets reduced in size at ends of each mesoseries. Larva to 4 cm. Middle instars elongate, mostly black, with bold markings (lower row). While Crumb’s (1956) account suggests that the larva of Citrus Fruitpiercer lacks the transverse white lines that ring each segment of its congener, G. unica, we have reared G. nutrix from caterpillars with prominent white rings. ! occurrence Orchards, hammocks, and scrub woodlands. Southern Florida. Continuously brooded with mature caterpillars throughout growing season. Common. ! common foodplants Pond apple (Annona glabra) and related Annona. ! remarks The genus is diverse in the Neotropics—Janzen and Hallwach’s website (http://janzen.sas.upenn.edu) illustrates more than a dozen species of these splendid moths. While the handsome black and orange caterpillars often attract attention from growers and the casual passerby, the larvae are rarely abundant enough to cause significant damage to pond apple or cultivated Annona species. Larval abundance peaks in the spring when new growth is burgeoning across south Florida. The insect is a more serious problem as an adult—both males and females have a stout proboscis that they insert into soft fruits to feed on juices. Feeding wounds sometimes spoil and render fruit unsalable. According to Jorge Pena, the tropical fruit entomologist who sent us many caterpillars, carambola (starfruit) sustains the most significant economic injury in southern Florida. The moth also feeds on oranges, tangerines, other citrus, kumquats, and probably additional species. Kimball (1965) notes that adult Lesmone (an erebine) sometimes feed at wounds left by the Citrus Fruitpiercer. Leaf fragments are tied into the cocoon (lower right).

82 Necklace Moths Subfamily Hypocalinae

Necklace Moths

Family Erebidae: Subfamily Hypocalinae This newly recognized subfamily includes seven North American species. Like the previous subfamily, the distance between SV1 and SV2 is only slightly longer than that between SV1 and SV3 on A1 (Crumb 1956). The clade is largely tropical with only two persimmon-feeding Hypsoropha resident northward.

Small Necklace Moth Hypsoropha hormos ! recognition Shape diagnostic. Late instars pale green, smoky black, or combination thereof. Green forms waxy green to lime green, and ranging from unmarked to heavily spotted and/or mottled in grays and black; often with subdorsal black spots. Dark form barklike with charcoal ground color faintly mottled with whites and grays (inset). All color forms often with conspicuous, white dorsal pinacula and abundance of minute white spots. Note shape and abbreviated stature of anal prolegs. Larva to 3 cm. Middle instars lime green, shiny, and translucent (lower right). Co-occurring on persimmon with Large Necklace Moth (H. monilis) during the spring and early summer. The larvae of the two are easily distinguished: the head of the Small Necklace Moth is green or mostly gray and mottled with black, whereas that of the Large Necklace Moth is pale orange with two pairs of black spots. ! occurrence Fields, woodlands, and forest edges from Kansas, Ohio, and New Jersey, south to Florida and Texas. Two or three generations over much of range with mature caterpillars from June onward; multiple generations from coastal North Carolina and southward. Common. ! common foodplants Persimmon; reports from sassafras likely in error. ! remarks Caterpillars of the Small Necklace Moth are easily beaten from foliage. They can also be readily located by turning shoots and examining leaf undersides—new leaves are preferred. At least in the spring, the solitary larvae eat back from a leaf tip. The black subdorsal spots present in some forms resemble the dark blight spots common to older persimmon leaves. Accordingly, dark-form caterpillars seem to be more common among second- and third-brood individuals. In contrast to the Large Necklace Moth (H. monilis), larvae are solitary, remain on the host through the day, and can be collected throughout the growing season.

Necklace Moths Subfamily Hypocalinae 83

Large Necklace Moth Hypsoropha monilis ! recognition Smoky green to black with broad, charcoal middorsal stripe and abundant white speckling. Head pale orange with large black blotch on either side of triangle and small black, lateral spot enclosed by eyes. Larva to 4 cm. Earlier instars dark and with numerous white spots, most of which are aligned into linear series (lower right). OCCURRENCE Fields, woodlands, and forest edges from central Missouri, southern Ohio, and southern New Jersey to Florida and Texas. One generation in North Carolina and New Jersey with mature caterpillars from early June into early July in New Jersey. Common southward. ! common foodplants Persimmon. ! remarks This is a boom and bust species that is decidedly more common in its “good” years. Small trees are commonly stripped of vegetation by the gregarious larvae. The female lays her eggs in rafts on the underside of new leaves. Caterpillars feed on spring foliage and appear to be especially fond of the purple-green, newly produced leaves. The larvae descend from foliage at or before dawn to pass the day in litter or uncompacted soil beneath the foodplant, often several centimeters from the stem base. At dusk, the caterpillars ascend, in line, back up the trunk. New shoots that have been stripped of adjacent leaves (including leaf petioles) are a sure indication of the species’ presence—even if no larvae can be located. The single, black “eyespot,” orange head, and gregarious nature make the caterpillar a credible sawfly mimic. If poked or pinched, the caterpillar reels around and simultaneously bites its attacker and vomits a clear orange liquid. The prepupal larva overwinters in a cocoon in the litter.

84 Scolecocampines Subfamily Scolecocampinae

Scolecocampines

Family Erebidae: Subfamily Scolecocampinae The taxonomic limits of this subfamily are in flux. Genera included by Lafontaine and Schmidt (2010), e.g., Abablemma, Arugisa, and Nigetia, that graze on algae and lichens are excluded here—we place these in Hypenodinae. Isogona also seems to be a poor fit because the caterpillars are arboreal leaf feeders. Scolecocampines are recognized by unique features of the adult tympanum and male and female genitalia (Fibiger and Lafontaine 2005). The nominate genus Scolecocampa has a mostly unpigmented larva that feeds in decaying logs.

Snow’s Owlet Isogona snowi

Southern Texas; breeds year-round. Common. Spiny hackberry (Celtis pallida).

Feeble Grass Moth Amolita fessa

Wisconsin to Nova Scotia, south to Florida and Texas. Evidently at least a partial second generation with mature caterpillars from June onwards. Locally common. Grasses.

Pink Sedge Moth Amolita roseola

Ohio to southeastern Massachusetts, south to Florida and Texas. One principal generation with mature caterpillars mostly in June in Connecticut; multiple broods through growing season in Florida. Local and generally uncommon. Pennsylvania sedge (Carex pensylvanica) in Connecticut.

Scolecocampines Subfamily Scolecocampinae 85

Dead-wood Borer Scolecocampa liburna ! recognition Shiny, whitish, somewhat translucent caterpillar with proleg on A3 absent (found boring in wood). Pinacula raised, dark brown, outer edge diffuse. Dorsal setae long, rusty, about as long as segment that bears them. Prothoracic shield shiny black with thin middorsal break; lower edge squared off, extending nearly to spiracle. Anal plate black, armored. Spiracle on A1–A6 about half size of that on A7; this spiracle in turn about half height of that on A8. Crochets long and slender. Larva to 4 cm. Superficially similar to larvae of ghost moths (Hepialidae) and carpenter moths (Cossidae), which also tunnel in wood; however, members of both of these families have a well-developed proleg on A3. Also, the crochets are arranged in circles rather than as a linear series as in all owlets. ! occurrence Woodlands and forests from southern Wisconsin and Michigan to central New England, south to Florida and Texas. Common over much of range. Evidently just a single protracted generation northward, but additional broods southward. In southern Ohio, adults have been collected from late May to late July and again in late August (Rings et al. 1992). ! common foodplants Caterpillar tunnels in fallen branches and trunks of chestnut, hickory, oak, sycamore, and other trees. Fallen and decaying tuliptrees often yield caterpillars (Rings et al. 1992 and DFS). ! remarks Like many borers, seasonal phenology is protracted. The larva tunnels under loose bark and in decaying wood, which is sometimes blackened. But we have also found caterpillars simply tunneling through moist, pulpy wood (with no discoloring and little hint of fungi), and we were wholly unsure how the caterpillars were managing nutritionally. The genus and its relatives are richly represented in low and midelevation rainforests of the Neotropics, where it is not uncommon to find 10–15 species at a single site. Costa Rica alone may have as many as 50 species of Scolecocampa. Evidently the larva of S. liburna overwinters.

86 Scolecocampines Subfamily Scolecocampinae

Thin-lined Owlet Isogona tenuis ! recognition Elongate, subtly striped bark mimic. Proleg on A3 a mere peg; that on A4 less than half size of those on A5–A6. A3 and A4 slightly swollen and often with vague brown patch over dorsum (especially on A3). Pale addorsal stripes narrower and straighter than five rows of subtending, wavy, frequently interrupted, subdorsal and supraspiracular stripes. Long setae borne from black or white warts. A1 long, approximately same length as combined length of T1–T3. Length of A2–A4 nearly twice that of A6; A7 shorter than A8. Dorsal setae on A8 and A9 very long; those on A9 extending beyond rear of A10. Venter frosty without markings. Head tan brown with black mottling, bearing long, forward-projecting black setae. Larva to 3 cm. Penultimate instar gray-green without stripes, and more subdued markings. Snow’s Owlet (I. snowi) of southern Texas is so divergent (page 84) in appearance that we wonder about its taxonomic placement as a congener of I. tenuis. ! occurrence Fields, woodlands, riparian and bottomland forests from Wisconsin, Ohio, and Connecticut, south to Florida and Texas. Two generations over much of East with mature caterpillars from June to early September in southern Ohio and New Jersey; additional broods southward. Locally common. ! common foodplants Hackberry. ! remarks The caterpillar is an excellent twig mimic—a person unfamiliar with the Thin-lined Owlet might even guess that hackberry was the host just by comparing the larva to an assortment of local twigs. The cocoon is a cigar-shaped construction often spun over a midrib; an elongate section of the leaf to either side of the cocoon is carefully cut away from the rest of the blade, wrapped over the top of the pupa, and tied down with silk. Each end is then sealed with a round leaf section (inset). The pupa overwinters.

Halflings Subfamily Hypenodinae 87

Halflings

Family Erebidae: Subfamily Hypenodinae Our treatment of the subfamily differs from that of Lafontaine and Schmidt (2010) who place some of these genera in the Scolecocampinae. Our concept of the subfamily, which leans more closely to that of Franclemont and Todd (1983), is based on our studies of Abablemma and Nigetia, along with published images and descriptions of European species of Hypenodes and Schrankia (Beck 1999, 2000; Ahola and Silvonen 2005; Wagner et al. 2008). The larva is elongate, lacks prolegs on A3 and A4, and the proleg on A5 is reduced in size and shifted rearward. A1–A4 are elongate. The D1 setae are tilted

Common Arugisa

Arugisa lutea (= Arugisa latiorella of Covell 2005 and others) Missouri to southern New Jersey, south to Florida and Texas. Breeds year-round southward. Common. Green and dead grasses (Crumb 1934); Tim McCabe (unpubl. data) reared figured individual ex ova on blue-green algae.

Fetching Halfling

Nigetia formosalis [Thin-winged Owlet] Ontario to Connecticut, south to Florida and Texas, but rare to absent over parts of range. One principal generation in Connecticut; two or more southward. Phenology unknown, presumably with mature larvae preceding adults in late spring and August over much of East. Locally common. Captive larvae reared to maturity on green algae (Protococcus) by Tim McCabe; presumably also consuming some lichens.

Braun’s Quandry Sigela brauneata

At least Kentucky, southern Ohio, and New Jersey, south to Florida and Texas. Two principal generations northward; multiple generations southward with mature larvae evidently present year-round. Captive larvae reared to maturity on green algae (Protococcus) by Tim McCabe.

forward and D2 setae rearward. In Hypenodes and Schrankia there are three SV setae on A1 but only two on A2. In Europe and elsewhere hypenodids have been reared from lichens. Arugisa feeds on green and dead grasses (Crumb 1934) as well as algae (Tim McCabe, unpubl. data); Tim McCabe has reared Abablemma and Nigetia on algae, and Dyspyralis on dead leaves. A few hypenodids are predaceous on scales and other Homoptera. Some Old World genera cover themselves with excised lichen fragments. Lichens and pieces of bark may be woven into the outer wall of the cocoon (see inset p.88); in Abablemma and some Palaearctic genera the cocoon is pedunculated. Larvae overwinter (Porter 1997, Sugi 1987).

88 Halflings Subfamily Hypenodinae

Double-eyed Halfling Abablemma duomaculata ! recognition Elongate, cryptically marbled in olive green, grays, white, and black with long, curved setae. Prolegs absent on A3 and A4. Dorsum with broad pale middorsal stripe with bulges at midsegment. Black subdorsal patch pushing up against middorsal stripe on A5–A7. Venter black, sharply contrasting with dorsum. Surface of body papillose. Dorsal setae over abdomen arising from warts. Larva under 1.5 cm. Other Abablemma occur in our region. ! occurrence Thorn scrub and palm woodlands. Southern Texas. Presumably multiple generations with mature caterpillars nearly year-round. Locally common. ! common foodplants Lichens; ours reared on Physcia. ! remarks The cocoon is a marvelous pedunculated construction (inset). The larva removes small fragments of lichen adjacent to the attachment site and meticulously weaves these into the wall of the cocoon. The caterpillar has a peculiar way of moving: the anterior end of the body is flopped to one side, then the caterpillar pauses before following with the rear of the body. The larva overwinters.

Brimley’s Halfling Abablemma brimleyana

New Jersey, south to Florida and Texas. Two generations in New Jersey where it established in late 1990s; more southward. Sometimes common in dry woods and xeric habitats. Captive larvae reared to maturity on green algae (Protococcus) by Tim McCabe; presumably also consuming lichens.

Fungus Moths Subfamily Boletobiinae 89

Fungus Moths

Family Erebidae: Subfamily Boletobiinae Most of the 17 North American species are classified in the genus Metalectra. The subfamily is closely related to Hypenodinae—characters listed there apply equally well to Metalectra. Metalectra can be reared on common algae (Protococcus) that

White-lined Fungus Moth Metalectra albilinea

At least Tennessee to North Carolina, south to central Florida, and west to Mississippi. Multiple generations with mature caterpillars throughout the growing season. Locally common. Green algae, fungus, and presumably other organic matter of forest floors. Small, mature larva less than 13 mm.

Four-spotted Fungus Moth Metalectra quadrisignata

Ontario to Maine, south to Florida and Texas. Presumably two or more generations. Common. Fungi, including dried fruiting structures. Eric Quinter reared ex ova larvae on a variety of bracket fungi and mushrooms, both living and dead—most preferred to feed on actively growing bracket fungi.

Black Fungus Moth Metalectra tantillus

Missouri to southeastern Massachusetts, south to Florida and Texas. Presumably two or more generations. Locally common. Kimball (1965) recorded larvae feeding on bark of a dead maple— presumably grazing on algae and lichens; Tim McCabe has reared a cohort on green algae (Protococcus) (Wagner et al. 2008).

grows on tree bark (Tim McCabe, unpubl. data; Wagner et al. 2008), although in the wild, larvae are usually found feeding on bracket fungi, mushrooms, and undifferentiated fungal mats. We have much success rearing Metalectra on fresh polypore fungi.

90 Fungus Moths Subfamily Boletobiinae

Common Fungus Moth Metalectra discalis ! recognition Waxy white to nearly black with conspicuous patterning and dorsal setae that alternate in their orientation: i.e., D1 setae directed forward and D2 setae rearward. Prolegs absent on A3 and A4. Long setae borne from pimplelike, often yellowed to pinkish warts. A1 with dark, oblique, subdorsal patch dropping down and forward from D1 pinaculum; oblique white patch running from brown middorsal spot to D2 pinaculum and beyond black spiracle on A1, A2, and other segments. D2 setae connected by ridge on A8 and A9. Larva to 2.5 cm. Other fungus moths (Metalectra and likely related boletobiine genera) occur in our region—identifications should be based on reared adults. ! occurrence Woodlands and forests from Wisconsin to Maine, south to Florida and Texas. Long flight period in Northeast extending from May to September, suggestive of two or more generations with mature caterpillars from late June onward. Common. ! common foodplants Bracket and other fungi; ex ova collections will mature on diet of green algae (Protococcus) (Wagner et al. 2008). ! remarks Adults are common at bait. Metalectra are most frequently found feeding on the underside of fleshy bracket fungi, but many mushrooms also serve as hosts. In captivity they can be reared on store-bought (Agaricus) mushrooms. We once found five larvae feeding on a mycelial (fungal) mat growing inside a waterlogged wooden compartment used for the storage of life preservers— evidently a gravid female had been drawn to the boat by volatiles emanating from the fungus. The prepupa weaves particles of soil, frass, and litter into the wall of the cocoon. The pupa overwinters.

Phytometrines Subfamily Phytometrinae 91

Phytometrines

Family Erebidae: Subfamily Phytometrinae This small group, previously classified with the hypenines (Hypeninae), was elevated to its own subfamily by Fibiger and Lafontaine (2005). Less than a dozen species occur in the East—adults are small to medium sized, usually under 35 mm in wingspan. We are dubious that the ten genera listed by

Lafontaine and Schmidt (2010) form a natural group. Larval prolegs are short and broad; in three species, the prolegs on A3 and A4 are absent or reduced; and at rest the anal prolegs are extended behind the body. The body wall is sometimes translucent and the setae are fine and of modest length. The taxonomic limits of the subfamily are in flux—our placement of Colobochyla and Hyperstrotia here is tentative.

Six-spotted Gray Spargaloma sexpunctata ! recognition Pale green, smooth, unpatterned, undistinguished. Proleg on A3 somewhat reduced. Anal prolegs stout, directed backward. Heart and tracheal trunks visible through body wall. Pale setae about half length of segment that bears them. Head small, pale green, and unmarked except for black eyes and waxy labrum. Larva to 2.5 cm. ! occurrence Dry fields, roadsides, woodland clearings, and other open habitats. Transcontinental across lower Canada, south in East to Florida, Mississippi, and Arkansas. One generation northward, two or more in Connecticut and southern Ohio with mature caterpillars from June onward; more generations southward. Locally common. ! common foodplants Dogbane. ! remarks The caterpillar feeds from and perches on the underside of leaf blades. Although they may be collected with beating sheets, we have had much success by simply turning shoots and examining leaf undersides on plants with conspicuous feeding damage—look for jagged leaf edges. Be advised that the caterpillar may rest on blades well away from those upon which it has fed—perhaps playing a “shell game” with predators and parasitoids that use leaf damage and volatiles released from feeding wounds, respectively, to locate prey. The pupa overwinters.

92 Phytometrines Subfamily Phytometrinae

Variable Tropic Hemeroplanis scopulepes ! recognition Smooth, brown above level of spiracles, with broad white lateral stripe. Prolegs absent on A3 and A4. Prominent white lateral stripe extending from antenna to anal proleg running immediately below level of spiracles, divided by and edged with fine reddish lines. Subventral area, below lateral stripe, cream to yellow, divided by three additional fine red lines on T3–A8; subventer mostly brown and otherwise unmodified on T1 and T2. Pale, dirty-yellow middorsal stripe best developed rearward, divided and edged with fine reddish lines; poorly differentiated over thoracic segments. Broad, pale midventral stripe bordered by brown adventral stripe. A9 short, only about a third length of A8. Head pale brown with pale snowflaking; clypeus mostly unpigmented. Larva less than 3 cm. ! occurrence Pastures, fields, roadsides, and other open habitats. Wide ranging throughout South, but straying north to Missouri, Ohio, and New York. Multiple generations with mature caterpillars through growing season. Common southward. ! common foodplants Our ex ova larvae were reared to maturity on clover. ! remarks The lethargic larva perches outstretched, often off the host with the anal prolegs extended rearward; the posterior end of the body is somewhat flattened. When alarmed the caterpillar releases its grip, falls to the ground, and folds its body in half (right). The caterpillar—grasping its prolegs with the thoracic legs—often remains motionless well after danger has passed. JBS has reared a second member of the genus from another legume, dwarf bristly locust (Robinia hispida nana), in North Carolina. A weak cocoon is spun in leaf litter. We suspect that the pupa overwinters.

Phytometrines Subfamily Phytometrinae 93

Pink-border Yellow Phytometra rhodarialis ! recognition Elongate, slender caterpillar with numerous, fine, red to smoky pinstripes. Prolegs absent on A3, those on A4 minute, peglike. Abdominal segments 1–4 elongate, nearly twice length of adjacent segments; body broadest about A5. Reddish middorsal stripe ending toward rear of T1 and front end of A10. Three rows of paired stripes above level of spiracle and two below. Setae dark, long, nearly length of segment that bears them. Body wall translucent, heart, tracheal trunks, and reddish testes of male visible through body wall. Head with numerous reddish stripes. Larva to 2.5 cm. ! occurrence Fens, grassy wetlands, flatwoods and moist pinelands, savannas and open woodlands from Missouri, southern Ontario, and New Hampshire (historic), south to Florida and Texas. Two or more generations in Ohio and New Jersey with mature caterpillars probably from June onward; more broods southward. Locally common. ! common foodplants Milkwort (Polygala). ! remarks At the suggestion of Tim McCabe, we confined females with leaves of milkwort and were soon rewarded with numerous maroon eggs. Our caterpillars accepted two milkworts in captivity: yellow polygala (Polygala lutea) and Maryland milkwort (P. mariana), with the latter favored. Larvae fed on flowers, chewed away the parenchyma and other green surface tissues of the square stems, and often ignored leaves. Alarmed caterpillars wriggle violently as do their kindred Hypeninae. Some of our larvae fashioned a cocoon by rolling up a leaf (inset). The male’s testes are visible through the body wall in the image above (behind the leading edge of A5). The pupa overwinters.

94 Phytometrines Subfamily Phytometrinae

Marsh Belle Colobochyla interpuncta ! recognition Small, elongate, yellow to lime green with deeper green heart stripe; faint, whitish addorsal and subdorsal stripes and creamy spiracular stripe. Incisure between T3 and A1 weakly expressed. Setae rusty to blackish, curved, rearward longer than segment that bears them. Areas of overlap between adjacent segments yellowish. Proleg minute on A3, smaller than basal segment of antenna; proleg on A4 half size of those that follow. Minute pale orange spiracles. Head pale green, unmarked, with long setae. Short anal prolegs splayed behind body. Larva to 2 cm. Easily confused with Black Snout (Hypena scabra), which occasionally can be found on willow, but lacks prolegs on A3. ! occurrence Wetlands from Wisconsin to Nova Scotia, south to Florida and Texas. Evidently with two generations in Ohio and Connecticut; more southward. Local and uncommon. ! common foodplants Willow. ! remarks Larvae shun older foliage. At rest, the body is somewhat flattened—the thoracic segments being the most obviously compressed. The larva is a looper. The working heart is readily observed through the transparent body wall. While Forbes (1954) notes that the cocoon is formed at ground level with dirt and sand interwoven, the norm for the Lesser Belle (C. salicalis), a European relative, is to fashion its firm cocoon along a twig of the host. Given the abundance of willow and frequency from which we beat larvae, it is curious that the moth is not more common in collections. The phylogenetic placement of Colobochyla is in doubt. Richards (1933) placed the genus in the Rivulinae, but molecular data support for its membership in the Hypeninae (Zahari et al. 2011). We leave the genus in Phytometrinae, near Phytometra, which shares similarities in habitus and proleg reduction.

Phytometrines Subfamily Phytometrinae 95

graylets Hyperstrotia

The caterpillars are small, elongate, pale to lime green, with prolegs on A3 absent and those on A4 reduced. The head is broad, and the body tapers rearward. Setal bases slightly raised, sometimes reddened or blackened. Larva under 2 cm. In dry woodlands and barrens, graylets are among the most common caterpillars in beating samples from beech and oak over many weeks during the summer. Early instars scrape away small

Yellow-spotted Graylet Hyperstrotia flaviguttata

Ohio (rare) to Massachusetts, south to Florida and Texas. Two generations over much of East with mature caterpillars from June to August, but with most in July and August after flight of summer brood; multiple generations in Florida. Local in barrens northward; common and widely distributed from New Jersey southward. Oak, both reds and whites (see Dotted Graylet).

Black-patched Graylet Hyperstrotia secta

Southern Ontario to Massachusetts, south to Florida and Texas. Two generations over much of East with mature caterpillars in June and then again in July and August. Locally abundant. Oaks, with some indication that white oaks are favored.

White-lined Graylet Hyperstrotia villificans

Great Lakes states to Nova Scotia, south to Florida and Texas. One generation in New England; two generations in New Jersey and Washington, DC area, perhaps more southward. Common and widespread. Chestnut and oaks mainly; also recorded from elm and possibly once from hickory (DLW).

patches of parenchyma, leaving the veins intact—the skeletonized “windows” are diagnostic. Later instars are large headed, and well suited to consume hardened summer foliage. They rest stretched out on the lower surfaces of leaves, and often can be recovered in number simply by scanning leaf undersides, especially at night. As they walk, the caterpillars lay down strands of silk. Graylets pass the winter as pupae in soil or duff. The confounding host associations of the genus are discussed in the Remarks section for the Dotted Graylet (H. pervertens).

96 Phytometrines Subfamily Phytometrinae

Dotted Graylet Hyperstrotia pervertens ! recognition General description supplied on previous page. We are unaware of species-specific characters for any of our eastern Hyperstrotia. Larva to 2 cm. The four widespread members of the genus in the East are illustrated in this work—additional species (H. aetheria, H. nana) are found in Florida and Texas. ! occurrence Woodlands and forests from Missouri to Nova Scotia, south to Florida and Texas. One principal generation with mature caterpillars from July to August in Connecticut; two generations in Washington, DC area with caterpillars mostly in June, then again in late July and early August (John Lill, unpubl. data); additional generations southward. Common. ! common foodplants Beech and oak. ! remarks The degree of red and black pigmentation about the pinacula varies considerably within species. Host records for our graylets are perplexing. The Dotted Graylet is reported to feed on both oak and elm (Forbes 1954, Prentice 1962), yet a cohort to which we offered beech, white oak, and American elm accepted only the first. In New England and elsewhere the abundance of the moth mirrors the range of beech more closely than either elm or oak. The dozens of Hyperstrotia larvae that John Lill has found in the Washington, DC area on beech have all yielded adults of the Dotted Graylet. We have a rearing of the Dotted Graylet from post oak in South Carolina, but the moth is smaller, brighter, and more sharply patterned than our H. pervertens collections from the North. Lill’s Hyperstrotia larval collections from white oak in the DC area have mostly yielded adults of the Black-patched Graylet (H. secta), fewer White-lined Graylets (H. villificans), and only one Dotted Graylet (H. pervertens). In New Jersey, we have reared 15 White-lined Graylets from larvae on post and black oaks, three Yellowspotted Graylets (H. flaviguttata) from post oak, one each from black and scrub oaks, and two Black-patched Graylets from white oak. This is yet another group where someone could make immediate contributions to lepidopterology. Wild larvae are easily collected (in large numbers by beating). Much could be learned by offering cohorts of larvae salads of potential host genera, although getting confined females to lay in captivity can be difficult—our egg clutches were obtained by sleeving females on bear oak (H. flaviguttata) and beech (H. pervertens).

Underwing Moths Subfamily Erebinae: Tribe Catocalini 97

Underwings, Zales, Witches, and Kin Family Erebidae: Subfamily Erebinae

This is the second largest owlet subfamily in North America with over 310 species listed by Lafontaine and Schmidt (2010). The subfamily is especially diverse in Florida and Texas (and the Neotropics). The 140 erebines figured in this work are grouped into six tribes: Catocalini, Euclidiini, Melipotini, Ophiusini, Poaphilini, and Thermesiini. Erebine caterpillars are often elongate with reduced or missing prolegs on A3 and a substantial proportion throw a loop into the body while walking, although exceptions are myriad. A few are even cutwormlike. There are three subventral setae on A1 and A2, and the SD1 seta on A9 is not thin and hairlike (except in some Euclidiini). The crochets are of a single length (uniordinal). Larval color and patterning may vary within a species—use coloration characters cautiously. Most are arboreal caterpillars that feed on woody plants, although many grass and forb feeders are included (for example, Euclidiini hosts include grasses and low-growing legumes). Later instars routinely remove themselves from the feeding site to seek shelter along a twig, stem, bark, or other site of safety; these, not surprisingly, are often colored in grays, browns, and other earth tones. The pupa (the overwintering stage common to most members of the subfamily) often has a waxy bloom, a characteristic that the Erebinae share with many tiger moths (Arctiinae), to which they may be related (see Mitchell et al. 2006). Most are foodplant specialists.

Underwing Moths

Family Erebidae: Subfamily Erebinae: Tribe Catocalini The tribe is represented by a single genus, Catocala, familiar to a broad spectrum of entomologists as well as the lay public.1 Catocala is the fourth largest genus of macrolepidopteran moths in North America with more than 110 species described and several as yet unnamed taxa awaiting formal recognition.

Approximately 80 of the named species occur in the East. Their size, beauty, and habits have made them a favorite with collectors and nature photographers. Diets are specialized: for example, all of our two dozen hickory feeders are associated with members of just one of the two North American hickory sections (i.e., either Carya or Apocarya2), and no less than six species appear to be specialists on a single species, shagbark hickory (Carya ovata). Foodplant preferences for many of the eastern North American Catocala were studied by Gall (1990a, b). Many ranges given here were provided by Larry Gall (unpubl. data). Species accounts and head capsule images are grouped by hostplant preferences as follows: Legume (Fabaceae) feeders Gale and sweet fern (Myricaceae) feeders Hickory and walnut (Juglandaceae) feeders Willow and poplar (Salicaceae) feeders Oak (Fagaceae) feeders Blueberry (Ericaceae) feeders Rose family (Rosaceae) feeders (apple, cherry, hawthorn, etc.) Underwing caterpillars are elongate with dark brown to, more commonly, purple-black spots along the venter; many species possess thickened setae (rootlet setae) that form a subventral fringe about the body. The “belly” of late instars is often tinted with pink, red, or yellow. Larval coloration can be exceedingly variable and is especially so among the gale feeders (pages 107–9). Many take on a waxy bloom in late instars—such is especially dense in Robinson’s Underwing (C. robinsonii). Head patterning is also variable, especially among the Rosaceae feeders, so use head coloration characters in Forbes’s larval key (and our plates) with caution. Forty eastern species are figured in Barnes and McDunnough (1918). We illustrate 78 species in this guide: 20 in full and an additional 58 in abbreviated accounts. Many of the latter are available as full species accounts on DLW’s website. The 90 head capsule images represent 76 species. Many diagnostic features in our species accounts were extracted from Forbes’s key.

Section Carya includes C. glabra, C. laciniosa, C. myristiciformis, C. ovata, C. ovalis, C. pallida, C. floridana, C. texana, and C. tomentosa; section Apocarya includes C. illinoinensis, C. aquatica, C. cordiformis, and C. palmeri, and C. myristiciformis in some treatments.

2

1 We exclude Spiloloma from the tribe and regard this moth to be better placed in Ophiusini.

98 Underwing Moths Subfamily Erebinae: Tribe Catocalini Genus-wide there is one generation a year. All overwinter as eggs—an attribute that attests to the decision by taxonomists to accord underwings their own tribe (other temperate Erebinae overwinter as pupae). The ova are laid singly, in small clusters, or in rafts (below left) in bark crevices, under flaps of bark, or in litter at the base of the foodplant. Females have an extensible abdomen that allows them to deeply insert the eggs into protected sites. The neonates are extremely active and routinely wander to locate their first meal. Most are new leaf specialists. Some, such as the Ilia and Scarlet Underwings (C. ilia and C. coccinata, respectively), will starve if offered older foliage, although the Girlfriend Underwing (C. amica), its kin, and many hickory feeders consume fully expanded leaves as late instars. Once leaves become fully hardened and leatherized they become unsuitable for many members of the genus. Of course, with over 100 North American species there are exceptions: Robinson’s Underwing (C. robinsonii), the last of our hickory feeders to complete its larval development, matures on midsummer foliage. Underwing caterpillars, and especially those that feed on trees, are strong, active crawlers that can move tens of meters to locate appropriate food or shelter. Five instars are typical of the smaller species, including the Epione Underwing (C. epione), The Consort (C. consors), and some of the willow feeders. Larger willow-feeding species and most hickory feeders have six instars, although both the Widow Underwing and The Bride (C. vidua and C. neogama, respectively) have seven (Larry Gall, pers. comm.). Early instars can be found resting on the underside of leaves or petioles. Middle and late instars, as their coloration suggests, spend most of the day resting on bark, positioning themselves along twigs, branches, and the trunk, often head down, sometimes with the face pushed up against the base of a leaf scar, petiole, or twig. Preferred resting sites differ markedly among instars (see Schweitzer 1982b; and White Underwing, Catocala relicta, page 125). The older instars of many select branches and stems that are approximately the same diameter as their own bodies. Start your search for these in the vicinity of the lowermost extent of the new year’s growth. Caterpillars that

descend to the main trunk frequently tuck themselves into crevices, cram their bodies under bark flaps, or descend into growth or other sheltered sites at the bases of their foodplant; a few hide in leaf litter and nearby vegetation by day (e.g., some legume feeders leave the host to shelter on nearby sticks or vegetation). At dusk, or sometimes earlier, the caterpillars move back onto foliage to feed. Some underwing caterpillars toss their fecal pellets by flicking their rear to one side. Catocala, especially those that shelter under hickory shags, are surprisingly muscular, and are capable of launching themselves from their purchase with great force. Once free of their perch, they continue to snap wildly; in nature, this behavior drives larvae beneath fallen leaves and other litter on the forest floor. Members of the serena-habilis-robinsonii group—all of which wedge themselves under shags of hickory by day—have an especially vigorous response, thrashing madly for many seconds, before coming to rest (and feigning death). Prepupal caterpillars thicken through the middle segments, sometimes take on a faint green to gray-green to gray-blue cast (especially visible about prolegs), lose patterning, and acquire a characteristic sheen. Such may wander from the host before spinning a sparse cocoon, usually in surface litter.

! Darling Underwing (Catocala cara) eggs. !

! Ilia Underwing (Catocala ilia) lichen form. !

Underwing Moths Subfamily Erebinae: Tribe Catocalini 99 Hunting Catocala caterpillars is good sport: challenging, engaging, and rewarding. Knowledge of a species’ preferred hosts, host size, seasonal phenology, and behavior will facilitate your search. For example, each species (and instar) may have a specific perching site—see account for White Underwing (page 125). Check leaves, petioles, and twigs for early and middle instars. Catocala that prefer larger trees (e.g., many of the shagbark hickory feeders) are not easily secured as early instars because few branches are at a height where leaves can be inspected or sampled. Nighttime flashlight searches can be fruitful. Watch for signs of feeding damage: late instars are sizeable, ravenous caterpillars—their feeding damage is conspicuous and diagnostic, although some (for example, Yellow-banded Underwings, C. cerogama) are reported to be leaf clippers (Heinrich 1993). For all but the hickory, walnut, and a few oak feeders, restrict your searches to new spring foliage. Beating sheets can be effective especially for early and middle instars. To acquire late instars from trees, lay down a white bed sheet(s) under low branches and use a baseball bat to rap branches and stems of targeted hosts. Consider working in teams with a person at each end or corner of one or two king-sized bed sheets; this also allows the sheet(s) to be quickly moved, held above vegetation (poison ivy), tilted, etc. The last two instars can be collected under burlap skirting (see page 16). Don’t expect many caterpillars: one or two for every 20 bands, on good days. Late instars of the shagbark feeders, e.g., C. angusi, C. habilis, C. judith, C. obscura, C. residua, C. retecta, C. robinsonii, and C. serena, can be found by checking under shags along the trunk. (You may want to position a beating sheet below the shags in case the larvae “jump” once exposed.) The best shagging is on young trees that are large enough to have shags at or below chest height, but not so large that larvae have abundant opportunities to shelter far above the ground. A significant proportion of the larvae that you will find under shags will be undergoing a molt and thus will be vulnerable to handling damage. The pupal stage is three to four weeks in Catocala; work backward from the beginning of a species’ flight season to time your searches. Baiting is a time-honored means of securing adult underwings (see below, and Sargent 1976). Another is “tree tapping.” On hot days, adult underwings perched on tree trunks can be flushed with a branch or stick. Roused individuals will often settle on nearby trees. Relocating these individuals and successfully photographing or capturing them is good sport— the moths have better than even odds of eluding you. Adult Catocala tend to perch on oaks, walnuts, and especially hickories; most shun smooth-barked plants and smaller trees where their coloration would make them apparent. Some of the shagbark hickory (Carya ovata) feeders will be found almost exclusively resting on their host, nestled under shags. Trees with vines growing up the sides are always worth checking— these and hickory shags should be lightly tapped with either the handle of one’s net, a stick, or brushed with a limb with a few leaves still attached at its tip. Once you get a feel for what

size and kinds of trees the moths are favoring, try locating the moths before they fly—individuals that have not yet flushed are more easily approached. Moths tend to move lower down on boles on the hottest days. Calm, humid, uncomfortably hot days in excess of 32° C (90° F) from the middle of July through the end of September will be the most productive. A good place to start your search is in a bottomland forest with an abundance of shagbark and/or shellbark hickories; shaded hollows or ravines with big trees may yield adults in numbers. It is also worth checking the underside of ledges, bridges, and culverts for adults; many of the willow feeders gather in such places, especially towards the end of summer. For additional information on tapping see Sargent (1976), Willis (1991), and Gibson (2008). Another sporting means to locate females is to go out to stands of the hostplant at dusk and look for ovipositing females—you will need a headlamp, and a long-handled net. This method can be especially effective for the hickoryfeeding underwings. Catocala caterpillars do well in captivity if offered acceptable foliage and ventilation, and if their containers are not crowded. What constitutes acceptable foliage is a matter of much mystery. We have lost many larvae through the years: some straggled for days and days before their demise on what appeared to be adequate foliage. On suitable foliage, vernal species grow fast. Watch for copious frass production and daily increases in body size, except over the two days of a larval molt (when the larva should not be handled). Offer straggling larvae salads of leaves and make note of what (aged) leaves are being eaten. Domestic apple and wild and ornamental crab apples can be used to rear many of the apple, hawthorn, and chokeberry feeders. Carya feeders will often accept walnut in captivity, especially in middle and later instars. Scrub or bear oak, and European cork oak (Quercus suber), are widely accepted by the oak feeders (Larry Gall, pers. comm.). For more information on this popular genus consult Sargent’s (1976) Legion of Night. The first chapter of the book has an engaging account on how many underwings came to be named after wives, lovers, and girlfriends.

BAITING FOR UNDERWINGS There is a long and lore-ridden tradition of using sugary baits to attract underwings. One of the most famous pieces of lepidopterological prose is Holland’s (1968: 146) account of a memorable night of baiting in western Pennsylvania. Bait yields can be spectacular. On 13 July 1981, DFS and Larry Gall encountered some 500 Catocala representing 21 species in about three hours along a bait trail through mixed pine-oak woods and swampland near Atsion, New Jersey. Be forewarned that moth activity at bait is notoriously unpredictable and often disappointing—a few years later, on a night of seemingly favorable weather conditions, the Atsion bait trail yielded no Catocala, yet adult underwings were seen flying in the woods. We have had much success during periods of dry weather when there have been few natural feeding resources available. Also, the nights following a hurricane or extended hard rains can be particularly productive—perhaps because naturally occurring sources of sugar, including the accumulation of aphid and scale honey dew, have been washed away.

100 Underwing Moths Subfamily Erebinae: Tribe Catocalini OBTAINING EGGS FROM CATOCALA Catocala that feed on willows, poplars, oaks, and members of the apple family (Rosaceae) usually oviposit readily in brown paper bags. While some females will lay on the first night of their captivity, most will need to be held for several nights, and in some cases weeks, before they lay. Food can be supplied on sponge pieces or cotton balls. Moisten the feeding station before dark with a plant mister and mix new solution about every three or four evenings. Recently emerged females may be unmated—this seems to be the case particularly among the late-season flyers, such as the Joined Underwing (C. junctura) and Robinson’s Underwing. If circumstances allow, wait until middle to late September to put females of these and other late-season Catocala up for eggs. John Peacock houses females in lunch bags. If available, a cool, damp basement is ideal for holding adults but any shaded outdoor or humid indoor situation will suffice. Another option, especially if moth-feeding rodents are likely to be a problem, is to suspend the bags from a span of clothesline. The lunch bags can be slipped into plastic bags left open at the top to keep the humidity higher and discourage the rapid evaporation of supplied food. Alternatively, bagged females may be placed in foam or plastic boxes (e.g., coolers) to reduce drying. Captive females frequently will push their eggs into what seem to be impossibly tight places, such as under the inside seam of a paper bag. For the larger oak feeders, make multiple small slits (2 cm in length) about the bag—females will push their ovipositor through the slits and lay their eggs on the outside of the bag. Some Rosaceae feeders oviposit more readily if a leaf of the foodplant is placed with the female. Likewise, pieces of bark from the foodplant will often stimulate egg laying in hickory and walnut feeders. Some, such as The Consort (C. concors)

and Epione Underwing (C. epione), and the Myricaceae and heath feeders, apparently oviposit in litter. For these, put twigs and dried leaves of the foodplant at the bottom of the bag. Both adults and their eggs are sensitive to desiccation, which may be caused by prolonged periods of air-conditioning. Hold eggs outdoors over the winter months and then check daily for the hatchlings beginning about the time of budbreak. Wet the eggs and allow them to air dry once a month (it is best if temperatures are above freezing when this is done). (Eggs will desiccate if held in air-conditioned or heated rooms or selfdefrosting refrigerators without misting.*) Soaking the eggs and exposing them to high temperatures (but not exceeding 32° C, 90–95° F) sometimes helps stimulate hatching in the spring, especially for southern taxa. Some species of underwings are exceptional among moths in having a protracted period of eclosion: some Catocala eggs from a single female may hatch over a five- to seven-week period. Egg clutches of willow feeders can still be yielding first instars at the time the earliest-hatching individuals are nearing pupation; larvae and adults of the Sad Underwing (C. maestosa) from the same population will sometimes overlap. Willow and walnut feeders—whose hostplants have indeterminate growth—are among those that have a notably protracted emergence period. However, there is little risk to holding eggs in the refrigerator for short periods of time, e.g., if you fear eggs will hatch while you are away traveling or before suitable foliage is available locally, you can refrigerate the eggs for two to four weeks with little risk. Do wet and air dry them first.

*

Yellow-banded Underwing Catocala cerogama

Saskatchewan to Nova Scotia, south to northern Georgia and Arkansas. One generation with mature caterpillars from late May to early July. Sometimes common. Basswood (Tilia). Color highly variable, but always with minute black spots arranged in lines; high warted hump on A8.

! CATOCALA HEADS. Species are arranged alphabetically by foodplant group. The degree to which patterns vary within species is unknown. We have found them to be more constant among the hickory feeders than among the Rosaceae and Myricaceae feeders. ! continued overleaf

101

102 Underwing Moths Subfamily Erebinae: Tribe Catocalini ! CATOCALA HEADS. ! continued

Underwing Moths Subfamily Erebinae: Tribe Catocalini 103 ! CATOCALA HEADS. ! continued

104 Underwing Moths Subfamily Erebinae: Tribe Catocalini

Three-staff Underwing Catocala amestris ! recognition Lavender, black, and yellow with six fine black pinstripes down each side of body. Ground color lavender above level of spiracles, contrasting with canary-yellow pinacula and white to cream subventral area. Venter as in lower right. Head with black stripes and spots; each lobe capped with yellow above. Larva to 5 cm. Caterpillars of the Married Underwing (C. nuptialis)—an uncommon species of Upper Great Plains—are similar, but less boldly patterned and peppered with more black spots. Search for its caterpillar (page 106) on indigo bush (Amorpha fruticosa), especially along watercourses. ! occurrence Prairies, limestone glades, savannas, especially along swales and river courses; also pinelands in Southeast. Eastern North Dakota and Wisconsin to North Carolina, south to central Florida and Texas; absent from Northeast. One generation with mature caterpillars in April to June. Locally common. ! common foodplants Desert false indigo (Amorpha fruticosa) and leadplant (A. canescens); works citing locust are in error. ! remarks Our Amorpha-feeding underwings (see also pages 105–6) are nocturnal creatures that frequently rest off the foodplant by day, at least as they near maturity. Last instar Three-staff Underwings are striking insects and rank among the most handsome of North American caterpillars. To find late instars by day, search around the base of plants, examining the trunk as well as nearby stems and grass blades. Jim Wiker has had success locating Amorpha-feeding Catocala at night by lightly tapping branches where he is searching—evidently, startled caterpillars lift the anterior end of the body and thereby reveal their whereabouts. If branches are abruptly jarred, the caterpillars release their grip and fall to the ground; unlike most underwings, the caterpillar remains motionless. The belly of the Three-staff Underwing is marked in black and yellow (right), which likely serves as a flash coloration that disappears once a caterpillar rights itself. Such bold ventral coloration seems to function in a fashion analogous to the orange and black hindwings of the adults (which are concealed when the adult is at rest).

Underwing Moths Subfamily Erebinae: Tribe Catocalini 105 Legume-feeding Underwings (Catocala) Caterpillars tend to be smooth and lack rootlet hairs except for the Little Underwing (C. minuta), which is unrelated to the others.

Abbreviated Underwing Catocala abbreviatella

Manitoba to Illinois, south to Texas. Mature caterpillars from April to June. Locally common. Leadplant (Amorpha canescens). Pinstripes of nearly continuous brown lines (not composed of spots); little yellow in dorsal pinacula (compare with Whitney’s Underwing). See Borth and Barina (1991).

Magdalen Underwing Catocala illecta

Nebraska to Wisconsin and southern Ontario, east to Pittsburgh, south to western South Carolina and Georgia, Gulf states (excluding Florida), and Texas. Mature caterpillars in May into early June. Common east to Ohio. Honey locust (but evidently not water locust). Distinctively ringed in black and white.

The Betrothed Catocala innubens

South Dakota, southern Canada, and Connecticut, south to Florida and Texas; more common west of Appalachians, where its foodplant is native. Mature caterpillars in June and early July. Common. Honey locust. Rootlet setae absent; bulge (low ridge) over A5; often with saddle; venter with bright lemon yellow patch at base of prolegs; pale waxy forms occur.

106 Underwing Moths Subfamily Erebinae: Tribe Catocalini

Little Underwing Catocala minuta

South Dakota east to southern Michigan and Massachusetts, south to Florida and Texas. Mature caterpillars in May and early June. Locally common. Honey locusts, including water locust. Small, highly variable; white-capped, rounded ridge over A5; prolegs on A3 much reduced; rootlet setae present (these absent in The Betrothed, C. innubens).

Married Underwing Catocala nuptialis

North Dakota through Ohio River valley to Kentucky, south to Arkansas and northeastern Texas. Mature caterpillars in May and June. Locally common. Amorpha species, especially indigo bush or desert false indigo (A. fruticosa) along watercourses. Similar to Whitney’s Underwing but more boldly marked: e.g., more prominent yellow-orange pinacula, and pinstripes more spotted; venter with purple-black spots on all segments.

Whitney’s Underwing Catocala whitneyi

Manitoba to Ohio, south to Tennessee and Kansas. Mature caterpillars in May and June. Locally common. Leadplant (Amorpha canescens). Dorsal yellow pinacula smaller than those of C. nuptialis; pinstripes composed of black spots; bold spiracular stripe; head with yellow spot over each lobe. See Borth and Barina (1991).

Underwing Moths Subfamily Erebinae: Tribe Catocalini 107

Sweetfern Underwing Catocala antinympha ! recognition Exceptionally variable, medium-sized Catocala. Setal bases white. No saddle or raised humps. Most individuals brown and undistinguished above, but some handsomely patterned with rust and cream and contrasting pinstriping. Venter yellow with dark spots and lines. Larva to 5 cm. Middle instars more obviously pinstriped (lower right). Coloration is exceptionally variable in the Myricaceae-feeding underwings (see also page 109), so much so that we are unsure if we can reliably identify larval collections without associated foodplant information. C. antinympha is the only underwing that feeds on sweet fern. In our examples there tends to be greater contrast between adjacent stripes, the white dorsal pinacula tend to be smaller, and the venter is more lemon yellow than in related species. In addition, the black markings on the venter may be less pronounced: purple-black spots on A3 and A4 are dark, the one on A5 is paler, those on T3 and A6 are diffuse and brown, and the linear midventral spots on T1, T2, and A8 are reduced or absent. Other characters are given in Crumb (1956). Larva under 5.5 cm. ! occurrence Acidic, sandy communities with host: barrens, fields, roadside banks, powerline right-of-ways, coastal strand habitats; also inland wetlands with sweet gale. Western Ontario to Nova Scotia, south to New Jersey, Virginia (mountains), and Great Lakes states. Mature caterpillars from June to early July. Often locally common, especially eastward. ! common foodplants Sweet fern (Comptonia peregrina) and sweet gale (Myrica gale). ! remarks These bold caterpillars, which rest exposed on leaves and twigs in early and middle instars, are conspicuous during the day. Beating samples from the first three weeks of June routinely yield this beautiful insect. The caterpillar’s wriggle response is violent and sustained. The middle instar illustrated (right) has been attacked by a braconid wasp; within an hour the smoky wasp larva spun a whitish cocoon about its body, and remained attached to the caterpillar until it emerged. In the Great Lakes region (and presumably elsewhere) larvae use sweet gale as well as sweet fern.

108 Underwing Moths Subfamily Erebinae: Tribe Catocalini

Bayberry Underwing Catocala badia ! recognition Polymorphic. All forms with bright white setal bases. Head orange with vague longitudinal streaking; inverted black “swooping pterodactyl mark” sometimes present, with its “feet” nearly reaching top of triangle. The triangle with orange medial line and pale lines to either side. Yellow venter with dark spotting. Larva to 5 cm. Co-occurring with The Little Wife (C. muliercula) on bayberry in coastal New England and Long Island. The white spotting, lines, and pinstripes tend to be more extensive in its various forms; additionally, The Little Wife tends to have less brown pigmentation above the level of the spiracles (Larry Gall, pers. comm.). ! occurrence Heathlands, powerlines, shrublands, dunes and sand plains, and coastal strand communities from southern Maine to Long Island (historic records from northeastern New Jersey); if range of C. badia coelebs is included, west through Great Lake region. One generation with mature caterpillars from late June into July. Locally common. ! common foodplants Bayberry (Morella (= Myrica) pensylvanica) for C. b. badia, usually Myrica gale for subspecies C. b. coelebs. ! remarks Our larvae were beaten off bayberry as middle instars by Ben Williams. Two hours of beating spread over a two-week period yielded numerous early and middle instars but no late instars, leading Ben to wonder if larger larvae moved to the base of bayberry during the day. Myricaceae-feeding Catocala are extremely polymorphic in coloration. Barnes and McDunnough (1918) provide paintings of five different C. badia forms—no other underwing species was illustrated with multiple forms. The Bayberry Underwing is closely associated with Morella pensylvanica. In Connecticut it occurs somewhat farther inland than The Little Wife (C. muliercula). Forbes (1954) associated the Bayberry Underwing with “Myrica barrens,” and regarded it as common “in wet areas along the coast.” Gall and Hawks (2002) regard the Bayberry Underwing to be conspecific with The Old Maid (C. b. coelebs) (next page). The two are southern (C. badia) and northern (C. coelebs) segregates of a single entity, with each being somewhat tied to a different foodplant, the former to Morella pensylvanica and the latter to Myrica gale.

Underwing Moths Subfamily Erebinae: Tribe Catocalini 109 Gale- and Sweet Fern-feeding Underwings (Catocala) Coloration is exceptionally variable across the group, which includes the Sweetfern Underwing (Catocala antinympha); the Bayberry Underwing (C. badia) and its northern segregate, The Old Maid (C. badia coelebs); and The Little Wife (C. muliercula). All bear numerous fine stripes that are particularly conspicuous in middle instars; venters are yellow, and dorsal pinacula white. Hosts and ranges are important clues to identity.

The Old Maid

Catocala badia coelebs Ontario to Nova Scotia, south to New Hampshire, northern New York, and upper Great Lakes states. Mature caterpillars from June into early July. Locally common. Sweet gale and occasionally bayberry.

The Little Wife Catocala muliercula

Mostly near coast from southeastern Massachusetts to Florida and Texas; inland records to Kansas, western North Carolina, Pennsylvania, etc., presumably strays. Mature caterpillars from May (Florida) to July. Common along coast. Mostly wax myrtle (Morella (= Myrica) cerifera), but bayberry (M. pensylvanica) northward. Usually with more extensive white spotting and pinstriping than Bayberry Underwing (C. badia).

110 Underwing Moths Subfamily Erebinae: Tribe Catocalini

Epione Underwing Catocala epione ! recognition Smooth, medium-sized Catocala, alternately striped in blacks, browns, and grays, without warts or ridges; sometimes beige and less strongly patterned than other hickory-feeding underwing caterpillars. Some mature individuals take on a waxy bloom. Brown to charcoal subdorsal and spiracular stripes, often broken in midsegment. Subventral rootlet setae lacking. Dorsal setae borne from small whitened pinacula. Venter with lance-shaped black spots (these often more rounded in other hickory feeders). Top of head with prominent, black, upside down “W”; triangle streaked with reddish-brown lines. Larva to 6.5 cm. Closely allied to The Consort (C. consors) (page 118) whose venter bears a broad, continuous black stripe; in C. epione the stripe is composed of a series of spots as in other underwings. While they co-occur in the South, The Consort is only rarely, and not recently, encountered above the Mason-Dixon line. ! occurrence Dry woodlands from Wisconsin, southern Canada, and Maine to Florida and Texas. One generation with mature caterpillars in May and June over most of range. Common. ! common foodplants Hickories, such as mockernut, pignut, sand, and shagbark; but not bitternut (Carya cordiformis). Often on small trees. ! remarks Caterpillars perch on twigs, branches, and trunks by day, or less commonly, under shags. The Epione Underwing is among the most commonly encountered hickory-feeding Catocala at light and bait across much of its range. Larvae, which pass through five instars, grow rapidly and mature in late spring—it and The Consort (C. consors), are the first of our hickory feeders to be seen on the wing. The phenology, forewing coloration, large rounded (not flattened) eggs, and number of instars of these two underwings attest to an evolutionary origin separate from that of most of our other hickory-feeding underwings (e.g., those that follow).

Underwing Moths Subfamily Erebinae: Tribe Catocalini 111

Mourning Underwing Catocala flebilis ! recognition Gray bark mimic with dark saddle over A5 and A6 (which may be weakened or broken middorsally). Dorsal setae from pink-orange warts; those on A8 enlarged, and connected by low ridge edged in black. Dark spiracular stripe best developed rearward on abdomen, e.g., below saddle and on A8. Rootlet setae essentially absent, occasionally with 1–3 very short rudimentary setae per segment (visible with lens). Venter with rose to red areas, bounded with greenish flush, on T1–T3 and A3–A6; medial black spots on A1–A4 and A7–A8; A9 with small V-shaped black spot. Head with rays of white snowflake spotting extending to vertex; black mask stretching between eyes and wrapping around and extending through lateral eyes. Larva to 6 cm. ! occurrence Woodlands from Illinois and southern Michigan to Massachusetts, south to Georgia and Texas. One generation with mature caterpillars in early summer. Locally common at times; both its range and abundance have been increasing in New England over the past decade. ! common foodplants Pignut, shagbark, and other section Carya hickories. Captive larvae, especially late instars, will accept walnut. ! remarks Mourning Underwing caterpillars can be collected by searching under shags as well as in and along bark fissures on pignut and related hickories. Search from chest height down to within a few centimeters of the ground. In Connecticut we find caterpillars at the same time of the season that we encounter Gypsy Moth (Lymantria dispar) caterpillars resting on trunks by day. Burlap banding is effective for the collection of Catocala caterpillars that feed on tree species with smooth bark. It and other hickory-feeding underwings, if dislodged from their perch, will writhe violently for several seconds. Hickory-feeding underwings will accept and mature on older foliage than our oak- and Rosaceaefeeding Catocala, which are largely new leaf specialists. Robinson’s Underwing (C. robinsonii) (page 122) is especially tolerant of older leaves and is among the last of the eastern Catocala to mature, feeding well into July in most years. Catocala do not tolerate handling well—they are quick to bite and frequently regurgitate a clear yellow-green fluid. They are easy caterpillars to photograph, settling quickly if offered bark or a twig upon which to perch.

112 Underwing Moths Subfamily Erebinae: Tribe Catocalini

The Bride Catocala neogama ! recognition Large, mottled in charcoal and gray. Spiracular area raised, most noticeably along anterior abdominal segments, giving caterpillars a corrugated appearance from above; thoracic segments much flattened (at rest); A8 with posterior dorsal setae (D2) on backward-directed protuberances connected by ridge. Dorsum with pale intersegmental addorsal spots between segments A1–A5; frequently with dark saddle on A5 and A6. Dark spiracular stripe often interrupted by oblique pale patches. Head with broad, dark horizontal bar above antenna; often with coronal bar. Venter pinkish red with conspicuous purple-black spots on A1–A4, A7, and A8—those on A1 and A2 large and somewhat squarish; spot on A8 teardrop shaped to elongate, longer than broad. Rootlet setae nearly absent, except for few, very short setae towards anterior end of some abdominal segments. Larva to 6.5 cm. Similar to and often found side by side on walnut with the Sad Underwing (C. maestosa). In our examples of the Sad Underwing, the caterpillars tend to have more tan or brown in the ground color, a narrower black line above the antenna, narrow black lines running below the D2 seta, orange plates on A3–A6 prolegs, and the black ventral spots reduced in size and limited to A1–A4; also the red ventral coloration is more subdued, and A6 and A7 in particular lack the bright red of The Bride. ! occurrence Parks and yards, thickets, woodlands, forests from Wisconsin, southern Canada and Maine, south to Florida and Texas. One staggered generation with mature caterpillars from early June through mid-July in New Jersey. Locally common. ! common foodplants Walnut and butternut (Juglans). ! remarks Most of our larvae have come from the trunks of large trees, resting one to three meters above the ground in bark furrows. Caterpillars wriggle violently when handled—the spasmodic contortions continue seconds after larvae have been released. Some occasionally came to rest belly up, exposing the bright pink-red underbelly and purple-black spots, although the propensity, across the genus, is to right the body. The Bride has a protracted period of eclosion from the eggs; ova from a single female will hatch over a period of a month or more: larvae can be found well into July, after nearly all other Catocala species have pupated or are on the wing. The extended eclosion period of The Bride (and Sad Underwing, C. maestosa) are made possible by the phenology of the foodplant, walnut (Juglans), which produces new foliage throughout much of the growing season.

Underwing Moths Subfamily Erebinae: Tribe Catocalini 113

Obscure Underwing Catocala obscura ! recognition Finely mottled in browns, grays, and charcoal, and rather even in color; lacking strong humps or warting; somewhat flattened. Most striping subdued; densely peppered with black dots. Rootlet setae absent. Poorly developed black subdorsal stripe, best developed rearward of D2 pinacula on A1–A8. Pinacula pale, whitish to pinkish or yellow-orange. Low hump connecting D2 setae on A8, darkened across its posterior face. Weak to strong black spiracular stripe best expressed on T1 and A8. Head with black line above eyes extending nearly half way to vertex; coronal bars sometimes expressed. Venter with bright red coloration; strong black midventral spots on A1–A4, A7, and A8; dark spots on thorax and A5 and A6 essentially absent. Larva to 6 cm. Caterpillar of the Residua Underwing (C. residua) closely similar but with black line above eyes shorter, not extended rearward of last eye (stemma 1). ! occurrence Wisconsin, southern Canada, and Massachusetts to northern Georgia and Arkansas. One generation with mature caterpillars from June into July. Common. ! common foodplants Shagbark hickory; generally on larger trees. ! remarks Nearly 20 underwing species feed on the leaves of shagbark hickory in our region, with many locations supporting a dozen or so species. It makes one wonder if there is a limit to the number of species that can share a single host. Across southern New England, the Obscure and Residua Underwings are among the most commonly encountered underwings, both as caterpillars and as adults, under shags. In Connecticut, many of the caterpillars found under shags in midJune are those of the Residua Underwing; by late June and early July, they are largely replaced by caterpillars of the Obscure Underwing. Like other underwings, if the caterpillar is alarmed, it is quick to launch itself into the air. Disturbed caterpillars may thrash the head and thorax vigorously from side to side, three to four times in each direction. The action is violent; we guess it would shake loose most invertebrate enemies, and stun a fly or wasp if it were hit by a direct strike.

114 Underwing Moths Subfamily Erebinae: Tribe Catocalini

Oldwife Underwing Catocala palaeogama ! recognition Highly variable, ranging from gray-green to charcoal, but always with pale yellow venter embellished by handsome set of midventral red and purple-black spots (inset). Dorsal pinacula whitened on T2 and T3 but orange to pink or red over abdominal segments, these often encircled by pale spot. D2 setae borne from conical warts on A8. Often with pale patch above spiracle on A1 and narrow, black saddle over A5–A6. Gray to rusty red head with numerous fine orange-brown lines; no black line above antenna. Rootlet setae present, of variable lengths. Venter pale lemon yellow with red spots (T1–T3, A5–A9), purple-black spots (A1–A2), or both (A3–A4). Larva up to 7 cm. Closely allied to the more southern Tearful Underwing (C. lacrymosa) (page 119) with which it occasionally hybridizes. We are unsure of larval characters that will reliably separate the two. ! occurrence Woodlands and forests from South Dakota, southern Canada, and Maine to northern Georgia and Texas. One generation with mature caterpillars mostly in June and early July. Very common in Northeast. ! common foodplants Mockernut, pignut, shagbark, and probably other section Carya hickories, but not pecan and related section Apocarya hickories; several sources also list walnut, based on lab rearings. ! remarks Numbers of the Oldwife fluctuate considerably from year to year (see Sargent 1976). Catocala caterpillars are quick to strike at objects that touch their abdomens. The “bite” is utterly harmless to humans but it comes with such rapidity and force that we are inevitably startled. The strike is so fast that we are guessing the insect is using its head as a mace to knock away the intruder, and that the mandibles may not be engaging in a bite. This innate behavior has been exploited by at least one lineage of braconid wasps that lay their eggs inside the early instar. Upon reaching maturity the wasp exits its (still much alive) host, crawls under the rear of the larva’s body, and then spins its cocoon. Ironically, the caterpillar is bound to its sentinel position by the wasp’s silk, and will animatedly strike at objects that touch the rear of its body; in so doing, it is fated to protect the very parasitoid that doomed it to a protracted death. The individual shown in the image above has been attacked by a tachinid fly—the black spot on the side of the seventh abdominal segment is the breathing spiracle of a maggot that will soon kill it.

Underwing Moths Subfamily Erebinae: Tribe Catocalini 115

Serene Underwing Catocala serena ! recognition Brown, shiny, flattened, widened through abdominal segments; rootlet setae absent. Small, often pale, wart rearward of each spiracle on first eight abdominal segments. Dark spiracular stripe often present but sometimes weakening through midabdominal segments and rearward. Head with strong coronal bars and broad diffuse black bar above each antenna with short spurs that reach upward towards vertex and back beyond eyes. Venter pale, lacking dark medial spots common to many Catocala. Larva to 5.5 cm. The larvae of Angus’s, Habilis, and Robinson’s Underwings also are smooth, shiny, flattened, and thus similar in appearance. The Serene Underwing is the only one of these that lacks black spots on its venter; head patterns also distinguish the four from one another. ! occurrence Woodlands from Wisconsin, southern Canada, and Massachusetts, south to South Carolina, Mississippi, and Oklahoma; absent from Atlantic Coastal Plain and most of Piedmont. Mature caterpillars in late May and June. Locally common. ! common foodplants Shagbark hickory. ! remarks The Serene Underwing has been expanding its range northeastward. It first appeared in eastern Connecticut in the 1980s and now ranks among the most common middle-season species there. Darryl Willis, an underwing aficionado who has reared nearly all eastern species of the genus, refers to caterpillars of the Serene Underwing as the ugly ducklings of the Catocala world—indeed, they surpass many cutworms in their homeliness. Larval coloration in the hickory feeders is a clue to the caterpillar’s perching habits. Underwing caterpillars that are handsomely mottled in grays and browns, or have raised ridges, prominent dorsal warts, and/or bear abundant rootlet setae, commonly perch on limbs or the trunk; whereas those that are flattened, smooth, dark brown, and comparatively featureless (as is true of the Serene Underwing), position themselves under shags during the day. But even members of the former group take shelter under shags in late instars if and when such are available (especially during a molt, which is drawn out over two days in underwings). The Serene Underwing and other “flattened-type” Catocala that roost under shags are muscular caterpillars with a powerful and prolonged alarm response. The integument of prepupal Catocala often becomes shiny, and that of the Serene Underwing particularly so (right).

116 Underwing Moths Subfamily Erebinae: Tribe Catocalini

Widow Underwing Catocala vidua ! recognition Large, comparatively smooth, pale gray, with subdued striping and variously expressed saddle over A5 and A6; Forbes (1954) notes that some caterpillars have a very subtle greenish aspect. A8 without hump, sometimes with faint oblique line extending down from D2 pinaculum. Pinacula minute, somewhat tan to pinkish in our examples. Rootlet setae numerous, fine, of moderate length. Head with bold black line from mandible to above antenna, mostly ending or becoming appreciably paler rearward of stemmata; often with vague yellow to rusty spots (below each P setae) over each lobe. Venter frosty white with handsome wine to purple-black spots; typically wine red (without dark spots) on T1–T3, A5, and A6; purple-black spots on A1, A2, and A7–A8 (sometimes on A9), and both colors present on A3 and A4. Larva to 6.5 cm. Similar to some forms of Yellow-gray Underwing (Catocala retecta) and no doubt others—its black eye line extends half way to vertex. ! occurrence Woodlands and forests from Wisconsin, southern Ontario, and lowermost Maine (historic) to northern Florida and Texas. One generation with mature caterpillars from late May to July. Often common. ! common foodplants Hickory; most or all section Carya hickories (e.g., pignut, mockernut, sand, and shagbark). Walnut accepted by captive larvae, but has not been verified in field. ! remarks The Widow Underwing is a common and widely distributed hickory feeder. Typically there are seven larval instars, in part a reflection of the small eggs produced by females and the large size of the last instar. To a greater extent than many other hickory feeders, the Widow Underwing will use small trees, sometimes with chest diameters of only three to five centimeters. Although eggs hatch about the same time as other hickory feeders, the last instar is protracted, sometimes lasting a month in New Jersey. The larvae mature in summer, completing their development on mature foliage. While collection records indicate that this underwing was once common, it virtually disappeared from New England beginning around the 1960s. Evidently it began rebounding in the late 1980s, and since 1990 has been common in eastern and coastal New England. The ranges of several other members of the genus also appear to be in flux. Angus’s (C. angusi) and Robinson’s (C. robinsonii) Underwings have contracted southwestward out of the Northeast. Others such as Mourning (C. flebilis), Judith’s (C. judith), and Serene (C. serena) Underwings became more common and widespread than in past times—we are unaware of any New England records for the last two of this threesome prior to 1961 (e.g., Farquhar 1934), but hundreds since then (e.g., Sargent 1976).

Underwing Moths Subfamily Erebinae: Tribe Catocalini 117 Hickory-feeding Underwings (Catocala)

Agrippina Underwing Catocala agrippina

Kansas to Carolinas, south to Florida and Texas, but straying northward. Mature caterpillars in May and June. More common southward. Pecan, water pecan, and perhaps other Apocarya hickories. Rootlet setae present; A5 with low knob; D2 pinacula on A8 greatly enlarged; dorsal pinacula of abdomen red-orange.

Angus’s Underwing Catocala angusi

Missouri to southern Michigan, and formerly New Jersey and southeastern New York, to north Georgia and Texas. Mature caterpillars from June into July. Locally common. Shagbark hickory; also walnut in captivity. Shiny, D1 and D2 often from minute white spots, head with somewhat less black mottling than Habilis Underwing (C. habilis).

Brou’s Underwing Catocala atocala

Southern Illinois to Mississippi, west to Texas and Oklahoma; old records to Florida. Mature caterpillars in May and June. Generally scarce and local. Nutmeg hickory (C. myristiciformis). Mousy gray with redorange setal warts, those over A8 enlarged; long rootlet setae. See Gall et al. (2002) and account in Schweitzer et al. (2011).

118 Underwing Moths Subfamily Erebinae: Tribe Catocalini Hickory-feeding Underwings (Catocala)

The Consort Catocala consors

Formerly more widespread in East; now Kansas and Illinois to Georgia, south to central Florida and Texas. Mature caterpillars mostly in April (southward) to June. Uncommon. Hickory, at least black and scrub; sand hickory likely. Smooth, plump; one common form tan with pinstripes; head with thick black line framing face; venter bearing broad, continuous black stripe. See account in Schweitzer et al. (2011).

Dejected Underwing Catocala dejecta

Missouri, southern Michigan, and Massachusetts to northern Florida and Texas. Mature caterpillars mostly May to June. Locally common northward. Section Carya hickories. Rootlet setae long; face with transverse black line above antenna; raised pinkorange dorsal pinacula; often with A1 frosted; venter with intense scarlet patches on T1–T3 and A3–A6.

Inconsolable Underwing Catocala insolabilis

Mostly Iowa to Ohio, south to Florida and Texas (formerly east to coastal Connecticut); rare strays north to Ontario. Mature caterpillars from April (Florida) into June. Uncommon over most of range; locally common in Florida. Hickory, especially pignut in Florida. Pale, frosty blue-white to gray; red-orange head with dark bar above antennae but otherwise lacking strong black lines; saddle obscure to absent.

Underwing Moths Subfamily Erebinae: Tribe Catocalini 119 Habilis Underwing Catocala habilis

South Dakota, southern Canada, and southern Maine to Georgia (mountains) and Arkansas; absent from Atlantic Coastal Plain and much of Piedmont. Mature caterpillars from June into July. Common. Shagbark hickory. Black lateral stripe well developed; D2 often from white spot; head with considerable black patterning; dark brown midventral spots on A1–A4.

Judith’s Underwing Catocala judith

Wisconsin, southern Canada, and New Hampshire to Georgia and Oklahoma; absent from Atlantic Coastal Plain and much of Piedmont. Mature caterpillars in June over most of range. Locally common. Shagbark hickory. Nearly black above; head with enlarged black spot above antenna, running to labrum; midventral spots red-brown.

Tearful Underwing Catocala lacrymosa

Mostly Missouri to Delaware, south to central Florida and Texas, but straying northward. One generation with mature caterpillars from April (Florida) into June. Common to abundant southward. Section Carya hickories. Head heavily mottled but without black lines; venter yellow with purple-black spots on A1–A4 and red patches on T1–T3, A5–A8.

120 Underwing Moths Subfamily Erebinae: Tribe Catocalini Hickory-feeding Underwings (Catocala)

Yellow-fringed Underwing Catocala luctuosa

Mostly eastern Kansas to Ohio, south to Mississippi and Georgia. Mature caterpillars probably from June into July. Local and uncommon. Captive larvae accept shagbark, shellbark, and pignut hickory (Gall 1990a, Darryl Willis, pers. comm.). Orange to pink dorsal pinacula; head with broad black band from antenna to vertex; rootlet setae long; venter of A5 and A6 salmon to orange and without purple-black spots.

Sad Underwing Catocala maestosa

Resident from Kansas to southern New Jersey to Florida and Texas, recently spreading northward, e.g., into Pennsylvania. Mature caterpillars from late June into August in New Jersey. Common southward. Pecan, other section Apocarya hickories, and walnut. Large, somewhat corrugated, often with black line extending back from SD2 pinacula, head with thin black line over antenna (only), and red reduced ventrally (relative to The Bride, C. neogama).

The Penitent Catocala piatrix

South Dakota, southern Canada, southern Maine, to Florida and Texas (subspecies to California). Mature caterpillars from June to July in much of range. Locally common. Mostly black walnut, pecan and other section Apocarya hickories; Sam Jaffe has collected caterpillars on shagbark hickory. Fine pinstripes composed of minute brown or black spots; head orange-red, framed by blurry, black “W”; venter with dull yellow hues.

Underwing Moths Subfamily Erebinae: Tribe Catocalini 121 Clouded Underwing Catocala nebulosa

Mostly South Dakota, southern Ontario, and Long Island, south to Florida and Texas; rare northeast of Delaware. Mature caterpillars mostly in June. Uncommon to locally common. Bitternut hickory (Carya cordiformis); possibly also pecan. Dorsal pinacula white, low over thorax, pink-red and wartlike over abdomen, forming cones on A8; low ridge on A5.

Residua Underwing Catocala residua

Wisconsin, southern Canada, and southern Maine to Georgia (mountains) and Arkansas; absent from Atlantic Coastal Plain. Mature caterpillars from June into July. Common. Shagbark hickory. Rootlet setae absent; black line ending behind eyes; venter with purple-black spots on A1–A4, A7, A8, with salmon tint to A3–A6.

Yellow-gray Underwing Catocala retecta

Wisconsin, southern Canada, and Maine to Georgia and Texas; absent from much of Atlantic Coastal Plain and Piedmont south of Delaware. Mature caterpillars in June and July. Very common. Shagbark and other section Carya hickories. Rootlet setae; head with black lateral stripe reaching well behind eyes, nearly joining reddish coronal bar; midventral spot on A1 squarish.

122 Underwing Moths Subfamily Erebinae: Tribe Catocalini Hickory-feeding Underwings (Catocala)

Robinson’s Underwing Catocala robinsonii

Mostly Iowa to Ohio, south to Georgia and northeastern Oklahoma. While now extirpated in Northeast, was moderately common eastward to New Jersey and Connecticut from late 1800s into 1960s. Mature caterpillars in July; among latest-maturing Catocala. Locally common to abundant. Shagbark and perhaps shellbark hickories. Head with thick black frame, not as shiny as C. habilis.

Sappho Underwing Catocala sappho

Missouri to southwestern Virginia (mountains), south through Gulf states. Mature caterpillars from April (Florida) to June. Scarce outside of Florida and southern Appalachians. Pignut and probably other section Carya hickories. Greenish-white lichen mimic; rootlet setae short, sparse; head with black mask between eyes. See also Slotten (1992).

Youthful Underwing Catocala subnata

Southern Minnesota and southern Canada to Georgia (mountains) and Arkansas; absent from Atlantic Coastal Plain and much of Piedmont. Mature caterpillars mostly in June. Local and uncommon. Bitternut hickory. Large, smooth and shiny; venter with yellow flush; head relatively unmarked, without black lines.

Ulalume Underwing Catocala ulalume

Mostly Missouri, Illinois, and southern Delaware to northern Florida and Texas. Mature caterpillars mostly in June. Usually uncommon to rare. Hickory; Slotten (1992) records mockernut hickory (Carya alba (= C. tomentosa)). Rootlet setae present; orangebrown coronal bar and black line from antenna through eyes; dorsal pinacula pink-orange.

Underwing Moths Subfamily Erebinae: Tribe Catocalini 123

Darling Underwing Catocala cara ! recognition Distinguished from related willow feeders by its buff to orange, forward-projecting cranial lobes (inset). Ground color orange to dark brown to gray or charcoal; striping vague. A5 with rounded middorsal lobe that may share coloration of cranial lobes. Dorsal pinacula red-orange, conspicuously wartlike over anterior abdominal segments. Head with thin black and orange lateral line ending above antenna; line largely broken across top of head; elongate, unpigmented patch to either side of triangle. Pinkish rootlet setae slender, elongate. Venter pinkish with wide oval purple-black spots on A3 and A4, smaller lens-shaped spot on A6; other segments lacking defined dark spots. Larva to 8 cm. Treasured Underwing (C. carissima) (lower right), a sister species of the Darling Underwing, occurs locally from Missouri and Kentucky to Delaware, south into Florida and eastern Texas. Its ground color is consistently paler than that of the Darling Underwing. Caterpillars of other related willow-feeding Catocala, all with proportionately smaller cranial bulges, are figured on pages 124–7. ! occurrence Wetlands and riparian corridors from Wisconsin, southern Canada, and New Hampshire, south to Florida and Texas. One generation with mature caterpillars in June and July over much of East. Sometimes common. ! common foodplants Willow, including nonnative species like weeping willow (Salix babylonica); especially black willow (S. nigra) southward, but apparently mostly on arborescent, smooth-leaved willows. ! remarks Darling Underwing caterpillars are among the more commonly collected, Salicaceae-feeding Catocala, in part because the females oviposit on black willow, which often grows as a small tree or shrub that is easily searched. As noted in the introductory box, eclosion from a single clutch of eggs is greatly staggered in our willow feeders, with siblings sometimes differing by as much as six to seven weeks in their “birth” dates. Presumably the indeterminate growth of willow shoots—and access to new foliage season long—makes such a bethedging strategy possible. Adults of C. cara and related willow feeders perch on the underside of ledges and fallen trees. Caves, porch ceilings, and culverts will sometimes yield individuals from late July through September. The willow feeders as a group are more likely to be seen at bait than at light, perhaps because they are active late in the season when baiting seems to work well across a spectrum of moths.

124 Underwing Moths Subfamily Erebinae: Tribe Catocalini

Pink Underwing Catocala concumbens [Sleepy Underwing] ! recognition Charcoal, dark gray, brown, tan, or reddish brown with slight hump on A5 that often bears some orange. Many forms with rusty band around posterior of A5. Pink-orange D2 pinacula pimplelike, partially encircled by black ring. Pinkish to orangish brows of head capsule bulging but not as much as in Darling Underwing (C. cara). Triangle with three faint black vertical lines. Rootlet setae, well separated, mostly of single length, shorter than height of spiracle. Venter red-pink with strong purple-black spots (see lower right). Larva to 7 cm. ! occurrence Wetland and riparian areas from Saskatchewan to Nova Scotia, south to Georgia (mountains) and Nebraska. One generation with mature caterpillars mostly in July. Common northward. ! common foodplants Willow and poplar. ! remarks This is the most commonly encountered willow- and poplarfeeding underwing in beating samples from southern Canada, the Northeast, and Great Lakes states, primarily because females oviposit on shrubby willow species that are shunned by other eastern Salicaceae-feeding Catocala (which tend to be tree feeders). In our rearing containers, late instars moved off foliage during daylight hours, suggesting that caterpillars commonly leave the host to pass daylight hours in cover at the base of their host. The figured individual was recovered after a deliberate effort to sample roadside willows the day after a torrential summer squall had filled roadside ditches and swales. We surmised that the flood waters would force larvae up onto stems and shoots where they could be effectively located by eye or sampled with a beating sheet—a half hour spent beating yielded a wealth of caterpillars, but only one Catocala. Salicaceae-feeding Catocala tend to be encountered in low numbers and require some effort to secure as larvae. As noted under the Three-staff Underwing (C. amestris), the bellies of Catocala are frequently boldly colored, particularly relative to the cryptic colors of the dorsal and lateral body surfaces. Presumably, the bellies serve in the same capacity as the bright hindwings of the adults, providing would-be predators with flash coloration that immediately disappears once an individual rights itself.

Underwing Moths Subfamily Erebinae: Tribe Catocalini 125

White Underwing Catocala relicta ! recognition Large, pale, and smooth with dark saddle over A5–A6 that reaches to prolegs on both segments. Low ridge over posterior half of A5. Rootlet setae long and numerous. Head with dark band over top that gives way to black mottling at about level of triangle; each lobe with white to orange spot above. Midventral black spots present on T2, T3, and A3–A7. Larva to 7.5 cm. Larva of Grote’s Underwing (C. grotiana), a Rocky Mountain species that feeds on aspen, is similarly colored. On the few that we have seen, the ridge on A5 was less swollen and marked with orange, and the black pigment on the head was confined principally to a black line across the top of each lobe. ! occurrence Forests across Canada, south in East to northern New Jersey, Virginia (mountains), and Missouri. One generation with mature caterpillars from late June to early July. Common northward. ! common foodplants Aspen and other poplars; also willow. ! remarks The host ranges for most of the eastern willow- and poplar-feeding underwings are poorly known, presumably because females preferentially lay their eggs on older trees that are largely inaccessible from the ground. Someone (or even better, a group) committed to a burlap skirt survey could contribute greatly to our knowledge of host utilization patterns across the Salicaceae feeders. The White Underwing’s pale coloration, both as a larva and as an adult, and its distribution suggest that the species is closely tied to quaking aspen. Natural selection has finetuned the larval coloration and behavior of each instar (Sargent 1976). The shiny greenish-brown first instar usually rests along a leaf edge. The pin-striped second instar rests on the underside of leaves, often with its body stretched out along a midrib. The third instar, reddish brown with white pinstripes, extends itself along the length of a petiole (which in aspens and many poplars is also reddish brown). The last three instars leave the foliage after feeding, perching along twigs and branches. As it grows, the larva rests on progressively larger twigs and branches. The final instar—the striking insect figured above—descends to the lower trunk and frequently seeks shelter in leaf litter at the base of its host. The markings on the caterpillar’s head are a good match for a leaf or bud scar.

126 Underwing Moths Subfamily Erebinae: Tribe Catocalini Willow- and Poplar-feeding Underwings (Catocala)

The Sweetheart Catocala amatrix

Idaho to Nova Scotia, south in East to northern Florida and Texas. Mature caterpillars in June and July. Sometimes common. Poplar (including cottonwoods and Lombardy), perhaps willows. Often with dark subdorsal and spiracular stripes; low middorsal wart on A5; lobes projecting somewhat.

Briseis Underwing Catocala briseis

Transcontinental in Canada, south in East to Long Island (rarely) and Great Lakes states. Mature caterpillars in June and early July. Common northward. Willow and probably poplar. Black stripes on head connecting; A5 with rusty to purplish hump.

Joined Underwing Catocala junctura

Western New York and Pennsylvania (as strays), south to Arkansas, westward to Pacific. Mature caterpillars in June and early July. Common westward. Willow and poplar. Dorsal pinacula orange; D2 pinacula modestly differentiated; low medial hump on A5; orange projecting brows on head.

Marbled Underwing Catocala marmorata

Ohio River valley to Delaware, south to South Carolina and Arkansas (formerly to southern Connecticut). Mature caterpillars in June and July. Generally rare. Principally swamp cottonwood (Populus heterophylla) but eastern cottonwood and willow suspected. Head coloration diagnostic (page 102)—see also Peacock and Gall (2000) and Schweitzer et al. (2011).

Underwing Moths Subfamily Erebinae: Tribe Catocalini 127 Meske’s Underwing Catocala meskei

Alberta to Maine to New Jersey (formerly), Ohio, and Arkansas in East. Mature caterpillars in June and July. Generally uncommon, rare eastward. Primarily poplars: mostly cottonwoods, including Lombardy in towns. Venter brownish, not pale as in other underwings, contrasting with grayish dorsum.

Mother Underwing Catocala parta

Saskatchewan to Nova Scotia, south to Virginia (mountains), Kentucky, and Kansas. Mature caterpillars in June and July. Uncommon to locally common. Poplar and willow. Tan to gray-brown, vaguely banded; black lines on head not meeting over top; lobes only modestly projecting.

Semirelict Underwing Catocala semirelicta

Transcontinental in Canada, south in East into northern New England and upper Great Lakes states. Mature caterpillars in June and July. Uncommon southward. Poplar, especially balsam poplar eastward. Gray with dark saddle extending from dorsal ridge on A5 to proleg on A6.

Once-married Underwing Catocala unijuga

Manitoba to Newfoundland, south in East to Virginia and Missouri; mostly as strays further south. Mature caterpillars in June and July. Common northward. Poplar, especially quaking aspen. Low reddish hump on A5; head smoothly rounded, not “browed,” and lower portion of face with three black lines.

128 Underwing Moths Subfamily Erebinae: Tribe Catocalini

Girlfriend Underwing Catocala amica ! recognition Small, pale gray to charcoal. Dorsal setal bases mostly black over thorax, becoming red-orange rearward. A8 humped with conical orange pinacula. Body broadest at A5; this segment bears low dorsal hump. Variously developed dark saddle over A5–A6 that reaches to proleg on both segments. Larva to 4 cm. The Girlfriend Underwing is part of a species complex that includes the Steely Underwing (C. lineella), Barrens Underwing (C. jair), and others whose taxonomy is still under study. Identifications for members of this group should be based on adults. ! occurrence Mostly woodlands and forests but also barrens from South Dakota, southern Canada, and Maine to Florida and Texas. One generation with mature caterpillars from May to early July. Very common. ! common foodplants Oak, especially white and related oaks, but also scrub, live, and others. ! remarks Members of this species group are among our most common underwings and usually make up a significant fraction of the adults seen at light, bait, or while tapping. Likewise, they were the most frequently encountered underwing caterpillar under the burlap bands that we used to monitor Gypsy Moth larval populations in the Appalachian foothills near Goshen, Virginia (Wagner et al. 1995). Over much of the East, the Girlfriend and related species begin emerging in late June or early July—a week or two later than our other small, yellow-hindwinged Catocala—and fly through much of the summer. In the Northeast, well-worn adults will be encountered through the month of September. Unlike other eastern oak feeders, caterpillars of this group can mature on fully expanded, somewhat hardened leaves.

Underwing Moths Subfamily Erebinae: Tribe Catocalini 129

Scarlet Underwing Catocala coccinata ! recognition Distinguished by prominent middorsal horn on A5 and paired dorsal horns on A8. Ground color gray to brown and charcoal with subdued striping. Ochre, rust, or black arching patch connecting prolegs on A5 and A6. Dorsal setae from orange pimplelike pinacula, especially D2 pinacula on abdominal segments. Rootlet setae numerous, especially rearward of A6 proleg. Head with thin, black (and sometimes orange) line framing face; short, black, vertical line above antenna, inwardly edged by white spot. Venter with purple-black spots on A3–A6; subtle, small, vague, purple-black spots on T2 and T3; spots essentially absent on A1, A2, and A7–A9 (lower right). Larva to 5 cm. Hoary-edged Underwing (C. herodias) is the only other oak-feeding Catocala with a middorsal horn on A5, although the horn is much smaller. Over much of its range, Hoaryedged Underwing is a species of barrens and balds and ridge tops with scrub oak (page 131). ! occurrence Woodlands and mixed forests from southern Canada to Florida and Texas. One generation with mature caterpillars from late May into June over much of East. Common some years, at least northward. ! common foodplants Oak; many records from red oaks northward (e.g., black, red, scarlet), but also post, southern red, and other oaks southward. ! remarks The caterpillars are specialists on young foliage, always maturing before oak leaves harden—it is among the first of our Catocala to complete development each spring. In the Northeast, the Scarlet Underwing is most common in dry oak and oak-pine woodlands. Like many Catocala, its population numbers vary considerably from year to year.

130 Underwing Moths Subfamily Erebinae: Tribe Catocalini

Ilia Underwing Catocala ilia ! recognition Exceptionally variable but always with rather corrugated texture: D2 pinacula connected by low ridge on A1, A2, A5, and A8. Recognizable by its large size, bright pink-red underside, and row of elongate, midventral black spots on A1–A8 (inset). Most forms with thin dark shading (saddle) over A5–A6. Body rather roughened but without conspicuous warts or humps; A5 with raised dorsal ridge. Black spiracular stripe commonly evident rearward of A7. Head with thick black band enclosing face. Subventral rootlet setae short and stubby. Larva to 7.5 cm. Caterpillars of its sister species, the Shadowy Underwing (C. umbrosa) (lower right), may not always be separable: in our examples, the ground color tends to be paler. The Shadowy Underwing is a species of dry, sandy woodlands, savannas, and barrens from at least Oklahoma (and probably North Dakota) to southeastern Massachusetts, south to Florida and west to Texas, but absent from many states and areas within this range. It is associated with scrub oak in the Northeast. ! occurrence Barrens, woodlands, and forests from southern Canada to Florida and Texas. One generation with mature caterpillars mostly from April (Gulf states) into early June. Very common. ! common foodplants Oak; most of our records from red oaks (subgenus Erythrobalanus), but also from white oaks. ! remarks Although adults are often underrepresented at lights, this is one of the East’s most commonly encountered underwings at bait and during the daytime. While most caterpillars are gray and essentially bark colored, occasionally one will encounter spectacular lichen mimics (page 98). Such striking forms are sometimes “discovered” nestled among foliose lichens, suggesting that the larvae are able to assess their immediate surroundings as they descend trunks to shelter for the day. Similar “lichen forms” also occur (rarely) in the genus Zale. Despite their large size, Ilia caterpillars are among the first underwings to mature each spring—the larvae require young leaves, and straggle and fail if fed older foliage.

Underwing Moths Subfamily Erebinae: Tribe Catocalini 131 Oak-feeding Underwings (Catocala)

Connubial Underwing Catocala connubialis

Wisconsin to Nova Scotia, south to Florida and Texas. Mature caterpillars in April (southward) to June. Rare to locally common. Oak. Obscure saddle and striping; numerous (pinkish) rootlet setae; often with white patch over each lobe.

Delilah Underwing Catocala delilah

At least formerly, Kansas and Illinois, south to Florida and Texas. Mature caterpillars mostly from April to early June. Rare east of Texas except in Florida. Oak. A5 with low hump; dorsal pinacula raised, reddish orange; head with black cheek line edged anteriorly with orange-yellow; ours with angled black line above triangle. See account in Schweitzer et al. (2011).

Hoary-edged Underwing Catocala herodias

Nominate subspecies (C. h. herodias) in Oklahoma and Texas; Gerhard’s Underwing (C. h. gerhardi) from eastern New York and Massachusetts, south to New Jersey; also mountains of North Carolina and Virginia. Mature caterpillars mostly in late May. Locally common in barrens. Scrub and blackjack oak primarily, and perhaps other red oaks westward. A5 with low, rounded horn; venter rosy red with purple-black spots on A3–A6. See account in Schweitzer et al. (2011).

132 Underwing Moths Subfamily Erebinae: Tribe Catocalini Oak-feeding Underwings (Catocala)

Barrens Underwing Catocala jair

Long Island, south mostly along coastal plain to south central Florida, west to Oklahoma and Texas. Mature caterpillars from April (southward) to June. Generally rare except in oak scrub and barrens of Long Island, Florida, and Texas. Scrubby oaks, especially blackjack, scrub, and turkey. Usually pale gray with weak saddle.

Coastal Plain Underwing Catocala near lineella

Arkansas to New Hampshire, south (mostly along coastal plain) to Florida and Texas; absent from Ohio River valley. Mature caterpillars from April (southward) to June. Common and widespread in south; local northward. Laurel, scrub, water, and other oaks. Gray with weak saddle. Compare with others in Girlfriend Underwing (C. amica) group (page 128).

Steely Underwing Catocala lineella

Southern Canada, south to Florida and Texas. Mature caterpillars from April (southward) to June. Common. Oak, especially those of red and black oak group. Gray with dark saddle. Sometimes distinguishable from Girlfriend Underwing (C. amica) by its grayer color, especially anteriorly.

Underwing Moths Subfamily Erebinae: Tribe Catocalini 133 Little Nymph

Catocala micronympha Minnesota to southern Maine, south to Florida and Texas. Mature caterpillars in April (southward) to June. Common and widespread. Oak; especially those of white oak group. Head framed with black above; dark saddle on A5–A6; dorsal diamonds often present; A1 and A4 commonly with faint, pale subdorsal patch. Very close to Similar Underwing (C. similis).

Similar Underwing Catocala similis

Minnesota, southern Canada, and Maine, south to Florida and Texas. Mature caterpillars in April (southward) to June. Locally common. Oak, often scrub and blackjack. Similar to Little Nymph (C. micronympha). We are unaware of characters that reliably separate the two; Forbes (1954) suggests C. similis tends to be darker with more obscure patterning. Associated with sandy, dry oak woodlands with small oaks; more restricted in habitat than Little Nymph.

134 Underwing Moths Subfamily Erebinae: Tribe Catocalini

Andromeda Underwing Catocala andromedae ! recognition Pinkish gray to tan, medium-sized Catocala with closely set striping. A5 with medial dorsal wart. Second dorsal seta (D2) borne from minute red-orange wart on T2–A9; warts on thoracic segments about half size as those along abdomen. Approximately eight dark wavy pinstripes along each side of body. Rootlet setae mostly of one length, numerous. Venter with wine-purple spots on T1–A5, those on A3 and A4 largest. Head conspicuously patterned with two prominent red-orange stripes: one passes along outer reach of triangle and the second extends to the eyes; frons white with medial dark line. Larva to 5 cm. Larva closely similar to Louise’s Underwing (C. louiseae) (page 135). Also resembling Graceful Underwing (C. gracilis), which tends to have darker ground coloration and lack the double striping of the Andromeda Underwing; in addition, the head capsule markings are more discrete, and there is often a dark spot over each lobe in the Graceful Underwing. ! occurrence Swamps, bottomlands, thickets, woodlands, and forests from Great Lakes states to Maine, south to Florida and Texas. One generation with mature caterpillars from late April (Florida) through June. Common. ! common foodplants Blueberry and pink azalea or (certain) other azaleas. ! remarks While blueberry is the principal host across much of its range, the Andromeda Underwing also uses Rhododendron. In late May 2004, Bryan Connolly was able to locate several larvae on pinxter-flower or pink azalea (Rhododendron periclymenoides), near Mansfield Reservoir in eastern Connecticut, by searching stems in the vicinity of shoots with feeding damage. Caterpillars had fed from the leaf tip back towards the petiole, leaving ragged edges that revealed their whereabouts. By day, the larvae rested head down, appressed to a stem. Rhododendrons are also used in the Smokies. In the Southeast, sparkleberry or farkleberry (Vaccinium arboreum) is the most frequently used host. The species name suggests that the moth was once reared from one of the heaths formerly known as Andromeda, most likely a Leucothoe or Lyonia species.

Underwing Moths Subfamily Erebinae: Tribe Catocalini 135 Blueberry and Other Hosts (Catocala)

Graceful Underwing Catocala gracilis

Michigan to Maine, south to Florida and Texas. One generation with mature caterpillars from late April (Florida) to June. Common eastward. Blueberry, especially highbush, and swamp doghobble (Eubotrys racemosa) (DFS). Sparse rootlet setae; A5 with low rounded hump; head with strong black lateral line; wine-red ventral spots on T1–A8.

Sordid Underwing Catocala sordida

Alberta to Nova Scotia, south to Georgia and Texas, but sporadic and local southward, e.g., mostly absent from Gulf region, Piedmont, and parts of Midwest. One generation with mature caterpillars from late May into early July. Common northward. Blueberry, especially lowbush (Vaccinium angustifolium and V. pallidum) over much of East. Rootlet setae absent.

Louise’s Underwing Catocala louiseae

Principally Florida to Oklahoma and Texas, but absent from many portions of range; isolated records northward to Tennessee and North Carolina. Mature caterpillars mostly in April. Locally common in parts of Florida. Farkleberry (Vaccinium arboreum) in the Southeast, and perhaps other blueberries elsewhere. Larval appearance close to Andromeda Underwing, C. andromedae.

136 Underwing Moths Subfamily Erebinae: Tribe Catocalini

Hawthorn Underwing Catocala crataegi ! recognition Smooth, mottled brown or smoky to mouse gray, with obsolescent middorsal stripe sometimes edged with darker pigment. Horn absent (usually) or present, but in latter case as dark rounded knob (see Remarks). Charcoal to russet saddle over A5–A6 present or absent. D1 pinacula minute and obscure; D2 small, dark, and comparatively undifferentiated relative to other Catocala. Rootlet setae varying from short and sparse to long and tightly packed (as in Wonderful Underwing, C. mira); many rootlet setae branched, sometimes pinkish. Head with discrete black band framing face (page 103); lobes on head not projecting forward. Venter with faint bluish cast; strongest purplish spots on A1–A4, especially those on A3 and A4; spots more poorly expressed on T2–T3 and A5–A9. Larva to 5 cm. The Hawthorn Underwing is a member of a poorly understood complex of species (or subspecies) in the Southeast. There is disagreement as to whether some populations found mostly along southern rivers are this species. ! occurrence Overgrown fields, woodlands, and bottomland forests from Manitoba to Nova Scotia, south to Florida and Texas. One generation with mature caterpillars in May and June over most of range. Locally common. ! common foodplants Apple and hawthorn. ! remarks The caterpillar(s) illustrated as C. crataegi by Barnes and McDunnough (1918) is either atypical or erroneously identified and has led to confusion in the literature (e.g., Forbes 1954, Schweitzer 1982a). In the 20 or so wild-collected larvae that we have seen, none possessed a high horn on A5 as in Barnes and McDunnough’s drawing on Plate XII (Fig. 18). Our caterpillars lacked a horn, or possessed a dark, rounded, knoblike projection (smaller in size than is typical for Wonderful Underwing, C. mira). On Plate X, Barnes and McDunnough show a dark face with protruding lobes. In our material, the face of the Hawthorn Underwing tends to be flatter and not darkly pigmented save for the black line that frames the face. The Wonderful Underwing is the most common member of this complex that has projecting lobes and a dark face (and a high horn on A5).

Underwing Moths Subfamily Erebinae: Tribe Catocalini 137

Wonderful Underwing Catocala mira ! recognition Tan, brown, gray, or charcoal to frosty blue-white, usually with middorsal horn over A5. Horn of last instar tends to be curved or sinuate, ranging from mere nub to prominent thornlike projection, sometimes orange or red, at least in part. D2 pinacula orange-red and knoblike on abdominal segments. Darkened saddle over A5–A6 present or absent. Occasionally bearing broad, dark middorsal stripe as seen in forms of several other Rosaceae-feeding Catocala. Face with upper portion of each lobe often orange-red and projecting forward (lower right); head of some individuals almost completely blackened. Rootlet setae long and branched, often with pinkish tint. Venter frosty with bluish or greenish hue and strong purple-black midventral spots; strongest purplish spots on A3 and A4, those on A1 and A2 only slightly smaller, spots vague on T2–T3 and A5–A9. Larva to 5 cm. At least a dozen Catocala feed on hawthorn in eastern North America—pay close attention to the features given above, and rear to adulthood any individuals in which the characters leave room for doubt. ! occurrence Thickets and woodlands from Manitoba to Quebec, south to Florida and Texas; more common westward. One generation with mature caterpillars from late April (Florida) through June. Locally common. ! common foodplants Hawthorn; less frequently reported from crabapple and plum (e.g., Prunus angustifolium) (Schweitzer 1982a). ! remarks The size and the coloration of the horn on A5 are variable in both the Wonderful and Hawthorn (C. crataegi) Underwings (page 136). The Wonderful Underwing is often the most common Catocala on hawthorn, especially from Ohio westward and across the South. In the Northeast, the Hawthorn Underwing is more common, in part because of the widespread abundance of domesticated apple. Across much of central Florida, where ten or more Catocala species share the spring flush foliage of hawthorns, the Wonderful and Alabama Underwings (C. alabamae) are the two most frequently encountered.

138 Underwing Moths Subfamily Erebinae: Tribe Catocalini

Ultronia Underwing Catocala ultronia ! recognition Medium-sized, gray to reddish brown or even charcoal with pinkish venter, and elongate horn on A5. Often with pale subdorsal patch anterior to horn and russet lateral patch connecting prolegs on A5 and A6 (lower right). Dorsal pinacula raised, especially over A8; upper portions orange-red. A8 with ridge connecting D2 pinacula; oblique black to rust line running from lower edge of ridge to vicinity of spiracle. Rootlet setae numerous; setae and venter of body pink with strong spots; those on A1–A4 purple-black. Horn elongate, and often somewhat sinuate. “Face” framed by solid black band. Larva to 5.5 cm. ! occurrence Late successional fields, woodlands, and forests from southern Canada to Florida and Texas. One generation with mature caterpillars from April to July. Very common. ! common foodplants Principally cherry and plum (Prunus) species, but also reported from apple and hawthorn. Black cherry and beach plum are favorites in Northeast. Our neonates refused domestic apple and crabapple. ! remarks The best way to locate underwing caterpillars is to search the bark of twigs and branches back from new spring growth. Use feeding damage, especially that of the last two instars, to anchor your larval searches. In some underwings, the caterpillar will be located near the leaves on which they have fed, e.g., the Yellow-banded Underwing (C. cerogama) rests on the underside of branches just back from the recent year’s growth. Most, including the Ultronia Underwing, will descend meters from the feeding site; look for caterpillars perched along the trunk. Start with smaller trees, which will have smoother bark, because larvae will be easier to locate. In contrast to most other Catocala that feed on members of the rose family (Rosaceae), egg hatch in the Ultronia Underwing is staggered—caterpillars can be found into July northward. Presumably the species takes advantage of the flushes of new growth produced by cherry and its relatives over the summer.

Underwing Moths Subfamily Erebinae: Tribe Catocalini 139 Rose Family-feeding Underwings (Catocala) The species in this plate and the next are difficult to distinguish both as adults and larvae. Moreover, the number of species involved is still unresolved. Horn development can vary, e.g., see Alabama and Hawthorn Underwings (C. alabamae and C. crataegi, respectively). Until reliable species characters are known, identifications should be based on adults, which can be challenging enough. Some of the hawthorn feeders use just one or a few Crataegus species at a given locality.

Clinton’s Underwing Catocala clintonii

Manitoba to southern Canada, south to Florida and Texas; formerly New York. Absent from New England. Mature caterpillars from late March to early June. Locally common. Apple, cherry, hawthorn, and plum (especially Chickasaw plum); often eating flowers. Chunky in aspect. Active early in season, maturing two to three weeks before other small underwings.

Sweet Underwing Catocala dulciola

Michigan to Ohio, south to North Carolina and Missouri, but absent from much of this range; formerly east to New York. Mature caterpillars in May. Generally uncommon and local. Hawthorn, closely associated with Crataegus margarettiae in Great Lakes region; one from wild plum (Prunus americana) (John Peacock record). Head pale, ringed with solid black; horn usually absent; pale middorsal stripe edged with black. See Schweitzer et al. (2011).

Woody Underwing Catocala grynea

South Dakota, southern Canada, and Maine, to Florida and Texas. Mature caterpillars in May and June. Common. Apple, crabapple, plum, and related plants, including European mountain ash. A5 with prominent horn; underside waxy white (not red); upper portion of head orange or white.

140 Underwing Moths Subfamily Erebinae: Tribe Catocalini Rose Family-feeding Underwings (Catocala)

Chokeberry Underwing Catocala praeclara

At least Wisconsin to Nova Scotia to South Carolina and Tennessee. Mature caterpillars in May and June. Locally common. Most records from chokeberry and serviceberry; also hawthorn. D1 pinacula inconspicuous; pink to red D2 pinacula minute on T2–A7; low ridge on A8 with conical D2 pinacula; horn short; head rimmed with red or black; some individuals with broad, brown middorsal stripe. Hawthorn-feeding populations from Ohio (shown here to left) thought by some to represent another species.

Precious Underwing Catocala pretiosa

Plymouth County, Massachusetts and southern New Jersey; subspecies C. p. texarkana from North Carolina south to northern Florida, and west to Missouri and Texas. Mature caterpillars in April (Florida) to June (Massachusetts). Locally common from New Jersey southward. Red chokeberry; less often apple, crabapple, hawthorns (especially for C. p. texarkana), and serviceberry. Short, rearprojecting horn; long, branched rootlet setae; lobes of head pale above; compare with Charming Underwing (C. blandula). See account in Schweitzer et al. (2011).

Underwing Moths Subfamily Erebinae: Tribe Catocalini 141 Alabama Underwing Catocala alabamae

Missouri and Illinois to New Jersey, south to Florida and Texas (our range includes C. alabamae titania). Mature caterpillars from April to May. Locally common. Hawthorn. Pale; A5 horn very reduced to elongate; lobes protruding; dense rootlet setae; dark to rusty arc connecting A5 and A6 prolegs.

Charming Underwing Catocala blandula

As currently understood, Alberta to Nova Scotia, south to Florida and Texas; however, coastal plain populations from North Carolina to Florida and Texas may represent a distinct entity. Mature caterpillars April to June. Common northward. Apple, crabapple, and hawthorn; oviposition observed on serviceberry (Larry Gall, pers. comm.). In our figures: warted dorsal pinacula; without horn on A5; saddle russet to light brown; head with pinkish raised lobes. Broad middorsal stripe (right) sometimes present. Compare with Precious Underwing (C. pretiosa). See also Muller (1981).

142 Underwing Moths Subfamily Erebinae: Tribe Catocalini Rose Family-feeding Underwings (Catocala)

Grisatra Underwing Catocala grisatra

Eastern North Carolina to northern Florida. Mature caterpillars in late spring, after other hawthorn feeders. Hawthorn, especially yellowleaf hawthorn (Crataegus flava) (Jeff Slotten, pers. comm.). Local and rarely encountered. Highly variable; we are not aware of diagnostic characters. See account in Schweitzer et al. (2011).

Lincoln Underwing Catocala lincolnana

North Carolina south through Gulf states, west to Arkansas and Texas. Mature caterpillars in April or May. Uncommon. Hawthorns, especially parsley hawthorn (Crataegus marshallii), also littlehip hawthorn (C. spathulata), growing in river bottoms (Jeff Slotten, pers. comm.). Larvae mature very early in season. See account in Schweitzer et al. (2011).

Miranda Underwing Catocala miranda

Southeastern Ohio to northeastern Pennsylvania (but historic north of Virginia), south to Florida panhandle and Mississippi; mostly in southern mountains and northern Florida, absent from Atlantic coast and most of Piedmont. Mature caterpillars from late April to mid-June. Rare over most of range. Hawthorn, especially littlehip (Crataegus spathulata) and green (C. viridis) in Florida (Jeff Slotten, pers. comm.). Note even color, horn shape, rootlet setae, and dark head with slightly protruding lobes. See account in Schweitzer et al. (2011).

Orba Underwing Catocala orba

Coastal Carolinas south to northern Florida, west across extreme northern parts of Georgia and Alabama to Arkansas and Texas. Mature caterpillars in April or May. Local, usually uncommon. Parsley hawthorn (Crataegus marshallii) and probably other lowland hawthorns (Jeff Slotten, pers. comm.). Head extensively blackened, and green cast to body.

Lesser Underwings Subfamily Erebinae: Tribe Melipotini 143

Lesser Underwings

Family Erebidae: Subfamily Erebinae: Tribe Melipotini Most of our 56 species are western. Eastern melipotines tend to occur in woods, scrublands, and other dry to xeric habitats. Two genera, Drasteria and Melipotis, contain 80% of the North American fauna. Significantly, four melipotines, the Black-dotted Brown (Cissusa spadix), Common Oak Moth (Phoberia atomaris), The Darter (Melipotis acontioides), and Indomitable Melipotis (Melipotis indomita), constitute more than half of the owlets in this book known to defoliate trees. It is unclear what aspects of melipotine life history or physiological makeup allow their population numbers to explode and temporarily break free from their local natural enemy complexes, and enjoy such extraordinary abundance. Characters given in Crumb (1956) are diagnostic: the labial palpus is less than half the length of the spinneret; stemma 4 and 5 are closely situated with the distance separating them about equal to the diameter of either stemma; and the V seta on A1 and A2 is replaced by a sclerotized ring (inset). The midventer is sometimes marked with purplish lines and spots (though faint in Cissusa and Phoberia). Females insert the eggs in clusters into bark crevices. Foodplants are mostly trees,

! Ventral setae are replaced by rings in melipotines !

shrubs, and low perennial plants. Like underwings (Catocala), melipotines are muscular creatures that range widely in search of suitable foliage. By day, late instars shelter on or near the ground, in bark fissures, or under loose bark, and then ascend to find foliage after nightfall. Development can be very rapid with the larval stage lasting less than three weeks. A sparse cocoon is spun underground or in duff. Pupae overwinter.

Blueberry Drasteria Drasteria occulta

Maine to New Jersey and Pennsylvania. Mature caterpillars in June and July (northward). Local. Lowbush blueberry. Occasional pest of cultivated blueberries in Maine (Forbes 1954). Image of middle instar.

Barrens Oak Moth Phoberia ingenua

Core range along Atlantic Coastal Plain from Long Island southward to Florida, and from Colorado and Missouri to Texas, but also Wisconsin and isolated populations in Pennsylvania and Albany, New York. April (Gulf states) to June on spring foliage. Locally common to abundant. Oak. Compare with Common Oak Moth (next page). Additional details in next account.

144 Lesser Underwings Subfamily Erebinae: Tribe Melipotini

Common Oak Moth Phoberia atomaris ! recognition Variegated pale to deep chocolate brown, smooth, shiny, somewhat flattened with tapered posterior. Beaded or wavy, black subdorsal stripe best expressed on abdominal segments. Obscure middorsal “stripe” outwardly edged with wavy addorsal stripe that may form diamond-shaped spots over abdominal segments. Abdominal proleg on A3 about third size of that on A4, which in turn is half size of those on A5 and A6; all with 16–22 crochets. Dark spiracular stripe, which includes black spiracles, prominent rearward. Often with whitish spot above and behind spiracle (visible with hand lens), with that on A7 frequently enlarged. Head with extensive mottling, especially notable to either side of paler triangle; many individuals with white comma-shaped spot over each lobe. Larva to 4 cm. Barrens Oak Moth (P. ingenua), a closely related species mostly of sandy pine oak barrens, tends to have a less pronounced subdorsal stripe, darker spiracular stripe, and often bears a waxy bloom over the body (Lafontaine et al. 2008) (page 143). Caterpillars of the Black-dotted Brown (Cissusa spadix) are closely similar in coloration (see opposite). ! occurrence Barrens, woodlands, and forest edges from Wisconsin, southern Canada, and Massachusetts, south to Florida and Texas. One generation with mature caterpillars in late spring. Very common. ! common foodplants Oak. ! remarks The eggs are laid in bark fissures. Though abundant in oak woodlands across much of the East, caterpillars of the Common Oak Moth are not apt to be encountered unless you search foliage by night or closely examine bark in late spring. It is one of the most common caterpillars under burlap bands used to census Gypsy Moth caterpillars in late May and early June (Wagner et al. 1995). Populations occasionally outbreak and cause local and regional defoliation events—Leucobalanus (white oaks) appear to be the most severely affected (e.g., see account of defoliation event in Ross County, Ohio in 2002, http://www.dnr.state.oh.us/DNN/health/oakmoth/tabid/5214/Default. aspx). The muscular caterpillars are quite arboreal. DLW once left the lid ajar on a Common Oak Moth rearing container in his garage and upon returning later found two larvae upside-down on the ceiling. When mildly alarmed the caterpillar plays dead or draws the head under the body. Agitated caterpillars hurl themselves from their perch and forcibly writhe and snap their bodies about. The pupa overwinters in soil.

Lesser Underwings Subfamily Erebinae: Tribe Melipotini 145

Black-dotted Brown Cissusa spadix ! recognition Elongate, somewhat flattened, smooth, shiny, with barklike coloration and patterning. Dorsum darkened between subdorsal stripes. Wavy middorsal stripe (when present) most pronounced over A8 and A9. Supraspiracular area with broad pale “stripe” made up of four thin, wavy lines. Black spiracular stripe edged with fine white line above and below; black spiracles touch along lower edge. Often with white spot above and posterior to spiracle on abdominal segments; spot on A7 largest. Proleg on A3 about a third size of those that follow, with 16–22 crochets (proleg more reduced than that of Merry Melipotis, M. jucunda, a related oak feeder). Prothoracic shield dark and shiny, essentially black where addorsal stripe passes through the plate. Head with broad dark line over each lobe and with heavy mottling. Larva to 4.5 cm. ! occurrence Barrens and woodlands from Missouri to Ohio and New Jersey to Florida and Texas; strays seen northward to Wisconsin, southern Canada, and Massachusetts. One generation with mature caterpillars in late spring. Locally common southward. ! common foodplants Oaks, including red, white, and live; evidently also hickory and others during outbreaks. ! remarks This is a remarkably active caterpillar capable of covering great distances quickly—presumably the larva ranges widely in search of young foliage. Both thoracic and abdominal prolegs are proportionately long. The muscular caterpillar is quick to release its grip and launch itself from its perch. When disturbed, the larva folds the head under the abdomen and tucks the head between the prolegs on A5 and A6—a habit shared with many other erebines. We reared Black-dotted Brown caterpillars on young red and white oak foliage. The two figured individuals, reared in crowded quarters, may be appreciably darker than those that normally occur in the wild—dark outbreak phenotypes occur in a kindred melipotine, the Common Oak Moth (Phoberia atomaris). Cissusa overwinter as pupae belowground or in duff. An expanded species account, noting a widespread outbreak in Georgia in the spring of 2010, is posted on the website associated with this guide.

146 Lesser Underwings Subfamily Erebinae: Tribe Melipotini

melipotises Tribe Melipotini: Melipotis

This genus is best represented in Florida, Texas, and especially the American Southwest. Several migrate or stray long distances—no doubt, some of the dozen species recorded from the United States represent nonresident vagrants from the Caribbean and Central America. Head patterning, microscopic sculpturing of the integument, spotting along the venter, and armoring of the anal plate can be useful in species identification. Markings are highly variable—keep this in mind when using our images and descriptions. All are striped, mostly in browns and other earth tones, and somewhat flattened, especially rearward. A few have forms with bright orange patches. To what extent dark or brightly colored forms are inducible (i.e., represent crowding or outbreak phenotypes) is unstudied. Janzen’s figures of Costa Rican melipotis amply demonstrate the variability that can be seen within a single species (http://janzen.sas.upenn.edu).

By day, late instars shelter under bark flaps, in litter, or even under their own frass during outbreaks. All are strong, active climbers capable of roaming many meters when seeking out suitable food. Pupation occurs in loose soil or duff in a slight cocoon. Most melipotis feed on legumes, although the Merry Melipotis (M. jucunda) uses willow and oak, in addition to various legumes. We have had trouble getting wild-collected melipotis caterpillars to accept salads of legumes—whether this is because their diets are more specialized than is generally appreciated or because the caterpillars prefer foliage of a certain age is not known. This is a group where a dedicated individual could make significant contributions: several of our species have yet to be reared and the life histories of all are incompletely known. Moreover, such information could prove to be of economic significance given that two Melipotis are defoliators. Adults are strongly attracted to both light and bait, and many can be flushed in the daytime.

The Darter

Melipotis acontioides Occasionally caterpillar bears two bright supraspiracular stripes as in individual shown here. Typically pattern comprised of subtle earth tones. A full species account appears opposite.

Bookend Melipotis Melipotis cellaris

Florida westward through American Southwest, but straying northward to Minnesota. Continuous broods throughout growing season. Common southward. Acacia and presumably related legumes.

Lesser Underwings Subfamily Erebinae: Tribe Melipotini 147

The Darter Melipotis acontioides ! recognition Highly variable, striped caterpillar that ranges from brightly colored to rather muddied and undistinguished. A8 with low ridge over posterior half. Proleg on A3 half-sized. Broad addorsal stripes flank broad middorsal stripe that in many forms flares outward into crude diamond-shaped spots, especially over A2–A6; middorsal stripe best expressed over A7–A9; obscured over thorax. Some individuals with two orange supraspiracular stripes (page 146). D2 and L1 pinacula often embedded in small white spot on first seven abdominal segments. Venter pale with dark spots on A1–A4 and A7–A9, with spot on A8 elongate and much narrowed anteriorly. Head heavily patterned with brown and prominent brown-black patches to either side of triangle (lower right). Mature larva to 3.5 cm. ! occurrence Mostly Florida and Texas in our region, westward to California and south to Argentina. Continuously brooded with mature caterpillars throughout growing season. Common to occasionally abundant. ! common foodplants Royal poinciana (Delonix regia), various Parkinsonia, and Poeppigia. ! remarks A principal host in southern Florida appears to be the royal poinciana tree, an ornamental native to Madagascar. Confined larvae, held by Doug Caldwell over a two-day period, refused Lysiloma and Acacia. In the fall of 2006, caterpillars of The Darter defoliated mature royal poinciana trees in the Naples area (Collier County) of south Florida. By day the caterpillars abandoned foliage and descended as much as 15 m out of canopy foliage to shelter along root flares, in nearby vegetation, or under duff, rocks, sloughed bark, and surface litter. During outbreaks, larvae will even take refuge beneath their own accumulated feculae; in the case mentioned above, the feculae reached depths of two to three centimeters! Approximately 20 minutes after sunset, the Naples population “erupted out of the ground and raced up the trunk to seek out foliage” (Doug Caldwell, in litt.). One homeowner described the mass movement as “a river of caterpillars that moved up the trunks at dusk.” Caterpillars may leave an unsightly, dark (fluid) stain on houses and other structures in the vicinity of infested trees— we do not know if these stains are left by all larvae, prepupae evacuating their gut, or only by those succumbing to starvation and disease. The caterpillar wriggles violently when disturbed, hurling itself from its perch. Well-fed larvae also regurgitate a sticky green vomit if handled roughly. Pupation occurs belowground in an earthen cell, thinly lined with silk.

148 Lesser Underwings Subfamily Erebinae: Tribe Melipotini

Indomitable Melipotis Melipotis indomita ! recognition Smooth, dark, striped, with minute prolegs on A3 and reduced prolegs on A4. Dorsum with numerous, alternating, fine wavy stripes; body pale below dark spiracular stripe. Spiracular stripe darkest about spiracles, often with white spot above and posterior to each spiracle. Many forms with orange flush above prolegs on A3–A6. Venter waxy green to yellowish and unpatterned. Integument with minute cobblestonelike plates (under high magnifications). Head dark reddish brown to almost black with obscure patterning. Dark anal plate thickened and roughened. Larva to 4 cm. Other legume-feeding Melipotis occur in Texas; determinations should be based on adults. ! occurrence Deserts, scrub, and thickets. American Southwest; thought to be resident east to Oklahoma and Texas, but commonly straying into northern Great Plains states and infrequently to Connecticut. Now established in Hawaii. Multiple generations in southern Texas with mature caterpillars throughout growing season. Very common southward. ! common foodplants Mesquite. ! remarks Regard our description as provisional given the extreme variability in coloration exhibited by Melipotis and related genera. For example, Crumb’s (1956) larvae were “uniformly pale” below the spiracles, lacking both the pale (subspiracular) stripe and the diffuse orange spots above the prolegs that we mention above. At rest the caterpillar is somewhat cutwormlike: stout with a broadly rounded rump. Michael Singer wrote to us about a widespread outbreak in southern Arizona in 2003, “You could see large stands of defoliated mesquites from the road all along the west slope of the Santa Rita Mountains during August. I saw the same thing in some other mountain ranges. Initially, I was perplexed because there were no caterpillars on the mesquites—I figured they had finished up. But when we went out blacklighting and poked around with a flashlight, we found dozens of Melipotis caterpillars on each tree!” To what extent outbreaks in the Southwest are linked to those years with captures of adults far out of range (e.g., from the Great Lakes states and New England) is unstudied. Note the three tachinid eggs on the left side of T2 in the inset.

Lesser Underwings Subfamily Erebinae: Tribe Melipotini 149

Merry Melipotis Melipotis jucunda ! recognition Smooth, striped in gray, brown, and other earth tones. Proleg on A3 about half-sized; that on A4 about three-quarter-sized with 24–34 crochets. Subdorsal and spiracular stripes darkest; former darkened along its lower (outer) margin. Head with dark coronal bars with inward bulge or separate spot near apex of triangle; lower portion of bar narrowing, reaching to antenna (lower right). Anal plate peach or tan, roughened. Venter with maroon stripe and vague purplish midventral spots on A1 and A2, larger spots on A3 and A4, and purplish line on A8. Larva to 3.5 cm. ! occurrence Scrub and thickets southward; sandy areas and other xeric openings in New Jersey (at northern extent of range). Missouri to western Long Island, south to Florida and Texas, but absent from many areas northward; straying to and occasionally breeding as far north as Canada. Three broods in New Jersey, multiple generations in Deep South with mature caterpillars throughout growing season. Common southward. ! common foodplants Hosts include catclaw blackbead (Pithecellobium unguis-cati) (Crumb 1956), oak (Ferguson 1975), and willow. We reared one ex ova cohort on willow that refused both young and mature foliage of willow oak (Quercus phellos), and found wild larvae feeding on turkey oak (Q. laevis) in North Carolina. ! remarks During a hot July period in southern New Jersey, most caterpillars completed the larval stage in 17–19 nights on a diet of young willow foliage. Late instars remove themselves from the foliage before settling for the day. Presumably in the wild they hide along the trunk, and more frequently, beneath strips of bark or in leaf litter. Their smooth body is somewhat cutwormlike, and lacks the complex texturing of erebines that rest exposed on bark. Like cutworms, they simply drop from their purchase and feign death when alarmed, but, being erebines, wriggle spasmodically and hurl themselves from their purchase if handled roughly. Disturbed caterpillars curl the head back and hold it under the abdomen. Ours were quick to produce a sticky green vomit. Females are not difficult to flush during the day and readily oviposit if held in a paper bag or jar with willow leaves and bark. Pupation occurs belowground or in duff in a weak cocoon.

150 Lesser Underwings Subfamily Erebinae: Tribe Melipotini

Figure-seven Drasteria Drasteria grandirena ! recognition Pale red to gray-brown with vague striping and “W”s over dorsum. Proleg on A3 absent and that on A4 minute. Body widest at A4 and A5, then tapering noticeably to A10. Head with complex patterning (inset). Setae short above; longer along sides and subventer. Larva to 4.5 cm. ! occurrence Woodlands and open forests from Wisconsin to Nova Scotia, south to Florida and Texas. At least a partial second generation as far north as Connecticut; two full generations southward with mature caterpillars from May to September. Locally common. ! common foodplants Witch hazel. ! remarks Caterpillars of the Figure-seven Drasteria feed on new leaves. Larvae chew holes in the lamina from the underside of the blade, leaving the leaf edges intact. Like some Catocala, the caterpillar throws its fecal pellets by rapidly flicking the abdomen to one side. We know of only one collection (in 2010) of this species in New Jersey in the past 30 years, even though the host is still common. We wonder if excessive grazing by deer, especially of basal sprouts, and a more limited supply of second-flush leaves, might have played a role in the species’ decline. The pupa overwinters.

Lesser Underwings Subfamily Erebinae: Tribe Melipotini 151

False Heather Drasteria Drasteria graphica atlantica [Atlantic Graphic Moth]

! recognition Smooth, elongate, nearly cylindrical, striped caterpillar

without ridges or warts. Mostly chocolate above subdorsal stripe and pale below. Dorsal abdominal segments with pale subdorsal line above supraspiracular-subdorsal stripe. Chocolate spiracular stripe includes the small black spiracles. Dark, shiny head capsule crossed by four prominent white lines that continue forward from thoracic (addorsal, subdorsal, supraspiracular, and subspiracular) stripes. Prothoracic shield somewhat differentiated from rest of thorax. Larva to 3 cm. ! occurrence Barrens, dunes, heathlands, and coastal strand communities from southeastern New Hampshire to Accomack County, Virginia (see below); also reported from Great Lakes region. Two generations north to at least Connecticut with mature caterpillars from June to early September. Locally common. ! common foodplants Mountain goldenheather (Hudsonia montana) and wooly beachheather (H. tomentosa); also beach pinweed (Lechea maritima) in Connecticut (all Cistaceae). ! remarks The description Forbes (1954) attributed to Abbot appears to belong to another species: “Green and yellow incisures and a broad yellow stigmatal line. Head orange.” The brown caterpillars of the False Heather Drasteria are cryptic and difficult to locate. DFS found larvae by carefully inspecting stems of Hudsonia—some were detected by touch before they were seen. The nominate D. graphica graphica is a southern moth that occurs along the coastal plain north to Wilmington, North Carolina, where only a single brood occurs. Adults are on the wing from the last week in March through April. The more northern double-brooded entity that we treat here may be a distinct species: Forbes (1954) identified several wing pattern characters that can be used to separate the two moths. The pupa overwinters in all Drasteria.

152 Grass Loopers Subfamily Erebinae: Tribe Euclidiini

Grass Loopers

Family Erebidae: Subfamily Erebinae: Tribe Euclidiini About 16 species occur in our region. The ventral setae on A1, A2, and A7 are conical, shorter than those on intervening segments (Crumb 1956); A3 proleg and sometimes that on A4 absent; SD1 sometimes hairlike on A9. While most erebines (and erebids more generally) feed on woody plants, euclidiines tend to be grass and/or legume feeders. We have reared most of the species treated here on lotus and clover.

Forage Looper

Caenurgina erechtea [Common Grass Moth] Southern Canada to Florida and Texas. Multiple generations with mature caterpillars from May onward. Locally very common. Clover and grasses.

Vetch Looper

Caenurgia chloropha Southern Michigan south to Florida and Texas, but only as a stray northward (e.g., Michigan and New Jersey). Two or more generations with mature caterpillars from May onward. Common southward. Vetch and other legumes.

Long-winged Doryodes Doryodes grandipennis

Prince Edward Island to North Carolina. Multiple generations with mature caterpillars from spring into fall. Common along coast. Smooth cordgrass (Spartina alterniflora). Southward the Long-winged Doryodes is replaced by D. spadaria, which also feeds on Spartina. Other species, including one associated with wiregrass (D. bistrialis), occur from North Carolina to Texas. Application of names across this genus is uncertain.

Grass Loopers Subfamily Erebinae: Tribe Euclidiini 153 Toothed Somberwing Euclidia cuspidea

Western Canada to New Brunswick, south to Florida and northern Arkansas. Probably one principal generation with one or more partial summer generations southward. Associated with open sandy places in New Jersey. Locally common. Clover and presumably other legumes (such as bush clover and wild indigo) and grasses. Small proleg on A4.

Yellow Mocis Mocis disseverans

North Carolina to Arkansas, south to Florida and Texas. Multiple generations with mature larvae throughout growing season. Common. Grasses.

Common Ptichodis Ptichodis herbarum

Missouri, Ohio, and New York, south to Florida and Texas; northern records represent strays—yearround range unclarified. Two or more generations southward with mature larvae from May onward. Locally common. Bush clover (Lespedeza) according to Franclemont in Forbes (1954).

DYAR’S RULE Harrison Dyar (1890) noted that head capsule widths differed proportionally between the successive instars of many butterflies and moths. In most Lepidoptera this ratio falls between 1.25 and 1.4, with the ratio remaining more or less constant across all instars of a given species. Hence, if the ratio for a particular owlet with six instars were 1.3 and first instars had an average head capsule width of 1 mm, then Dyar’s Rule predicts the head widths for the five successive instars would be approximately 1.3 mm, 1.7 mm, 2.2 mm, 2.9 mm, and 3.7 mm. This rule has great utility in circumscribing insect life histories, and applies across many insect orders and varied anatomical features. For example, if one had first instar head capsule widths and just a single final instar head capsule, e.g., that recovered from a pupal crypt or cocoon, it would be possible to infer the number of intercalated instars.

154 Grass Loopers Subfamily Erebinae: Tribe Euclidiini

Clover Looper Caenurgina crassiuscula [Range Grass Moth] ! recognition Coloration and patterning highly variable; often tan to pale brown, striped. Prolegs absent on A3 and A4. Pale stripe below spiracle, a composite of two pale outer stripes and beige inner stripe that runs forward to antenna. Thin, pale middorsal stripe edged with tan to reddish-brown stripes that in turn are outwardly edged with prominent white addorsal stripes. All stripes extending onto head (upper inset). Larva to 4 cm. Caterpillars of Forage Looper (C. erechtea) and Vetch Looper (Caenurgia chloropha) (both figured on page 152) are similarly marked. Characters given by Forbes (1954) for the separation of the three species are contradicted by our images. The Toothed Somberwing (Euclidia cuspidea) co-occurs with this trio over much of the East but is much more localized. Its caterpillar, striped in a similar fashion, is best recognized by the presence of reduced prolegs on A4. Larval variation within this group of loopers is considerable—identifications are best based on adults. ! occurrence Pastures, fields, and roadsides from northern Canada to Florida and Texas. Multiple generations over much of East with mature caterpillars from April onward. Abundant. ! common foodplants Legumes evidently preferred, but also grasses. ! remarks This caterpillar turns up in sweep samples from fields and meadows in the summer and fall. It and related loopers can be especially common in nighttime surveys. Alarmed caterpillars will roll the head under the abdomen (lower inset). The Clover Looper is sometimes reported as a pest of legumes. The pupa, heavily dusted in purplish-white wax, overwinters in a sparse cocoon in leaf litter.

Grass Loopers Subfamily Erebinae: Tribe Euclidiini 155

Striped Grass Looper Mocis latipes [Small Mocis] ! recognition Mocis caterpillars can be recognized by the intersegmental dorsal black spots between A1–A2 and A2–A3 (concealed in body fold when larva is at rest), conspicuously striped trunk, and absence of prolegs on A3 and A4. Ground color often yellow-brown with broad chocolate striping along subdorsum and subventer; wide, yellow-brown lateral area often divided by narrow, dark brown stripe that runs through spiracle on T1 to A8. Pinacula inconspicuous; mostly whitish, often with minute adjacent gray-black spot. Head orange to tan brown with numerous fine stripes and contrasting white lines: addorsal line flaring to either side of triangle, subdorsal line running to whitened antenna, and lateral line running below eyes to antenna. Larva to 4.5 cm. Four mocis occur in our region; all are closely related and not readily identifiable as larvae—determinations should be based on adults. A preliminary key to the four Floridian species was authored by Ogunwolu and Habeck (1979). The Withered Mocis (M. marcida) and Texas Mocis (M. texana) are discussed on pages 156 and 157. The caterpillar of the Yellow Mocis (M. disseverans), described by Crumb (1956), is figured on page 153. To complicate matters further, Mocis caterpillars are easily confused with those of Caenurgia, Caenurgina, and Ptichodis. ! occurrence Fields, grasslands, mesquite scrub, and other open, grassy habitats from Missouri to North Carolina southward; adults stray north to southern Canada in fall. Multiple generations with mature caterpillars from late May onward. Continuously brooded in southern Florida and Texas. Common. ! common foodplants Grasses, including corn, rice, sorghum, sugarcane, as well as lawn, turf, and pasture grasses. Occasional pest in Deep South (and especially so in Caribbean and Central America). ! remarks A detailed life history of the Striped Grass Looper was published by Reinhert (1975). Population numbers build through the growing season and the moth may become common by September. Some Japanese mocis eat legumes in addition to grasses (Sugi 1987) and thus share the unusual and seemingly incongruous diet choices of Euclidea, Caenurgina, and other euclidiines. Mocis overwinter as pupae, although in the Deep South caterpillars can be found year-round. Prepupal caterpillars sometimes fashion their cocoons in a folded grass blade, occasionally well above the ground—watch for doubled-over grass blades that have been silked together. Mocis adults come to both floral (e.g., phenylacetaldehyde) and sweet baits (Landolt 1995, Meagher and Mislevy 2005).

156 Grass Loopers Subfamily Erebinae: Tribe Euclidiini

Withered Mocis Mocis marcida ! recognition Similar to Striped Grass Looper. Diagnostic characters not yet known. Pale or tan to yellow-brown, with darker brown (chocolate) addorsal and subspiracular stripes. Larger stripes dissected by multiple, fine, reddish to brown, wavy stripes. Chocolate subventral stripe contrasting with whitish lower edge of lateral stripe. Larva to 4.5 cm. Middle instars colored much like the above, but with black intersegmental spots lacking or less conspicuous (inset). ! occurrence Fields, marshes, grasslands, and other open, grassy habitats from coastal North Carolina, south to Florida and west to Texas. Multiple generations with mature caterpillars from late May onward. Locally common. ! common foodplants Grasses. ! remarks Mocis, like many grass-feeding caterpillars, have large heads with powerful grinding mandibles—the teeth are very short and the grinding area comparatively enormous (line drawing). Mocis caterpillars are capable of feeding on robust grasses such as sugarcane and St. Augustine grass, and are occasionally pests of the former. Females readily lay eggs if held for a night or two in a container with a selection of grasses. Unlike most erebine larvae, Mocis do not wriggle when accosted, but simply drop to the ground and remain motionless—a habit they share with other grass feeders—their coloration is well suited to blend in with dead, withered grass blades. The caterpillars can be common in beating and sweep samples from tall grasses. They frequently co-occur with wainscots (Leucania).

Grass Loopers Subfamily Erebinae: Tribe Euclidiini 157

Texas Mocis Mocis texana ! recognition Gray to yellow-brown, or sometimes reddish brown with numerous, closely set stripes that run length of body and spill over head. Ground color often darkened between addorsal and subdorsal stripes and above thoracic prolegs. White pinacula especially obvious over dorsum. Larva to 5.5 cm. In the Striped Grass Looper (M. latipes) ground color paler, pinacula are white or same as ground color, and not as conspicuous; subdorsal spots between A1–A2 and A2– A3 less pronounced (Forbes 1954). ! occurrence Fields, marshes, grasslands, and other open, grassy habitats from Minnesota to Martha’s Vineyard, south to Florida and Texas. Three generations in New Jersey and Ohio with last instars mostly from June to October; additional broods in Deep South. Often common. ! common foodplants Grasses, including crabgrass; DFS also records Panicum virgatum, Andropogon virginicus, and Spartina patens in New Jersey. See also Remarks. ! remarks This is the only mocis that commonly breeds and overwinters in most of our coverage area. The subdorsal black spots on the anterior abdominal segments (exposed in the inset) are normally concealed. When disturbed, older larvae flash these spots, and if bothered further, flip themselves to the ground. (Less developed black spots occur in Caenurgina larvae.) Collection records and our observations suggest that the Texas Mocis is a breeding resident north at least to Ohio, central Connecticut, and southeast Massachusetts. The rounded fecal pellets are surprisingly small given the caterpillar’s large body size. In captivity our caterpillars wove grass blades, all running lengthwise, into the wall of their cocoons; in the wild, cocoons can be found by watching for folded-over blades or places where two or more overlapping blades have been silked together. Brief life histories of three mocis, including the Texas Mocis, were published by Ogunwolu and Habeck (1975).

158 Grass Loopers Subfamily Erebinae: Tribe Euclidiini

Black Bit Moth Celiptera frustulum ! recognition Striped with pale lateral abdominal patches. Prolegs on A3 absent and those on A4 very reduced. D2 pinacula on T2 and T3 embedded in black spot. Thin middorsal stripe sandwiched between pale addorsal stripes. Some forms with one or more pale lateral patches: these may include an oblique patch extending back from leg on T3; large pale patch on A4 with subspiracular ray extending back to above proleg on A6; and vague patch on A8–A10. Head conspicuously striped, mostly light tan with dark subdorsal stripes that extend to antennae. Pinacula minute, white, not appreciably raised. Larva to 4 cm. Two or three zales occur on locust in the East: these often have more distinctively patterned heads with a spot on each lobe, as opposed to, in the words of Forbes (1954), a “confusedly striate” patterning. The dorsum of A8 is without ridging or warts in the Black Bit Moth but is invariably raised in zales, especially about the posterior dorsal (D2) seta. ! occurrence Shrubby fields and woodlands from Wisconsin, southern Ontario, and Connecticut, south to Florida and Texas. Two generations over much of East with mature caterpillars from June onward. Locally common. ! common foodplants Our larvae and cohort reared by Franclemont accepted black locust; natural hosts unrecorded (see Remarks). ! remarks The natural hosts of the Black Bit Moth are unknown. While the species is commonly encountered in the vicinity of locust, adults also turn up where Robinia is absent, suggesting that the moth uses other hosts or that the adults are vagile. In New Jersey adults of the Black Bit Moth are usually seen in the vicinity of bush clovers (Lespedeza). Our description is based largely on images taken of a single larva. Forbes (1954) notes that coloration is variable. Overwinters as a pupa.

Chocolates and Kin Subfamily Erebinae: Tribe Poaphilini 159

Chocolates and Kin Family Erebidae: Subfamily Erebinae: Tribe Poaphilini

Lafontaine and Schmidt (2010) recognize the tribe based on similarities in mitochondrial DNA and male genitalia. The North American fauna comprises six genera and 22 species, nearly all of which had been previously placed in the Euclidiini. The tribe is currently under study by JBS. As in the previous tribe, the head capsule is always striped, and larvae sometimes have a waxy bloom. Larval diets are specialized. Most are multiple brooded and overwinter as pupae.

Erasa Chocolate Argyrostrotis erasa

North Carolina to Florida, west to Texas. One principal generation with smaller facultative broods, and with mature caterpillars from April to October. Leatherwood or titi (Cyrilla racemiflora). Locally common. Middle instar (upper) and last instar (lower). Waxy bloom in latter not always present.

Yellow-lined Chocolate Argyrostrotis flavistriaria

North Carolina to Florida, west to Texas. Multiple generations with mature caterpillars from April to October. Leatherwood or titi (Cyrilla racemiflora). Locally common. Coloration highly variable.

160 Chocolates and Kin Subfamily Erebinae: Tribe Poaphilini

Short-lined Chocolate Argyrostrotis anilis ! recognition Elongate, slender, frosty to charcoal gray, with prolegs on A3 missing and those on A4 half-sized. Some individuals uniformly colored while others heavily patterned; commonly with large, pale, lateral patch extending forward from spiracle on A1. Thin, black line extending along outer face of proleg on A5 and A6. Raised, reddish dorsal tubercles (bearing D2 seta) on A1 and A8; warts on A2 and A7 appreciably smaller, but larger than those on A3 and A6. Dorsal pinacula white. Head with gray to reddish-brown “eye patches” to either side of triangle and reddish-brown-black bar across each lobe (above lower right). Larva to 4 cm. Ten species of Argyrostrotis are listed for our region, although at least three of these are synonyms. ! occurrence Barrens, powerline right-of-ways, thickets, woodlands, and coastal strand communities from Wisconsin to southern Canada, south to Florida and Texas. Evidently two generations over most of East with mature caterpillars in late spring and early summer and then again with smaller brood in mid to late summer. Active throughout growing season in Florida. Locally common. ! common foodplants Beach plum and other plums, crabapple, and hawthorn. ! remarks In Connecticut this moth is scarce except along the coast where it is known to feed on beach plum (Prunus maritima). The description for A. anilis in Forbes (1954) of a Sabatia-feeding, pink-striped larva, with ridges over A5 and A8, appears to belong to a member of another genus. DNA barcodes suggest that two species are currently going under the name of A. anilis in collections. By day, the Short-lined Chocolate caterpillar rests stretched out along a twig, with both the anterior and posterior abdominal segments flattened. Several larvae were sent to us by Lyle Buss, most of which he found after nightfall with a flashlight: “Yesterday afternoon it took me an hour to find only one while it was still light out, but between 11 and 11:30 p.m. I found four more on the same plants.” His collections from hawthorn included the whitish-gray larva figured above, a color form perfectly suited for the light bark of hawthorn. The larvae seem to do fine on old foliage. Prepupal larvae take on a bluish-green cast along the venter. The cocoon, a comparatively open network of reddish-brown silk, is spun among leaves or in leaf litter. The pupa overwinters.

Chocolates and Kin Subfamily Erebinae: Tribe Poaphilini 161

Four-lined Chocolate Argyrostrotis quadrifilaris ! recognition Spindle shaped, charcoal with subtle striping above, and prominent creamy subspiracular stripe. Proleg on A3 missing and that on A4 reduced. Black spiracular stripe including black spot behind spiracle on first four abdominal segments. Conspicuous black subdorsal spot on A1. Venter black with obscured patterning. Head with three prominent creamy stripes. Larva to 4.5 cm. Earlier instars gray-brown with more prominent striping and black spotting (inset). Until the life histories of our southeastern Argyrostrotis fauna are clarified, identifications should be based on reared adults. ! occurrence Wet pinelands, pocosins, savannas, flatwoods, powerline right-of-ways, and other open, acidic wetlands from Albany Pine Bush (where it is historic) to New Hampshire (also historic), south to Florida and Texas. A single principal generation with mature caterpillars in late spring and early summer; sporadic activity thereafter. Locally common. ! common foodplants Our ex ova larvae accepted maleberry (Lyonia ligustrina); first instars also nibbled on staggerbush (Lyonia mariana) and swamp doghobble (Eubotrys racemosa), which they later rejected. Reports on cotton in error—see below. ! remarks Adults are easily flushed, and sometimes fly during the day in late spring when new foliage of maleberry is abundant. Adults also are taken in light traps. Presumably mating and oviposition occur at night. The moth is local, and often absent from Lyonia wetlands northward, but occurs regularly from New Jersey southward. Our captive larvae were specific in their feeding preferences— they favored young leaves (but often not the youngest) from plants growing in sun, and ignored both flower buds and older leaves. Foliage from forest understory plants was rejected by first instars, and adults were not found in such places. Early instars reside on leaves, whereas older larvae roost on stems, sometimes away from the foodplant; the thoracic and terminal abdominal segments are somewhat flattened, appressed to the perch. It is our guess that their tendency to rest off the foodplant accounts for the curious and almost certainly erroneous host records for the genus: e.g., cotton for A. quadrifilaris, rose gentian (Sabatia) for A. anilis, and skullcap (Scutellaria) for A. flavistriaria.

162 Chocolates and Kin Subfamily Erebinae: Tribe Poaphilini

False Underwing Allotria elonympha ! recognition Barklike, elongate, mottled in grays, pinks, and/or browns. Ridge over A8 with edge blackened posteriorly; black extended laterally as oblique line toward spiracle. Dorsum and sides of A1 often noticeably lighter than T3 and A2. Head usually with white spot over each lobe and fine black line running from eyes to antenna. Subventral fringe (rootlet) setae absent. Prominent brown spot on venter of A3, and less conspicuous midventral spots on A4 and A5. First two prolegs (on A3 and A4) reduced. Larva to 4 cm. ! occurrence Swamplands and mesic woodlands from Missouri to Michigan and southern Maine, south to Florida and Texas. Two or three generations through most of East with mature caterpillars from June onward. Common. ! common foodplants Tupelo and other Nyssa. ! remarks Middle and late instars rest on bark by day. Caterpillars can be located by visual searches, but beating will yield them in numbers. We recall an afternoon in early August along the shores of Lake Michigan where caterpillars of the False Underwing and The Hebrew (Polygrammate hebraeicum) were so common in beating samples that dozens had to be individually reattached during a hopeless search to find larvae of Cadbury’s Mystique (Comachara cadburyi), which we were later to find out had finished feeding weeks earlier. The pupa overwinters.

Chocolates and Kin Subfamily Erebinae: Tribe Poaphilini 163

Gordonia Darkwing Gondysia similis ! recognition Integument plum with waxy bloom; smooth skinned with small black subdorsal eyespot on A1. Middorsal stripe, a composite of three fine purple and four pale lines, flanked by dark addorsal area. Dorsal setae borne from minute whitish pinacula. Sides of body with numerous fine pinstripes. White subdorsal, supraspiracular, and spiracular stripes continue forward onto purple-black head; subdorsal stripe ends above triangle, supraspiracular line ends about eyes, and spiracular reaches to antenna. Prolegs on A5 and A6 with dark plate on outer face. Spiracles black, small; that on A8 more than twice height of other abdominal spiracles. Venter with pale blotching. Larva to 3.5 cm. Three other species occur in the Southeast—see below. ! occurrence Pine flatwoods, savannas, bay forests, and pocosins of coastal plain from North Carolina to Mississippi. At least three generations in coastal North Carolina with mature caterpillars from late May onward. Local and uncommon. ! common foodplants Loblolly bay (Gordonia lasianthus). ! remarks Mature caterpillars become engorged and movement causes the skin to ripple like that of a bull walrus. Like the chocolates (Argyrostrotis), older larvae (and pupae) develop a distinctive bluish waxy bloom. JBS has located caterpillars twice on loblolly bay, and captive females will lay eggs if confined with its leaves. Adults are occasionally found flying diurnally as are those of its more common and widespread congener, Smith’s Darkwing (G. smithii); the latter occurs from southeastern Virginia to Florida, west to Texas, and north to Missouri. John Abbot illustrated a larva on what appears to be beech (Fagus) (Sullivan 2010), although we regard this as an unlikely foodplant. The rare Consobrina’s Darkwing (G. consobrina) occurs from North Carolina to Florida westward to Texas. Like Smith’s Darkwing it ranges into the Piedmont; its early stages are unknown. A fourth sibling species, G. telma, was recently described by Sullivan (2010); it occurs from North Carolina southward at least to the Florida Panhandle and west to Texas with a single record from southern Indiana. We encourage efforts to “crack” the life histories of these three darkwings, because all strike us as worthy conservation targets.

164 Chocolates and Kin Subfamily Erebinae: Tribe Poaphilini

Maple Looper Parallelia bistriaris ! recognition Highly variable in coloration and patterning: usually some shade of tan, brown, red, or gray; occasionally with alternating dark and light coloration (upper left). Prolegs on A3 and A4 reduced. Ridge over A8 with edge often blackened over its posterior side. Head patterning diagnostic: black line extending from subdorsum of T1 to antenna; second, narrower, black lateral line reaching from vertex to antenna; often with dark coloration to either side of dorsal midline. Anal proleg with fine black line on outer face. Subventral fringe (rootlet) setae absent. Larva to 4 cm. Younger larvae greenish (lower right above). ! occurrence Woodlands and forests from Minnesota to Nova Scotia, south to Florida and Texas. Two or more generations with mature caterpillars from June onward over much of East. Very common. ! common foodplants Maple, especially red; other reported hosts, including yellow and white birch and walnut, may be erroneous. ! remarks All of our collections of the Maple Looper have come from red and silver maple. The waxy green, early instars sit on a large leaf vein and reach out to either side to feed on patches of tissue. The pupa overwinters in a distinctive tacoshaped cocoon, fashioned from an excised leaf section.

Distinguished Cypress Owlet Cutina distincta

Missouri to southern Indiana and Maryland, south Florida and Texas. Multiple generations with mature larvae throughout growing season. Common. Cypress.

Chocolates and Kin Subfamily Erebinae: Tribe Poaphilini 165

Gray Cypress Looper Cutina albopunctella ! recognition Elongate, bright, lime green with longitudinal striping. A8 slightly humped. Proleg absent on A3; proleg on A4 one-third size of those on A5 and A6. White subdorsal and spiracular stripes; the latter passing through lower edge of pale yellow-orange spiracles, dipping below spiracle on A8, then running down anal proleg, and frequently more yellowish than subdorsal stripe. Supraspiracular stripe on T1 continues onto head, but fades out rearward. Faint, very thin, white addorsal stripes. D2 setae on A8 from small warts. Larva to 3 cm. Pogue and Ferguson (1998) recognized four members of the genus, all evidently associated with cypress. Identifications should be based on adults. Distinguished Cypress Owlet (C. distincta)—the most common member of the genus over much of its range—is figured on the previous page. In our examples of C. distincta, the subdorsal stripe is broader, whiter, and more parallel-sided than that of C. albopunctella, and the spiracular stripe contains more yellow. ! occurrence Swamps, blackwater streams, and other cypress wetlands from Missouri and southern Illinois to Maryland, south to Florida and Texas. Multiple generations with mature caterpillars from May to October. Locally common. ! common foodplants Bald cypress (Taxodium). ! remarks Accurately identified images and preserved larval specimens are needed for all four of our Cutina. The individual figured above from Okefenokee Swamp in southeast Georgia was photographed and identified by Doug Ferguson. We have had much success beating Cutina larvae from sucker shoots and sapling trees, some less than a meter in height. As might be expected for a species living above waterways, disturbed larvae do not readily release and drop from their perch. When beating for Cutina, pass the beating stick back and forth across the foliage to dislodge caterpillars. The dozen or so larvae that we have secured have come from new foliage, as suggested by their bright, lime green coloration. Given the habitat and growth habit of the host, the best way to obtain unambiguously determined larvae of Cutina may be to light trap or bait for gravid females, and then rear broods. Pupation occurs in a nest of webbed foliage, often along a shoot or twig. As in many other erebines, the pupa takes on a bluish, waxy bloom after a few days.

166 Wavy-lined Owlets Subfamily Erebinae: Tribe Ophiusini

Wavy-lined Owlets

Family Erebidae: Subfamily Erebinae: Tribe Ophiusini This is a large tribe with nearly 100 species north of Mexico, some 40% of which are zales (introduced separately below) (Lafontaine and Schmidt 2010).1 Generic diversity is greatest in subtropical Florida, Texas, and the Southwest. The cylindrical caterpillars, usually variegated in earthen tones (a few conifer-feeding zales are green and white), rest on twigs, bark, or in litter by day; the head is roughly the same diameter as T1. The first two pairs of prolegs are absent in Selenisa sueroides and Tyrissa multilinea, but present in the other genera that we treat here. Most are Catocala-like with an elongate tapered

body, slightly humped eighth abdominal segment, and dark ventral spotting. (In Zale these spots often join rearward to form a line; spots remain separated in Catocala.) A fringe of subventral setae (more slender than the rootlet setae of underwings, Catocala) is present in a few genera, e.g., Coxina. Abdominal segments five and six are roughly the same diameter as those that follow (these segments are often swollen in Catocalini). Coloration and patterning varies considerably among individuals. Apply names with caution and rear adults to confirm identities. Because of their cryptic coloration and habits, ophiusines are most efficiently collected with a beating sheet. Adults are strongly attracted to baits. Most come to lights, others (e.g. Confused Zale) rarely do so.

Black-waved Owlet Coxina cinctipalpis

Southern Florida and Texas. Active year-round. Very common. Legumes; often Acacia and Pithecellobium in Texas.

Live Oak Metria Metria amella

Southeast Virginia to Florida, west to Texas. Multiple generations. Common in Deep South. Our records from live and turkey oak. Head coloration diagnostic.

1

The recent inclusion of Amolita by Lafontaine and Schmidt in the Ophiusini, based principally on the position of the ostium (female copulatory orifice), radically changes our concept of the tribe. Characterization of the tribe above is traditional and does not accommodate Amolita.

Wavy-lined Owlets Subfamily Erebinae: Tribe Ophiusini 167 Tailpipes

Selenisa sueroides (= S. monotropa) [Pale-edged Selenisa] Florida and Texas, becoming common in subtropical areas where it is active year-round. Common. Acacia, blackbead (Pithecellobium), senna, and other legumes (our records from woody species). Larvae tunnel into wood and pithy stems to form their pupal cells. See Miller et al. (2006) for an interesting account of this species.

The Bravo

Tyrissa multilinea [Multi-lined Tyrissa] South Florida and Texas, south into Caribbean and Neotropics. Our caterpillars came from October collections in the Rio Grande Valley. Common. Acacia.

Detracted Owlet Lesmone detrahens

Kansas to Long Island (as a stray northward), south to Florida and Texas (and Neotropics). Mature caterpillars throughout growing season southward, probably July to October in New Jersey. Common in Deep South. Cassia.

168 Wavy-lined Owlets Subfamily Erebinae: Tribe Ophiusini

Moon-lined Moth Spiloloma lunilinea ! recognition Elongate with tapering posterior end. Prolegs on A3 and A4 about half size of those that follow. Trunk with alternating grayish stripes. Pale stripes alternating with somewhat darker (addorsal, supraspiracular, and spiracular) stripes; spiracular stripe especially evident on A8. Russet middorsal brand over A4 and A5. Middorsal stripe constricted between segments. Forbes’s (1954) description suggests ground colors may include yellows and tans. A8 with enlarged, white, pink, or orange D2 pinaculum, borne from elevated ridge. D2 pinacula on A7 and A9 also pimpled. Venter white with dark medial spots with those on A3–A7 largest. Head patterning diagnostic although often less boldly marked than individual figured here. Larva to 5 cm. ! occurrence Floodplain forests and woodlands, often over limestone soils from Missouri to western Pennsylvania, south through Appalachian region and Texas; more frequent west of Appalachians, and scarce to absent over much of Atlantic Coastal Plain. Evidently two generations over most of range with mature caterpillars from mid-June onward. Local and uncommon. ! common foodplants Honey locust. ! remarks The sienna saddle over A4 and A5 and smoky patch between A1 and A2, both evident in the individuals that we figure, are not mentioned by Forbes (1954) or Crumb (1956); this suggests that coloration varies markedly among individuals. The Moon-lined Moth was at least briefly established in central New Jersey where Joseph Muller collected larvae. It is not known if the moth feeds on water locust (Gleditsia aquatica) in the Southeast. Honey locust has been widely planted in cities and yards throughout much of New England and the Eastern Seaboard—perhaps one day the moth will establish more widely. It overwinters as a pupa. Lafontaine and Schmidt (2010) place the Moon-lined Moth in the Catocalini. We transfer it here to the Ophiusini because the caterpillar resembles a zale and because Spiloloma differs from our concept of Catocalini which all overwinter in the egg stage, are strictly univoltine, and usually bear rootlet setae. In addition, Moria Robinson (in litt.) notes that Spiloloma pupates in duff, as do Zale; Catocala fashion cocoons in leaf litter.

Wavy-lined Owlets Subfamily Erebinae: Tribe Ophiusini 169

zales Zale

Zale is a large and taxonomically challenging genus with nearly two dozen named, and a few still unnamed, species in our region. The many pine feeders can be bewilderingly difficult to identify both as larvae and adults. Because of the abundance of misidentifications of the pine feeders in collections, and even otherwise reliable state lists, we base our ranges only on specimens examined by DFS. Our use of names closely follows Forbes (1954). Ranges are most reliable from Wisconsin to Quebec, and south to Ohio, New Jersey, and North Carolina, but are incomplete south and west of North Carolina, especially through the Gulf Coast states. The Z. squamularis-obliqua complex is particularly problematic taxonomically. We group the angiosperm feeders and pine feeders separately. The green or brown caterpillars are elongate with a gradually tapering abdomen. The (D2) setae on A8 are perched atop two closely set protuberances (these are reduced in the green pine-feeding species, e.g., Z. buchholzi, Z. helata, and Z. curema). The caterpillars resemble those of underwings (Catocala), whose caterpillars also bear dark spots along the midventer of the trunk. However, the spots of zale caterpillars tend to form a midventral line on A1, A2, and A7; zales never have a fringe of subventral rootlet setae (page 97) as is commonly the case for our underwings. Often the prolegs on A3 are half the size of those on A4, which in turn are half the size of those on A5 and A6. Several have a pale, inverted V-shaped spot over each lobe (to either side of the triangle). The labrum may be deeply cleft, or in a few pine feeders, slightly emarginate. A fine black line runs along the outer face of the anal prolegs. We use subdorsal stripe in two different senses: unambiguously in the green, pine-feeding zales (e.g., Z. buchholzi and Z. helata), and for the dark stripe bordering the middorsal stripe (that includes the D2 pinacula) in several of the gray, pine-feeding Zales (e.g., Z. confusa, Z. metata, Z. metatoides, and Z. submediana). Some are reasonably constant in coloration and identifiable as larvae (e.g., Z. buchholzi, Z. curema, and Z. bethunei), while others are highly variable in appearance (e.g., Z. helata, Z. metatoides, and Z. lunata). It is our impression that colors and patterning are more pronounced in the penultimate and early last instars—especially among those zales that feed on deciduous plants—and that coloration becomes more subdued over the course of the ultimate instar. Identifications should be based on reared adults or, in some cases, foodplant associations. Others are more constant and identifiable as larvae (e.g., Z. galbanata, Z. phaeocapna, Z. buchholzi, Z. curema, and Z. bethunei). Crumb (1956) provides a key to 11 eastern species. Nine zales are illustrated in Wagner (2005); eight of the eastern conifer-feeding zales are figured by Maier et al. (2004). Where coverage overlaps with

these two publications, we selected images of caterpillars that differ in appearance, to convey a better understanding of color variation within a given species. Female zales lay about 200 ova singly over their three- to four-week adult lifespan. Zale eggs are comparatively large, rounded, and pale green; they hatch after 5 to 15 days. Larvae mature in about four to five weeks, feeding mostly at night. About half are pine feeders, and about half of these are specialists on only one or a few species of Pinus. Cherry, locust, and oak are used by several of those that feed on broad-leaved plants. Most zales prefer young leaf tissue, especially in early instars; they then consume older leaves and needles in late instars, although there are important exceptions, particularly among the pine feeders. Univoltine, spring-active species are often fully reliant on young leaves, and will straggle and perish if offered fully expanded, newly hardened foliage. Middle and late instars of most species rest on branches by day, and are cryptically rendered in browns and grays; the few pine feeders that perch along needles by day are green and, predictably, bear longitudinal white striping. According to Heinrich (1993), some are leaf clippers that chew through the petiole, dropping evidence of feeding activity to the forest floor; the chewed leaves might otherwise be used by birds to locate a caterpillar’s whereabouts. Zales are strong, active caterpillars that wander widely in search of appropriate foliage. Forbes (1954) notes zale caterpillars as being exceptionally muscular, capable of hurling themselves from their perch when alarmed. Keep this in mind if you plan to handpick your quarry. All overwinter as pupae in weak cocoons spun in leaf litter, or (and probably more often) in surface soil or humus, but not really underground. Several spring-flying species (Z. phaeocapna, Z. intenta, Z. lunifera, Z. unilineata, and the green pine feeders) are univoltine range-wide; but many species have two or more generations. Even in those Zale with more than one brood, some pupae from the first brood enter diapause in June or early July and overwinter, but we have not had any overwinter more than once. The adults of most come to lights but much more so to baits (see below), with the latter being an efficient means of securing females. One of our species (Z. lunata) is migratory. Additional life history information is given in the species accounts. To obtain eggs, we place each female in a half-gallon ice cream container with a mesh or other airy lid. Then we add a twig of host foliage (in a vase or water pick), paper toweling, and a sponge square moistened with sugar or honey water. Well-fed females will lay eggs over a period of up to four weeks on toweling, netting, and other surfaces (most pine feeders will lay on needles). Five-gallon paint filter bags make good sleeves for rearing small numbers of larvae. Because springflying zale are new-leaf specialists, move sleeves frequently so that the caterpillars will have access to young foliage.

170 Wavy-lined Owlets Subfamily Erebinae: Tribe Ophiusini

Green-dusted Zale Zale aeruginosa ! recognition Pink-brown, orange- to red-brown zale with welldifferentiated middorsal stripe outwardly edged with thin white border. Dark addorsal stripes most evident over thoracic segments and A8. Posterior half of A1 with darkened, modestly swollen saddle. D1 seta from minute white pinaculum on A1–A7. A8 humped with orange to black wart-bearing D2 seta. Often with interrupted, thin, black lateral stripe running under spiracles on abdominal segments. Venter of A3 and A4 with large purple spots; smaller linear lines on A1 and A2. In our examples, inverted V-shaped spot over each lobe of head indistinct; brown lateral line above eyes. Larva to 4 cm. ! occurrence Barrens, pocosins, woodlands, and forests from Ontario to Nova Scotia, south to Florida and Texas. One generation northward; two or more generations in South. In Connecticut mature caterpillars occur in June and July. Evidently, trivoltine in New Jersey, but both second and third broods appear to be partial. Generally uncommon, except in sandy and acid soil areas. ! common foodplants Kimball (1965) records white and live oak. Eric Quinter (pers. comm.) and two of us have taken larvae from blueberry. We have reared ex ova broods on oak. ! remarks The Green-dusted Zale could be polyphagous, much like the Colorful Zale (Z. minerea) and Lunate Zale (Z. lunata), yet it is decidedly less common than either of these two general feeders. We have beaten caterpillars from sparkleberry in North Carolina. On three occasions Eric Quinter has taken larvae on blueberry (and not other hosts), an observation consistent with where one most often sees adults of this handsome moth. We have images of zale caterpillars from blueberry in New Jersey and North Carolina that appear to represent the Green-dusted Zale. However, we also note that some Piedmont localities for the Green-dusted Zale lack blueberry and, moreover, have few native heaths (Ericaceae) of any kind.

Wavy-lined Owlets Subfamily Erebinae: Tribe Ophiusini 171

Dixie Zale Zale declarans ! recognition Longitudinally striped, gray to yellow-brown zale with dark middorsal, subdorsal, and spiracular stripes; often heavily salted with minute white spots. Three oblique black patches, when enlarged, may fuse over the dorsum on A1, A4, and A8 to form “saddles”; posterior margin of each patch often followed by diffuse white subdorsal or supraspiracular patch. Thicker longitudinal stripes edged above and below with thin tan to white stripes. Occasionally with small white spot before and following each abdominal spiracle. Head tan to orange with linear arrays of spots and other mottling; usually with black line above eyes. Ventral spotting dark maroon on white background: small spots on T1–T3; A1 and A2 with longitudinal bar; A3 and A4 with large, nearly round spot. Spiracles tan, circled with black. Larva to 5 cm. Middle instars (inset) smoky green with more obscure striping. ! occurrence Live oak groves, especially along coast, from North Carolina south through Florida, and west to Louisiana. At least three broods with last instars from May through September in North Carolina. Locally common. ! common foodplants Live oak. ! remarks Early-hatching larvae may consume oak catkins. More typically first and second instars feed on the underside of new leaves, grazing away patches of tissue. Second- and third-generation caterpillars presumably must locate and feed on scarce new shoots. The Dixie Zale is sometimes common in dunes and maritime forests of the southeastern coastal plain. The first brood is the most numerically abundant; later summer broods appear to be partial. The pupa overwinters, presumably in a cocoon fashioned in the duff and surface litter.

172 Wavy-lined Owlets Subfamily Erebinae: Tribe Ophiusini

Lunate Zale Zale lunata ! recognition Highly variable, usually gray, brown, or black, with slight swelling around D2 seta on A1 and prominent ridge over A8. Beautiful gray-green lichen-mimicking forms (McCabe 1990) occasionally encountered. Intersegmental area between A1 and A2 yellow (visible when a loop is thrown into body). Subdorsal and spiracular stripes darkest rearward. Posterior dorsal seta (D2) often at upper end of small elongate white spot on A1. Head with dark mottling over each lobe and with pale brown to white lines over sides and triangle. Frons high: onethird again as high as its width (see Colorful Zale, Zale minerea, next page). The venter bears black midventral dots that fade to red-brown (Forbes 1954) (lower right). Larva to 5 cm. ! occurrence Yards, parks, thickets, woodlands, and forests from southern Canada to Florida and Texas, but uncommon northward. One or two generations in New England and Upper Midwest, evidently three or four in New Jersey; mature caterpillars from June onward over much of East. Very common. ! common foodplants Apple, birch, blackberry, cherry, chokeberry, hawthorn, plum, raspberry, sensitive plant (Cassia), willow, and wisteria, but also oak and many other woody plants, and occasionally forbs, especially among the pea family (Fabaceae). Black cherry (Prunus serotina) and willow commonly used. ! remarks As noted by Ferguson (1991), this species is migratory. In southern New Jersey, worn migrants usually appear in April and May and a large local brood of adults appears in late June, at about the same time as early Catocala are seen on the wing. Successive broods are produced through the rest of the season with adult numbers building into October, November, and December. The sequence is similar in eastern North Carolina but starts earlier, and more often the early season adults are fresh and appear to have eclosed locally. We do not know if there is a southward return flight in autumn, but suspect so based on the abundance of adults along the Atlantic Coastal Plain most falls. (Annual, highaltitude, southward migrations missed by generations of scientists were recently documented in the hawk moth genus Autogramma by Chapman et al. (2008).) DFS collected a teneral male in early January and two fresh adults in a mid-March heat wave in 1990, indicating that some pupae overwinter at least as far north as New Jersey; however, most individuals are believed to emerge by December (and fly south or fail to reproduce).

Wavy-lined Owlets Subfamily Erebinae: Tribe Ophiusini 173

Colorful Zale Zale minerea ! recognition Highly variable and easily confused with Lunate Zale and other zales that feed on broad-leaved shrubs and trees. Ground color gray, tan, brown, to nearly black; patterning ranges from almost uniformly colored to striped or strongly mottled. Intersegmental area between A1 and A2 yellow over dorsum. Often with a pale lateral saddle extending between A4 and A5. Dorsal and often lateral setae borne from white pinacula. Crumb (1956) notes that frons is shortened, scarcely as high as it is wide at its base. Ventral coloration as in lower right. Larva to 4 cm. In the Lunate Zale the frons is a third again higher than broad. ! occurrence Woodlands and forests. Transcontinental in Canada, south to Florida and Texas. One principal generation over much of East with mature caterpillars in June and July; a few individuals eclose as a second (summer) generation from Pennsylvania southward. Common in much of its range, except along Atlantic Coastal Plain. ! common foodplants Prentice (1962) lists alder, American hornbeam, aspen, basswood, birch, elm, Douglas-fir, hazel, hickory, hophornbeam, maple, oak, poplar, spruce, and willow. If our slide images of wild-collected caterpillars are identified correctly, then we have numerous records for chestnut and oak, and one from hawthorn. See Remarks. ! remarks Adults of this species and the Lunate Zale are commonly confused in collections. Hence, we have some doubt about published foodplant records, although both species appear to be generalists. Until reliable characters can be identified to distinguish the caterpillars of these two zales, determinations (and host associations) based on larval identifications should remain suspect. Perhaps a careful study of their larvae, especially the labrum, mouthparts, frons height, head patterning, crochets, and spots along the venter, will reveal features that can reliably distinguish the two. Last instars of the Colorful Zale sometimes hide under loose bark rather than rest on branches. Like some other erebids, prepupal zales take on an aqua or blue-green aspect.

174 Wavy-lined Owlets Subfamily Erebinae: Tribe Ophiusini

One-lined Zale Zale unilineata ! recognition Mostly yellow-brown to brown; some individuals with considerable charcoal coloration laterally. Distinct yellow dorsal intersegmental patch between A1 and A2. A8 elevated and humped with prominent, widely spaced, D2 warts directed away from midline. Most forms with vague pale patch between A4 and A5. Usually with poorly defined, somewhat yellowish, middorsal and supraspiracular stripes. White pinacula conspicuous. Proleg on A3 much reduced. Venter somewhat yellowish with “dark center line, except on segments with prolegs” and “A3 and A4 only with dark patches” (Forbes 1954). Larva to 4.5 cm. Penultimate instar decidedly more yellow with obscure pinstriping often evident (inset). The eighth abdominal segment is not as strongly humped and the D2 warts are proportionately smaller in the Black Zale (Z. undularis), another legume feeder. In addition, caterpillars of the Black Zale usually have a more subdued white lateral patch over A4 and A5, less contrast between adjacent stripes, and the yellow intersegmental band rearward of A1 is not as evident. Additionally, the venter bears a nearly continuous purplish line that is much widened on the thorax and between the prolegs (Forbes 1954) (page 184). Other Zales such as the Lunate Zale (page 172) and Colorful Zale (Z. minerea) (page 173) may also use legumes. Until host ranges and diagnostic characters for the genus are better understood, the identity of legume-feeding zales is best confirmed through rearing of the adults. ! occurrence Overgrown fields, woodlots, fencerows, parks, and woodlands from Manitoba to New Brunswick, south to Georgia and northern Arkansas. One generation with mature caterpillars mostly in May and June. Locally common. ! common foodplants Black locust (Robinia pseudacacia). ! remarks The One-lined Zale and Black Zale (Z. undularis) have benefited greatly from the planting of black locust through much of the East. It is unclear if other locusts are used southward and westward.

Barrens Zale Zale lunifera

Maine to Florida, west to at least Mississippi. One generation with mature caterpillars early in season, mostly in April to June. Local. Scrub oak northward and blackjack and other scrubby oaks southward. Schmidt’s (2010a) new application of the name Z. lunifera is discussed under Z. intenta on the next page.

Wavy-lined Owlets Subfamily Erebinae: Tribe Ophiusini 175

Bold-based Zale Zale intenta (= Zale lunifera of previous authors)

! recognition Yellow- or red-brown zale with elevated and strongly angulate

eighth abdominal segment; D2 protuberances reduced, relative to elevation of entirety of A8. A1 with white oblique subdorsal patch behind darkened, modest swelling that includes D2 seta. Second, more ill-defined, pale, lateral patch extending between A3 and A4. Middorsal stripe vague, edged by more conspicuous, thin, white addorsal stripes. Dark, interrupted dorsal-subdorsal stripe; especially dark over T2, T3, A3–A5, and A8. Setae borne from white pinacula. Dark lateral patch extending from posterior half of A7 down outer face of anal proleg, broadest around spiracle on A8. V-shaped spot on each lobe of head small and narrow relative to other zales. Spiracles dark (see also Crumb 1956). Larva to 4 cm. Earlier instars are more yellow-green. The Bold-based Zale is a member of a complex of species. The Barrens Zale (Z. lunifera), an oak feeder associated with barrens, is figured on page 174. The westward reach of the species and what names apply to populations in Texas are in need of study. ! occurrence Coastal strand communities, shrubby fields, woodlands, and forest edges from southern Canada to Florida and Texas. One generation range-wide with mature caterpillars from late May to July in New England, April and May southward. Covell’s (2005) treatment indicates a second generation (presumably southward), although we are unaware of evidence for such in this complex. Common. ! common foodplants Cherry, especially black cherry, and beach plum (DFS); one adult reared from caterpillar on willow oak (DFS). ! remarks Schmidt (2010a) recently revised the species taxonomy for the Bold-based Zale (Z. intenta) and the closely allied Barrens Zale (Z. lunifera). Previously the name lunifera had been widely applied to the former entity. Where the two species fly together, there is little seasonal overlap, with Z. intenta flying two to three weeks earlier. All members of this species group are thought to be univoltine, and new leaf specialists: caterpillars of the Barrens Zale straggle and eventually die if fed any but the youngest oak leaves (Mike Nelson, pers. comm.). The larva attributed to Bold-based Zale by Crumb (1956) matches our concept of Brown-spotted Zale (Z. helata) (page 179), which is consistent with his recorded host of white pine. The last two instars of both the Bold-based and Barrens Zales are dark, cryptic, rest on branches by day, and lack the prominent longitudinal striping (characteristic of pine feeders) described by Crumb (1956).

176 Wavy-lined Owlets Subfamily Erebinae: Tribe Ophiusini BAITING FOR ZALES Baiting for zales in barrens and dry pine-oak woodlands can be engaging and productive. April and May are the best months, beginning after red maples have gone to seed and are no longer providing nectar. DFS recalls a phenomenal late April evening in southern New Jersey following a warm spring day. He baited about 35 trees and then walked the bait line for three hours after dark. Over 700 individuals representing 13 zale species were seen along the bait trail—a typical rate of visitation in April for southern New Jersey might be 0.5 to 5 zales per bait patch per hour. (The three blacklight traps deployed in the same woodland collectively yielded fewer than 20 zales.) It is our experience that some zales only occasionally come to light under normal weather conditions, the Confused Zale (Z. confusa) being a case in point among eastern species. Baiting should be included in biotic inventory efforts where zales are among the target species, and requisite in surveys for state-listed species. Connecticut, Massachusetts, New York, and Ohio, and perhaps other states, have at least one pine-feeding Zale included on their rare and endangered species lists.

Confused Zale Zale confusa

Missouri, Ohio, and New Jersey, south at least to northern Georgia and Texas. Two or three generations range-wide; perhaps four southward, but later broods partial. Mature caterpillars from late May to September in New Jersey, starting earlier southward. Locally common. Shortleaf and perhaps occasionally other pines.

Fawn Virginia Pine Zale Zale metata

Southern Ohio, extreme south central Pennsylvania and southern New Jersey to central Florida and Texas. Two broods in New Jersey with mature caterpillars from late May to August; more broods southward, but later ones partial. Locally common. Virginia pine, probably sand pine (Pinus clausa) in Florida.

Wavy-lined Owlets Subfamily Erebinae: Tribe Ophiusini 177

pine-feeding zales The genus Zale has radiated on pines. We treat 11 species here, but suspect that others will be recognized, e.g., from the Z. obliqua complex. Adults are among our most frequently misidentified Erebidae in collections and in the literature. Reliable species-level characters for larvae are unknown for

most: identifications should be based on reared adults. (The genitalia of both sexes are usually diagnostic as are many wing pattern characters with practice.) Foodplant identity, plantcommunity association, and seasonal activity provide good clues to a zale caterpillar’s identity.

Buchholz’s Zale Zale buchholzi ! recognition Green to burnt sienna (but with at least some green ventrally); prominent, white subdorsal stripe narrowed along thoracic segments, with lower edge broadened and ill-defined on A6–A8. Orange-brown spiracular stripe edged with white, although only sparsely so above. Almost no hump on A8; D2 pinacula scarcely raised. Smoky green middorsal stripe, edged with white, sandwiched between broad orange-brown addorsal stripes. Each lobe with two short white lines. Spiracles tan to white with narrow black ring. Larva to 4 cm. Early instars green with black pinacula (lower inset); ground color of middle to penultimate instar about evenly divided between green and orange brown (upper inset). ! occurrence Pinelands of Atlantic Coastal Plain from New Jersey to Florida westward along Gulf coast to Texas. One generation with mature caterpillars as early as late May in New Jersey, earlier southward. Common (at bait) in much of its range. ! common foodplants Pitch (and pond) pine in New Jersey; apparently also associated with loblolly and longleaf pines southward. ! remarks There is an unnamed entity with similar-appearing adults that occurs from the Delmarva Peninsula southward into Florida that is associated mostly with native loblolly pine stands. Since the male genitalia are not particularly similar, we do not assume its larva is similar. Its flight period is mostly about a month later, with larvae presumably maturing in June in North Carolina and July on the Delmarva.

178 Wavy-lined Owlets Subfamily Erebinae: Tribe Ophiusini

Banded Zale Zale duplicata ! recognition Remarkably variable: greenish, reddish, maroon-gray or brown with or without conspicuous striping. Closely set protuberances over A8; somewhat smoother and glossier integument than that of other Zale. It also strikes us as having a more slender aspect. Most individuals with vague patterning, but strongly mottled individuals (Ives and Wong 1988) as well as striped forms occur (Forbes 1954) (see Remarks below). Dorsal pinacula minute, often white. D2 seta sometimes set in dark oblique to triangular spot over abdominal segments, borne on prominent protuberances on A8, and smaller raised warts on A9. A1 commonly with obscure, darkened saddle that includes D2 setae; less commonly with pale subdorsal patch between A4 and A5. Intersegmental area between A1 and A2, and A2 and A3, occasionally pale or rarely yellowed. Head with gray to white inverted “V” over each lobe; labrum with only shallow notch, and no black lateral line above eyes. Striped forms may not be separable from Brown-spotted Zale (Z. helata). Forbes (1954) described Z. duplicata as “striped in rusty and yellow, shading into white; dorsal stripe less regular than in Z. helata, shading into yellow on midsegments; the bordering brown stripes darkened at rear of segments; lateral darker zone more even in color, finely edged below with white; substigmatal area yellow; midventral stripe normally interrupted, much broadened between prolegs.” Larva to 4 cm. Middle and penultimate instars greenish to greenish yellow. ! occurrence Woodlands and forests from at least Manitoba to Nova Scotia, south to Georgia (mountains). One generation with mature caterpillars in June or July. Often common to locally abundant. ! common foodplants Eastern white pine; records for lodgepole and jack pine listed by Ives and Wong (1988) presumably refer to Z. duplicata largera; Prentice (1962) also lists Scotch pine, red pine, and larch, but these too may apply to other species. ! remarks The larva is widely known as the Pine False Looper. All zales in which the final instar is green have a single annual generation with springemerging adults. The Banded Zale is associated with eastern white pine from Canada to northern Georgia. Northward and westward it is reported to use several hard pines (two- and three-needled species). But there is reason to believe that these records are referable to Z. d. largera, a known jack pine feeder—a segregate that some believe warrants full species status, distinct from the partially sympatric Banded Zale.

Wavy-lined Owlets Subfamily Erebinae: Tribe Ophiusini 179

Brown-spotted Zale Zale helata ! recognition Numerically more prevalent green form with bright white middorsal, subdorsal stripes, and thin, poorly differentiated subspiracular stripes. Latter two stripes enclose brick-red to orange-brown spiracular stripe that may be infused with green. Middorsal stripe begins on T2 and ends over A9, nearly parallel-sided; subdorsal stripe runs from head to top of proleg on A10. Middorsal and subdorsal stripes delimiting greenish addorsal stripe commonly edged with black over back half of each segment. Other forms with green ground color partially or almost completely replaced with reddish brown, but otherwise similarly marked. A range of intermediate phenotypes link the green and orange- to red-brown extremes. A8 without distinct ridge or raised setal warts. Prothoracic plate green or tan. Head often with white to yellow “V” over each lobe and white line rearward of eyes. Labrum only shallowly emarginate. Larva to 4 cm. One form of the Banded Zale (Z. duplicata) purportedly resembles the Brown-spotted Zale. Caterpillars of the two are easily distinguished: the dorsum of A8 is always strongly humped with the D2 protuberances in the Banded Zale (page 178). ! occurrence Barrens and pine woodlands from Manitoba to Maine, south at least to northern Alabama and reportedly to Texas. One generation with mature caterpillars from June to July in most of range, earlier southward. Common. ! common foodplants Both hard and soft pines; rarely larch. ! remarks The ground color of most of our pine-feeding zale caterpillars is gray or brown, but this species apparently does not have a form with barklike coloration. Caterpillars of the Brown-spotted Zale rest on needles with the head wedged near the base of a needle fascicle (Maier et al. 2004) in the same fashion as many of our angles (Macaria). Larvae, especially in early instars, consume young needles. Across much of its range, the Brown-spotted Zale is among the most widespread and commonly encountered pine-feeding zales, although it is apparently uncommon in the Southeast. Adults have a rather uniform appearance across the southern portion of the range (Rings et al. 1992: Plate XII, figs. 26, 27, the later as curema), roughly from New Jersey southward; populations from mountainous areas of Pennsylvania northward can be highly variable (Rings et al. 1992: Plate XII, figs. 21, 22?, 25) and difficult to recognize. The degree to which larval appearances differ geographically is unknown.

180 Wavy-lined Owlets Subfamily Erebinae: Tribe Ophiusini

Washed-out Zale Zale metatoides ! recognition Caterpillar variable in color and among more difficult members of genus to characterize: larva usually orange-brown, brown, or gray but more greenish forms occasionally occur (Maier et al. 2004). Striping usually vague, but conspicuous in some green forms. Usually with pale subdorsal patch toward rear of A1 and A4. D2 setae from larger warts on A8. Most consistent stripes include beaded middorsal, dark subdorsal (that includes D2 setae), and pale supraspiracular stripe. Oblique subdorsal patches, darker forward and paler rearward, covering back third of A1–A7. Head with large white oval to lens-shaped spot, capped with black spot over each lobe; considerable dark mottling; black lateral line, edged with white, extending back from antenna. Forbes (1954) noted that the proleg on A3 is “much reduced,” but to our eye the first set of prolegs appears close to the norm for the genus. Larva to 4 cm. Caterpillars of Bethune’s Zale, Confused Zale, Fawn Virginia Pine (Z. metata), and Gray Spring Zale (Z. submediana) may be indistinguishable—all have pale subdorsal patches on A1 and A4, a small proleg on A3, and large warts on A8. Identifications in this group are best based on adults. ! occurrence Barrens and pine woodlands from at least Manitoba to Maine, south to Georgia (mountains); range in Gulf states unclarified. One generation with mature caterpillars from late June into July in New Jersey; into August northward. Locally common. ! common foodplants Hard pines such as jack, pitch, red, and Virginia. ! remarks The Washed-out Zale is in one of a complex of some dozen eastern pine-feeding zales that vex all but seasoned owlet enthusiasts. Over much of the Northeast and south through the Appalachians, it is second only to the Brownspotted Zale (Z. helata) in abundance. In the Appalachian foothills in late May and early June, this and the Esther Moth (Hypagyrtis esther) will be among the most common caterpillars in beating samples from Virginia pine.

Wavy-lined Owlets Subfamily Erebinae: Tribe Ophiusini 181

Oblique Zale Zale obliqua ! recognition Mostly gray-brown caterpillar with black and white speckling, and low protuberances over A8. Middorsal stripe pale and often broken over abdomen, including white Y-shaped spots that are best defined toward rear of each segment on A3–A6. Dorsal seta from minute, whitish pinaculum; D2 seta embedded in oblique blackish spot. Supraspiracular area pale, becoming more clearly defined as stripe rearward of A6. Spiracles towards bottom half of dark spiracular stripe. Inverted V-shaped spot over head not as strongly whitened as in many other pine-feeding zales; black line extending behind eyes; labrum shallowly cleft. Larva to 4.5 cm. Third instar slightly greenish brown with little pattern; dorsum paler. Gray-banded Zale (Zale squamularis) caterpillars may be indistinguishable from those of the Oblique Zale: most individuals of that species are more boldly marked, average a bit smaller, and occur both earlier and later in the season. Because other pine-feeding zales also have similar caterpillars, identifications are best based on adults. ! occurrence Barrens, plantations, pinelands, and mixed-pine woodlands. At least Ohio to southern Maine, south to northern Florida, Mississippi, and Texas, apparently not reaching Canada or the northern Great Lakes states. From New Jersey northward, one brood with mature caterpillars mostly in late July and August. From eastern Maryland to northeastern North Carolina, two broods with larvae probably maturing in late June and July, then again from late September to October. Locally common. ! common foodplants Primarily pitch pine northward; southern hosts probably include loblolly, pond, and longleaf pine. ! remarks Adults fly mostly in June and July from New Jersey northward. They are the last of the pine-feeding zales to appear; in New Jersey only third-generation Z. squamularis and Z. confusa fly later. Young larvae consume newer terminal needles; last instars feed on older, hardened needles. The larval stage lasts about a month. The Z. obliqua-squamularis complex needs taxonomic study. The entity that we illustrate from New Jersey is the northern member of a species complex that ranges from Maine south, at least into eastern North Carolina. Farther south there may be additional species. The adults are sometimes confused with those of Z. submediana (page 182), a spring flyer.

182 Wavy-lined Owlets Subfamily Erebinae: Tribe Ophiusini

Bethune’s Zale Zale bethunei

From at least Ohio to southern New Jersey, south to Carolinas and Tennessee. Two generations with mature caterpillars in May and June, then again in July and August in most of range; small third brood in North Carolina. Locally common. Virginia pine. Often with yellow aspect to middorsal and supraspiracular stripes.

Black-eyed Zale

Zale curema [Northeastern Pine Zale] Albany, New York, to Maine, south to western North Carolina; we have not examined specimens from Gulf states and Texas identified as this species in collections. One generation with mature caterpillars from July into August in the Northeast. Locally common northward. Pitch and perhaps other hard pines.

Gray-banded Zale Zale squamularis

Ohio to Long Island, south to Florida and Texas; range limits are uncertain as the “species” may represent more than one taxon southward. Three generations in New Jersey with mature caterpillars from June onward; both later generations partial. Locally abundant. Pitch, pond, and perhaps other hard pines.

Gray Spring Zale Zale submediana

Wisconsin to Maine to New Jersey and North Carolina. A later-flying entity also goes under this name in Ontario, Michigan, and Wisconsin. One generation with mature caterpillars mostly in June in New Jersey. Locally common over much of range. Jack, pitch, and other hard pines.

Wavy-lined Owlets Subfamily Erebinae: Tribe Ophiusini 183 Double-banded Zale Zale calycanthata

Ontario to New Jersey, south to Florida and Texas, apparently as transient resident northward. One generation with mature caterpillars in April and May. Locally common. Oak; captive larvae accept apple and cherry; Forbes (1954) doubted the moth’s association with allspice or sweetshrub (Calycanthus).

Boxelder Zale Zale galbanata

At least Manitoba to New Brunswick, south to northern Florida and Texas. Two or more generations with mature caterpillars from late May to October. Common. Boxelder and perhaps other maples.

Scaled Zale Zale horrida

Canada to Florida and Texas. Two or more generations with mature caterpillars from May to October. Common. Viburnum; our collections mostly from arrowwood (V. dentatum) and related viburnums.

184 Wavy-lined Owlets Subfamily Erebinae: Tribe Ophiusini

Okefenokee Zale Zale perculta

Southern Georgia and northern Florida. One generation with mature caterpillars in late spring. Very localized. Climbing fetterbush (Ampelothamnus phillyreifolius). See account in Schweitzer et al. (2011) and cover of this book.

Hazel Zale

Zale phaeocapna Southern Canada south to Florida and Texas. One generation in spring with mature caterpillars from late April (Florida) to June and July (much of range). Locally common; scarce in collections before 1970s. Hazel and hophornbeam (Ostrya), as well as witch hazel. Bright, yellow-form caterpillars frequent on latter host. More typical brown form shown below. See McCabe (1987).

Black Zale

Zale undularis Minnesota and Manitoba to New Brunswick, south to Florida and Arkansas. Evidently one principal generation with mature caterpillars mostly in late June and July; partial second and third broods southward. Locally common. Black and honey locust.

Wavy-lined Owlets Subfamily Erebinae: Tribe Ophiusini 185

Locust Underwing Euparthenos nubilis ! recognition Stout, more cutwormlike than most erebines. Broad, crenulate, tan, yellow-brown, or brown stripes alternating with darker stripes. Often with pale spot behind each spiracle on A1–A7. Easily recognized by subdorsal, crescent-shaped cluster of four creamy spots on A1, when present. Beaded middorsal stripe, often edged with black, over posterior abdominal segments. Prolegs on A3 reduced. No black spots on venter. Larva to 5.5 cm. ! occurrence Abandoned fields, roadsides, and woodlands from Wisconsin to southern Canada, to northern Florida and northern Arkansas. Two generations over much of range with mature caterpillars from June onward. Common. ! common foodplants Locust, especially black locust. ! remarks The coloration of the Locust Underwing caterpillar suggests that it spends its days on the lower trunk or ground. Early instars are common in beating samples from the foodplant. According to Forbes (1954), the genus represents a basal offshoot of the lineage leading to true underwings (Catocala), but as pointed out by our colleague Richard Peigler (in litt.), one would be hard pressed to find evidence of a close association when comparing the larvae of the two genera. The pupa overwinters.

186 Witches Subfamily Erebinae: Tribe Thermesiini

Witches

Family Erebidae: Subfamily Erebinae: Tribe Thermesiini Witches are largely tropical owlets and include some of the World’s largest insects, e.g., the wingspan of the White Witch (Thysania agrippina) may exceed 30 cm. More familiar is this

behemoth’s little “sister,” the Black Witch (Ascalapha odorata), a large, powerful flyer that sometimes strays as far north as southern Canada. The generic limits of the tribe are in flux—as presently circumscribed, our fauna includes six species (Lafontaine and Schmidt 2010). We are unaware of larval characters that are unique to the tribe.

Black Witch Ascalapha odorata ! recognition Very large, robust, striped, smooth, brown caterpillar with pale dorsal patches over thorax, A2, A5, and A7; body foreshortened. Venter pink to magenta with purple-black spots on A1–A4; spot on A1 nearly round while those on A3 and A4 more barlike. Pale, bat-shaped spot over A2; pale spot over A5 transversely rectangular. Darkened elliptical spiracles, very large. Legs long, often pinkish red; planta greatly flared, flukelike. Prothoracic and anal shields deep brown-black above; former reddened laterally. Head very dark above; triangle pale except for vague medial spot; gena (cheek) with squiggly reddish lines. Approaching 8 cm. ! occurrence Thorn scrublands, palm woodlands, hardwood hammocks, and other subtropical habitats in southern Florida and Texas, with range contracting southward in years with hard freezes; adults stray north to Canada and Alaska. Larval populations highest from September to November. Seasonally common in southern Texas and Florida. ! common foodplants Various woody legumes. We have reared larvae on acacia and collected wild larvae from blackbead or ebony (Pithecellobium); also recorded from cassia, locust, and mesquite. ! remarks The Black Witch is our largest breeding owlet. It is well established in southern Florida and Texas (and now, in the Greater Los Angeles area on planted acacias). In Texas, the moth populations are highest during the wet season, beginning with the summer monsoons and continuing into November when the state’s legume-rich scrublands are producing new foliage. The moth is especially apt to be encountered before and after hurricanes and tropical storms, sometimes far out of range. Adults are familiar to all southern collectors who employ bait to attract moths. The caterpillar secrets itself under litter or bark by day and emerges at nightfall to feed. No stages are believed to tolerate prolonged freezes.

Eulepidotines Subfamily Eulepidotinae 187

EULEPIDOTINES

Family Erebidae: Subfamily Eulepidotinae This is a diverse assemblage of moths that were formerly classified as catocalines (in the broad sense). Most of the 27 North American species derive from tropical lineages that reach their

northern limit in subtropical Florida and Texas. Only two of our 15 genera are widespread in the East: Anticarsia and Panopoda. We are too unfamiliar with the larvae and natural histories of the subfamily to offer a diagnosis. Placement of Litoprosopus here is provisional (Lafontaine and Schmidt 2010).

Orange Panopoda Panopoda repanda ! recognition Pale to emerald green, peppered with abundant, minute, dark green to purple-black spots; body notably tapered rearward. Proleg on A3 reduced, often held above substrate. Faint yellow subdorsal stripe often weakening rearward. Middorsal stripe weakly developed. T2–A8 with oblique yellow lateral lines that start on previous segment (above spiracle) and run down to proleg. Dorsal setae borne from small, yellowish pinacula. Head large, rounded, with numerous smoky spots. Elongate pinkish antenna. Large, orange, elongate spiracles. Larva to 4 cm. ! occurrence Woodland and other coastal plain communities from southeastern Virginia to Florida, west to Texas. At least two generations in North Carolina with mature caterpillars from early summer onward; continuously brooded in southern Florida and Texas. Common in Deep South. ! common foodplants Live oak. ! remarks The early instars skeletonize patches of leaf tissue from below. Reports of the larvae feeding on para grass (Panicum purpurascens) are certainly in error. Panopodas are midsummer caterpillars that feed on mature foliage. Not surprisingly, their heads are comparatively large relative to owlets that feed on spring foliage. Such is especially apparent in recently molted late instars (inset). The pupa overwinters.

188 Eulepidotines Subfamily Eulepidotinae

Red-lined Panopoda Panopoda rufimargo ! recognition Lime to blue-green with variably developed subdorsal stripe and abundant, minute, smoky to blue spots. Middorsal stripe weak and broken or absent. Oblique cream to yellow lines running under spiracles on A1–A8. Prolegs on A3 and A4 only slightly reduced in size; prolegs with ends splayed forward and rearward, reddened. Anal prolegs large and directed backwards. Head rounded, green with abundant dark speckling, usually with transverse yellow line over triangle, and red antennae. Larva to 4 cm. Penultimate instar with reduced spotting. Caterpillar of Hickory Panopoda (P. carneicosta) similar but feeds on Carya (inset); the characters given in Forbes (1954) for the separation of Panopoda larvae vary within species and thus are unreliable for the separation of eastern panopodas. The range of the Hickory Panopoda follows that of Carya, its principal host. Kimball (1965) also lists basswood, oak, and willow, but these hosts seem unlikely given our understanding of the genus and the life histories of other panopodas. The Hickory Panopoda has one generation in Connecticut, a partial second in New Jersey, and is active over much of the growing season in Florida. ! occurrence Barrens, woodlands, and forests from Minnesota to New Brunswick, south to Florida and Texas. One generation in Connecticut with small second brood in New Jersey; evidently two or more broods in South with mature caterpillars from June onward. Very common. ! common foodplants Beech and oak. ! remarks Caterpillars commonly perch on the upper side of leaves. The subdorsal stripes are a credible match for the midrib and lateral veins of the red oak leaves on which the caterpillars are commonly found. Larvae sometimes throw a low hump into the anterior half of the body. Disturbed larvae “jump” from their perch and continue to thrash wildly for seconds. This same behavior assures that beating samples will contain essentially all the larvae from any tapped limb or branch. The pupa overwinters in a thin, cocoon spun in leaf litter, the walls of which may contain detritus and much of whatever else is in the prepupa’s vicinity.

Eulepidotines Subfamily Eulepidotinae 189

Cabbage Palm Caterpillar Litoprosopus futilis [Palmetto Borer Moth]

! recognition Mature larva somewhat shiny brown to pink with variously

developed brown to red striping. Prothoracic shield, anal plate, and pinacula black and hardened. D1 and D2 fused onto a single pinaculum on T2 and T3; D2 setal base on mesothorax often ringed with white. D2 seta of T2, A8, and A9 extremely long. Broad, brown to red middorsal stripe edged with white. Broad, orange-flushed spiracular stripe also edged with white. Head rounded, shiny, deep reddish brown above and nearly black below level of triangle. Larva to 4.5 cm. Middle instars with long white setae, browner ground coloration, and more conspicuous striping (inset). ! occurrence Coastal strand communities and scrublands from North Carolina to Florida, west to Texas; breeding range unclarified as adults are infrequently sampled at light. Active nearly year-round in southern Florida with mature caterpillars from March to November. Common in Florida. ! common foodplants Saw palmetto, sabal, and related palms. ! remarks The caterpillar tunnels in and about palm inflorescences, consuming unopened and newly opened flowers (Dekle 1968). Occasionally the larvae are abundant enough to consume most of the flowers in a population, thereby depressing local honey production. The caterpillars feed beneath a web of silk that is fortified with reddish frass and fecal pellets. Larvae are highly cannibalistic. Both early and late instars disperse on silken threads. Charles Bordelon reports that caterpillars commonly visit and consume sugar bait, especially during times when flowers are in short supply. He reared one late instar through to the adult stage solely on a banana-beer mixture that he was using in his bait traps. Interestingly, it is not the caterpillar’s feeding damage that draws the most complaints from affected landowners, but rather the exploits of the wandering prepupal larvae, which may enter homes and other places where they are not welcome. Larvae release reddish liquid that stains walls and floors. In addition, cocooning larvae will tear up rugs, clothes, furniture pillows, and other fabrics if chosen as a pupation site. (Normally the cocoon is fashioned in palm fibers and other surface litter.) Prepupal Cabbage Palm Caterpillars are pink to red in color. Adults visit bait—they are wary and difficult to approach, continuing to beat their wings while feeding and quick to move if illuminated (Jim Vargo, pers. comm.).

190 Eulepidotines Subfamily Eulepidotinae

Velvetbean Caterpillar Anticarsia gemmatalis ! recognition Exceedingly variable in coloration: normally green, but ranging from yellow-green to olive gray, green-brown, and occasionally black (lower right). Usually with broad, white to orange spiracular stripe, edged with white above and below, that continues down outer face of anal proleg. Spiracular stripe decidedly thinner through thoracic segments. Thin, whitish middorsal, subdorsal, and supraspiracular stripes. In darkly pigmented forms, ground color frequently darkened below subdorsal and above supraspiracular stripe. Prolegs on A3 and A4 reduced. Height and width of abdominal segments greatest through A4 and A5, then smoothly tapering to A10 (especially evident in resting caterpillars). Setae black, long; dorsal setae greater than half length of segment from which they arise. Head green, pale yellow, or orange, sometimes with dark reticulations; setal bases on head sometimes darkened. Larva to 4.5 cm. ! occurrence Agricultural fields and other open habitats. A regular breeding immigrant in Gulf states, but establishing northward temporarily during late summer, rarely as far north as Maryland, but usually arriving too late to breed successfully. Continuously brooded in southern Florida and, in mild years, south Texas. Abundant in Deep South. ! common foodplants Most records from alfalfa, kudzu, peanut, soybean, and velvetbean; also hairypod cowpea (Vigna luteola), locust, snoutbean (Rhynchosia), and other legumes. ! remarks At rest the anal prolegs are directed straight backward, almost parallel to the body axis. During warm periods development can be exceeding fast, taking only 21 days from egg hatch to adult emergence (Anawondu and Johnson 1986). When handled, the caterpillars wriggle violently. The Velvetbean Caterpillar is an infrequent pest in South and Central America and across our southern states. Late instars are green at low densities but take on greater amounts of black as densities climb, with nearly black caterpillars occurring during outbreaks (as shown in image to right). Physical contact with other larvae is the principal trigger that induces the transformation, although the hostplant, and perhaps other environmental conditions, also affect expression (Anawondu and Johnson 1986). Dark-form caterpillars reach the pupal stage more quickly, in part because they stop feeding and pupate at a smaller body mass (Anawondu and Johnson 1986). All of this suggests that the inducible black forms in Velvetbean Caterpillar and other owlets, e.g., the Cotton Leafworm (Alabama argillacea) and Eight-spot (Amyna axis), are

Eulepidotines Subfamily Eulepidotinae 191 an adaptive response, perhaps because darker larvae enjoy thermal advantages that allow them to better compete for dwindling food supplies during outbreaks. This largely tropical moth is a strong migrant (Johnson 1995) that commonly moves into our Mid-Atlantic and Midwestern states, with adults showing up as far north as southern Canada by autumn. We have had no success obtaining eggs from latesummer (Georgia) and fall (New Jersey, North Carolina) migrants—even when held and fed for weeks—and suspect that such individuals are in reproductive diapause. No stages are known to be cold hardy. Kudzu is an important “wintering” host for larvae (Buschman et al. 1977) in those areas of the Deep South that escape freezing temperatures. The larva’s shape, proleg complement, green-to-black polyphenism, wriggling response, and migratory nature suggest that the moth would be better classified in or near the Scoliopteryginae.

Curve-lined Owlet Phyprosopus callitrichoides ! recognition Fantastically bizarre and wholly uncaterpillarlike: long tentacular appendage over A2, and smaller recurved extension from dorsum of A3. Ground color rusty with darker brown markings, and Y-shaped spot extending up from anterior prolegs. Body and head densely covered in fine setae. Prolegs missing on A3 and A4. Larva less than 3.5 cm. ! occurrence Barrens, shrubby fields, powerline right-of-ways, woodlands from Missouri to Ohio and New Hampshire, south to Florida and Texas. Probably two broods in Maryland and New Jersey with mature caterpillars from late June onward; perhaps three broods in southern Missouri, and presumably more southward. Common. ! common foodplants Greenbrier. ! remarks The caterpillar is a dead leaf mimic. It quavers from side to side when disturbed, much like a leaf would do in a breeze. Similar dorsal tentacles occur in the Filament Bearer, an inchworm (for example, see Wagner 2005: 194), as well as other unrelated taxa in the Neotropics and Old World. A cocoon, with “measured” and excised stems woven into the outer walls, is shown in the inset. The pupa overwinters.

192

Elves Family Euteliidae

T

his tropical family includes seven widespread eastern moths; additional species occur in Florida and Texas. The family shares similarities with Baileyas (Nolidae: Risobinae): the facies of the larvae are alike, with prepupae turning pinkish red, both incorporating leaf fragments into the walls of their cocoon, and the pupae lacking cremasters. Caterpillars tend to be green, stocky, and thickest about the thoracic segments, with a flurry of creamy to white spots. The large head is partly withdrawn into the thorax. Legs are small, at least in proportion to the body. On A1 the SV group is bisetose or trisetose. SD1 is located forward of spiracle on A8 (Forbes 1954). Crochets are of one or two lengths (e.g., in Paectes the end crochets are reduced in size). Prolegs of Paectes include a fleshy lobe that curls upward both fore and aft.1 All but two of the species treated here feed on members of the sumac family (Anacardiaceae); subtropical and tropical euteliids feed on a number of additional plant families. Development is rapid, and both eutelia and paectes caterpillars can pass through five instars in just three weeks. Larvae are cannibalistic (at least in captivity). Elves overwinter as pupae, usually in a cocoon spun in leaf litter.

Dark Marathyssa Marathyssa inficita J recognition  Yellow-green with swollen thorax, body appearing smooth and hairless. Speckled with creamy spots, especially below subdorsal stripe that begins over T2. Larva to 2.5 cm. J occurrence  Shrubby fields, woodland edges, and other open habitats from Manitoba to Nova Scotia, south to Florida and Texas. Two or more generations, with mature caterpillars from May to October; three or more broods in Missouri. Common. J common foodplants  Sumac (Rhus); our collections from skunkbush (R. trilobata), smooth (R. glabra), staghorn (R. typhina), and winged (R. copallinum). J remarks  Search for caterpillars by turning twigs and branches and examining the underside of leaves, especially in late summer. Euteliids are easily taken with beating sheets. Many have narrow periods of seasonal activity, so expect caterpillars to be common for only a few weeks over a given brood. At rest the head is partially retracted into the first thoracic segment.

1 We coin the name elves for the family because the prolegs curl upward in much the same way as elfin boots in many renderings; moreover the moths are small relative to most noctuoids.

Elves Family Euteliidae 193

Light Marathyssa Marathyssa basalis J recognition Stout, boldly-marked sawfly mimic: waxy white with bold black and yellow spotting. Pale orange head with large black spot to either side of triangle withdrawn into thorax. T2–A8 with large anterior subdorsal spot and much smaller, black posterior subdorsal spot; black (spiracular) lateral spots on T1 to A8. Yellow supraspiracular patch over anterior half of thoracic and abdominal segments. Black thoracic shield and anal plates. Larva to 2.5 cm. J occurrence Coastal scrub, woodlands, and forests from southern Canada, south to Florida and Texas. One generation over much of East with mature caterpillars in late spring. Common, but usually less so than Dark Marathyssa (M. inficita). J common foodplants Poison ivy. J remarks The single black spot on the head, robust build, waxy coloration, and rounded rear are all suggestive of a sawfly. Its lethargic behavior and propensity to drop from the host and coil into the shape of a C are also consistent with the demeanor of many sawflies. Larvae are new leaf specialists. In rearing containers few larvae rest on foliage; more commonly caterpillars perch on stems, paper toweling, and the sides of the container. It is noteworthy how different our Marathyssa larvae are in appearance, given the close resemblance of their adults. Predictably, they also differ in habit: caterpillars of the Dark Marathyssa (M. inficita) are cryptically colored, cling more to their host, and do not coil into a C shape when molested.

194 Elves Family Euteliidae

Beautiful Eutelia Eutelia pulcherrimus J recognition  Elongate, rather cylindrical, olive green caterpillar, with dirty brown-green and white agateware mottling. Middorsal stripe broken into irregular brown-green spots; markings over A2–A6 broadly keyhole shaped. A8 modestly swollen. Setae short, somewhat bristlelike, often borne from contrasting white pinacula. Spiracles orangish. Upper portion of head smoky black; frons and lower portion of head pale and largely unpigmented. Larva to 3 cm. Middle instars markedly different: lime green, unmarked, translucent, and sawflylike (lower right). J occurrence  Swamps, marshes, wet meadows, rich woodlands and forests from eastern Missouri, southern Michigan and Ontario, and Massachusetts, south to Florida and Texas. One generation with mature caterpillars in April and May southward; June (mostly) to early July in Northeast (but see below). Generally scarce but sometimes common locally. J common foodplants  Poison sumac and poison ivy (but see Remarks). J remarks  Our images are from a cohort of ex ova caterpillars reared by Tim McCabe. The female was taken at light in Albany, New York: her eggs, laid about the 1st of June, hatched on the 15th, and yielded fully mature larvae during the first week of July. Initially, the larvae accepted and established on poison ivy, but many straggled and mortality was high; after the larvae were switched to poison sumac, the caterpillars developed quickly with only minimal loss. Nevertheless the moth occurs regularly in habitats with no poison sumac and it has been successfully reared on poison ivy (Ferguson 1975). Age of the foliage may be important, and we note that its Arizona counterpart, which goes under the same name but is specifically distinct, feeds only on the youngest leaves of poison ivy. Contrary to the supposition in Forbes (1954) that the caterpillar of the Beautiful Eutelia “presumably resembles that of E. adulatrix, its European congener,” the Beautiful Eutelia is far more sawflylike in appearance. The adult is a fine animal, well deserving of both its common and scientific appellations. We cannot explain Forbes’s (1954) statement that larvae have “been beaten from Rhus as early as February in the vicinity of New York City.” Phenologies given above are based on the assumption that larvae follow the adults, and on our belief that pupae overwinter.

Elves Family Euteliidae 195

Large Paectes Paectes abrostoloides J recognition  Bright green with prominent yellow spotting and well-developed, yellow spiracular stripe. Prolegs reminiscent of elfin boots, curling upward, forward, and rearward. Leading edge of prothorax yellow. Larva to 2.5 cm. J occurrence  Woodlands and forests from Missouri to New York, south to Florida and Texas. At least two generations with mature caterpillars from May to November. Very common. J common foodplants  Sweet gum. J remarks  Paectes are easy to find as caterpillars because they perch exposed on the underside of leaves and their feeding damage is conspicuous. Early instars chew characteristic “windows,” removing patches of tissue almost to, but short of, the upper leaf surface. Larvae are also easily sampled with a beating sheet. On two occasions we have inadvertently collected Paectes larvae by bagging terminal foliage (for other caterpillars). The Large Paectes migrates out of range. Mark Mello (unpubl. data) has collected nearly a dozen specimens from southeastern Massachusetts, spanning dates from July to October, yet the nearest native stands of the larval foodplant occur no closer than the western end of Long Island and New Jersey—and all moths taken by Mello appeared well flown. The moth’s seasonal phenology in New Jersey is consistent with that of other migrants: adults occur from April (infrequent) into December and are usually common by July, peaking (at least at bait) in October and November. Prepupal larvae become reddish (right).

Clouded Paectes Paectes nubifera Central and southern Florida. Multiple broods with mature larvae through much of growing season. Locally common. One larva reared from live oak— late instars fed on new growth, carving out cavities along stem (Lyle Buss, unpubl. data).

196 Elves Family Euteliidae

Eyed Paectes Paectes oculatrix J recognition Like other paectes: bright lime with conspicuous yellow spotting and bullet-shaped body, tapering rearward of A6. Yellow spiracular stripe mostly broken into spots forward of abdomen. Head sea green and unmarked, partially drawn into T1. Spiracles orange. Larva to 2.5 cm. The early instars are pale and elongate—it is not until the last two instars that the peppering of yellow spots is evident. Three other eastern paectes are believed to feed on sumacs: the Pygmy Paectes (P. pygmaea) is a widespread species that we have reared from winged sumac (see Remarks, next page); the Blue-gray Paectes (P. near pygmaea) is a western species that enters our coverage area in the Great Plains states (see next page); and P. abrostolella, which occurs from southern Ontario to at least Missouri and Kentucky (Chris Schmidt, pers. comm.). The larva and life history of P. abrostolella are unknown, although we suspect it is using Rhus aromatica and perhaps other low sumacs. J occurrence Woodlands and forests from southern Canada to Florida and Texas. At least two generations with mature caterpillars from June to September over much of East. Common. J common foodplants Our rearings have come from poison ivy and poison sumac. J remarks Paectes eggs are distinctive: flattened with a central raised area surrounded by a clear ring of colorless shell. The feeding damage of late instars is conspicuous: large patches of tissue are removed and telltale jagged edging flag a caterpillar’s whereabouts. In our experience the larvae are especially likely to be found on climbing vines growing in the shade, 0.25–2 m above the ground. The shiny black spots in the background of the figure are dried poison ivy latex droplets. Larval development is rapid, lasting less than three weeks. One cohort of five larvae from New Jersey proved highly cannibalistic—only two survived to pupation. Prepupae of all euteliids excise dozens of small leaf fragments and meticulously weave these into the cocoon wall (inset). Adults are infrequent at light or bait relative to the numbers of larvae that can be gleaned from nearby poison ivy vines.

Elves Family Euteliidae 197

Pygmy Paectes Paectes pygmaea J recognition  Similar to the preceding Paectes. Larvae under 2 cm. Other than its hostplant associations, we are unaware of characters that reliably separate the larva from that of the Eyed Paectes (P. oculatrix). It lacks the yellow prothoracic edging of Paectes near pygmaea (see below). Presumably the caterpillar also resembles P. abrostolella, a prairie species whose early stages and life history have, vexingly, eluded us (see also previous page). J occurrence  Fields, powerline cuts, coastal strand communities from Michigan, southern Ontario, and eastern Massachusetts, south to Florida and eastern Texas. One principal generation in the Northeast with mature caterpillars in July; two or three generations over much of East. Locally common. J common foodplants  Winged sumac (but see below). J remarks  The Pygmy Paectes purportedly feeds with the Large Paectes (P. abrostoloides) on the leaves of sweet gum (Forbes 1954, Covell 2005). The use of both sweet gum and winged sumac by a euteliid seems exceptional to us and may indicate that the sweet gum records are based on erroneous identifications. We had a clutch of P. pygmaea larvae from Cosby, Tennessee refuse sweet gum. Our records of the Pygmy Paectes in Pennsylvania and all of those from New England are from locations outside areas where sweet gum grows. The matter could be laid to rest by setting up (or sleeving) gravid females for eggs, and then transferring newly hatched larvae onto leaves of sweet gum, various sumacs, or preferably both. Both sexes come to light; curiously, the sex ratio is highly skewed towards females, so much so that we wonder if the mating flight is occurring at dusk.

Blue-gray Paectes Paectes near pygmaea

Wyoming to central Texas, west to California. Presumably multiple broods with mature larvae through growing season. Our records from prairie sumac (Rhus lanceolata), skunkbush sumac (R. trilobata), and evergreen sumac (R. virens). Anterior of prothorax edged with yellow and bearing four minute black dots.

198

Nolas and Punkies Family Nolidae

T

his heterogeneous family is represented in North America by five subfamilies: the Nolinae, Chloephorinae, Collomeninae, Risobinae, and Afridinae. Including undescribed taxa, about two dozen species occur in our region. The family, and in particular the Nolinae, is richly represented in lowland rainforests of the Neotropics, where it is not uncommon to find 20 or more species flying at a single station. Larvae frequently bear an exceptionally long seta from the D and SD1 setal warts at either end of the body (T1–T3 and A7–A9). At least three subfamilies (Nolinae, Risobinae, and Afridinae) painstakingly weave numerous, small pieces of “plant” tissue into the walls of their cocoons—a complex and fascinating process to watch. Nolid pupae lack cremasters. All are foodplant specialists, mostly on shrubs and trees, although a few use forbs and grasses. Afrida feed on algae and lichens. Because nolids are so diverse in form and habit, we introduce each subfamily separately.

Nolas and Punkies Family Nolidae: Subfamily Nolinae

This is the largest of the North American subfamilies with nine or ten species in the East. Their small, hairy caterpillars lack prolegs on A3. Larvae bear numerous, barbed secondary setae, clustered on verrucae (setal warts). The D1 and D2 setal clusters are fused. Warts on thoracic and posterior abdominal segments (especially A9) often include a long seta. Larvae of some Old World species retain the head capsules after a molt, with older caterpillars carrying a concatenated set of previous

instar head capsules over the thorax (McFarland 1978). Caterpillars sometimes spin a belay line to escape disturbances and to move to new feeding sites. While the caterpillars are commonly found inside leaf shelters, it is unclear to what extent they seek out the constructions of other lepidopterans, or if noline larvae can “roll their own.” Dyar (1899) records six instars for two eastern species. The prepupal larva excises small elongate strips of bark from the vicinity of the cocooning site and then weaves each piece into the wall of its cocoon (see that of Nola clethrae on page 202).

Confused Oak Punkie Meganola minuscula

J recognition  Dorsum mostly smoky gray and white with dark saddle over T2–T3, A3, and A7–A8. Thin, gray middorsal stripe. Larva to 2 cm. J occurrence  Barrens, woodlands, and forests from extreme southern Canada to Florida and Texas. Two generations over much of East with mature caterpillars from late May into September. Common. J common foodplants  Oak. J remarks  Dyar (1892) noted that the larvae often hide in dry, curled leaves or along bark. Certainly their coloration is well suited for the latter. Evidently, the pupa overwinters in a gall-like cocoon spun along a twig. There is considerable variation among nolines as to which stage overwinters: the Sharp-blotched Nola (Nola pustulata) is thought to overwinter as an egg; the Once-charred Punkie (M. spodia) and Blurry-patched Nola (Nola cilicoides) overwinter as partially grown larvae; other eastern nolines are believed to overwinter as pupae.

Nolas and Punkies Subfamily Nolinae 199 Sharp-blotched Nola Nola pustulata

Central New York to Massachusetts, south to Georgia (mountains) and Texas. One generation with mature caterpillars from mid-May to mid-June over much of range, earlier southward. Locally common. Maleberry (Lyonia ligustrina). Evidently overwinters as an egg. Caterpillar feeds on newly expanding leaves in spring. See McCabe (1996) for details on its life history. Penultimate (right) and final instar (below).

Live Oak Punkie Meganola new species

At least central Florida; taxonomy and range unclarified. Figured larva reared by Lyle Buss from live oak. Confused with M. phylla in many collections (Hugo Kons, unpubl. data).

Once-charred Punkie Meganola spodia [Ashy Meganola]

Wisconsin to southern Quebec, south to Georgia. Single generation with mature caterpillars in early spring (Butler 1989). Locally common. Red and white oaks.

200 Nolas and Punkies Subfamily Nolinae

Two-spotted Oak Punkie Meganola phylla [Coastal Plain Meganola]

J recognition Tan, yellowish to pinkish, often with dark saddle over A3 (and

A7). Middorsal stripe pale, made up of two, closely set stripes. Setal warts large, often pinkish. Larva less than 1.5 cm. A number of other oak-feeding nolines occur in the East: the larva of M. spodia is yellowish, pinkish to pale green with a broad, creamy, middorsal stripe (page 199). It overwinters as a partially grown larva, and then matures on early spring foliage (Butler 1989). Lyle Buss sent us images of a Meganola that he reared from Gainesville, Florida on live oak that appear to be M. phylla (inset). Nola ovilla is also reported from oak. We suspect its larva would be less dorsoventrally flattened than those of a Meganola, and would have prominent dorsal warts bearing clusters of pale secondary setae as do its congeners. J occurrence Barrens, woodlands, and forests from Wisconsin, southern Ontario and New England, south to Florida and Texas. Two generations over much of East with mature caterpillars from June to October, although the second generation may be partial; additional broods likely southward. Locally common. J common foodplants Beech, and red and white oaks. We have records from dwarf chinquapin oak, blackjack oak, and white oak; our record of beech is based on collections made by John Lill. J remarks A Two-spotted Oak Punkie female confined with white oak laid her eggs on leaf undersides, placing them along midribs. Look for caterpillars on the undersides of leaves, on or near necrotic or scraped patches of tissue, or perched along the midrib. They also seek refuge in the shelters, usually abandoned, of other caterpillars. Like other nolines, the prepupal caterpillar labors over its cocoon for hours, weaving minute, individually excised strips of bark from a nearby petiole or twig into the outer wall of the cocoon. The finished cocoon, higher at one end and spun along a petiole or twig, is highly cryptic. The pupa overwinters.

Nolas and Punkies Subfamily Nolinae 201

Blurry-patched Nola Nola cilicoides J recognition Smoky yellow-orange with black crescents beneath the dorsal, and sometimes subdorsal, setal warts. Head prominently marked with black. Larva to 1 cm, or just exceeding that length. A small undescribed Nola, closer to N. pustulata (page 199) than N. cilicoides in appearance occurs in seepage areas and other wetlands in savannas and other coastal plain areas of the Southeast. Its early stages are unknown. J occurrence Wet meadows, marshes, edges of watercourses, mesic woodland edges, and forest clearings from southern Canada to northern Florida and Arkansas. In Connecticut with small spring brood and larger summer brood; mature caterpillars in spring and then again in July; three broods in Georgia (James Adams, pers. comm.) with adults into September. Locally common. J common foodplants Fringed loosestrife (Lysimachia ciliata) and perhaps related species, but not whirled loosestrife or swamp candles. J remarks This insect has a fascinating life history. In the spring, (approximately third instar) larvae that have overwintered initiate feeding by boring a tiny hole into the side of a fringed loosestrife shoot; after coring the shoot, they then move distally to feed on flower buds, opened flowers, and young leaves. Wilted tops are, so to speak, a dead giveaway for the presence of larvae. Early instars were not observed to engage in “pithing,” but instead tunneled into leaves, forming full depth blotch mines (inset). Larvae from August collections in Connecticut exited their mine, fashioned a weak circular cocoon between two overlapping leaves, and then entered a nine-month state of diapause. To have both pithing and mining behaviors comingled in the life history of a single species, qualifies N. cilicoides for recognition as one of the East’s most remarkable moths. A more detailed life history is given in Wagner and Connolly (2009).

202 Nolas and Punkies Subfamily Nolinae

Sweet Pepperbush Nola Nola clethrae J recognition Dusky green, gray, brown, or charcoal with dark middorsal stripe. Body with mottling above level of spiracles; ground color paler and essentially unmarked below. Setal warts pale, each with dark ring about base. Prothoracic shield with dark brown to black, bat-shaped plate. T1, T2, and A9 with especially long setae. Head with dark splotch over each lobe. Larva to 1.5 cm. The middle instars greenish (upper inset). J occurrence Floodplains, riparian corridors, wetlands in pinelands, wooded swamps, and mesic woodlands from central Massachusetts, south to Georgia, Alabama, and Texas; most records from Atlantic Coastal Plain, although numerous records from mountains. At least two generations with mature caterpillars from May to October; spring brood numerically the largest. Often abundant. J common foodplants Coastal sweetpepperbush (Clethra alnifolia) and presumably mountain sweetpepperbush (C. acuminata). J remarks In New Jersey swamplands and pinewoods, adults may account for more than 10% of the individual moths in light trap samples for the year, and occasionally account for more than 40% on spring nights. The prepupal larva individually attaches strips of bark to the outer surface of the elongate cocoon (lower inset). The larva frequently enters and feeds within the shelters of other lepidopterans; in Connecticut, caterpillars of the Sweet Pepperbush Nola can be found by unrolling the abandoned shelters of a Herpetogramma (pyralid leafroller) that are seemingly omnipresent on sweet pepperbush. The fortunes of the Sweet Pepperbush Nola may even be linked to those of the pyralid as nearly every shelter is occupied by the noline inquilines in some years—one wonders what the predation rate on the Nola caterpillars would be without the foliar engineering efforts of the pyralid. The pupa overwinters.

Nolas and Punkies Subfamily Nolinae 203

Three-spotted Nola Nola triquetrana J recognition Small, hairy with yellow-orange ground color. Orange dorsal and subdorsal setal warts issuing from black spot. Thoracic shield black, shiny, divided by pale middorsal line. Lateral setal tufts, especially those towards end of body, with one extremely long seta. Larva to 1.5 cm. Middle instars smoky green and lack black spotting. J occurrence Mesic woodlands and forests from southern Canada to Florida and Texas. One generation with mature caterpillars from May to June over much of East. Common. J common foodplants Witch hazel. J remarks Caterpillars live in leaf rolls, sometimes with other larvae if population numbers are high. Presumably the caterpillars do not spin their own shelters but instead enter the rolls and leaf shelters of other lepidopterans. The abandoned shelters of Olethreutes and other tortricid leafrollers are especially likely to yield larvae of the Three-spotted Nola. The caterpillars are skeletonizers, removing all but the vascular tissue over sections of an occupied leaf (upper inset). Where the moth is abundant, such as in the foothills of the Appalachians, the caterpillar’s feeding damage may significantly affect the appearance of witch hazel. Larvae sometimes spin down on a thread of silk when alarmed, or when searching for a new feeding site. The pupa overwinters in a highly cryptic, spindle-shaped cocoon, spun along a twig (lower inset).

Sorghum Nola

Nola cereella (= N. sorghiella) Resident in Deep South; northern limit of yearround range unclarified. Adults commonly occur north to Missouri, Ohio, and southern New England as summer and fall migrants. Multiple generations throughout growing season. Common southward. Caterpillars feed gregariously in heads of grasses (Franclemont 1960). An occasional pest of sorghum; known in applied literature as Sorghum Webworm.

204 Chloephorines Subfamily Chloephorinae

Chloephorines

Family Nolidae: Subfamily Chloephorinae This largely Old World tribe is represented in our area by three genera: Garella, Iscadia, and Nycteola. Larvae of Nycteola are semitranslucent green with long, thin setae, some considerably longer than the segment that bears them. The prolegs on A3–A6 are equally developed, and there are two SV on A1. The apex of the spinneret bears a flap to either side (Beck 1999). Our species feed on members of the willow family (Salicaceae). The cocoon is diagnostic—right. The larvae of Garella, also with hypertrophied setae, are sometimes internal feeders in nuts and shoots (Sugi 1987) and galls (see below).

Ash-colored Nycteola Nycteola cinereana

Boreal species; transcontinental in Canada, south in East to New York (mountains only) and northern New England. One generation with mature caterpillars in late spring and early summer. Uncommon as an adult. Poplar. Note silk webbing below caterpillar.

Sienna-dotted Nycteola Nycteola metaspilella J recognition  Lime to yellow-green, smooth, elongate, with long thin setae; longer dorsal setae greater than twice length of segment that bears them. Setae especially long rearward. Body semitranslucent with thin, spiracular stripe that is actually the internal tracheal trunk (and accentuated in flash images). Spiracles pale brown, minute. Larva to 2.5 cm. Two other Nycteola occur in our region. The Frigid Nycteola (N. frigidana) is believed to be principally a willow feeder; it is a boreal, transcontinental moth that ranges south in the East into mountainous areas of New York and New England. In Canada and northern New England, it is joined by the Ash-colored Owlet (N. cinereana), a poplar feeder (above). We are unaware of characters that will distinguish the caterpillars of our three eastern Nycteola. Fletcher (1959) was the last to revise the genus. J occurrence  Edges of watercourses, fields, and forest edges from southern Canada, south to Florida (Don Lafontaine, pers. comm.). At least two generations over much of East with mature caterpillars in June and July, and again in August and September. Infrequently encountered as adult. J common foodplants  Our records from big-toothed aspen, balsam poplar, and various willows. J remarks  Early instars live in small groups beneath a loose webbing of silk spun over the upper surface of a new leaf (opposite page), or in a shelter made

Chloephorines Subfamily Chloephorinae 205

by weaving together two or more leaves. Our captive larvae (sleeved on aspen) preferentially chewed into leaf petioles even when leaves were available. Last instars tend to be solitary, although occasionally two caterpillars will be found together sharing a single leaf. We suspect that the exceptionally long setae and webbing work in concert to provide the larva with an extended sensory network for the early detection of natural enemies in a manner analogous to that for shelter-dwelling metalmark moths (Rota and Wagner 2008). McCabe (2001) observed five Frigid Nycteola (N. frigidana) adults probing the remains of dead bugs on the windshield of his car on a night when no members of the genus came to traditional moth baits or light. Adults of the Oak Nycteoline (N. revayana) in Britain are occasionally seen at flowers (Porter 1997). The white, flocculent cocoon, higher at one end, is reminiscent of a spider egg “nest” (see top of page 204). Adults overwinter.

206 Chloephorines Subfamily Chloephorinae

Merchant Punkie Garella nilotica J recognition Resembles a microlepidopteran in size and aspect. Blackish prothoracic shield sometimes bearing dark T-shaped pattern with middorsal and posterior portions more darkly pigmented. Mostly cream or greenish to yellowbrown or red-brown with darkened supraspiracular area and peppering of minute brown to reddish spots, again mostly above spiracles. Setae long, especially rearward where they exceed length of two adjacent abdominal segments. Head black-brown. Larva to 1 cm. J occurrence Orchards, parks, city streets, riparian corridors, and open second-growth areas. Southern Canada, south through our region into Caribbean and Mexico, but presumably as a migrant northward. Present throughout growing season in parts of Florida, but especially numerous from April to June. Common in south Florida and Texas; frequent summer migrant as far north as Missouri and Long Island. J common foodplants Pest on black olive or gregorywood (Bucida buceras) in south Florida; also almond, button mangrove (Conocarpus erectus), rhododendron, willow, and many other woody plants. J remarks G. nilotica is a nearly cosmopolitan insect, recorded from six continents and many remote islands. Its diet and feeding habits are extraordinarily varied. The larvae may feed externally on foliage, web up flowers, feed internally in developing fruits, or tunnel through galls. The caterpillar is a common pest on black olive in southern Florida and through the Caribbean, that sometimes defoliates entire trees. During infestations, falling debris, and especially larval fecal matter, can stain homes, sidewalks, cars, and other surfaces (Caldwell 2008). Our image shows a caterpillar and its feeding damage inside a “string bean” gall—induced by an eriophyid mite—on the developing fruits of Bucida. During outbreaks, larvae are commonly encountered spinning down on silk threads, and as a consequence, they have acquired the name of “Bungee Caterpillar” in parts of Florida (Caldwell 2005). An ovoid, silky white cocoon—devoid of distinctive design elements common to other nolids—is spun among debris in or about the feeding site (right). The common name Black Olive Caterpillar, used by entomologists in Florida, is a misnomer given that the insect is polyphagous—we suggest the Merchant Punkie as an alternative, reflecting our belief that its global distribution relates to early sea trade.

Baileyas Subfamily Risobinae 207

Baileyas

Family Nolidae: Subfamily Risobinae This subfamily is represented in the East by seven species. The green larvae are somewhat dorsoventrally flattened, possess a strong subdorsal line, short midabdominal prolegs that are rolled at their ends, and moderately long anal prolegs that splay out behind the body. The SV group is bisetose on A1. Prior to pupation the larvae turn olive to reddish brown. Su-

Sleeping Baileya Baileya dormitans

Wisconsin, southern Canada, and Maine, south to Florida and Texas. In Northeast, mature caterpillars mostly in July and August with small second generation caterpillars in fall; two generations in coastal North Carolina. Common. Hickory and walnut for certain and probably also butternut and pecan.

Doubleday’s Baileya Baileya doubledayi

Across southern Canada to northern Florida and Texas, but local and scarce southward. Two generations over much of East with mature caterpillars in late June and July, then again in fall; at least three generations in coastal North Carolina. Uncommon. Alder.

perficially, Baileya caterpillars resemble those of some euteliids—so much so that even seasoned lepidopterists sometimes confuse the two. Reported foodplants include members of the birch and walnut families; hosts for the two recently described species from the Southeast are unknown. The pupa overwinters in an elongate cocoon into which dozens of excised leaf fragments are interwoven (see image of Doubleday’s Baileya, below).

208 Baileyas Subfamily Risobinae

Pale Baileya Baileya levitans J recognition Pale green, stout, mostly parallel-sided with prominent, yellow subdorsal stripe that may be broken at midsegment as well as between adjacent segments. Head yellow-green to sea green, shiny, unmarked. Prolegs with faint pinkish cast, distal portion flaring outward, but not as pointed forward and rearward as some other baileya. Spiracles small, pale yellow-orange, and inconspicuous. Larva to 2.5 cm. We are unaware of characters that distinguish caterpillars of the Pale Baileya from those of the Sleeping Baileya (B. dormitans) with which it co-occurs on both hickory and walnut. J occurrence Woodlands and forests from southern Canada to Florida and Texas. Two generations through much of East, presumably more in Gulf states with mature caterpillars from May onward. Common, but absent from portions of Atlantic Coastal Plain. J common foodplants We reared a wild larva from shagbark hickory, and a clutch of ex ova larvae on pignut hickory and black walnut (all Juglandaceae). J remarks The body wall is transparent. With a lens one can watch hemolymph being pushed forward into the thorax and head by the heart, or make out the arrangements of major spiracular trunks that lead away from each of the spiracles. Baileya are lethargic or at least slow moving insects with no obvious escape strategies. Even when molested they merely amble away, seemingly without conviction. Dozens of excised plant fragments are woven into the outer wall of the cocoon. In Connecticut the second brood is partial, with some pupae from the first generation holding over until the following spring.

Small Baileya Baileya australis

Michigan, southern Ontario, and southern New York, to Florida and Texas. Two generations northward, at least three in South with mature caterpillars from May onward. Common. Black walnut.

Baileyas Subfamily Risobinae 209

Eyed Baileya Baileya ophthalmica J recognition Stout lime green caterpillar with prominent subdorsal stripe. Supraspiracular stripe represented by two or three white to yellow spots on many segments; often with an additional spot or two below and behind spiracle. Head green, shiny, rounded, with subtle mottling. Larva less than 2.5 cm. Easily distinguished from other baileya by its choice of foodplant; however, we caution that there are two recently described Baileya that occur through the Gulf states north to the Carolinas—their hosts, life histories, and caterpillars are unknown. J occurrence Woodlands and forests from Wisconsin to Nova Scotia, south to Florida and Texas. Two generations, with second apparently partial over much of range; mature caterpillars from June to September; in Carolinas with caterpillars mostly in late May and June and again in July. Locally common. J common foodplants American hornbeam, hophornbeam, American hazel, and beaked hazel; records from butternut presumably apply to another species. Covell’s (2005) record of beech probably refers to blue beech (= American hornbeam). J remarks Adults of this species occasionally show up dozens of miles from their foodplants. Baileya caterpillars are easily collected with beating sheets. The resemblance of the caterpillars to euteliids has been mentioned elsewhere. Adult wing venation in the two groups of moths is also similar (Forbes 1954). Furthermore, the two groups of moths both weave leaf fragments into the walls of their cocoons. The caterpillars are sometimes mistaken for those of a Hypena (= Bomolocha) with which they frequently co-occur because of overlapping host preferences, especially during late summer and early fall. The two are easily distinguished by many features: e.g., the complete complement of prolegs on A3–A6 in Baileya (the proleg on A3 is greatly reduced or absent in Hypena). The animated wriggling response of all our snouts will immediately distinguish the two. Prepupal larvae chew free elongate strips of leaf tissue and layer these into the cocoon wall.

210 Concanas and Kin Subfamily Collomeninae

Concanas and Kin

Family Nolidae: Subfamily Collomeninae This tropical tribe is represented in our area by three genera and four species. The first two sets of prolegs are greatly reduced and the SV group is trisetose on both A1 and A2. Several New World species have been reared from Malpighiaceae,

Locust Berry Concana Concana mundissima

South Florida into Caribbean. Year-round, but most common when locust berry is producing new foliage. Uncommon. Locust berry (Byrsonima lucida).

Barbados Cherry Concana Concana near mundissima

South Texas. Our larval collections taken over fall and winter months; presumably active year-round. Locally common. Dwarf Barbados cherry (Malpighia glabra).

Buttonwood Punkie Motya abseuzalis

Florida south into Caribbean. Active throughout growing season. Uncommon. Button mangrove (Conocarpus erectus).

including two of the three species figured here. Recent molecular studies suggest Concana and related genera should be reclassified near Bagisarinae (Reza Zahira unpubl. data).

Lichen Punkies Subfamily Afridinae 211

Lichen Punkies

Family Nolidae: Subfamily Afridinae This tropical subfamily is represented in our region by a single genus with three species. The minute larva bears distinctive droplet-bearing setae. Caterpillars feed on lichens and algae.

Gulf Lichen Punkie Afrida ydatodes [Dyar’s Lichen Moth] J recognition  Very small, gray to brown, mottled with black. Long setae borne from raised pinacula. Dorsal as well as some lateral setae glandular, with secretion accumulating at apex. Body thickest through thorax and A1; dorsum of T3 and A1 bearing black V-shaped mark. Black middorsal patches over A7–A8. Relatively thick setae borne from raised, white, conelike warts. Larva to 1 cm. An even smaller lichen punkie occurs in south Texas—A. minuta. J occurrence  Mesic woodlands and tropical hardwood hammocks. North Carolina south to Florida, and west to Texas. Records in Kimball (1965) suggest year-round generations in southern Florida. Common. J common foodplants  Lichens. J remarks  The blue-green cast to the larva figured here is indicative of the caterpillar’s prepupal status. While lichen punkies (Afrida) have been classified as arctiids by some workers (e.g., Franclemont 1983), Kitching and Rawlins (1998) regarded them to be nolids and proposed a new subfamily for the moths. The cocoon—the hemispherical swelling along the twig—is phenomenally cryptic. The caterpillar painstakingly incorporates pieces of lichen, moss, and algae that it has chewed free from adjacent areas of bark into the wall of the cocoon. A similar cocooning behavior is exhibited by nolines and risobines, both of which add bark strips to the walls of their cocoons. The individual shown here was beaten from an ash tree in Houston, Texas, in the latter half of April. The caterpillar fed on lichen for only a day before it began construction of its cocoon. We do not know which stage overwinters, but it is likely to be as a half-grown larva, given the early date that the figured individual was collected.

212

Trifine Owlets Family Noctuidae1

A

s currently circumscribed, the Noctuidae includes 20 subfamilies in North America, all but two of which are represented in this guide. More than 2500 species have been recorded north of Mexico, and thus trifines account for nearly 70% of the Nearctic Noctuoidea. No larval characters uniquely unite the members of the family. In most there are only two SV setae on A1 and A2, and SD1 is hairlike (fine) on A9, although some (early diverging) subfamilies have three SV setae and an undifferentiated SD1 seta on A9 as in erebids. The latter character is somewhat variable both within the Erebidae and Noctuidae, and should be used in combination with other features. The first two pairs of abdominal prolegs are usually absent in five basal trifine subfamilies (Acontiinae, Bagisarinae, Cydosiinae, Eustrotiinae, and Plusiinae). More detailed diagnoses and life history synopses are given in the boxes that introduce each subfamily of this varied and extraordinarily successful lineage of moths.

Loopers

number, and mouthparts. Godfrey’s (1987) treatment based on mandibles and chaetotaxy is reliable, but includes only pest Approximately 50 plusiines occur in the East. Contrary to the species. Lafontaine and Poole (1991) provide larval descripdiversity trends seen across most animal and plant groups, tions for nearly all of the eastern species. Be forewarned that plusiine diversity increases with latitude, due mostly to the ground coloration and expression of the stripes varies among richness of two boreal genera, Autographa and Syngrapha. individuals—if crowded, many plusiines develop dark or meAs caterpillars, plusiines are distinguished by the absence or lanic forms (Long 1953; see discussion under Rachiplusia). rudimentary nature of the prolegs on A3 Conversely, plusiine larvae reared on artiand A4 (with the exception of Abrostola ficial diets can be largely devoid of green and Mouralia). Crochets appear biordinal pigments. Even the lateral black stripe on (inset) except in Plusia.2 In several genera, the head comes and goes within a spedeep intersegmental incisures give the abcies (contrary to keys that we have seen). domen a scalloped profile. Most are green Charles Bordelon reports that caterpillars with two to three (sometimes four) pairs of commonly visit and consume sugar bait, white longitudinal stripes. Commonly the especially during times when flowers are rump is arched or swollen and the anal proin short supply. He reared one late instar legs are comparatively short. The integuthrough to the adult stage solely on a bament is frequently spinulose: in the Bilobed nana-beer mixture that he was using in his Looper (Megalographa biloba) the cuticle is bait traps. Plan to rear out adults if certain so densely set with spinules that the cateridentity is required. pillar appears fuzzy. The head is small, at Many plusiines, including all migratory least in proportion to the girth of the thotaxa, are generalists that consume herbs, rax. Thoracic legs are long, and sometimes grasses, and low-growing woody plants. blackened. Most rest with a loop thrown Loopers that feed on plants with milky into the body and the head pushed down to latex and other kinds of toxic sap, such as ! Looper proleg showing the substrate (see Gray Looper page 217). wild lettuce and dandelion, are able to do so SD1 on A9 is thin and hairlike (Abrostolini apparently alternating by severing the midrib or girdling the stem, and Argyrogrammatini) or undifferentiat- crochet lengths—see text. ! thereby circumventing the plant’s ability ed (Plusiinae); there are either two or three to deliver latex and sap to the site of feedSV setae on A1. Yellowish testes are visible in the later instars ing injury (see Common Looper, page 224). Rather than feed of male caterpillars. from an edge, most remove patches of host tissue from central Plusiines are frustratingly undistinguished as caterpillars: portions of a leaf. Look for caterpillars by examining leaf uneven identification to genus and tribe may require careful dersides in the vicinity of recent feeding damage, which will examination of microscopic features. Characters for distin- often be found on newer foliage. They are slow, deliberate, and guishing among species, especially in larger genera like Syn- seemingly vulnerable in nature, seldom moving far from their grapha, will require examination of the chaetotaxy, crochet natal site. Some, e.g., Plusia, are cannibalistic (Long 1953).

Family Noctuidae: Subfamily Plusiinae

1 In most trifines, the cubital vein in the hindwing appears three branched because of reduction of the second medial vein (M2). In erebids, euteliids, and nolids, the cubital vein appears four branched, and hence the common name, quadrifines (see Lafontaine and Fibiger 2006 for discussion and several helpful figures). 2 While the hooks appear to alternate in length, in the caterpillars that we examined the alternating “lengths” were due (primarily) to differences in alternating insertion and deflection angles.

Loopers Subfamily Plusiinae 213 Except for Abrostola, which overwinters as pupae, plusi- Among these are a number of crop and garden pests. Adults, ines diapause as middle instars (commonly as third instars). especially of the migratory species, sometimes come to bait, They are difficult to overwinter: try placing caterpillars in a particularly in the fall after natural sources of nectar have covered flowerpot sunk partially into become scarce. Plusiines account for the ground where they will receive many of the strong-flying moths that exposure to rainfall. A light covering one sees coursing through gardens of leaves (over their container) will and visiting flowers at dusk; even temper exposure to low temperatures. while gathering nectar they are wary, Their white to yellow silken cocoon— and quick to fly off when approached. somewhat reminiscent of a spider nest Butterfly bush and other “butterfly (right)—may be spun well above the flowers” are eagerly visited. Blooming ground, commonly on the underside milkweed and viper’s bugloss (Echium of a host leaf. Among their natural vulgare) patches are a good place to enemies are polyembryonic encyrtids, observe nectaring adults. Many speminute wasps whose eggs clonally recies are more reliably observed at produce themselves once inside the flowers than at light (see species achost’s body—thousands of eggs will count for Wavy-chestnut Y, Autogdevelop from a single “parent” egg. rapha mappa, on page 223). High Seemingly the whole of the caterpillatitudes not only hold the promise of lar’s body is replaced by the bodies greater species richness but also offer of the next generation of microscopic an extended period of twilight when wasps (see page 29, top row, center). the adults can be seen without the aid Several plusiines are migratory, of a flashlight. Adults of both sexes are moving northward each spring and ! This cocoon, with caterpillar visible attracted to the floral scent phenylacsummer, and southward in the fall. within, is only half complete. ! etaldehyde (Meagher 2002).

Golden Looper

Argyrogramma verruca (Tribe Argyrogrammatini) South and Central America, Caribbean, and Gulf states but straying northward in fall. Multiple generations with mature caterpillars year-round in southern Florida and Texas. Common southward. Many herbaceous plants including arrowhead, dock, and tobacco; minor pest of field and garden crops. A3 and A4 with minute vestigial prolegs; SD1 pinaculum smaller than spiracle of same segment; pinacula flat. Easily confused with Sharp-stigma Looper Moth (Ctenoplusia oxygramma) (below).

Sharp-stigma Looper Ctenoplusia oxygramma (Tribe Argyrogrammatini)

Southern Canada to Florida and Texas, but only as an occasional summer- and fall-breeding migrant over much of East. Multiple generations with mature caterpillars year-round in southern Florida and Texas. Common southward. Many herbaceous plants; Crumb (1956) lists asters, goldenrod, horseweed (Conyza), and tobacco. Similar to above but with pimplelike (raised) pinacula.

214 Nettle Loopers Subfamily Plusiinae: Tribe Abrostolini

Nettle Loopers

Family Noctuidae: Subfamily Plusiinae: Tribe Abrostolini This small tribe of loopers bears functional prolegs on A3 and A4. The SV group is bisetose on A1, and the SD1 seta on A9 is thin (more slender than dorsal setae). The tribe is represented by two genera in North America: Abrostola and Mouralia. Foodplants include members of the nettle (Urticaceae) and spiderwort (Commelinaceae) families for the two genera, respectively.

Nettle Looper Abrostola urentis

[Speckled Nettle Moth or Variegated Brindle]

! recognition Bright lime green to olive, with prolegs on A3 and A4, and

oblique white subdorsal lines. In darker forms, smoky pattern elements dominate (as in inset image of A. ovalis). A8 strongly humped with raised shelf that bears whitish wart with D2 seta at apex. A1 and A2 often with dark dorsal patch sandwiched between oblique subdorsal lines. Head with dark addorsal and lateral lines; area between sometimes filled with dark reticulations. Spiracles pale with darkened rim. Setae borne from small white pinacula. Larva to 3 cm. Whiteshouldered Nettle Moth (A. ovalis) is distinguished by its bright white to yellow subdorsal spots on A1 and A2 (inset). In our images the smoky green ground color is darker and the white pinacula are larger and brighter than those of A. urentis. A. ovalis occurs from the Great Lakes states to southern Quebec, south to Tennessee and North Carolina (mountains). It feeds on both nettle (Urtica) and woodnettle (Laportea). ! occurrence Fields, wetlands, roadsides, and woodland and forest edges. Transcontinental in Canada, south in East to North Carolina (mountains) and Kansas. One generation northward; two generations in Connecticut with mature caterpillars from June onward. Common. ! common foodplants Stinging nettle, probably woodnettle as well. ! remarks The caterpillar often rests with the first two abdominal segments looped. Individuals reared at high density often darken appreciably in color.

Golden Loopers Subfamily Plusiinae: Tribe Argyrogrammatini 215

Golden Loopers

Family Noctuidae: Subfamily Plusiinae: Tribe Argyrogrammatini Four of the six North American members of the tribe are figured in this volume. The prolegs on A3 and/or A4 are minute (vestigial) and SD1 on A9 is thin (more slender than the dorsal setae). A few are crop pests.

Soybean Looper Chrysodeixis includens ! recognition Green with minute peglike vestigial prolegs on A3 and A4 (only visible with lens) that bear three setae. Dorsal pinacula usually black or white; SD1 pinacula black, often twice size of spiracle of same segment. Integument with minute spinules. Spiracular stripe white, not edged with black above, often continuing to anal proleg. Dorsal pinacula on T2 more closely situated to one another than D2 is to SD1. Head green, shiny; black lateral line (if present) thin, and not including lateral eyes (stemmata). Setae on head arising from black spots or rings. Larva to 3.5 cm. Many identification resources fail to account for the substantial variation in the species and thus are unreliable—rear out adults if certain identification is required. ! occurrence Fields, gardens, greenhouses, agricultural fields, waste places, and other open and early successional habitats from southern Canada to Florida and Texas. Two to many generations with mature caterpillars from May onward in South, mostly late summer and autumn in Northeast. Very common. ! common foodplants A wide array of herbaceous plants, including many crops; e.g., alfalfa, bean, cotton, kidney bean, lettuce, mustard, soybean, tobacco, and tomato. ! remarks Across our region, the Soybean Looper is one of the most commonly encountered plusiines at light, especially in the fall. It is one of several eastern loopers that are migratory, flying northward each spring and summer and fall, sometimes in great numbers. We have observed adults suddenly appear in great numbers at flowers in late September, coincident with storm episodes. On occasion the insect is a pest of soybeans and other legumes. Reverse migration in the fall has not been documented. Like many plusiines, the Soybean Looper is a trencher: the caterpillar can circumvent the vascular systems of latex- and resin-containing foodplants by chewing through the veins that would otherwise deliver sap to the caterpillar’s feeding site—see Remarks for Common Looper (Autographa precationis) on page 224. The insect is thought to be freeze intolerant, capable of overwintering successfully only in the deep South.

216 Golden Loopers Subfamily Plusiinae: Tribe Argyrogrammatini

Cabbage Looper Trichoplusia ni ! recognition Green with minute peglike vestigial prolegs on A3 and A4 (difficult to see without strong lens) each of which bears three setae. Integument covered with microscopic granules (not elongate spinules). Setal base above spiracle (SD1) may be black. Spiracular stripe white or yellow, not edged with black above, fading rearward of A7. Three uppermost setal bases on T2 equidistant. Head green, shiny, without black lateral line. Larva less than 3.5 cm. Antepenultimate instar (upper inset) with more prominent striping and yellow-orange head. Penultimate instar (lower inset). Our main image features a mature caterpillar with subdued markings. (Our three images are of same individual.) ! occurrence Fields, gardens, agricultural lands, and other early successional habitats from southern Canada to Florida and Texas. Three or more generations with mature caterpillars from April onward, at least in South. Common southward. ! common foodplants Many herbaceous plants; crop hosts include asparagus, beans, broccoli, cabbage, cauliflower, corn, lettuce, pea, parsley, tobacco, tomato, watermelon, and many others. ! remarks One of our most widely distributed moths, present in both the New and Old Worlds. In recent years it has been curiously scarce in the Northeast. Although the caterpillars are polyphagous, they are most apt to be encountered on cultivated crucifers; latex-containing composites are also favored. The caterpillar is an occasional greenhouse pest. Most years this species does not overwinter north of the Carolinas (Lingren and Green 1984), and migrants then reestablish the upper midwestern and northeastern populations. The pale Cabbage Looper caterpillar shown to right was reared on artificial diet. In the two insets the caterpillar’s testes are visible.

Loopers Subfamily Plusiinae: Tribe Plusiini 217

Loopers

Family Noctuidae: Subfamily Plusiinae: Tribe Plusiini This tribe contains some 68 North American species. The prolegs are wholly absent on A3 and A4 and the SD1 seta on A9 is setose.

Gray Looper Rachiplusia ou ! recognition Yellow to smoky or lime green with three white to yellow

stripes above; thin, wavy addorsal stripes; somewhat thicker addorsal stripe enveloping D1 pinacula; thin subdorsal stripe running below D2. (This striping subdued in main image, perhaps because larva is prepupal.) Spiracular stripe running along upper edge of spiracles on A1–A7, then dropping down anal proleg. Integument spinulose (visible with magnification). Straw-colored to white spiracles with darkened rims (note contrast especially on A8). Head with black lateral line. Pinacula white. Three SV setae on A2 borne from separate pinacula. Spinules over venter of body usually darkened; distance between V1 setae on A2 nearly equal to distance between V1 and SV2 (Lafontaine and Poole 1991). Larva to 3.5 cm. ! occurrence Fields, gardens, croplands, and other early successional habitats from southern Canada, south through Florida and Texas into South America; principally as stray or autumn migrant north of Gulf states. Mature caterpillars occur year-round along Gulf and in southern Florida and Texas. Common southward. ! common foodplants Many herbaceous plants, including a wide array of crops: clover, corn, cosmos, mallow, mint, ragweed, tobacco, wheat, and Mexican tea (Chenopodium ambrosioides) (see also Eichlin and Cunningham 1978). ! remarks The Gray Looper is among the most commonly encountered loopers in the South. We include it here, in part, to underscore how confoundingly similar our eastern plusiines are in appearance, across genera, and even tribes. Certain identification of many plusiines requires careful examination of microscopic characters. The Gray Looper Moth is migratory and in some years will be found far northward of the states where it is resident. The darkly pigmented individual figured in the inset likely represents an inducible crowding form— several like it turned up among a cohort that was reared in rather dense quarters.

218 Loopers Subfamily Plusiinae: Tribe Plusiini

Splendid Brass Looper Diachrysia balluca [Green-patched Looper]

! recognition Handsome bright green and white larva, with long setae

arising from whitish pinacula that cap conelike swellings. Integument shiny. Broad frosted addorsal stripe broken over each abdominal segment by an oblique green line that runs between dorsal setae. Subdorsal stripe well defined along thoracic segments but fusing with addorsal stripe over abdomen. Setae white and comparatively long, especially on A8. Head shiny green, unmarked. Thoracic legs with black outer line. Larva to 3.5 cm. ! occurrence Fields, wetlands, riparian corridors, woodlands, and forest clearings from Manitoba to Nova Scotia, south to northern Delaware, North Carolina (mountains), and Great Lakes states. Two generations north to southern Quebec with caterpillars in late spring, then again in midsummer. Mostly local and uncommon. ! common foodplants Often reported from hops. Also anise or blue giant hyssop (Agastache foeniculum), aspen, raspberry, and woodnettle (Laportea); but presumably eating many herbaceous and woody plants, perhaps with preference for Urticales (such as hops and nettles). ! remarks Initially, we regarded this moth to be a creature of Canadian zone forests, and only infrequently encountered it in Connecticut and northern New Jersey. More recently, our impression is that the Splendid Brass Looper is becoming more widely distributed across the Northeast and may be expanding its range southward. The moth was recorded from Delaware for the first time in 2005—in association with recently invasive Japanese hops. Diachrysia adults are strongly attracted to flowers. The image of a Splendid Brass Looper caterpillar in Lafontaine and Poole (1991) appears to be of an early (recently molted) last instar—the pinacula are not nearly so pimplelike in larvae that we have encountered. The ground color of plusiine caterpillars is much influenced by diet. Larvae reared in captivity are often more yellow to lime green than those found in the wild; the latter tend to be more blue-green and, in some species, more heavily frosted with white.

Loopers Subfamily Plusiinae: Tribe Plusiini 219

Unspotted Looper Allagrapha aerea ! recognition Green to blue-green caterpillar with four wavy stripes above level of spiracles. Dropping away from midline, each stripe thinner, more crenulate, and broken; lowermost supraspiracular stripe sometimes absent (only three stripes mentioned by Lafontaine and Poole 1991). Profile somewhat corrugated with transverse creases towards posterior half of each segment. Spiracular stripe passing through top of spiracle on A1–A4 and A8, running above spiracle on A5–A7, weakening anteriorly and posteriorly, and often absent on A7 and A8. Dorsal and lateral pinacula porcelain white, raised, diameter about half that of spiracle; SD pinacula green, and dorsal pinacula on A8 especially elevated. No vestigial prolegs on A3 and A4. Head green, shiny, smooth, and unmarked. Larva to 3.5 cm. Caterpillar very similar to that of Lined Copper Looper (Diachrysia aereoides), which differs in having the head integument roughened (granulate) in texture. ! occurrence Fields, gardens, croplands, and open habitats from Wisconsin and southern Canada, to Alabama and Louisiana (uncommon to absent along lower Atlantic Coastal Plain). Two or three generations in Connecticut with mature caterpillars from May onward; three or more broods southward. ! common foodplants Aster, nettle, soybean, and probably many other herbaceous plants; our larvae were reared on goldenrod. ! remarks The yellow subdorsal area in the fifth abdominal segment of the figured individual above is a testis. Testes are visible in the latter instars of male plusiine caterpillars. Allagrapha and Diachrysia are thought to be sister genera; as noted above, their caterpillars are quite similar.

220 Loopers Subfamily Plusiinae: Tribe Plusiini

Pitcherplant Looper Exyra fax [Pitcher Plant Moth] ! recognition Dark brown to maroon with deep pale constrictions between

segments. Integument covered with minute spinules. Thoracic setae borne from raised, darkened warts. Head with black transverse cheek bar and large black spot capping each lobe. Larva to 3 cm. Two other Exyra, both with greatly extended lateral warts on A1–A4, occur in the Southeast. In the Pale Shoulder Pitcherplant Looper (E. semicrocea), the raised pinacula along sides of thorax are about the same size as those over the dorsum, and the integument is densely set with spinules (book cover). It occurs from coastal North Carolina to eastern Texas, and is associated principally with Sarracenia flava and S. leucophylla, but also using S. minor, S. alata, and others (Jones 1921; Rymal and Folkerts 1982; Debbie Folkerts, pers. comm.). In Riding’s Pitcherplant Looper (E. ridingsii) the raised subdorsal warts are longer than the dorsal warts on the thoracic segments, and sometimes approach those of abdomen in height (following page). Most frequently it is found feeding in S. flava. All three species of Exyra have been reared from S. purpurea. Folkerts and Folkerts (1996) and Stephens and Folkerts (2012) provide much useful information on our three species. ! occurrence Bogs and acid wetlands from Manitoba to Nova Scotia, south through Great Lakes states, New England, and Atlantic Coastal Plain to North Carolina. Evidently two generations in Northeast with mature caterpillars from late May into June, and at least a partial second generation in July. Additional generations in Southeast. Locally common. ! common foodplants Pitcherplant (Sarracenia purpurea). ! remarks The female lays her eggs in the inside of the pitcher near the water line. The caterpillar grazes the inner surface of the blade and leaves the outer epidermis largely intact; a hole is chewed into the lower side of the blade that drains the pitcher. The entrance of the pitcher is sealed off by the larva, either by silking the edges together or spinning a silken sheet across the mouth of the pitcher; in so doing many potential enemies are excluded. Middle instars overwinter within the sealed pitchers. In the spring, the caterpillar completes its development or moves to a second pitcher plant if its host leaf begins to senesce. The characteristic damage and copious feculae left inside the pitchers can be used to quickly assess whether Exyra larvae are present at a given locality. See Jones (1921), Folkerts and Folkerts (1996), and Stephens and Folkerts (2012) for detailed notes on the biology of our pitcherplant moths.

Loopers Subfamily Plusiinae: Tribe Plusiini 221 Riding’s Pitcherplant Looper Exyra ridingsii

Coastal North Carolina south through the Florida panhandle. Multiple generations with mature caterpillars from late spring into late summer. Locally common. Typically associated with Sarracenia flava. See above.

Bilobed Looper Megalographa biloba (= Autographa biloba)

[Stephen’s Gem]

! recognition Immediately recognizable by fine hairlike spinules over dorsum of body; setae more spinelike below spiracular stripe. Dorsal and subdorsal areas with three thin, white stripes, with integument between uppermost stripes sometimes filled with white. Thin, wavy, white spiracular stripe beginning on T3 or A1 and usually fading rearward of A7, often edged with darker green above; T1, A9, and A10 mostly devoid of striping. Head with broad black lateral stripe that extends through eyes. Thoracic legs shiny black. Spiracles nearly white with dark rim thickened forward and rearward. Larva to 3.5 cm. ! occurrence Fields, gardens, croplands, waste places, as well as woodlands and forests. Resident in Deep South but migrating northward into Canada most years. Two or more generations in Connecticut with mature caterpillars from early June onward. Sometimes common. ! common foodplants Many herbaceous plants, including alfalfa, banana, barley, cabbage, clover, cranesbill, dandelion, hedge nettle, kidney bean, larkspur, lettuce, plantain, sage, tobacco, vervain, and many others. ! remarks The Bilobed Looper is our only representative of this largely tropical genus. The moth is a strong migrant that occurs throughout much of the New World as well as Hawaii. In England it is known as Stephen’s Gem. Occasionally, it is common enough to cause damage in gardens. In Connecticut and New Jersey, adults wing into the state as early as May or June in some years. They are common backyard visitors—at dusk, watch for nectaring or ovipositing adults. The garden is as good a place as any to search for this looper—larkspur and lettuce are among its favorites.

222 Loopers Subfamily Plusiinae: Tribe Plusiini

Climbing Looper Autographa ampla [Broken-banded Y] ! recognition Translucent yellow-green, green, or blue-green caterpillar with two wavy dorsal stripes: diffuse, thickened addorsal stripe that runs through D1 pinacula, and fragmented, yellow stripe (running between two dorsal setae). Latter stripe often broken into oblique lines that more strongly expressed toward anterior end of each abdominal segment. Spiracular stripe cream to yellow, often thinning across thoracic segments, and passing through upper edge of spiracles on A1–A4 and above spiracles on A5 and A6. White speckling reduced. Pinacula inconspicuous, pale white or green. Head green, shiny, unmarked. Straw-colored spiracles without darkened rim. Integument with minute spinules. SV group trisetose on A1. Larva to 3.5 cm. ! occurrence Wetlands, shrub swamps, meadows, edges of watercourses, woodland and forest edges and clearings especially of boreal zone. Transcontinental in Canada, south in East to Long Island, Georgia (mountains), and Great Lakes states. One generation with mature caterpillars in spring and early summer. Common northward. ! common foodplants Alder, aspen, birch, cherry, poplar, serviceberry, soapberry, wild raisin, and presumably other woody plants. ! remarks This species is also known as the Brown-patched, Large, and Raspberry Looper. While the loopers that range into warmer parts of the United States favor nonwoody plants, A. ampla and other univoltine northern Plusiinae prefer shrubs and small trees. All overwinter as larvae and mature on the nutritious, new growth of late spring and early summer.

Loopers Subfamily Plusiinae: Tribe Plusiini 223

Wavy-chestnut Y Autographa mappa ! recognition Light green with frosty white dorsum. Addorsal stripe diffuse, broad (usually), reaching almost to D2 pinaculum; in some forms, addorsal and dorsal stripe distinct, i.e., area between two stripes not filled (as described above and in figured individual). Subdorsal stripe thin, wavy, broken. Yellow spiracular stripe passing below spiracle on T1, through middle of spiracle on A1, along top edge on A2–A4, above spiracles on A5–A7, and again dipping down to touch top edge on A8. Deep incisures between A1–A7. SD pinacula and those on T1 and head black. Head green, shiny, without black bar or short inconspicuous bar through eyes. Integument with sharp-tipped granules. A1 with SV group trisetose. Larva to 3.5 cm. ! occurrence Woodland and forest edges and clearings. Transcontinental in Canada, south in East to northwestern Connecticut and Great Lakes states. One generation with mature caterpillars in May and June. Common northward. ! common foodplants Nettles, blueberry, and presumably other low plants. ! remarks A. mappa is the only member of the “jota species complex” in eastern North America—its closest relatives are in Europe, Asia, and Alaska (Lafontaine and Poole 1991). Given its taxonomic isolation we suspect that the diagnosis given above will distinguish the caterpillar from all other plusiines in our fauna. A British relative, the Plain Golden Y (A. jota), often spins its cocoon on the underside of a host leaf; its pupal stage lasts about four weeks (Porter 1997). Adult plusiines are avid flower visitors. Nielsen (1981) records 20 species of plusiines from his early evening surveys in upstate Michigan. Four of his collections were state records and several others previously were thought to be strays or extremely rare in Michigan. Blooms of fireweed proved to be the most attractive—19 of the 20 species found during his seven-year survey were taken while feeding at Epilobium flowers.

224 Loopers Subfamily Plusiinae: Tribe Plusiini

Common Looper Autographa precationis ! recognition Green with roughened integument densely covered with minute spinules. Scalloped profile. In many individuals, first 4–6 abdominal segments with prominent, shiny black SD1 setal base above spiracle; that on A3 largest. A8 humped with dorsal setae perched on raised bases. Head green, shiny, with prominent black bar down each side. A1 with SV2 seta absent (SV group bisetose). Larva to 3.5 cm. Bilobed Looper (Megalographa biloba) has much larger integumental spinules (page 221). Caterpillar of Soybean Looper (Chrysodeixis includens) similar, but head stripe thinner, and with minute (vestigial) prolegs on A3 and A4 (page 215). ! occurrence Fields, gardens, agricultural fields, waste places, and other open and early successional habitats from southern Manitoba to Nova Scotia, to northern Georgia and Mississippi. At least two generations in Connecticut with mature caterpillars from June onward, three or more broods southward. Very common. ! common foodplants Many herbaceous plants, including beans, cabbage, clover, dandelion, fleabane, hollyhock, sunflower, thistle, vervain, wild lettuce, and some grasses. ! remarks Also known as Plantain Looper. This is one of many plusiines known to “trench.” When feeding on plants with milky latex, such as dandelions and wild lettuce, the Common Looper incapacitates the pressurized vascular system of its foodplant by chewing a trench through the veins (Dussourd and Denno 1991, Dussourd 1993), thwarting the plant’s ability to deliver sap to the feeding site. Upon severing the midrib and other veins, usually near the leaf base, the caterpillar moves beyond the trench to feed. Captive larvae will eat, and may even prefer, slightly wilted foliage. Plusiines have rapid development times; summer generation caterpillars mature in three to four weeks. The individual in the inset was reared indoors from an egg and lacks some of the bright green pigments of wild larvae. In November 2005, DFS collected a larva in New Jersey and reared it outdoors. Except for a 23-day cold spell in December, it remained intermittently active all winter with molts on 1 January and 28 February, spun its cocoon on 27 March, pupated on 5 April, and eclosed on 1 May. It is a well-known migrant; we recall occasions, especially in late summer, when numerous adults suddenly appeared in our gardens and at light. We are uncertain how far north the species is able to overwinter.

Loopers Subfamily Plusiinae: Tribe Plusiini 225

Celery Looper Anagrapha falcifera ! recognition Green, yellow-green, or blue-green with prominent white to cream spiracular stripe that continues to anal plate, and down anal proleg in some forms. Upper edge of spiracular stripe passing through spiracle on A1–A4 and A8; spiracles on A5–A7 sometimes falling below stripe. Three thin, wavy dorsal and subdorsal stripes evenly partitioning D1 from D2 setae. Stripes obscure on T1. Shield and head green without markings in our examples; Lafontaine and Poole (1991) describe a form with black lateral line that passes through eyes. Sparse, elongate spotting above and below spiracular stripe. Integument appearing smooth, but bearing minute spinules; subventer and venter with smoky spinules. Dorsal pinacula white; SD pinacula green. Three SV setae on A1. Spiracles white with dark rim, especially forward and rearward. Larva to 3.5 cm. Superficially similar to many other plusiines, especially Syngrapha species. Most Anagrapha larvae can be distinguished from those of Syngrapha by their host (more likely to be found on forbs), and southward by range. ! occurrence Fields, gardens, croplands, waste places, and other open and early successional habitats across southern Canada; in East south to Georgia (mountains) and central Texas. Three generations in Connecticut with mature caterpillars from May onward, more broods southward. Often very common. ! common foodplants Generalist. Field crops include bean, cabbage, carrot, celery, lettuce, sugarbeet, tobacco. Less commonly on woody plants such as blueberry, large cranberry, and viburnum. Crumb (1956) also mentions corn. ! remarks The caterpillar is an occasional pest of celery and other vegetables. While there is a dearth of literature mentioning Anagrapha as a migrant (but see Lingren et al. 1993), its presence at light in the advance of storm fronts, asynchronous phenology over the summer months (Rings et al. 1992), and frequent adults in November and early December in New Jersey (DFS) suggest that at least a fraction of midlatitude populations represent continuously breeding migrants from the South. The March adults in Ohio (Rings et al. 1992) are probably migrants since pupal overwintering seems unlikely; however, more numerous late April records in Ohio, and the presence of vernal form adults in April and May in southern Pennsylvania and New Jersey, suggest that Celery Looper larvae overwinter successfully in colder climates. See also Schweitzer and Roberts (2007).

226 Loopers Subfamily Plusiinae: Tribe Plusiini

Abstruse Looper Syngrapha abstrusa ! recognition Dorsum with broad cream to white addorsal stripe and thin bluish-white subdorsal stripe nearly fusing with addorsal stripe over T1 and T2. Broad yellowish spiracular stripe. According to Lafontaine and Poole (1991), SD1 pinaculum small, approximately half height of spiracle on abdominal segments; D2 pinaculum narrowly ringed with black. Three SV setae on A1. Tan spiracles along upper edge of spiracular stripe. Larva to 3 cm. Three other conifer-feeding Syngrapha occur sympatrically over much of Canada and the northeastern United States, and have similar caterpillars (additional species also occur westward). In both the Salt and Pepper Looper (S. rectangula) (see page 229) and Spruce Looper (S. alias) (see next page), the white addorsal stripe is pure white in color, the subdorsal stripe is thicker, and the diameter of the SD1 pinaculum on A1 is about the same height as the spiracle. In S. rectangula the SD1 pinaculum on A1 is about twice the height of the spiracle. A fourth member of the genus, Green-sigma Looper (S. viridisigma), looks like S. rectangula, but has only one SV seta on T2 and T3; in addition, its addorsal and subdorsal stripes are more slender (page 229). ! occurrence Forests especially in dry, well-drained habitats. Transcontinental in Canada, south in East to New Jersey and Great Lakes states. One generation with mature caterpillars in May and June. Common northward. ! common foodplants Eastern hosts include jack pine and white spruce (Lafontaine and Poole 1991), and presumably other conifers. ! remarks The 24 North American species of Syngrapha are largely northern and montane in distribution, and best represented in coniferous forest habitats. A few fly north of the Arctic Circle. Their caterpillars commonly feed on members of the pine, fir, and spruce (Pinaceae), heath (Ericaceae), and willow (Salicaceae) families. Ten members of the genus are figured in this work. All overwinter as third instar caterpillars on foliage, exposed to the harshest temperatures of winter. On a number of occasions DLW has found Syngrapha larvae, 6–8 mm in length, on the family Christmas tree (usually a Fraser fir).

Loopers Subfamily Plusiinae: Tribe Plusiini 227 Beautiful Looper Chrysanympha formosa

Manitoba to Prince Edward Island, south to Georgia (mountains) and Great Lakes states. One generation with mature caterpillars in May and June. Often common in bogs, barrens, and acidic swamps. Blueberry. A8 strongly humped.

Spruce Looper

Syngrapha alias [Hooked Silver Y] Transcontinental in Canada, south in East to North Carolina (mountains) and northern Great Lakes states. One generation with mature caterpillars from early June to July. Common northward. Spruce, fir, western hemlock, and perhaps other conifers. SD1 pinaculum on A1 nearly equal to height of spiracle.

Inscribed Looper Syngrapha epigaea

Transcontinental in Canada, south in East to northern Pennsylvania, Ohio, and northern Great Lakes states. One generation with mature caterpillars from June into August. Locally common northward. Blueberry, sheep laurel, and presumably other heaths. Also known as Pirate Looper.

Little Bride Looper Syngrapha microgamma

Much of Canada, south in East to southern Maine, northern Massachusetts, and Great Lakes states. One generation with mature caterpillars from late May into July. Locally common northward. Labrador tea, leatherleaf, and blueberry; willow in Europe. Closely associated with bogs in United States. Ground color dark green or red-brown.

228 Loopers Subfamily Plusiinae: Tribe Plusiini

Labrador Tea Looper Syngrapha montana

Transcontinental across much of Canada, south in East to extreme northern New England and Great Lakes states. One generation with mature caterpillars from May into June. Locally common. Labrador tea: favoring new growth and feeding from upper leaf surface. Note prominent white pinacula.

Boreal Looper

Syngrapha octoscripta [Dusky Silver Y] Transcontinental across much of Canada, south in East to northern Pennsylvania, Ohio, and Great Lakes states. One generation with mature caterpillars in June and July. Common northward. Blueberry. Often olive green, with white speckling below yellow spiracular stripe. One SV seta on T2 and T3.

Loopers Subfamily Plusiinae: Tribe Plusiini 229 Salt and Pepper Looper Syngrapha rectangula

Eastern portion of range: Manitoba to Newfoundland, south to North Carolina (mountains) and Great Lakes states. Mature caterpillars in June and July. Locally common. Firs and spruce, with former evidently preferred (Lafontaine and Poole 1991). Broad addorsal stripes; SD1 pinaculum on A1 twice height of spiracle; two SV setae on T2 and T3.

Chosen Looper Syngrapha selecta

Northwest Territories to Newfoundland, to northern Michigan. Mature caterpillars in June and July. Rare and local. Caterpillars accept birch, blueberry, and willow (Lafontaine and Poole 1991); natural foodplant unknown. Crochets numbering 21–24 on midabdominal prolegs; D2 seta equidistant between D1 and SD2 on T2.

Golden Looper Syngrapha u-aureum

Manitoba, Labrador, and Cape Breton Islands, south to northern New England (alpine summits). Mature caterpillars from late June to August. Locally common. Blueberry. Crochets numbering 27–30 on midabdominal prolegs; two SV setae on T2 and T3. Closely similar to S. interrogationis and S. epigaea.

Green-sigma Looper Syngrapha viridisigma

Transcontinental in Canada, south in East to western Massachusetts and Great Lakes states. Mature caterpillars mostly July to early August. Locally common. Spruce, fir, and Douglas-fir. Crochets numbering 25–28 on midabdominal prolegs; D2 seta closer to D1 than SD2 on T2; and only one SV seta on T2 and T3.

230 Loopers Subfamily Plusiinae: Tribe Plusiini

Putnam’s Looper Plusia putnami [Lempke’s Gold Spot] ! recognition Smooth, translucent, gray-green to blue-green with three thin, white addorsal and subdorsal stripes. Spiracular stripe pure white, thick, parallelsided, running length of body, sometimes edged with dark green above, along thorax. SD pinaculum on A1 black, small, less than half height of spiracle. Head green, smooth. Setae black on head and above spiracular stripe. Thoracic legs green. White spiracles embedded along upper edge of spiracular stripe. Crochets appearing uniordinal. Larva to 3.5 cm. Plusia caterpillars are the only North American loopers (plusiines) in which the crochets appear to be all of a single length (uniordinal); in addition Plusia caterpillars have three SV setae on A1, a smooth integument, and they lack the conspicuous intersegmental incisures of other loopers. The Connected Looper Moth (P. contexta) is broadly sympatric with P. putnami; its caterpillar has only two thin addorsal and subdorsal stripes above the level of the SD pinacula, and the spiracular stripe is more apt to be yellow. The recently described Large-spotted Looper Moth (P. magnimacula) occurs in bogs and mesic woodlands from Alberta to Newfoundland, south to Massachusetts and Nebraska (Handfield and Handfield 2006). Its larva, figured by Landry et al. (2002) (as P. putnami), purportedly differs from P. putnami in having the spiracular stripe more apt to continue down the anal proleg and more weakly developed subdorsal and addorsal stripes; however, these characters show some variation across the genus. The larva of a fourth eastern member of the genus, P. venusta, is unknown. ! occurrence Wet meadows, fens, floodplains and riparian corridors, marshes, and other open grassy habitats. Transcontinental from Alaska to Newfoundland, south in East to Pennsylvania and Great Lakes states. Two generations from southern Canada southward with mature caterpillars in spring and early summer, then again in July and August. Locally common. ! common foodplants Grasses and sedges. ! remarks The larva can sometimes be found by sweeping in wet meadows. Nighttime searches also yield caterpillars, especially in the spring (Porter 1997). The caterpillar rests with the body fully extended along a leaf blade, not arched upward as is typical for other loopers (plusiines). In Britain P. putnami is known as Lempke’s Gold Spot.

Loopers Subfamily Plusiinae: Tribe Plusiini 231 Variegated Violet Looper Autographa rubidus

Transcontinental in Canada, south in East to northwestern Connecticut and northern portions of Great Lakes states. One generation with mature caterpillars in June and July. Uncommon. Natural foodplants unknown; lab-reared caterpillars accept dandelion and artificial diet (Lafontaine and Poole 1991).

Pink-patched Looper Eosphoropteryx thyatyroides

Much of Canada, south in East to Georgia (mountains) and Missouri. Two generations with mature caterpillars in spring and then again in midsummer. Uncommon. Meadow-rue, columbine, and possibly lousewort.

Connected Looper Plusia contexta

Central Saskatchewan to Nova Scotia, south to New Jersey and Nebraska. Two generations over much of East with mature caterpillars in spring and again in summer. Locally common. Grasses. Dorsal stripe between addorsal and subdorsal absent; spiracular stripe yellowish.

Straight-lined Looper Pseudeva purpurigera

Manitoba to Newfoundland, south to Georgia (mountains) and Missouri. One generation with mature caterpillars in spring and early summer. Uncommon. Meadow-rue.

232 Tumblers Subfamily Bagisarinae

Tumblers

Family Noctuidae: Subfamily Bagisarinae Bagisarines were formerly classified as bird-dropping moths (Subfamily Acontiinae), but were removed from that subfamily by Crumb (1956), and more recently by Kitching and Rawlins (1998). We place Amyna here, because of its shape and posture, the subapical tooth on the crochets, and because its animated alarm response resembles that of Bagisara (see Wagner and Binns 2010). Twelve Bagisara and two Amyna species occur north of Mexico (Ferguson 1997, Lafontaine and Schmidt 2010). Like acontiines, the first two pairs of abdominal prolegs are wanting and the D2 setae are thickened and upcurved on A10 (right). The body setae are long and conspicuous (especially rearward). The SV group is trisetose on A1 and A2. In addition to the structural characters noted above, bagisarines and acontiines share the habit of flicking their feculae, and both

include many mallow feeders among their ranks. Bagisarine caterpillars also share numerous structural and life history characters with the scalloped moths (Anomis and kin) (see Wagner and Binns 2010). Larvae are quick to hurl themselves from their perch and wriggle violently when alarmed, hence the name Tumblers. Prepupae take on maroon hues. Pupation occurs in a soil-cocoon belowground or in a sparse cocoon spun in duff and leaf litter.

! Thickened d2 setae of anal plate !

Hazel Tumbler Bagisara rectifascia

What we regard to be true B. rectifascia occurs from at least Manitoba (Forbes 1954) to Massachusetts and Pennsylvania (Grote 1874); range confused with that of Bagisara near rectifascia southward (see page 233). One principal brood with mature larvae mostly in July into early August in Northeast. Generally rare, but sometimes locally common. Hazel.

Sandstorm Tumbler Bagisara buxea

South Texas to California. Multiple generations with mature caterpillars from April to October, but mostly in wet season. Locally common. Ours reared from globemallow (Sphaeralcea) and Texas Indian mallow (Abutilon fruticosum).

Rayed Tumbler Bagisara tristicta

South Texas to southern Arizona. Multiple generations with mature caterpillars from May to November, but especially during wet season. Local and uncommon. Bladdermallow (Herissantia crispa).

Tumblers Subfamily Bagisarinae 233

Common Sida Tumbler Bagisara repanda [Wavy-lined Mallow Moth]

! recognition Slender, elongate, lime green with faint, dark green addorsal,

subdorsal, and supraspiracular stripes. Prolegs absent on A3 and A4. Pale, thin spiracular stripe, beginning on T2 and ending on A10 above proleg, usually better developed anterior to spiracle. Setae dark, long; those over dorsum about as long as segment that bears them; pinacula black, minute. Green head peppered with tiny black dots, those over vertex arranged in lines. Prolegs with pink cast at their apex; anal prolegs splayed out behind body. Mature larva under 2.5 cm. Middle and penultimate instars with somewhat more developed stripes. Its relative, the Hazel Tumbler (B. rectifascia), appears to represent two species. Along the Gulf coast, a moth going under the name of B. rectifascia feeds on hibiscus and Turk’s cap (Malvaviscus) (Bottimer 1926; John Tveten, pers. comm.). What we regard to be true B. rectifascia feeds exclusively on hazels (page 232). Two hypenines that sometimes feed on sida in Florida and Texas, the Mallow Snout (Hypena minualis) (page 63) and Black Snout (Hypena scabra) (page 70), are superficially similar to the Common Sida Tumbler—both are missing the first set of prolegs (on A3), but bear prolegs on A4. ! occurrence Fields, roadsides, waste lots, and other open areas in Deep South. Continuous broods in southern Florida. Common to abundant southward. ! common foodplants Sida and false mallow (Malvastrum). ! remarks Larva often perches with a loop thrown into the anterior portion of the body and the head cocked to one side. If poked, the head is rolled under. When threatened further, caterpillars launch themselves from their perch and writhe wildly. The alarm response is violent, the wriggling so animated that caterpillars can instantly hurl themselves 15 cm (6 in.) to a new point of purchase. Larvae are readily sampled with a beating sheet (but be careful not to bump target foliage beforehand). The caterpillars flick away their fecal pellets with a snap of the abdomen—the walls of our rearing containers will sometimes be peppered with adherent feculae. Pupation occurs in a sparse cocoon in litter.

234 Tumblers Subfamily Bagisarinae

Eight-spot Amyna axis (= A. octo) ! recognition Elongate, shallowly constricted between segments, with long, dark setae borne from blackened pinacula. Prolegs absent on A3 and A4. Pale green to black, with thin, usually broken, addorsal, subdorsal, supraspiracular, and spiracular stripes running length of body. Spiracular stripe somewhat thickened, weakening rearward of A8. Caudal segments flattened with prolegs directed behind body. Head shiny, pale green, with setae borne from dark bases. Larva to 3 cm. A. bullula co-occurs with the Eight-spot throughout much of the East—its early stages and foodplants are unknown. We wonder if it occurs principally as a migrant to our area, because most records are for late in the season, even in Florida. ! occurrence Fields, roadsides, waste lots, agricultural lands, and other open habitats. Gulf states with strays north to Great Lakes states and southern Canada. Multiple generations with mature caterpillars through growing season in southern Texas and Florida, but even there most activity late in season, perhaps due to influx of migrants from Caribbean and Mexico, respectively. Common in Gulf states. ! common foodplants Our collections from south Texas all from amaranth and croton; Ferguson et al. (1991) list amaranth, pigweed (Chenopodium), Parasponia, and balloon vine (Cardiospermum) in Bermuda. ! remarks At low densities caterpillars tend to be pale green. At higher densities larvae take on darker patches, and in the extreme, the ground color is given over to black (right). Caterpillars can be abundant in south Texas in October—to the extent that they sometimes defoliate small stands of amaranth. Most of our larvae have come from mature amaranth plants, especially those that were well into seed, turning red, and senescing. Like Bagisara, they are quick to release their grip and thrash and snap wildly, sometimes for several seconds after contact has been made. A cocoon is spun belowground or in debris at the soil surface. A. axis (formerly known as A. octo) is a pantropical moth that has been named nearly 20 times from 15 different countries (and at least four continents); if nothing else, its list of aliases is a testament to the insect’s exceptional migratory abilities. See Wagner and Binns (2010).

Resplendent Moths Subfamily Cydosiinae 235

Resplendent Moths

Family Noctuidae: Subfamily Cydosiinae While historically Cydosia were thought to be Acontiines, Lafontaine and Schmidt (2010) regarded the larvae to be so exceptional that they placed the genus in its own subfamily. The group is essentially Neotropical with three species in our region, mostly confined to Florida and Texas. The larvae are immediately recognizable by the enormously elongated lobes that bear the paddlelike D2 setae. New World hostplant records include members of the Loganiaceae and Verbenaceae.

Regal Cydosia Cydosia nobilitella

South Florida. Multiple generations over growing season. Local and uncommon. Robert Pilla found and photographed the caterpillars shown here on West Indian pinkroot (Spigelia anthelmia). Adult shown below.

Tricolored Angel

Cerathosia tricolor (Subfamily unclarified) Central Texas and westward. One generation in Austin area with mature caterpillars in May. Valerie Bugh has twice photographed the caterpillar, once on rainlily (Cooperia) and once on milkweed. Uncommon and local. (We place Cerathosia here because it too possesses elongated lobes bearing black, spatulate setae—but its full proleg compliment and some adult features suggest it belongs elsewhere or warrants its own subfamily.)

236 Eustrotiines Subfamily Eustrotiinae

Eustrotiines

Family Noctuidae: Subfamily Eustrotiinae Members of this subfamily were formerly classified with the Bird-dropping Moths (Subfamily Acontiinae). All are small moths, generally with wingspans under 2 cm. The North American eustrotiine fauna—a heterogeneous (and perhaps unnatural) assemblage—consists of some 43 species and 11 genera (Lafontaine and Schmidt 2010). The two largest gen-

era, Tripudia and Cobubatha, are largely southwestern. The life histories (and larvae) are unknown for most. The halfdozen, widespread eastern species (all formerly in the genus Lithacodia) are mostly grass or sedge feeders. The proleg on A3 is missing or reduced; in some, the A4 proleg is also reduced or wanting. The SV group on A1 is usually trisetose in our species (although Kitching and Rawlins 1998 state that most bear only two SV). The pupa overwinters.

Burnished Bog Moth

Deltote bellicula [Bog Deltote] Illinois to Newfoundland, south to Florida and Texas. Single generation over much of East with mature caterpillars in July and August. Very common. Tim McCabe reared a cohort on upright sedge (Carex stricta); likely feeds on other sedges as well because C. stricta is absent from many sites where moth occurs.

Wild Petunia Seed Moth Tripudia rectangula

Iowa to southern New Jersey, south to Florida and Louisiana (record from Martha’s Vineyard likely a stray or mislabeled). Multiple generations with mature larvae from July into October over most of range. Common. Larvae seed predators in pods of wild petunia (Ruellia). Mature larva less than 9 mm.

Brass-dotted Grass Moth Maliattha synochitis [Black-dotted Maliattha]

Dakotas to Nova Scotia, south to Florida and Texas. One principal generation over much of East with mature caterpillars from middle to late summer. Very common. Grasses; ours reared on crabgrass (Digitaria). Tim McCabe reared a cohort on Hordeum.

Eustrotiines Subfamily Eustrotiinae 237

Black-bordered Lemon Moth Marimatha nigrofimbria

(= Thioptera nigrofimbria)

! recognition Small, bright green with white longitudinal striping and reduced prolegs on A3 and A4. Thin, white addorsal and supraspiracular stripes; the latter weakening rearward. Subdorsal stripe white, prominent, twice width of other stripes. Body bright green and unmarked. Setae darkened. Prominent, thickened, darkened, upward-curving seta arising from upper portion of anal proleg just below anal plate. Head, legs, prolegs with pale orange cast. Spiracles straw colored. Larva to 2 cm. ! occurrence Grasslands, fields, and waste lots from Missouri to Massachusetts, south to Florida and Texas. At least two generations in much of East, with mature caterpillars nearly year-round in Deep South. Common to abundant over much of range. ! common foodplants Grass, including crab (Kimball 1965); in captivity ours consumed both panic grass and bluegrass. ! remarks This is an abundant moth from New Jersey southward. The small anal prolegs extend from the rear of the body when the caterpillar is at rest. The caterpillar may wriggle violently and regurgitate a yellow-green fluid if handled roughly. The force of the wriggle is sufficient to throw the caterpillar centimeters from its perch. We observed one caterpillar attach itself to the end of a blade then violently throw the anterior portion of the body from side to side, causing the blade to which it was attached to move rapidly to and fro. Feculae are “tossed” with a whip of the terminal abdominal segments. One of our larvae, perched 8 cm above the ground on a potted clump of grass, launched its pellets 55 cm from its feeding site.

238 Eustrotiines Subfamily Eustrotiinae

Large Mossy Grass Moth Protodeltote muscosula (= Lithacodia muscosula)

! recognition Small, striped, tan to brown with prominent transverse creases

toward back half of each segment. Proleg on A3 absent and that on A4 very reduced. Broad, brown-olive middorsal stripe, sometimes edged with red, bounded by wide pale stripes that may be divided by beige to reddish stripe anteriorly; these too may be edged with red. Thin, reddish-brown stripe, somewhat thickened forward and rearward, passing through black spiracles. Anal plate with dark medial line. Anal proleg with dark bar down outer face. Setal bases black with that above spiracle (SD1) somewhat enlarged, especially so on A8. Head with brown reticulations that coalesce to form coronal bar and two less conspicuous lines behind eyes. Larva under 2.5 cm. One other Protodeltote occurs in our region, P. albidula. Crumb (1956) describes its caterpillar as having a purple cast to many of the stripes, the supraspiracular area as being more dusky than the middorsal area, and the venter as “sordid tinged with purple.” ! occurrence Meadows, marshes, and other mesic to wet, often wooded, grassy habitats. Transcontinental in Canada, south in East to Florida and Texas. Evidently at least two generations in Northeast with mature caterpillars from June to September. Uncommon to locally common. ! common foodplants Grasses; Crumb (1956) reported sawgrass (Cladium) as a favorite. ! remarks According to Porter (1997), a good method for collecting British relatives of the Large Mossy Grass Moth is to sweep, especially at night. The pale brown coloration of the caterpillar suggests that it is largely nocturnal in habit and does not perch on green leaves by day. Poole (1996) transferred a number of eastern moths that had been formerly been classified in Lithacodia into other genera, e.g., the Pink-barred Marvel (Pseudeustrotia carneola) (page 379) and Brass-dotted Grass Moth (Maliattha synochitis) (page 236). The caterpillars of both feed on members of the dock family (Polygonaceae). Life histories of other eustrotiines and the early flight season of the Large Mossy Grass Moth suggest that the pupa overwinters.

Bird-dropping Moths Subfamily Acontiinae 239

Bird-dropping Moths Family Noctuidae: Subfamily Acontiinae

So named because the adults are often a combination of white, gray, and black, and patterned in such a way as to resemble bird excrement. The wings, held tentlike over the body, help in the ruse, as does their tendency to perch on the upper side of leaf surfaces, especially of their foodplant. The scope of the subfamily has been reduced considerably by recent taxonomic decisions—all five tribes from Hodges et al.’s (1983) Moths of North America checklist have been given full subfamily status (Kitching and Rawlins 1998, Fibiger and Lafontaine 2005). In addition, the generic classification of the subfamily was revised by Lafontaine and Poole in 2010, e.g., many of the eastern species of Acontia were transferred to Tarache and our Tarachidia moved to Ponometia. Bird-dropping moths are represented by about a dozen widespread eastern species. Their numbers and species richness rise rapidly in the American Southwest—Knudson and Bordelon (2000) record 68 species from Texas. The subfamily is often associated with dry, sunny habitats including waste lots, fields, prairies, sand plains, thorn scrub, and desert environs. However, at least three of our widespread eastern species of Tarache also occur in wetlands, though still in open, sunny habitats. Acontiine caterpillars lack prolegs on A3 and A4. The scalelike spinneret scarcely projects beyond the labium. In most bird-dropping moths there are two SV setae on A1. North American species fall into the nominate tribe, the Acontiini, which often have one or two pairs of stout setae on the anal plate (Crumb 1956). (Presumably these assist in flicking frass away from the body, although Ponometia and perhaps others, which do not have especially differentiated anal setae, are also able to flick their fecal pellets many body lengths.) Coloration can be confoundingly variable within species. Many species of Ponometia come in both a green and brown form. Head capsule patterns are intricate and can assist in distinguishing species; unfortunately they too tend to be variable—a thoughtful photographic study of variation in and across species (and genera) would no doubt yield numerous taxonomic characters. Until the Acontiinae receives more study, identifications should be based on reared adults, especially in Texas and wherever the subfamily is represented by many species. Eusceptis, Ponometia, Spragueia, Tarache, and others perch with a loop thrown into the abdomen. Tarache and kin sometimes rest with a high loop so that caterpillars seen in profile resemble the Greek letter omega. A few hold the head and legs above the substrate at rest. Adults are thought to be principally nocturnal, although they can be flushed during the day. Acontiines are avid flower visitors; at least one is known to puddle (Adler 1982) (see text essay on puddling, page 447). Females are reluctant to oviposit in captivity, necessitating larval searches for those wishing to see or study the early stages. The caterpillars are quick to release their grip (and some even jump) when disturbed, and thus can be readily sampled with beating sheets; care should be taken not to jostle target vegetation beforehand. On low plants over sandy or rocky substrates it is also possible

to rake one’s fingers through a hostplant and then examine the ground below for larvae; these will be motionless and folded in half, or wound up into a ball. Early and sometimes late instars perch fully exposed on upper leaf surfaces—they are easily located once a search image is formed. Once sated, last instar Tarache and related genera may move away from the feeding site and come to rest along a stem: follow shoots to the soil, especially when searching low-growing foodplants, and look for caterpillars with a loop thrown into the abdomen, facing head down. While most, if not all, acontiines will consume and mature on vegetative tissues, many are partial to flowers, young fruits, and other reproductive tissues. Henning von Schmeling and DLW have observed pollen predation in two species, and suspect the behavior is widespread. Those from arid regions can be tolerant of older or damaged foliage that would be rejected by forest-dwelling caterpillars. Eastern acontiines feed mostly on members of the mallow (Malvaceae) and sunflower (Asteraceae) families. Many use just one or two foodplants at any single location, but exploit a range of hosts across their range. Two Spragueia have been recorded from bindweeds (Convolvulaceae).1 Ponometia, Tarache, and others sometimes feed by scraping patches of tissue from the upper side of a mature leaf. Older larvae frequently chew holes in the middle of the blade, rather than feed from a leaf edge. At least spragueia and presumably others shoot a clear fluid several body lengths when squeezed, presumably from the cervical neck gland. Prepupae take on a red or blue-green cast, or in some cases both; and to a much greater degree than most Lepidoptera, they shrink in size. Under normal conditions, acontiines spin a sparse cocoon below the soil surface, and then secrete considerable fluid that dries and hardens the cocoon-crypt, providing a cell that prevents desiccation, among other things. Denied of soil, Ponometia and Spragueia weave slight cocoons in paper toweling or pupate without spinning a cocoon. Expect to lose a high percentage of acontiines if you fail to supply prepupae with adequate substrates and moisture regimens. Tim McCabe places last instars over dry, loose loam (collected from beneath the foodplant at the time the larvae are discovered). We have had success with a mixture of slightly dampened sand and silt that allows the prepupa to construct a “mud pot” cocoon. Acontiine pupae may overwinter for more than one year.

! Spotted Spragueia moth (Spragueia guttata). ! The veracity, or at least the primacy, of bindweed as a common spragueia host is in need of confirmation because acontiines may retire away from foliage when not feeding, and on occasion, come to rest on nonhosts.

1

240 Bird-dropping Moths Subfamily Acontiinae

Exposed Bird-dropping Moth Tarache aprica (= Acontia aprica)

! recognition Chunky, green to brown mottled, often with prominent black

spot around spiracle on A1. Small swellings above spiracles on abdomen (evident when larva is at rest). Face flattened, heavily mottled with black, appreciably darker than posterior half of head; jet black spot above each antenna, lower half of triangle mostly black (lower right). Prothoracic shield strongly pigmented; some forms with hint of orange spiracular line passing through first two thoracic segments. Dorsal pinacula pimplelike, small, embedded in black spot; enlarged over A8. Vague, zigzagging lateral stripe dropping down prolegs on A5 and A6. Hump over A8 rounded. Spiracles black with white halo. Thoracic legs with black bands. Larva to 3 cm. A number of other Tarache occur east of Texas, four of which are discussed below. ! occurrence Swamps, marshes, gardens, fields, and waste lots from Kansas, southern Canada, and Long Island to Florida and Texas. Two generations over much of East with mature caterpillars from May onward; nearly year-round in Florida. Locally common. ! common foodplants Hollyhock (Alcea rosea), hibiscus, and presumably related mallows—see below. ! remarks Our caterpillars were collected from hollyhock in southern Pennsylvania by Valerie Giles. Three larvae were found and reared to maturity on flowers; a fourth larva was found and reared on leaves. Hollyhock is an introduced species: presumably the caterpillar also feeds on hibiscus and other native members of the mallow family. The Curve-lined Bird-dropping Moth (T. terminimaculata) occurs widely throughout our region. It is reported to feed on both basswood and mallows; DFS has collected larvae on seashore or Virginia saltmarsh mallow (Kosteletzkya virginica), and has seen adults flying in the vicinity of marsh mallow (Hibiscus) in New Jersey. The rarely encountered Delightful Bird-dropping Moth (T. delecta) is believed to feed on swamp rose mallow and other Hibiscus. Its caterpillar bears prominent white and yellow spots about the abdominal spiracles and spotting on the head is sparser than on the Exposed Birddropping Moth (Crumb 1956). The Brown Cotton Leafworm or Cotton Birddropping Moth (T. dacia), reported from Louisiana and Texas, is a pest of cotton (Todd 1955). The Four-spotted Bird-dropping Moth (T. tetragona) (next page) occurs throughout southern Florida and Texas; it is believed to feed on abutilon, bladdermallow, false mallow (Malvastrum), and sida.

male

female

Bird-dropping Moths Subfamily Acontiinae 241

Four-spotted Bird-dropping Moth Tarache tetragona ! recognition Yellow or tan to dark brown with detailed white, black, and orange markings. Dorsal and subdorsal setae and spiracles often surrounded by black spot. Spiracular stripe best developed rearward, often infused with orange. Blackened spiracles atop lateral swellings on abdominal segments. A8 humped. D2 pinacula borne from swollen black wart. Head patterned with black and white and orange patches over each lobe, through triangle, and rearward from eyes (lower right). Larva to 2.5 cm. Middle instars with pinacular warts, especially following a molt (above lower right), and thus resembling a spragueia, which feed on the same hosts. ! occurrence Fields, agricultural lands, waste lots, and other open habitats in southern Florida and Texas. Presumably active through growing season with mature caterpillars in all months. Common. ! common foodplants False mallow (Malvastrum) commonly; abutilon and bladdermallow (Herissantia crispa) (both Berry Nall, unpubl. data), sida, and presumably other members of mallow family. ! remarks In the late summer and fall, this species is widespread and common throughout the Rio Grande Valley. Larvae prefer flowers over leaves. Over the weeks they were held in the lab, we had a cohort of caterpillars become progressively bolder in coloration, with elaboration of the black and orange pattern elements. We are unclear if the darkening and other phenotypic changes were induced by our rearing conditions. Disturbed caterpillars drop from the host, fold the body in half, and remain motionless.

Turk’s Cap Bird-dropping Moth Eusceptis flavifimbriata

Texas, south into Mexico. Mature caterpillars from at least April into November—larvae common in October. Locally common. Turk’s cap or wax mallow (Malvaviscus).

242 Bird-dropping Moths Subfamily Acontiinae

Yellow Spragueia Spragueia apicalis ! recognition Ground color yellow and tan-green to brown with black markings; subspiracular spots on A1–A4. Spots on A2 and A3 often larger than those on A1 and A4. Doubled, yellow lateral stripe enclosing orange spot on thoracic and abdominal segments. Often with additional orange spot above proleg on A4 and A5. White, lateral patch on head tends to be globular, not as longitudinally elongate as some spragueia. Larva under 2 cm. Middle and penultimate instar (inset) appreciably darker and more warted. Species-specific characters for spragueia have yet to be worked out: photograph and rear out adults for certain identification. Head capsule patterns appear to hold promise for sorting out Spragueia, Ponometia, and other acontiines. ! occurrence Fields, gardens, waste lots, and other open habitats from Kansas and Illinois to Maryland, south to Florida and Texas (John Wilterding, unpubl. data). Multiple generations with mature larvae through growing season, but becoming more common as season progresses. Locally common. ! common foodplants Sida (multiple records) with questionable records from other hosts, such as broom snakeweed (Guteirrezia sarothrae); but see remarks under Southern Spragueia (S. dama). ! remarks Look for spragueia larvae resting on the upper side of leaves or along the stem of a shoot producing flowers or new leaves. Like other acontiines, older larvae will also be found perched along a stem, head down, near the base of their host plant. While larvae will consume leaves, they have a special penchant for flowers and especially pollen and anthers, which presumably are rich in nitrogen, lipids, and other nutrients. Their dark coloration (especially in middle instars) and conspicuous perching habits are perplexing, and immediately give them away once a search image is developed. If disturbed, the caterpillar is quick to release its grip and drop, or in some cases jump from its perch. At least in early instars, it will drop on a belay line. Alarmed caterpillars may wriggle violently. In any case, it is best not to jostle potential host plants when searching for larvae.

Bird-dropping Moths Subfamily Acontiinae 243

Southern Spragueia Spragueia dama ! recognition Variegated with brown, white, orange, and yellow stripes; ground color smoky. Dorsal pinacula on A1–A2 wartlike. Black lateral spots on A1–A4. Head usually with white cheek and crown (vertex) separated by broad black patch. White patch on top of head enveloping black spot (P1 seta). Upper portion of triangle white; lower portion shiny black. Larva under 2 cm. Middle instar shown in inset. In our examples, both the ground and head capsule coloration are considerably darker than those in other eastern spragueia, but until the genus is better known, identifications should be based on adults. ! occurrence Fields, waste lots, and other open habitats from Oklahoma and Missouri to Virginia, south to Florida and Texas (John Wilterding, unpubl. data). Multiple generations with mature caterpillars from May onward. Common. ! common foodplants Evidently bindweed (Ipomaea); possibly also sida. ! remarks We have found and reared two singletons on bindweed, but found others (possibly conspecific caterpillars) on prickly fanpetals (Sida spinosa). Host-use patterns for spragueia are perplexing given that acontiines are usually specialized in diet. Covell (2005) lists chinquapin and field corn for the Blackdotted Spragueia (S. onagrus); we have reared the species in appreciable numbers from annual ragweed (Ambrosia artemisiifolia) in Florida and South Carolina (page 245). The Common Spragueia (S. leo) is recorded from bindweed, but our collections (totaling more than 60 wild-caught larvae) have come from annual ragweed, A. artemisiifolia (next page). The Yellow Spragueia (S. apicalis) (page 242) is reported from broom snakeweed (Guteirrezia sarothrae), but we have taken it only on prickly fanpetals (Sida spinosa) (on seven occasions). Interested persons are urged to search for and rear out any spragueia and to challenge larvae with alternative hosts, especially those listed in this guide, to help clarify which foodplants are acceptable to larvae. The oval cocoon, spun belowground, incorporates soil and other nearby debris into the wall. Spragueia prepupae secrete a fluid which hardens the walls of their soil-cocoon. All members of the genus overwinter as a pupa.

244 Bird-dropping Moths Subfamily Acontiinae

Common Spragueia Spragueia leo ! recognition Pale green or tan to brown mottled with spots and narrow

stripes; white spiracular stripe with downward-dipping spurs. Often with blurry subspiracular spots on A1–A4, with that on A1 largest. Dorsal pinacula pimplelike over A1, A2, and A8; small white spot often subtending black second (D2) pinacula on A1 and A2. Head shiny, dark brown to black with two white lines over each lobe (below right). Prominent white cheek patch longitudinally elongate; lowermost portion largest and roughly parallel-sided. Larva under 2 cm. Middle and penultimate (inset) instars smoky green to mostly black, with warted dorsal pinacula. Very close to and perhaps inseparable from Black-dotted Spragueia (S. onagrus) (next page), which also uses ragweed (Ambrosia) as a host. Last instars of Common Spragueia resemble those of the Small Bird-dropping Moth (Ponometia erastrioides) (page 247), but larvae of the latter species lack the black subspiracular spots on A1–A4, lack the white lines over each vertex, and the spots making up the white cheek patch tend to be less longitudinally elongate than those of S. leo. ! occurrence Fields, waste lots, roadsides, and other open habitats from Kansas, Illinois, and southeastern Massachusetts; south to central Florida and eastern Texas. Multiple generations with mature caterpillars from May onward. Common. ! common foodplants Our numerous collections all from ragweed (Ambrosia); also recorded from bindweed. ! remarks As noted previously under the Southern Spragueia (S. dama), there is much uncertainty about spragueia hosts, and diagnostic larval features have not been identified for members of the genus. Take images, record notes, and preserve voucher specimens; according to John Wilterding, the taxonomic authority for the genus, reliably determined larval and pupal specimens are all but lacking for the genus.

Bird-dropping Moths Subfamily Acontiinae 245 Spotted Spragueia Spragueia guttata

Mississippi to Texas (John Wilterding, unpubl. data). Multiple generations with mature caterpillars from May onward. Locally common southward. Parthenium and ragweed; we also have beaten (possibly wandering) singletons from pelotazo (Abutilon incanum) and woolly croton or hogwort (Croton capitatus).

Confounding Spragueia Spragueia jaguaralis

Texas westward to southern Arizona. Multiple generations with mature caterpillars from May onward. Common. Ours from false mallow (Malvastrum), ragweed, and toothleaf goldeneye (Viguiera dentata); recorded from lantana (L. camara) in Mexico (John Wilterding, pers. comm.). We suspect that we have more than one species sorted under this name. That shown to right is from false mallow in southern Texas and that shown below is from ragweed in Arizona.

Black-dotted Spragueia Spragueia onagrus

North Carolina to Florida, west to Mississippi. Multiple generations with mature caterpillars from May onward. Common southward. Ours beaten from annual ragweed (Ambrosia artemisiifolia); other reported hosts, i.e., chinquapin, corn, and goldenrod, strike us as erroneous or, at best, exceptional.

246 Bird-dropping Moths Subfamily Acontiinae

Olive-shaded Bird-dropping Moth Ponometia candefacta

! recognition Elongate, slender, green and white with numerous white and

darker green pinstripes. Variable in coloration, ranging from essentially green forms to those that bear prominent dark subdorsal and lateral patches on A1–A4, with those on A3 and A4 largest. Abdominal spiracles occasionally embedded in russet to purplish or black spot. A8 humped. Head with white stripe to either side of triangle and prominent white lateral marbling. Larva under 2.5 cm. Green forms of the Small Bird-dropping Moth (P. erastrioides) are easily confused with those of the Oliveshaded Bird-dropping Moth. Head capsule coloration for each is diagnostic: the former is black and white with a white cheek patch that is divided by a black line running back from the eyes. The head of the Olive-shaded Bird-dropping Moth is mottled in green and/or reddish brown and white, and lacks black pattern elements. The early and middle instars of Ponometia resemble various Spragueia species—see notes under the Common Spragueia (Spragueia leo). ! occurrence Fields, waste lots, roadsides, and other open habitats across much of southern Canada, to Florida and Texas (west to California). Evidently two or three generations over much of East with mature caterpillars to be expected from May (southward) until first frosts. Abundant. ! common foodplants Annual, giant, and other ragweeds. ! remarks Bird-dropping moths do poorly in containers with saturated atmospheres—they are creatures of arid environs. Keep the caterpillars supplied with fresh leaves, flowers, and young fruits, and provide some ventilation. Across much of the East this is the more numerous of our two common ragweed-feeding bird-dropping moths. Come fall, look for the caterpillars with their heads buried in flowers. The larva to the right has a minute clump of pollen affixed to its frons. Beating samples will sometimes yield caterpillars in large numbers. When alarmed, the caterpillar either throws a high loop into the abdominal segments or releases its grip and drops to the ground, feigning death. The Olive-shaded Bird-dropping Moth has been introduced into Russia as a biocontrol agent of introduced ragweed. All Ponometia overwinter as pupae, and some of ours have done so for more than a single year.

Bird-dropping Moths Subfamily Acontiinae 247

Small Bird-dropping Moth Ponometia erastrioides ! recognition Elongate, slender, green to tan, brown, or dark brown. Dorsum

of A1–A3 with ground color darkened to either side of midline. A8 humped. Fine, white spiracular stripe zigzagging along abdomen with spurs extending along outer face of prolegs on A5 and A6. Head with extensive black patterning; cheek with divided white patch. Larva under 2.5 cm. A green-form larva (penultimate instar?) shown in upper inset. Early and middle instars with warted dorsal pinacula, especially over A1 and A2 (lower inset), approaching appearance of some spragueia. The similarly patterned Olive-shaded Bird-dropping Moth (P. candefacta) (page 246) remains lime green through the last instar and lacks the black patterning over the head capsule. The caterpillar may also be confused with those of various spragueia, and especially that of the Common Spragueia (S. leo), with which it cooccurs on ragweed. The Common Spragueia is a bit smaller, lacks brown pattern elements, bears two white lines over each lobe of the head, and the pale cheek patch is more longitudinally elongate than that of the Small Bird-dropping Moth. ! occurrence Fields, waste lots, roadsides, sand plains, and other open habitats from Minnesota to Maine, south to northern Florida and Texas. Two or three generations over much of East with mature caterpillars from April (southward) until first hard frosts. Common. ! common foodplants Ragweed. ! remarks Caterpillars are common in late summer and fall. Beating will yield caterpillars in short order, typically in lesser numbers than those of the Oliveshaded Bird-dropping Moth (P. candefacta). Larvae are quick to regurgitate a deep green vomit if handled. Ponometia rest with a high loop thrown into the anterior abdominal segments, and will sometimes elevate the head (right). In nature, caterpillars fashion a cocoon in soil, but will pupate naked on the floor of their rearing containers if denied sand or soil. Adults are attracted to lights; they are also readily flushed during the day.

248 Bird-dropping Moths Subfamily Acontiinae

Half-yellow Bird-dropping Moth Ponometia semiflava ! recognition Elongate, brown (usually) or green (lower right), with

pale pinstriping; A8 strongly humped. White spiracular stripe broken into thin, overlapping lines. Ground color becomes increasingly mottled with white lines and stripes in later instars. Dorsum of A1–A4 sometimes with dark saddles. A8 with black, pointed warts (D2 pinacula). Venter well marked with white striping, differing little from that of patterning over dorsum. Thoracic legs alternately banded with white and shiny black. Head handsomely marked with white and brown; base of antennae porcelain white. Larva to 2.5 cm. Early and middle instars bear comparatively conspicuous dorsal warts (inset). Related species that could be confused with Half-yellow Bird-dropping Moth occur from central Texas southward and westward, where identifications should be based on reared adults. ! occurrence Roadsides, fields, prairies, coastal strand communities, and other open habitats from Manitoba to southeastern New York, south to Florida and Texas. Two or three generations over much of East with mature caterpillars from June until end of October. Locally common southward. ! common foodplants Maryland goldenaster (Chrysopsis mariana) in New Jersey and North Carolina; camphorweed (Heterotheca subaxillaris) in parts of North Carolina and Texas. Also sunflower, toothleaf goldeneye (Viguiera dentata), and awnless bushsunflower (Simsia calva) in Texas (Berry Nall, pers. comm.), and presumably other composites. ! remarks First-brood larvae mature on leaves—look for them resting on stems, leaves, or nearby litter. They can also be sampled with a sweep net. The tough cocoon can occasionally be found just below the soil surface, near the foodplant. Our collections from along the Eastern Seaboard have all been brownform caterpillars like that shown in the main image. Populations of what we are calling the same species from Texas and Arizona often have both brown- and green-form larvae. Ponometia caterpillars do poorly in closed, moist containers; provide ventilation and a layer of sand, into which the prepupal larvae can tunnel and spin their cocoons. Ponometia is a large genus with 36 United States species, mostly denizens of prairies, deserts, and other arid communities. The biology of all but a handful remains unknown. If you find caterpillars that do not match those figured above (or from different hosts), take images and notes, get the foodplant identified, and save reared adults. The six species that we have collected as larvae have all come from members of the sunflower family.

Pantheas and Yellowhorns Subfamily Pantheinae 249

Pantheas and Yellowhorns Family Noctuidae: Subfamily Pantheinae

Pantheines are robust moths that share similarities with daggers (Acronictinae). Only five of the 17 North American species are eastern. The larvae possess numerous secondary setae on the head and body; the latter are often borne from fleshy warts. All have a wart that bears numerous setae anterior to the spiracle on

T1; a small wart follows the spiracle on A8. The second stemma is often elongate. The spinneret pore opens on the dorsal surface, sometimes well back from the apex (Crumb 1956). Panthea and yellowhorn caterpillars are large headed, well equipped to feed on fully hardened summer leaves (Charadra and Colocasia) and conifer needles (Panthea). They overwinter as pupae in cocoons, usually in the litter. Adults do not feed.

The Laugher Charadra deridens ! recognition Pale green to smoky (or rarely black), with abundant, long, white wispy setae not grouped into fascicles or lashes. Ends of longer setae often somewhat curled. Setal warts usually pale. Head shiny black with bright, lemonyellow triangle and yellow crescent to either side of triangle (lower inset). Some forms with sparsely intermixed black setae at ends of body. Larva to 4 cm. ! occurrence Barrens, woodlands, and forests from southern Canada to Florida and Texas. Two generations over much of East with mature caterpillars from June to October. Common, although absent from parts of Atlantic Coastal Plain. ! common foodplants Beech and oak most commonly; also reported from birch, elm, and maple. ! remarks The Laugher larvae fashion a shelter or use those of other caterpillars. The pre-molt caterpillar in the upper inset has had the upper leaf of its shelter pulled free. Caterpillars do well on tough summer foliage. The yellowish testes of last instar male caterpillars can be seen through the body wall of A5. Forbes (1954) notes that the larva spins a light cocoon on the foodplant.

250 Pantheas and Yellowhorns Subfamily Pantheinae

Tufted Spruce Panthea Panthea acronyctoides

[Black Zigzag]

! recognition Ground color charcoal, gray, or black, with pale spots along sides. Dorsal setae mostly white and (usually) not organized into fasciculated lashes over T1, T2, A1, and A8 as in Tufted Pine Panthea (P. furcilla). Paddle-shaped setae absent from dorsal setal clusters (especially that over T2). This description, consistent with most of our images of Panthea acronyctoides caterpillars from the eastern United States, is contradicted by the Panthea images provided by Ives and Wong (1988: 18) (e.g., longer setae in their panel 18H are fasciculate). See text for Tufted Pine Panthea below. Larva to 3.5 cm. Until this group receives more study, identifications are best based on adults. ! occurrence Forests from Canada south to Great Lakes states and Georgia (mountains). Common northward and in southern Appalachians. ! common foodplants Fir, hemlock, larch, pine, and spruce. ! remarks Although generally regarded as rare in the South, the Tufted Spruce Panthea is common in hemlock stands throughout the Great Smoky Mountains, down to 550 m. It is, apparently, a glacial relic in the southern Appalachians; northward, at least through West Virginia, Pennsylvania, the Adirondacks, and New England, it is more local and occurs at higher elevations, keeping company with the Green Comma (Polygonia faunus) and other northern species. Our caterpillars from Tennessee reared on hemlock and pine (in separate years) had different color forms—an intriguing study would be to rear caterpillars from a single clutch of eggs on different conifers (or on foliage of different ages) to see if such color variants are inducible. Caterpillars tuck the head under the body and/ or are quick to release their grip and tumble from their perch if molested. Panthea overwinter as pupae in tough cocoons spun in leaf litter or humus.

Pantheas and Yellowhorns Subfamily Pantheinae 251

Tufted Pine Panthea Panthea furcilla (= P. pallescens)

[Eastern Panthea]

! recognition Variegated in orange, red, white, and black. Conspicuous subdorsal black hair pencils on A1 and A8; sparser fascicles (lashes) on T1 and T2. Longer lashes, usually with some paddle-shaped setae. Setal warts often orange. A1–A8 in many forms with oblique, pale patch below and forward of white spiracle. Larva to 4 cm. P. furcilla australis occurs from southern Missouri to Virginia south to Florida and Texas (Anweiler 2009). Presumably its caterpillars are indistinguishable or at least very similar to those of the nominate subspecies figured above. ! occurrence Woodlands, pine plantations, and forests from southern Canada to Florida and Texas (but see Remarks). One generation northward and two in Connecticut and New Jersey with mature caterpillars from May to November (but see below); farther southward there are additional broods—moths fly year-round in Florida. Adults common to locally abundant. ! common foodplants Larch, pine, spruce, and probably other conifers. ! remarks The middle instars crudely resemble the shed male strobili (flowers) of their conifer hosts. The thin subdorsal black tufts on T1 and T2 are absent in the images of the Tufted Pine Panthea in Ives and Wong (1988: 18G, J). Perhaps these and other setal tufts vary in their development, are deciduous and sometimes lost, or the image of Ives and Wong is misidentified. Here is another case where an individual with a knack for rearing could help unravel a perplexing tangle of variation (or erroneous literature). Cohorts of ex ova siblings from southern New Jersey reared by DFS in the fall of 2003 and 2006 yielded 19 and 29 adults, respectively, through the month of May, and another 31 and 27 adults from mid-August to early September of the following years—no adults emerged in the intervening months. Offspring from the May-hatching cohort matured by the end of June, diapaused as pupae for about a month, and also emerged in late summer (with the unhatched pupae from the preceding year). In Florida and Georgia, where Panthea furcilla flies into November, larvae may occur all winter. Larvae are rather tolerant of freezing temperatures. The cocoon consists of a loose outer layer and a sturdy, densely woven, inner one.

252 Pantheas and Yellowhorns Subfamily Pantheinae

Yellowhorn Colocasia flavicornis ! recognition Color presumably variable as in other pantheines. In our examples, ground color black with rusty orange hairs over thorax that make a transition to paler setae over abdomen. Other forms more uniformly rendered (upper inset). Dorsal abdominal setae with rusty tinge; those along sides often lighter. T2 with yellow-orange to black hair pencil (setal bundle). Black middorsal tuft over A1 and A8 nearly as broad as high. White, flatly oblique line under spiracle on A2 to A8, getting stronger rearward. Spiracles white. Head solid black. Larva to 3.5 cm. Penultimate (lower inset) with continuous subspiracular stripe and pale patches over A3–A6. Antepenultimate mostly yellow green over abdominal segments A2–A7 and below spiracles. Very close to, and perhaps inseparable from, the Close-banded Yellowhorn (C. propinquilinea)—see discussion below. ! occurrence Woodlands and forests from Missouri to Canada, south to Georgia and Arkansas. Two generations over much of East with mature caterpillars from late May to October. Locally common, but absent from regions of Atlantic Coastal Plain and Piedmont. ! common foodplants Captive larvae readily consume American hornbeam (Carpinus caroliniana), beech, and birch. Our field-collected larvae have all come from beech. ! remarks The caterpillars are shelter formers, usually drawing two leaves over one another and silking them together. The shelters are crude and readily abandoned. The setae bear minute branches, perhaps revealing pantheines’ phylogenetic affinity with the daggers (Acronictinae). Our images of C. flavicornis caterpillars represent a single cohort reared from a female sent to us by Steve Johnson; the female came from a rich, deciduous woodland in eastern Pennsylvania where both the Yellowhorn and Close-banded Yellowhorn (C. propinquilinea) fly together. Over much of the East, the Yellowhorn tends to be encountered in northern hardwood forests or in wetlands, whereas the Close-banded Yellowhorn is more generalized in habitat, turning up in a variety of woodland and forest types.

Pantheas and Yellowhorns Subfamily Pantheinae 253

Close-banded Yellowhorn Colocasia propinquilinea ! recognition Ground color variable, ranging from white to rust or black, with spreading clusters of white to orange setae. Subdorsal hair pencils on T2 and unpaired middorsal lashes present on A1 and (usually) A8. Middorsal tufts often rusty but sometimes black. Head capsule orange or black. Larva to 3.5 cm. Antepenultimate instar darkened at either end (lower right). Co-occurring with Yellowhorn (C. flavicornis) over much of East. Characters suggested by Crumb (1956) for the separation of these two moths appear to be variable within each species. In our examples, as well as those of Tim McCabe’s, the middorsal tuft on A8 is more likely to be orange in C. propinquilinea and black in C. flavicornis. ! occurrence Forests from Canada south to north Georgia, northern Alabama, and Louisiana. Two generations with mature caterpillars mostly from late May into June and again in fall over much of East. Common, although largely absent from Atlantic Coastal Plain. ! common foodplants American hornbeam and birch commonly; also reported from beech and oak, and less commonly elm, maple, and walnut. ! remarks The caterpillar is a shelter former that silks together adjacent leaves or cuts deep incisions into a leaf edge and then draws the freed flap over itself. In Connecticut the Close-banded Yellowhorn is common and widespread, while the Yellowhorn (C. flavicornis) tends to be locally common, but curiously scarce to absent from much of the state. The Yellowhorn is also local in Louisiana, Missouri, and elsewhere; yet in the Appalachian foothills, and much of Maryland (John Glaser, pers. comm.), it is often the more common of our two Colocasia. Our confirmed rearings for the Yellowhorn are from beech. Perhaps beech is a principal host of the Yellowhorn while the birch family includes the principal hosts for the Close-banded Yellowhorn. Larval color varies within both the Yellowhorn and Close-banded Yellowhorn, so much so that we cannot reliably discriminate between the two. We encourage others to collect wild larvae, take pictures, and save reared adults. The pupa overwinters in a cocoon.

254 Hieroglyphic Moths Subfamily Diphtherinae

Hieroglyphic Moths

Family Noctuidae: Subfamily Diphtherinae This newly recognized subfamily (Fibiger and Lafontaine 2005) is represented in our region by a single tropical species. The gregarious, boldly-marked caterpillar can be confused with no others in our fauna.

Hieroglyphic Moth Diphthera festiva ! recognition Handsome black and white caterpillar with bright red-orange head and anal plate. Ground color white to blue-white with three to four narrow, incomplete black rings on T2–A7; ring anterior to spiracle thickest. Rings ending below level of spiracles, and sometimes broken by black middorsal and/or subdorsal black stripes. Prothoracic plate shiny black. Red to orange subventral spots, especially over prolegs on A3–A6. Prolegs with shiny orange and black plate. Larva to 4.5 cm. ! occurrence Tropical scrub, hammock edges, disturbed sites, and other open habitats from coastal North Carolina south through Florida, west through much of Texas, and southward at least to Bolivia and Brazil. Adults occasionally straying northward. Multiple generations with caterpillars through growing season in southern Florida. Common southward. ! common foodplants Most records from chocolateweed and pyramidflower (both Melochia) and sleepy morning (Waltheria) (all cacao family, Sterculiaceae); also soybean, bush clover (Lespedeza), and Mimosa (all legumes), and members of mallow family. Covell (2005) gives “pecan, and other trees; sweet potato” but these records seem dubious—perhaps older larvae wander to alternate foodplants, especially after having defoliated their original host. ! remarks The boldly colored caterpillars feed openly in the daytime in clusters. Their coloration and gregarious behavior leaves little doubt that they are chemically protected. Birds feeding on Hieroglyphic Moth larvae sometimes reject their prey and spit the caterpillars back out (Collins and Watson 1983). Four of the Hieroglyphic Moth caterpillars collected by Dan Janzen’s “gusanero team” in Costa Rica proved to be free of parasitoids (http://janzen.sas.upenn.edu/), a feat more species wish they could boast of. In the wild, cocoons are often found along stems and in the crotches between twigs (above, lower right). New molecular data collected by Reza Zahiri (pers. comm.) suggest that Diphthera is quite divergent and may be better classified outside of Erebinae. Presumably overwintering, at least northward, as a pupa.

Brothers Subfamily Dilobinae 255

Brothers

Family Noctuidae: Subfamily Dilobinae The subfamily is represented in the East by only a single genus with two closely related species. The stocky caterpillars bear small, dorsal processes on T2. The SV group on A1 is bisetose. Brothers possess characteristics in common with both daggers (Acronictinae), and pantheas and yellowhorns (Pantheinae). Our species feed on aspen, poplar, and willow.

The Brother Raphia frater ! recognition Blue-green to bright lime green with yellow spotting and transverse yellow lines running over A1, A5, and A8. Yellow lines edged with red or white anteriorly. T2 with rose to red dorsal conical projections, yellowish at their bases and rose-pink apically. Larva to 3 cm. We are unaware of characters that will distinguish larvae of the closely related Abrupt Brother (R. abrupta) (inset). In our limited material the dorsal projections average higher in R. abrupta. The Abrupt Brother is found mostly in the Midwest and Gulf states; it occurs north to Maryland, New Jersey (once), and Indiana. It is unreported from peninsular Florida, Connecticut, Ohio, Wisconsin, among others. ! occurrence Edges of watercourses, late successional fields, woodlands, and forests. Transcontinental in Canada, south in East to Virginia, Missouri, and Kansas. At least two generations with mature caterpillars from late June to October. Common, northward and inland. ! common foodplants Especially aspen and poplar, also willow; records from alder and birch strike us as erroneous. ! remarks The eggs are laid in a small group, often in a linear, partially overlapping array. The early instars are narrowly elongate, their appearance contrasting markedly with that of the stubby later instars. The first two pairs of prolegs are rudimentary even in the second instar—the larva, predictably, loops. The caterpillars rest on leaf undersides by day and are easily found by turning branches or examining leaves from below. Provide mature caterpillars with pulpy wood or a roll of tightly wrapped cardboard into which the larvae can tunnel and fashion their pupal cells. Like other noctuids that fashion pupal chambers in wood, the prepupa turns red. Winters are passed as a pupa.

256 Balsas Subfamily Balsinae

Balsas

Family Noctuidae: Subfamily Balsinae Balsas were recently placed in their own subfamily by Lafontaine and Fibiger (2006) because the SD1 seta on A9 is unmodified, i.e., not hairlike. We place the subfamily near the Dilobinae, in part because of the superficial similarities be-

tween the caterpillars of Raphia and the Many-dotted Balsa (Balsa malana), and because members of both subfamilies form pupal cells in soft wood and bark. Prepupae take on a pinkish-red cast. The prolegs on A3 and A4 are reduced. Balsas overwinter as pupae. We have had success obtaining livestock by sleeving well-fed females.

White-blotched Balsa Balsa labecula ! recognition Bright lime to yellow-green with conspicuous white to yellow spots, and yellowish spiracular stripe. Dorsal pinacula prominent, round to more commonly transversely oval, somewhat raised. Spots between setal bases more irregular in outline. Posterior half of A3–A8 with elongate, more or less parallelsided, middorsal spot that extends forward to posterior (D2) dorsal setae. Subdorsal stripe broken. Head pale green, unmarked, shiny. Larva less than 2.5 cm. ! occurrence Forests, swamps, barrens, and woodlands from Manitoba to eastern Canada, south to northern Florida and Texas. One principal generation in Northeast with mature caterpillars in June and July, and partial second brood in late summer. Locally common. ! common foodplants Our larval collections from chokeberry and hawthorn; we reared one cohort on serviceberry, a common shrub or small tree at sites where we have taken adults at bait and light in New Jersey. ! remarks The larva rests with the head curled to one side as shown. Balsas are semiloopers—only the prolegs on A5 and A6 (and A10) are used in walking. Caterpillars are readily collected with beating sheets. If available, prepupal larvae will tunnel into bark and soft wood, and construct a cocoon of silk and interwoven wood chips much like daggers (Acronictinae). Our pupae from a first-brood New Jersey female went into diapause (in early July) and did not yield adults until the following year, yet second-brood adults occur regularly in July in New Jersey.

Balsas Subfamily Balsinae 257 Three-lined Balsa Balsa tristrigella

Minnesota to eastern Canada, south to Georgia and Texas. Two generations over much of East with mature caterpillars in early summer, then again in fall. Common. Our collections from apple, hawthorn (Rockburne and Lafontaine 1976), and perhaps related members of rose family. A3–A8 with middorsal spotting ending rearward of D2 setae (compare with White-blotched Balsa, B. labecula).

Many-dotted Balsa Balsa malana ! recognition Small, stout, heavily spotted; somewhat spindle shaped, thickest through thoracic segments. Pinacula often white; other spots frequently with yellow cast. Head also heavily spotted. Prolegs short, inconspicuous from above; those on A3 and A4 slightly reduced. Spiracles orange. Anal prolegs directed backward. Larva less than 2 cm. Much shorter and stockier than our other eastern Balsa. ! occurrence Orchards, parks, and woodlands from Manitoba to Nova Scotia, south to Florida, Mississippi, and possibly Texas. Two generations in Ohio and Connecticut with mature caterpillars in June and early July, then again from August to October. Sometimes common. ! common foodplants Apple and crabapple (Rings et al. 1992) most commonly. Handfield et al. (1999) also list serviceberry. Cherry, elm, pear; Covell (2005) includes plum. Elm record erroneous (and may refer to misidentified caterpillar of Delightful Dagger, Acronicta vinnula). ! remarks The caterpillar is widely known as the Many-dotted Appleworm. The figured individual was beaten from apple in early August (in Goshen, Virginia). We suspect host records from cherry refer to domestic varieties— we have not encountered the Many-dotted Balsa in our collections from black (Prunus serotina) and chokecherry (P. virginiana), nor do we associate adults with either plant.

258 Daggers Subfamily Acronictinae

Daggers

Family Noctuidae: Subfamily Acronictinae Daggers derive their name from the sharp, black (sometimes knifelike) dashes present in the adult forewing markings. The eastern fauna consists of seven genera and about 60 species. More than 50 of these are Acronicta, an extraordinarily successful genus if evolutionary prosperity is measured by species diversity. Acronictines buck the species diversity trend for most animals, i.e., their richness drops southward; Costa Rica has but two and both of these are restricted to mountainous areas. We treat 37 Acronicta individually and another ten are figured in plates with abbreviated text. We present species in groups based on their relatedness (or appearance) and/ or their principal host associations. Chris Schmidt provided helpful distributional data and taxonomic insights. The genus had long captured the interest of Charles V. Riley, our nation’s first state entomologist, chief entomologist for the U.S. Department of Agriculture, and father of biocontrol in the New World. It was Riley’s hope to complete a systematic work on Acronicta, whose larvae were of special interest to him, but his untimely death in 1895 robbed him of the chance to complete his studies. His notes were subsequently passed along to John B. Smith and incorporated into Smith and Dyar’s (1898) monograph on the genus—a work rich in life history information on our fauna. Acronicta caterpillars are marvelous animals—the warblers of the temperate caterpillar world, colorful, diverse, and somewhat elusive—and once a hand lens is swapped for binoculars, they appear more beautiful still. Their behavior and natural history ranks among the more interesting and varied in our fauna, with marked behavioral and morphological changes sometimes occurring across instars, and in at least one species (Greater Oak Dagger, A. lobeliae), even within an instar (see Transformer essay, page 36). No larval (or adult) characters are known to be uniquely shared by members of the subfamily. Most acronictine caterpillars are endowed with conspicuous, long, hairlike setae, and in many these arise from tufts or warts. Secondary setae are

 Yellow-haired Dagger (Acronicta impleta). 

nearly always present, especially above the prolegs (although members of the genus Cerma have only primary setae, page 304). The integument is often granulose or microspinulose. The head has only primary setae (with the exception of Fragile Dagger, A. fragilis). The distance between the first and second stemmata is greater than that between the second and third; in many the second stemma is elliptical (Crumb 1956). Typically, the head crests well forward of its posterior margin. Most have two or more setae arising from the L3 pinaculum on A1–A8 (Crumb 1956; Beck 1999, 2000) (but again, Cerma has only a single seta). The dorsal pinacula are frequently fused over T2 and are even more likely to be fused over T3. Several Acronicta resemble lasiocampids, prominents, tussock moths, and tiger moths, and to our eyes, provide many compelling cases of caterpillar mimicry (see essay on page 78). The somewhat flattened eggs are laid singly. Larvae pass through six to eight instars. Many notes on morphological differences across instars are given in Smith and Dyar’s (1898) monograph. The caterpillars are grazers in the first two instars, skeletonizing irregular patches of tissue, usually from the lower leaf surface. Larvae consume their skins after a molt and may even scour into underlying and adjacent leaf tissue (Judy Semroc, pers. comm.), thereby eliminating chemical and physical clues of their whereabouts. Later instars of several daggers sometimes carve out deep cavities to either side of the midrib; such damage is diagnostic and can be used to locate larvae, e.g., see Fragile Dagger. In the middle to the penultimate and occasionally through the final instar, a large fraction feed and perch on upper leaf surfaces in full view, suggesting that they enjoy some level of protection from visual predators; the nature of this protection remains unstudied. Leaf perchers spin a slight sheeting of silk below their perch, into which the crochets engage. Some late instar daggers are quick to snap (their head and thorax) at would-be attackers. Presumably the long spatulate setae of some, and the retained head capsules of Harris’s Three-spot (Harrisimemna trisignata), provide additional defense, serving as swatters and maces (respectively) to strike away parasites and other small attackers. Cocooning habits vary greatly and no doubt are reflective, at least in part, of underlying phylogenetic relationships within the genus. The Fingered Dagger (A. dactylina) and a few others make a soft, arctiinelike cocoon: spacious, fashioned with generous amounts of silk and riddled with setae from the last instar. Stout, tough-walled cocoons—with interwoven frass, feculae, and soil bits—are produced by several woodland members of the genus (see page 260). Others make a slight cocoon among tied leaves, usually in duff or leaf litter at the soil surface. The lion’s share excavate a pupal cell in pulpy wood; although some silk may be used to tie up the shavings during the chamber’s construction, the cell itself tends to be mostly free of silk. The winter is passed as a pupa, sometimes for as many as five winters in some European species (Ahola and Silvonen 2005), although we have not observed protracted emergence in our collections.

Daggers Subfamily Acronictinae 259 Acronictines may undergo dramatic color, behavioral, and life history changes between the penultimate and ultimate instars; these changes may even take place over the course of the last instar as in The Hebrew (Polygrammate hebraeicum), Greater Oak Dagger (Acronicta lobeliae), and others. With the exception of Simyra and some (possibly related) Acronicta, members of the subfamily feed on shrubs and trees. While most are foodplant specialists, quite a number are generalists on woody broad-leaved shrubs and trees. The American Dagger (A. americana) and Narrow-winged Dagger (A. lanceolaria) also include conifers in their diets. Most dagger caterpillars prefer older foliage—their large heads and color patterns are testimony enough that this is normal fare. Exceptions include two spring fliers: Doll’s Dagger (A. dolli) and Narrow-winged Dagger are univoltine generalists that feed up on the new leaves of trees and shrubs. The

Long-winged Dagger Group Acronicta near longa

We have received images of this caterpillar on three occasions from Mississippi and Texas. Hosts include beautyberry (Callicarpa), dogwood, and wax myrtle (Morella cerifera). While closest in appearance to the Long-winged Dagger (A. longa), the caterpillars are consistently distinct, and so we wonder if an as yet unrecognized species is involved. Those encountering the caterpillar are encouraged to photograph, save, and rear examples.

Hazel Dagger Acronicta falcula

At least Manitoba to Massachusetts, south to northern New Jersey, southern Ohio, Iowa (Forbes 1954). Two generations with mature caterpillars in June and again in September. Generally rare (in part, because adults are only weakly attracted to light). American and beaked hazel.

The Agate Cerma cora

Eastern Canada, south to Florida and Texas, but rare or absent over much of this range. One generation with caterpillars in late spring (Florida) to August (Ontario). Generally rare; locally common in central Florida and Maine. Most records from hawthorn and pin cherry. See account in Schweitzer et al. (2011).

majority of eastern species are double brooded through much of their eastern range, although it is sometimes difficult to disentangle the broods (our overwintered full-sibling pupae of A. albarufa, A. lithospila, and A. lobeliae eclosed over a period of eight weeks or more). Second broods tend to be partial, and especially so northward. Offer prepupal larvae pulpy wood in which to tunnel. Access to wood or equivalent pupation substrates is particularly important to those species that turn red as prepupae. Adults of several species come to bait, although usually in modest numbers. To obtain eggs, hold females in smooth-walled containers. Acronictines can be recalcitrant about ovipositing in captivity, e.g., we have yet to figure out a means of obtaining eggs from The Hebrew. Members of the oak dagger complex (A. haesitata and kin, page 276) are also reluctant to lay in containers and are best fed and then sleeved on a suitable host.

260 Daggers Subfamily Acronictinae

American Dagger Acronicta americana J recognition  Large, densely vested with white or pale yellow setae from which emerge black, diverging, dorsal hair pencils on A1 and A3. Integument usually pale green. Unpaired, black hair pencil from dorsum of A8. Larva to 5.5 cm. Early and middle instars often more yellow. J occurrence  Woodlands and forests, especially mesic to swampy bottomlands, from southern Canada to Florida and Texas. Evidently one principal (extended) generation with mature caterpillars from July to October in Northeast; two or three broods from Missouri and New Jersey southwards. Common. J common foodplants  Alder, American hornbeam, ash, basswood, beech, birch, blueberry, boxelder, buckeye, elm, hazel, hickory, maple, oak, poplar, redbud, red cedar, sycamore, walnut, willow, and many other woody plants. J remarks  The American Dagger caterpillar rests with its head curled to one side. Larvae are commonly encountered in late summer and fall beating samples from woody vegetation, and especially so from basswood, red and silver maples, and members of the birch family. Drooz (1989) makes brief mention of an outbreak of the American Dagger in Maine. While harmless to the touch, roughly handled caterpillars, and particularly those that find their way into clothing, may produce a severe rash. The prepupal caterpillar excavates a pupation crypt in wood or bark. Gnawings are combined with silk and perhaps other oral secretions to fashion a tough-walled, domed-shaped cocoon over the caterpillar’s troughlike excavation. The cocoons sometimes turn up in odd places: e.g., in wood siding or firewood. In some situations, the caterpillar will spin a flimsy outer layer or veil (which we have detached and flipped up in the image to right); this affords the caterpillar modest protection during the long tunneling process. One of the parasitic wasps to attack the caterpillar is Aleiodes stigmator (Braconidae)— the female deposits numerous eggs in each host larva. Upon consuming the caterpillar, the wasp larvae pupate within the cadaver and chew their way out of their mummified victim as they emerge; they leave the corpse riddled with holes, nearly two dozen of which are visible in the Parasitoid Plate (row 3) on page 28.

Daggers Subfamily Acronictinae 261

Connected Dagger Acronicta connecta  recognition Blue-green to dark green with prominent yellow subdorsal stripes and blackened dorsal pinacula. Dorsal setae long: many black. Abundant white lateral setae below level of spiracles. Lemon subdorsal stripe edged with white extending from T1 to A10. Prothorax with elbowed to triangular spot bordering pale middorsal stripe. Head coloration diagnostic: each lobe with triangular red mark that narrows and darkens to eyes (antenna). Riley in Smith and Dyar (1898) describes a brown-form larva. Larva to 3.5 cm. Penultimate instar with less prominent dorsal setae, wine red spots about dorsal pinacula and D1 pinacula over A1 and A4 nearly joined by enlarged reddish spots; subdorsal stripe more irregular in expression (inset).  occurrence Wetlands and edges of watercourses from southern Ontario and Great Lakes states to central New England, south to Florida and Texas. Two generations with mature caterpillars mostly from June onward. Local and uncommon over much of East; more common southward.  common foodplants Willow.  remarks DLW once mistook a melanic female of the Connected Dagger for that of the Paddle Dagger (A. funeralis), and tried to rear the offspring on elm. Only a few A. connecta larvae survived. If available, larvae pupate in pulpy wood (Riley in Smith and Dyar 1898). The Connected Dagger has been reared from trap nests placed out for cavity-nesting bees (Godfrey 1987); evidently prepupal caterpillars occasionally enter holes, such as those fashioned by wood-boring beetles and bees, to seek suitable pupation sites. The adults are not common in collections; perhaps, like some other members of the genus, the Connected Dagger is only weakly attracted to light. Adults do show up at baits, sometimes half a mile or more from any willow. We are uncertain if the brown form described by Riley represents a (genetically) distinct morph, an inducible phenotype, or if larvae commonly turn cinnamon brown late in their development, as is the case with the Clear Dagger (A. clarescens), and to a lesser extent, the Witch Hazel Dagger (A. hamamelis).

262 Daggers Subfamily Acronictinae

Fingered Dagger Acronicta dactylina  recognition Densely hairy; chocolate brown setae over dorsum give way to frosty setae below level of spiracles. Black middorsal hair pencil over A1, A3, and A8. Integument black; spiracles white. Long, pale setae along sides; those over dorsum short and bristlelike. A few long setae extend from either end of body. Larva to 4.5 cm. Middle instars (inset) often more salmon or peach above.  occurrence Wetlands, edges of watercourses, and fields with woody growth from Manitoba to Newfoundland, south to Georgia (mountains) and Illinois. Evidently just one generation with mature caterpillars mostly in August and September. Common northward.  common foodplants Alder, aspen, birch, and willow; also dewberry (Rubus).  remarks This is one of the most frequently encountered daggers across our northern tier of states and Canada. The larva perches and feeds from either leaf surface, bears rather bold coloration, and its feeding damage is apparent—these are all indications of unpalatability. Evidence of chemical protection is all but lacking for members of the subfamily Acronictinae. Northward, the Fingered Dagger is routinely encountered in beating samples from alder. When alarmed the larva curls up and exposes jet black integumental membranes; the venter too is black and contrasts boldly with the animal’s frosty lateral setae. The prepupa tunnels into wood or fashions a spacious silken cocoon into which many of the larval hairs are interwoven. Cocoons spun in the open can be easily mistaken for that of a tiger or tussock moth.

Daggers Subfamily Acronictinae 263

Hoary Alder Dagger Acronicta hastulifera [Frosted Dagger]  recognition Densely hairy with hoary dorsal setae giving way to mostly orange setae along sides. Single, very long seta extending from dorsal and lateral warts at either end of body. Setae over dorsum mostly black with frosty tips (giving caterpillar its hoary appearance); orange subdorsal and lateral setae also white tipped. Black lashes absent. White spiracles contrasting with flat black ground color. Head shiny black. Larva to 4 cm. Penultimate instar with some setae over dorsum white to base, and short, closely set, paired, black lashes on A1, A3, and A8 (inset).  occurrence Swamps, wet meadows, and edges of watercourses from southern Ontario to Georgia. Our larval collections are from August and September, suggesting a single annual generation. Local; sometimes common in Appalachians.  common foodplants Alder, birch, and hazel, although nearly all our wild larvae from the first.  remarks While generally regarded as rare (Forbes 1954, Rings et al. 1992), the Hoary Alder Dagger is locally common in alder swamps at middle and high elevations in the Appalachians of West Virginia and Pennsylvania. Alarmed caterpillars will tenaciously engage the leaf on which they are perched. Laura Miller sent us our only record from birch, a singleton beaten from river birch (Betula nigra) in West Virginia. Literature for this species and the Fingered Dagger (A. dactylina) may be confused because the adults are almost inseparable. Prepupal larvae will tunnel into pulpy wood if available.

264 Daggers Subfamily Acronictinae

Paddle Dagger Acronicta funeralis [Funerary Dagger] J recognition  Flamboyant and unmistakable: black, white, and yellow with spatulate dorsal setae. Dorsum of each segment boldly marked with yellow and black spot—those over first eight abdominal segments reminiscent of bottomside-up cowry shells. Larva to 3.5 cm. Early instars are bird-dropping mimics with spatulalike setae only on first thoracic segment (upper right images). J occurrence  Fields and woodlands from southern Canada to Georgia, Mississippi, and Texas, although absent from many parts of this range (e.g., much of Atlantic Coastal Plain and Piedmont). Two to three generations in Missouri with mature caterpillars from June onward. Mostly uncommon. J common foodplants  American hornbeam, alder, apple, birch, blackberry, blueberry, cherry, hazel, hickory, mountain ash, oak, poplar, willow, witch hazel, and other woody plants. J remarks  Dietary staples in our area include oak and plants in the birch (Betulaceae) and rose (Rosaceae) families. Related Old World species, sometimes classified in their own genus (Jocheara), are associated principally with oaks. The differences between the last two instars of this stunning insect are nothing short of spectacular—one wonders how and why evolutionary forces have pushed the appearances of the two larval instars so far apart. The middle to penultimate instars, especially when curled to one side, resemble bird droppings. The adaptive significance of the final instar eludes us—and surely there is one as the caterpillar will perch on upper leaf surfaces, exposed in all its glory. Porter (1997) remarks that the related Alder Moth (A. alni) of Britain is warningly colored, and one is compelled to agree, but such a claim presupposes that the caterpillar is in some way protected. Although many Acronicta are brightly colored, conspicuous in habit, and sometimes foul-smelling (see Long-winged Dagger, A. longa, page 293), studies are needed to document that they are, in fact, manufacturing or sequestering toxic compounds and are unpalatable. A single prepupal Paddle Dagger reared by DLW on hazel released a detectable scent when squeezed, but the smell was neither especially unpleasant nor lasting. When touched the caterpillar is quick to snap its anterior end to the point of contact—presumably the paddles serve to knock away ants, flies, wasps, and other small enemies. Pupation occurs in soft pulpy wood, if available.

Daggers Subfamily Acronictinae 265

Interrupted Dagger Acronicta interrupta  recognition Red, white, and blue-gray, immediately recognizable by its paired middorsal red spots: A1–A7 with red dorsal spots that may fuse, especially rearward. Broad, frosty white to cream subdorsal stripe. Dorsum of A8 swollen, smoky, and bearing wartlike setal clusters. Subdorsal and lateral pinacula only modestly raised and frequently yellowed. Setae long; orange over dorsum and mostly white along sides. Head dark, reddish brown to black, with or without red patch over each vertex. Larva to 4 cm. Middle instar shown in inset. The larva of the Square-eyed Dagger (A. quadrata), a boreal species, is similar. Its caterpillar is more boldly marked, bears little red on A1, sports larger red patches over A5–A7, and possesses broader subdorsal stripes (see Ives and Wong 1988: 125). Like the Interrupted Dagger, its principal hosts are cherry, serviceberry, and other members of the rose family.  occurrence Woodlands and forests from Manitoba to Nova Scotia, south to Florida and Texas, although absent from many parts of this range and becoming scarce southward. Two generations over much of East with mature caterpillars from June to September. Locally common northward.  common foodplants Apple, cherry, chokeberry, mountain ash, plum, serviceberry, and related plants in rose family (Rosaceae) commonly; other reported hosts include basswood (Tim McCabe), birch, elm, and ironwood. In our experience pin cherry and apple preferred over black cherry; black birch frequently in central Connecticut.  remarks Rings et al. (1992) report the Interrupted Dagger as abundant in Ohio. In southern New England it tends to be local and scarce, and in some years goes undetected, although in 2003 the species was common over eastern regions of the state. In northern New Jersey and Pennsylvania adults are frequent in woodlands with apple or serviceberry. The dark spot between the fifth and sixth segments of the penultimate instar figured to right marks the location of a tachinid fly larva. In the center of the spot the fly maggot has pushed through its posterior spiracles so that it can breathe while it feeds within its caterpillar host.

266 Daggers Subfamily Acronictinae

Radcliffe’s Dagger Acronicta radcliffei  recognition Dark brown with prominent, yellow middorsal, subdorsal, supraspiracular, and thickened subspiracular stripes. Dark rump patch over A8. Thorax and abdomen with long white setae; those over dorsum greater than twice length of segment that bears them. Upper half of head cherry red. Larva to 3.5 cm. Penultimate instar blue-green with broad, red-brown dorsal stripe, edged with yellow, and extending length of body (inset).  occurrence Fields, swamps, barrens, and woodlands from southern Canada to mountains of Georgia and northern Arkansas (and northern California), but absent from many areas within this range. At least two generations with mature caterpillars from May through November. Locally common southward.  common foodplants Cherry and other members of rose family (Rosaceae), such as apple, chokeberry, hawthorn, mountain ash, and serviceberry.  remarks Yet another dagger with very different middle and last instars. As noted under the Yellow-haired Dagger (A. impleta), mimicry has rarely been reported as occurring in caterpillars. Last instar Radcliffe’s Daggers so closely resemble datanas (Notodontidae) that seasoned entomologists confuse them. The alarmed caterpillar even cranes the head back in the same fashion as do Datana (although not to the same degree). One wonders if these two insects, as well as others, such as the Orange-striped Oakworm (Anisota senatoria) and Turbulent Phosphila (Phosphila turbulenta), have somehow entangled their fates in a shared mimicry complex. All four taxa occur together in southern New Jersey and elsewhere, with the dagger, by far, the scarcest of the lot. Our image is of a recently molted last instar.

Daggers Subfamily Acronictinae 267

Clear Dagger Acronicta clarescens (= A. pruni)  recognition Last instar characteristically cinnamon to rusty brown with broken, gray middorsal stripe embedded in broader, brown stripe that widens over A4 and A5. Closely set paired knobs over T2–A3 and A8; dorsal knobs over A3 about half length of those over A2. First two thoracic segments bearing triangular, brown spot over dorsum, outwardly edged with white to yellow dashes (upper inset). Head bearing distinctive elongate black spots. Larva to 3.5 cm. Penultimate instars blue-green with red-brown middorsal stripe that widens on A4 and A5, and pale, whitish spots along sides of body; dorsal setae black, and setae along subventer downy white (lower inset).  occurrence Fields and woodlands (and yards) from Wisconsin to eastern Canada, south to Florida and Texas. Two generations over much of East with mature caterpillars from June to October. Uncommon over much of East, at least as an adult.  common foodplants Apple, cherry, chokeberry, cotoneaster, hawthorn, mountain ash, plum, and probably other woody members of rose family.  remarks The last instar, initially blue-green after the molt, transitions to the dramatically distinct, cinnamon-brown form shown above. The caterpillar is timid, quick to tuck its head under and arch the thorax upward—no other dagger in the East has such an exaggerated posture, dragonesque in profile. The Clear Dagger fashions its pupal chamber in soft wood if available, or failing that, in leaf litter. While we characterize the species as being uncommon above, such may be more true for the adult than the caterpillar, which turns up regularly across much of the Northeast, especially over late summer and early fall. Adults come well to baits in southern New Jersey.

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Fragile Dagger Acronicta fragilis J recognition  Bright green with broad, mostly reddish-brown dorsum from T2–A8 outwardly edged with broken, yellowish subdorsal stripe. T1 with subdorsal stripe absent. Long black setae from dorsal and subdorsal warts on T2 and T3, extending forward of head. Dorsal warts over T2–A8 deep burgundy, shiny, bearing numerous, black setae; those over abdominal segments relatively short (compare with Splendid Dagger, next page). Lateral setae dark, sparse (other daggers with many more lateral setae, especially above the prolegs). Head with secondary setae; reddish vertex giving way to black spot that narrowly reaches eyes. Larva to 3 cm. The larva of Heitzman’s Dagger (A. heitzmani) has not been described, although it likely resembles that of the Fragile Dagger, the species with which it had previously been confused. Its foodplant is unknown but if it is an evolutionary sister of the Fragile Dagger, one would suspect that it feeds on members of the rose family. It is a denizen of the Ozarks and Ohio River Valley. J occurrence  Edges of watercourses, woodlands, and forests. Transcontinental in Canada, south through New England and Kentucky, and in mountains to Georgia. Two generations north at least to southern Quebec with mature caterpillars from July to September. Common northward. J common foodplants  Alder, apple, beech, birch, cherry, mountain ash, plum, rose, serviceberry, and willow; Prentice (1962) also reports white spruce. See Remarks. J remarks  Larvae from a batch of Fragile Dagger eggs, sent to us by Gary Anweiler from Alberta, took well to gray birch; yet two clutches obtained from the Great Smoky Mountains straggled on leaves from the same plant. In hindsight we might have also offered our larvae apple or cherry, as many of the recorded hosts and most wild collections in the East are members of the rose family. Mitochondrial DNA from specimens taken in the Great Smokies differs somewhat from those found in Canadian populations of A. fragilis. The caterpillar of the Fragile Dagger feeds in from a leaf edge and chews its way through the blade down to the midrib—the damage is diagnostic. When such is done to either side of the midrib, the distal portion of the leaf (with larva aboard) becomes isolated, accessible only to those intrepid predators willing to traverse the still intact midrib out to the “refuge.” (Analogous feeding behaviors occur in limenitidine Nymphalidae and some Notodontidae.)

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Splendid Dagger Acronicta superans  recognition Beautiful blue-green caterpillar with broad, chocolate- to red-brown dorsal stripe outwardly edged with yellow. Long, black dorsal setae arise from black pimplelike pinacula. Numerous, white secondary setae below level of spiracles. A8 with sharp hump; middorsal stripe lighter brown, rearward from hump. Head distinctively marked: lobes orange-red above with bold black spotting, frons rosy, lower sides cream and unmarked. Larva to 4 cm. Co-occurring with the Fragile Dagger, A. fragilis, which differs in coloration and overall shape. In the Fragile Dagger, the dorsal markings never form a broad, solid stripe, and A8 is but little elevated and bears four cherry red warts. Dorsal warts in the Splendid Dagger are blackened and less wartlike. In addition, each dorsal wart in A. superans bears one very long blackened setae; in A. fragilis each dorsal wart bears two to six blackened setae, none of which approach the hypertrophied condition of the Splendid Dagger. Penultimate instars of Radcliffe’s Dagger (A. radcliffei) are also close in appearance. Its caterpillar lacks the prominent black snowflaking on the head capsule and the red-brown middorsal stripe on A9 and A10 that are present in the Splendid Dagger.  occurrence Shrubby fields, powerline right-of-ways, orchards, woodlands, and forest edges from Manitoba to Newfoundland, south to Pennsylvania, Great Lakes states, and North Carolina (mountains). Two generations in Connecticut with mature caterpillars in July, then again in August and September. Common.  common foodplants Woody members of rose family (Rosaceae): apple, cherry (e.g., pin but not black), hawthorn, mountain ash, plum, and serviceberry, with many reports for first and last of these; also reported from birch and hazel (both Betulaceae).  remarks Across much of Canada and the northern United States this is among the more common of the dozen or so daggers known to feed on cherry. The late instars rest with the thoracic and anterior abdominal segments arched upward; commonly, the large head, pulled well into the thorax, is appressed to the leaf. Prepupal larvae turn smoky green to cinnamon or red-brown (inset), and will fashion a pupal chamber in soft wood if available.

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Cherry Dagger Acronicta hasta (= A. furcifera) [Speared Dagger]

 recognition Last instar unmistakable: charcoal with broken bright red

middorsal stripe outwardly edged with black. Larva to 4 cm. Penultimate instar green with broad red and yellow middorsal stripe that narrows rearward and A8 somewhat humped (inset). Vertex and head to either side of triangle bright red.  occurrence Fields, powerline right-of-ways, dunes, woodlands, and forest edges from Canada to Florida and Texas. Evidently two generations in Northeast with mature caterpillars from June onward; two or three annual broods in Missouri and New Jersey. Second and third broods appear to be partial. Common.  common foodplants Cherry (especially black) and plums.  remarks Cherry and plum have a rich dagger fauna in the East: six Acronicta use Prunus as a primary foodplant and another half-dozen feed on it occasionally. Several of the cherry (Rosaceae) specialists have similarly colored middle and penultimate instars, but there will be no mistakes if you wait until the last instar to make your identification—all 12 are decidedly distinctive. Middle and penultimate instars of the Cherry Dagger perch on the upper side of leaves. We have never seen the black and red final instar on foliage except immediately following the last larval molt and at night while examining foliage by flashlight— by day, the last instars rest on bark, usually along the main trunk. We have had much success locating last instars on isolated trees in fields with lower trunk diameters in the range of 7–15 cm. A thin cocoon of brown silk is spun in leaf litter or upper soil layers.

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Gray Dagger Acronicta grisea  recognition Humpbacked, emerald to yellow-green (and tending toward latter in mature individuals) with reddish-brown “saddle” over A3–A8. First two thoracic segments with reddish Y-shaped spot over dorsum, connecting to abdominal saddle by nearly complete middorsal line over T3–A3. Abdominal saddle broadest over A4–A7, enclosing green middorsal “island” on A6 and A7. Dorsal setae borne from brown, shiny, pimplelike warts; these largest on A1 (D1) and A8 (D2). Head mostly red to brown except for green triangle and “cheeks.” Larva to 2 cm. Middle instar shown in inset. Similar to Hazel Dagger (A. falcula) (page 259) in color, form, and posture. Its dorsal pinacula are more warted, especially over A1, A4, A5, and A8; in addition the middorsal line over T3 to A3 is largely absent.  occurrence Edges of watercourses, wetlands, wooded swamps, woodland and forest edges and openings across Canada, south in East to Massachusetts and Great Lakes states. Curiously absent (or rare) from the Appalachians south of New York. One generation with mature caterpillars mostly in August and September. Common northward.  common foodplants Alder, apple, birch, cherry; also reported from American hornbeam, dogwood, elm, mountain ash, poplar, willow, and others. Frequently associated with mountain ash, paper birch, and pin cherry in the Adirondacks.  remarks Unlike the vast majority of caterpillars that hide by day or feed from the lower side of leaves, middle to penultimate instars of many daggers perch on leaf upper sides with their crochets enmeshed in a thin sheet of silk that they spin over the leaf surface. Prior to molts additional strands of silk are laid down. Because each silk fiber contracts as it is deposited, the molting platform may become elevated above the leaf surface (inset). Feeding damage of late instars is diagnostic: the caterpillar eats from a leaf edge down to the midrib and then does the same to the other side of the leaf, leaving the distal portion of the leaf as a “flag.” See also the Remarks section of the Fragile Dagger (A. fragilis). Our captive larvae incorporated small leaf fragments into the walls of their cocoons, or occasionally tunneled into pulpy wood when it was offered.

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Streaked Dagger Acronicta lithospila  recognition Blue-green with rosy to reddish-brown saddle over dorsum; the saddle is outwardly edged with cream or white. Saddle narrowing to white middorsal stripe over A3 and from there continuing forward over thorax to T2 where it flares outward; middorsal stripe absent on T1. D1 pinacula on A4 and A5 darkened. Setae above level of spiracles mostly long and dark; those below level of spiracles paler, shorter, and more numerous. Upper portion of head with reddish band that narrows to eyes; lobes with reddish to black snowflake spotting. Larva to 3.5 cm.  occurrence Barrens and dry oak woodlands from southern Canada to Florida and Texas; local and scarce north of barrens of Cape Cod. Two generations over much of East with mature caterpillars from June to October (but see below). Common southward.  common foodplants Principally oak and chestnut, but also reported from hickory and walnut.  remarks Like other daggers the first two instars are scrapers—they feed partway through the leaf, leaving vascular tissues in place. Upon cessation of feeding, prepupal larvae lose their blue-green coloration and gradually redden over a period of a day or two (inset). In our experience, daggers that turn red in the last instar are frequently the same species that prefer to tunnel into pulpy wood to pupate. The Streaked Dagger will wander some distance to locate a substance into which it can tunnel. Although specimen data is suggestive of two broods (e.g., see Rings et al. 1992), it is also possible that there is but a single protracted generation. A cohort of larvae that were reared from a clutch of eggs obtained from a female taken in early August 2004 by DFS in southern New Jersey produced adults over an eight-week period in June and July.

Daggers Subfamily Acronictinae 273

Triton Dagger Acronicta tritona  recognition Yellow green with dorsal reddish to purple-brown saddle, edged with yellow, and interrupted over A2. Prothoracic shield with oblique black bar that carries forward as reddish bar to eyes. Dorsal saddle enveloping green “island” over A4–A7. A8 strongly humped. Dorsal pinacula enlarged on A1 and A8. Setae above level of spiracles long, black, and shiny; those below mostly shorter and pale; numerous secondary setae above prolegs. Larva to 4 cm. Lime-green penultimate instar similarly marked, but with yellow coloration absent or more subdued (upper inset).  occurrence Heathlands, bogs, barrens, ericaceous woodlands, and acid swamps from at least Manitoba to Nova Scotia, south to Florida and Texas. Two generations in Connecticut with mature caterpillars in July, then again in August and September; additional generations southward. Locally common.  common foodplants Azalea, blueberry, and possibly other heaths.  remarks The anterior abdominal segments are often raised, giving the caterpillar a humpbacked aspect, especially when molested. The caterpillar’s markings suggest a relationship with the Fragile (A. fragilis) and Gray (A. grisea) Daggers. Prepupal larvae take on a wine to maroon cast (lower inset). Like many daggers, the larva pupates in old bark or dead wood, especially where these substrates are in contact with soil. Prepupae excavate a shallow area out of the wood and then spin a cocoon over themselves, incorporating soil and wood particles. Although Covell (2005), Rings et al. (1992), and Handfield (1999) regard the moth to be uncommon to rare, it is often common over acid soils of the Atlantic Coastal Plain with an abundance of heaths. Adults come to light and occasionally to bait. The few wild caterpillars that we have found have come from lowbush blueberries (Vaccinium angustifolium and V. pallidum) and swamp azalea (Rhododendron viscosum).

274 Daggers Subfamily Acronictinae

Witch Hazel Dagger Acronicta hamamelis (= A. subochrea)  recognition Last instar green or cinnamon with white to pink and dark brown dorsal markings. Dorsum with dark lens-shaped spots over A1, A4, A5, and A8. A1 and A8 with raised dark wart (bearing D1 seta) in lens-shaped spot, and pale wart (bearing D2 seta) just outside it. A8 bent strongly downward. Limegreen to cinnamon ground color most apparent along sides. Head with abundant dark snowflaking and long setae. Larva under 4 cm. Middle to penultimate instars bright green with contrasting smoky red patches over A1, A4, A5, and A8; dorsal pinacula on these segments bright, shiny red (upper inset).  occurrence Mesic forests and woodlands from Wisconsin to Nova Scotia, south to northern Florida and Texas. No longer found in Northeast (unless possibly in Nova Scotia and Maine) but still regular southward and westward from central Pennsylvania. Two generations with mature caterpillars from June onwards. Uncommon and local.  common foodplants Witch hazel.  remarks When alarmed the caterpillar hunkers down while raising the first two abdominal segments. If agitated further, the caterpillar swings around to bite its attacker. Fully mature caterpillars lose their bright lime-green coloration, and take on a smoky green to cinnamon coloration. A prepupal caterpillar that we reared wandered continuously about its container for more than three hours (and shrunk rather greatly in length), until we provided it with a piece of pulpy wood. It immediately began tunneling into the wood and within the course of the next hours managed to conceal itself. It is our guess that the caterpillar’s strongly angled eighth abdominal segment (and that of other wood-boring acronictines) facilitates tunneling by making it possible to generate a greater downward force while excavating the pupal crypt.

Daggers Subfamily Acronictinae 275

Afflicted Dagger Acronicta afflicta  recognition Last instar beige-orange or smoky orange to salmon with charcoal middorsal stripe (sometimes faint) and clubbed, black, dorsal setae about as long as segment that bears them. One pair of clubbed setae per segment (D2 setae). Pale spiracles ringed with black. Larva to 4 cm. Middle instars of Afflicted Dagger with bristly warts and pale middorsal stripe (upper inset) resemble those of the oak dagger complex (A. haesitata-increta). The Charred Dagger (A. brumosa) has two pairs of black, clubbed dorsal setae per body segment (both D1 and D2 setae) (lower inset). This oak specialist flies in xeric oak woodlands, barrens, and savannahs, from Colorado to Missouri, south to Texas; and in the East from New Jersey to peninsular Florida, west through the Gulf states. In the eastern portion of its range, it is a barrens species of the outer coastal plain. We associate the Charred Dagger with blackjack thickets in New Jersey. Moria Robinson collected wild larvae from turkey oak in North Carolina. Its yellowish middle instars are somewhat shiny and bear fewer secondary setae than the “spiny” middle instars of the Afflicted Dagger.  occurrence Barrens, woodlands, and forests from Wisconsin to Nova Scotia, south to Florida and Texas. Two or three generations over much of East with mature caterpillars from June to October. Common.  common foodplants Oak, both red and white groups.  remarks The heads of most daggers are large relative to the body size. Those of both the Afflicted and Charred Daggers (A. brumosa) are especially broad, allowing the caterpillars to grind up tough, fully sun-hardened oak leaves unmanagable to most caterpillars. Larvae can be found by scanning leaf undersides; nocturnal searches with illumination from below can be fruitful. Prepupal larvae of the Afflicted and Charred Daggers tunnel into bark or wood. Wood chips, laboriously interwoven into the wall of the overlying cocoons, render the pupation sites invisible.

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oak daggers

Acronicta haesitata-increta group This species complex is the most taxonomically challenging of eastern acronictines. Its members include A. albarufa, A. exilis, A. haesitata, A. increta (= inclara), A. modica, A. ovata, and A. tristis, and additional species still unrecognized. No two collections are likely to have the members of the complex sorted in the same way. What is recognized as A. increta in most collections includes more than one biological species, and we have never been confident about the limits of A. tristis. Rings et al. (1992: 76) provide a set of useful characters for the separation of the adults. Forbes (1954) regarded the caterpillars within the complex to be largely indistinguishable, although the larvae of the Lesser Oak Dagger (A. exilis) and Medium Oak Dagger (A. modica) differ substantially from the others in both form and habit. Our diagnoses and species accounts should be regarded as provisional until the taxonomy of the group has been reviewed. The caterpillars are large headed and thus well suited for feeding on hardened summer and fall oak foliage. All bear variously developed pale blotches over the dorsum of A2–A8. The dorsal spots fluoresce somewhat, under far UV blacklight (below). Members feed on beech, chestnut, oak, and presumably chinkapin (all Fagaceae). Northward “increta” also feeds on members of the birch family. Caterpillars perch exposed on lower leaf surfaces with the head curled back and held along the abdomen. They are easily mistaken for a leaf gall or blemish; their coiled bodies and markings also bear a resemblance to acorn caps. Larvae hold their purchase by laying down a thin film of silk about their perch into which the crochets are secured, but even without a silk lattice the caterpillars have a

tenacious grip. Members of the complex frequently seek refuge in leaf curls or in and about the shelters of oak-feeding microlepidopterans. The Lesser Oak and Medium Oak Daggers stand apart from all others in that they leave the foliage to seek out hiding places under bark flaps in the last two instars. And, like many other bark resters, their caterpillars can forcibly hurl themselves from their perch when exposed or otherwise alarmed. In nature the prepupal larvae excavate pupal chambers in softened wood or bark or make a cocoon of soil bits and litter; otherwise they will fashion cocoons in leaves and toweling if these are all that are provided. The prepupal larvae take on a caramel, orange, or more rarely, red-purple flush. There is a need to rear out cohorts from individual females to establish limits on the levels of variation displayed by families of larvae and/or adults. Captive females in this complex are often reluctant to lay eggs in vials; better success will be had by feeding females, then sleeving them on an appropriate oak. The larvae may be found by scanning the underside of oak foliage. When common, early instars can be quickly located by the distinctive windows (skeletonized patches) they chew into the underside of the leaf. If searching visually, be sure to examine protected areas in and about the shelters made by other species of Lepidoptera for later instars. Beating is also effective. Because some individuals take up residence in shelters, and two species hide on bark by day, nighttime searches with headlamps may be the most productive.  Eclipsed Oak Dagger (A. increta) larva. Some forms of this complex have bright white dorsal patches that fluoresce under far ultraviolet radiation. 

Daggers Subfamily Acronictinae 277 Barrens Dagger Acronicta albarufa

Widely scattered sites from Missouri to Cape Cod, south to North Carolina sand hills; westward reach unclarified due to confusion with A. exempta (Chris Schmidt, pers. comm.). At least formerly ranging north to Manitoba and Ontario; absent or historic from many states in this range. One principal (staggered) generation with mature caterpillars from July to October in New Jersey. Generally rare except in Ozarks and barrens of coastal Massachusetts. Oak: bur, dwarf, post, scrub, and presumably other oaks. See Schweitzer et al. (2011).

Beech Dagger

Acronicta increta “beech” This taxon is possibly an undescribed species. Range unclarified: at least Washington, DC area. Principal generation with mature caterpillars mostly in August and September. Locally common. Beech. Larva consistently smaller and more pale than Eclipsed Oak Dagger (A. increta), page 281.

Sad Dagger

Acronicta tristis Range unclarified because of confusion with melanic forms of Eclipsed Oak (A. increta) and Hesitant (A. haesitata) Daggers: at least eastern Canada, south to New England, Great Smokies, and Great Lakes states. Evidently two generations north to southern Quebec with mature caterpillars from July to October. Uncommon, or at least not recognized to be otherwise. Oak (mostly reds) and beech. (This image was borrowed from the LaPlante slide collection at the Canadian National Collection in Ottawa; we did not find the adult on which the determination is based.)

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Lesser Oak Dagger Acronicta exilis [Exiled Dagger]  recognition Small, darkly mottled, densely peppered with abundant minute black spinules; A8 slightly humped. Pinacula pimplelike. Dorsum of A8 pale above oblique, black, subdorsal line. A1–A6 with dark V-shaped middorsal spot rearward of dorsal setae. Head mostly beige with black coronal line that extends to triangle, and short black line directed back from eyes; triangle inwardly edged with black and lower portion outwardly bounded with black spot that extends only to antenna (lower right). Larva to 2.5 cm. Early to middle instars resembling those typical for oak dagger complex with long, stiff primary setae and pale, paired blotches over dorsum. Late instars are close in appearance to those of Medium Oak Dagger (A. modica), but smaller. In our examples, the black markings on the head, especially about triangle, are less extensive in the Lesser Oak Dagger, e.g., the black bar above the clypeus ends above the antenna (in A. modica the bar is thicker and extends outward to eyes).  occurrence Oak woodlands and forests from Wisconsin to southeastern Massachusetts, to Florida and Texas. Infrequent northward, but widespread and regular from Long Island to Florida panhandle. Two generations in southern New Jersey (second partial) with mature caterpillars in July, then again in August and September. Locally common.  common foodplants White oak and probably other Leucobalanus oaks, such as post and chestnut.  remarks The biology of the Lesser Oak Dagger and its kin, the Medium Oak Dagger (A. modica), is closely similar—comments made below apply to both. The first three instars feed from and rest on leaf undersides. The antepenultimate instar is mottled in yellows, reds, and brown, and is transitional in appearance between the pale early instars and the brown final instars (described above). In culture, the dark instars tend to seek shelter between two overlapping leaves, hide in abandoned leaf shelters of other caterpillars, or rest away from foliage. DFS found a wild larva of either the Lesser Oak or Medium Oak Dagger under white oak bark. The caterpillar is patterned and behaves much like those of the Common Oak Moth (Phoberia atomaris). Our first wild-collected larvae were lost because we failed to recognize that the caterpillars represented a species of Acronicta, and consequently we didn’t think it necessary to add wood or bark for pupation. A cohort of caterpillars that DFS sleeved on white oak in southern New Jersey matured in five weeks.

Daggers Subfamily Acronictinae 279

Medium Oak Dagger Acronicta modica  recognition Last two instars chocolate to smoky brown with pale mottling. Pale middorsal stripe most evident over thorax, and at leading and trailing edges of each abdominal segment. Subdorsal stripe comprised of broad, dark lower portion and thin, pale upper portion. Thin supraspiracular and spiracular stripes variously developed, but always broken and incomplete. Dorsal setae embedded in pale spot; D1 spots often meeting over midline rearward. Dorsal setae reduced in length; basal half whitish and contrasting with browner distal half. Integument densely peppered with minute black spines (visible with lens). Head shiny, broad, mostly beige with strong, black coronal bars that drop down and join prominent black “mask” extending between eyes, dodging up and over triangle. Thick black bar runs from lower edge of triangle to eyes (in A. exilis this bar ends above antenna). Larva to 2.5 cm. Similar in appearance and biology to caterpillars of the Lesser Oak Dagger (A. exilis) (previous page).  occurrence Oak woodlands and forests. Reported range from Minnesota, southern Canada, and southern Maine to Florida and Texas (Covell 2005), although many specimens in collections, and thus in literature, are confused with those of Hesitant Dagger (A. haesitata). One principal generation, probably with partial second generation southward, with mature caterpillars in July, then again in August and September (but see below). Common.  common foodplants Oak; our records from white oak.  remarks The larval coloration and flattened aspect suggest that the last two instars rest on or under bark. Late instars are quite motile and capable of covering considerable ground when moving between perches. If gently touched with a finger or brush, the middle instars are quick to turn and bite their attacker. Through the fourth instar, caterpillars may drop from their perch on a belay line— with the exception of its relative, the Lesser Oak Dagger (A. exilis), such behavior does not occur commonly in daggers. The last two instars are quick to tumble free from their perch; dislodged caterpillars roll about, usually making more net rearward movement—they do not snap violently to and fro like some underwings (Catocala), geometrids, and others. A southern New Jersey cohort reared from eggs laid in May yielded 26 adults the following May, all within 11 days of one another. However, adults are common in late July and August in southern New Jersey, which suggests that at least a partial second generation is produced.

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Hesitant Dagger Acronicta haesitata J recognition  Ground color of last instar ruddy, reddish, beige, tan, pinkish, salmon, orange, or yellowish, handsomely mottled with darker and paler splotches above. In our examples of A. haesitata, dorsal splotches, while contrastingly paler than adjacent coloration, are not as white or yellow as is commonly the case in Epauleted Oak Dagger (A. ovata). Middorsal stripe thin and broken. Integument minutely spined. Pinacula pimplelike, white to yellow. Head with reticulate patterning. Larva to 3.5 cm. Middle instars shown below right. We are unaware of reliable characters that will distinguish the caterpillars of the Hesitant Dagger from those of the Barrens (A. albarufa), Eclipsed Oak (A. increta), Epauleted Oak (Acronicta ovata), and Sad (A. tristis) Daggers. J occurrence  Woodlands and forests from southern Canada to South Carolina and Texas. May be partially double brooded in New Jersey where phenology is similar to that of Medium Oak Dagger (A. modica). John Lill’s collections from Maryland are suggestive of at least partial second and third broods from June into October. Very common. J common foodplants  Oak, mostly white oaks; reports from beech and chestnut require confirmation (see A. increta). J remarks  In our experience, this is the most common oak-feeding dagger in many mesic and northern woodlands and forests in the East. While beech, chestnut, and oak are listed as hosts, taxonomic uncertainty across the oak dagger group renders literature records suspect. From the Albany Pine Bush south through New Jersey and the Piedmont, A. haesitata appears to favor white oaks over reds. In southern New Jersey, DFS found 45 larvae in late June and July by checking the undersides of leaves on low branches of subcanopy white oak. Substantially more effort on southern red and black oak yielded no larvae. Late instars are frequently found inside curled, dead leaves or the abandoned shelters of other Lepidoptera (e.g., those of various microlepidopterans and the Confused Woodgrain, Morrisonia confusa). Substantial color changes may occur in the last instar: in one of our larval cohorts, the caterpillars were pale yellow to orange initially, but became more rusty orange as they aged. Prior to pupation, they took on the rose and red hues common to many wood-tunneling Acronicta. Caterpillars form pupal chambers in pulpy wood, or in bark and/or other organic litter near the soil line. Larval development is relatively slow, extending over six to eight weeks in some individuals; we have had final instars take nearly two weeks to complete their development.

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Eclipsed Oak Dagger Acronicta increta J recognition  Green, yellowish, pinkish, or salmon to orange with variously developed, paired, creamy splotches over dorsum of T2–A7. Extent of spots over T2, T3, and A7 often reduced. Larva to 3 cm. As noted above, the Eclipsed Oak Dagger is a member of a taxonomically difficult species complex—the only thing we are certain about is our uncertainty, both with regard to number of species involved and how the members can be separated from one another and the various forms of A. haesitata. J occurrence  Barrens, woodlands, and forests from southern Canada to Florida and Texas. Evidently a single protracted generation with mature caterpillars from late July to November in Northeast; small second brood near Philadelphia, and two full broods in Missouri. Often very common, especially northward. J common foodplants  Chestnut, oak (especially red and black oaks), and beech; also from hazel and birch northward (McCabe 1990; DLW, unpubl. data). J remarks  The species-level taxonomy of the Eclipsed Oak Dagger is in need of study. John Lill has sent us specimens of A. increta from Maryland and vicinity that we refer to as the Beech Dagger (A. increta “beech”); these may represent a distinct species. Adults are smaller on average, and the forewings are powder gray with a vague whitish field beyond the oblique antemedial line. The caterpillars are pale yellow or yellow-green with subdued coloration and markings; some bear subtle pinkish to reddish patterning over the trunk and head, and a well-developed middorsal stripe (page 277). Adults of A. increta from southern New Jersey and coastal North Carolina consistently stand out as being gray with extensive dark shading; we have not seen melanic adults in our samples despite their prevalence across the core of the Eclipsed Oak Dagger’s range. Caterpillars of this segregate are thought to feed principally on post and probably other white oaks; populations are bivoltine. What appears to be true A. increta is illustrated above. Where it flies with the beech-feeding entity that we are calling A. increta “beech,” adults average larger, have a darker ground color that ranges from smoky gray to charcoal, and have forewings that lack the contrasting field of whitish scales through the middle. A. increta is most often found feeding on red oaks, although caterpillars also can be found on white oak and beech.

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Epauleted Oak Dagger Acronicta ovata [Ovate Dagger] J recognition Ground color greenish, tan to pinkish or orangish, with paired dorsal patches down trunk. One form with bright creamy spots about dorsal setae; second form with spots greatly suffused with red or orange. Integument microspinulose. Pinacula white to yellow, slightly raised. Head patterning variable: weak, irregular, with brown-black coronal bars and dark line running back from lateral eyes, or without dark markings. In both cases, head is marked with abundant snowflake spots. Mature larva to 3 cm. Early instars more strongly mottled and prickly in appearance (inset). Because the Epauleted Oak Dagger is a member of a difficult species complex, identifications should be based on adults— see also below. J occurrence Woodlands and forests from southern Canada to North Carolina in mountains (but absent from many areas of coastal plain); southern limits elsewhere unclarified due to confusion with others in complex. Apparently one protracted generation over much of East with mature caterpillars from late July to October. Often common. J common foodplants Most of our records from white oak but several from red oak; also reported from beech, chestnut, and other oaks (but see below). J remarks We have adult vouchers that are unambiguously assignable to A. ovata for the larvae figured. No doubt, the diagnosis given above applies to other members of the oak dagger complex; this simply reinforces our plea for additional taxonomic study. The wild caterpillars that we have collected and reared to adulthood have come from white and red oak; all of those that John Lill and Tim McCabe have collected have come from white oak. Northward, the range of A. ovata roughly approximates that of white oak.

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Yellow-haired Dagger Acronicta impleta J recognition  Although variable in coloration, always recognizable by dense sets of pale, paired tufts over T3, A1, and A2 that envelope row of four taller, brown to black tufts across dorsum of A1. Broad, reddish spiracular stripe. A8 with pair of diverging brown to black setal lashes subtended by paler and shorter subdorsal setal tufts. Larva to 4.5 cm. Easily separated from all tussock moths (Lymantriinae), which it appears to mimic, by absence of unpaired middorsal glands on A6 and A7. J occurrence  Fields, woodlands, and forests. Transcontinental in Canada, south in East to Florida and Texas. Evidently at least two generations over much of range with mature caterpillars from June onward; three or more broods in Deep South. Common. J common foodplants  Hickory and walnut among favorites, but also alder, ash, birch, blueberry, cherry, dogwood, elm, hazel, maple, mountain ash, oak, rose, sassafras, sourwood, sweet gum, willow, witch hazel, and other woody plants. J remarks  The Yellow-haired Dagger provides a promising case for larval mimicry: its caterpillar looks remarkably like those of a tussock moth in the genus Dasychira, sharing the same overall shape, lashes, and tufts. It matches in detail; e.g., the setae that make up the shorter tufts are densely plumed and downy, like those of a tussock moth caterpillar. Behaviors too are copied; when alarmed, the Yellow-haired Dagger caterpillar will arch its first two abdominal segments upward, more fully displaying its tufts, in much the same fashion as true tussock moths. As further evidence of the resemblance, let it be known that two of the coauthors of this volume believed they had collected a tussock moth caterpillar at the time the larva was boxed up and sent to Connecticut to be photographed and reared. On the matter of mimicry, we have no doubts—our only issue is what kind of mimicry: Batesian or Mullerian? And if the latter, what is the basis of the Yellow-haired Dagger’s protection? See essay on page 78.

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Ochre Dagger Acronicta morula J recognition Last instar densely vested with brown, black, and white setae. Setae over dorsum short and mostly of one length; those along sides long, white, and of variable lengths. Black middorsal warts over A1, A4, and A8. Head black to either side of pale triangle with red over each lobe; lobes mostly pale with dark spots forming arcs (left inset). Larva to 4.5 cm. Penultimate instar (right inset) lime green with yellow-edged, purple-brown spots over A1, A4, and A8; head with reddish patches and darker snowflake spotting. J occurrence Fields, woodlands, and forest edges from Manitoba to Nova Scotia, south to northern Florida and Texas; increasingly local southward. At least two generations as far north as Quebec and likely with additional broods southward; mature caterpillars from early June through November. Locally common northward and in mountains. J common foodplants Elm; other reported foodplants, including apple, basswood, and hawthorn, need verification. J remarks This is yet another Acronicta with two stunningly distinct larval forms. In Connecticut, this moth is not common in many forested parts of the state, even where American elm is relatively frequent. In upstate New York, the Ochre Dagger does well in open habitats, e.g., alvars and along windbreaks and fencerows in agricultural landscapes. Young trees are especially likely to yield caterpillars. Southward, in the Appalachians, we associate it with mid-elevation forests as well as woodlands of the Atlantic Coastal Plain. The penultimate instar perches on the upper side of a leaf. The last instar passes the day perched on bark—it is best sought at night or sampled by beating.

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River Birch Dagger Acronicta betulae J recognition Last instar smoky green brown to reddish, purplish, or rusty brown, with long, whitish hairs and sprinkling of white spots along sides of body. Dorsal setae on T1 project well forward of head. Anterior dorsal seta (D1) arising from small gray spot; posterior dorsal seta (D2) enveloped by pale orange spot that may reach to D1. Another small, orange spot above each spiracle on abdomen (containing SD pinaculum). A1 and A8 scarcely enlarged, and more importantly, less so than in related birch-feeding species. Head with orange to reddish patch over each lobe and abundant black “snowflaking”; frons mostly pinkish brown and unmarked. Integument with abundant black spinules. Larva to 3.5 cm. Penultimate instar pale to blue-green with slightly humped A1 and orange-brown to reddish middorsal stripe (inset). J occurrence Wetlands, edges of watercourses, and floodplain forests from Wisconsin to New Hampshire, south to Florida and Texas (see Remarks). Two or three generations with mature caterpillars from June onward over much of range. Local and apparently uncommon. J common foodplants River birch (see Remarks). J remarks In our experience, this species is closely tied with river birch (Betula nigra). We wonder to what degree records from outside the range of this tree represent misidentifications of the Unmarked Dagger (A. innotata). As might be predicted by the caterpillar’s coloration, our green-form larvae (early to penultimate instars) rested on foliage, whereas the brown-form larvae (last instar) perched on twigs when such were provided. Adults come to both light and bait. Our prepupal larvae spun loose cocoons between leaves—they were not offered pulpy wood.

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Unmarked Dagger Acronicta innotata J recognition Chocolate to purple-brown or smoky brown; sides bearing numerous, grayish setae below level of spiracles. Lacking continuous orange middorsal stripe of related species (compare with Pleasant Dagger, A. laetifica, next page). Dorsal pinacula enlarged on A1 and A8; dorsal pinacula reduced on A7. Often with pale spot above and below each spiracle. Head with red patches over vertex, although these may be pulled into prothorax; below these patches each lobe with conspicuous dark spotting. Larva to 4 cm. Penultimate (left inset) and middle (right inset) instars pale yellow- to blue-green, with red dorsal spots edged with yellow over first eight abdominal segments; head pale yellow-green with diffuse red patches over lobes and above eyes. Closely allied to Pleasant and River Birch (A. betulae) Daggers. The latter occurs in association with river birch from areas largely to the south of those where the Unmarked Dagger flies. The ground color of its caterpillar usually has a decidedly pinkish or clay-colored cast, and smaller dorsal warts on A1 and A8. J occurrence Woodlands and forests from Manitoba to Newfoundland, south in mountains to Georgia. At least two generations with mature caterpillars from June onward. Common northward and in mountains. J common foodplants Birch; also reported from alder, aspen, hickory, mountain ash, pin cherry, poplar, serviceberry, and willow, but see below. J remarks The color and form changes that occur between the last two instars are extensive, and even the shape of the head is transformed. The middle instars, colored yellow, green, and red, reside on the upper sides of leaves, whereas the dark last instar rests on the bark of twigs and branches. We have had minimal success rearing larvae from eggs on hickory, a commonly listed host for the Unmarked Dagger (e.g., Smith and Dyar 1898, Crumb 1956). We have never found a larva feeding on hickory in the wild, and hickory is often absent from the forests where the Unmarked Dagger moth is common. In our experience, the species is more apt to turn up on members of the birch family (Betulaceae); we have found larvae on black or yellow birch on five occasions, and once each on paper birch, gray birch, and mountain ash (Rosaceae).

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Pleasant Dagger Acronicta laetifica J recognition Smoky to gray-black with subtle, broken, pale orange middorsal stripe running from T2–A8. Middorsal stripe edged with black at anterior end of each abdominal segment (from D1 seta forward). Sides of abdomen with sprinkling of pale spots. A1 and A8 slightly raised with enlarged black pinacula; on both segments, D setae flanked by diffuse pale patch that can be nearly white in some individuals. Setae above and below spiracle often arising from orange-yellow pinacula. Abundant pale setae along sides of body. Integument densely peppered with minute black spinules. Head with reddish patch over each lobe. Larva to 4 cm. Penultimate instar blue-green with orange-brown middorsal stripe that fades rearward; as in last instar, middorsal stripe with hint of black edging forward from D1, and pinacula distinctly warted over T1 and A8 (inset). J occurrence Woodlands and forests from southern Canada to Florida and Texas. Two generations over much of East with mature caterpillars from May through November. Locally common. J common foodplants American hornbeam and hophornbeam; Laura Miller sent us a caterpillar from hazel (Corylus americana) (though the caterpillar arrived dead, it appeared to represent this species). J remarks Forbes (1954) listed hickory as the host. A cohort we tried to rear on pignut hickory fared miserably, with all but two dying by the last instar. We have found wild larvae on American hornbeam on seven occasions and hophornbeam once. The prepupal caterpillar tunnels into pulpy wood to fashion its pupal cell.

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Nondescript Dagger Acronicta spinigera J recognition Last instar with peach to orange middorsal stripe frequently broken into spots (two per abdominal segment), running from T2–A7; spots weakening over thorax. Segments A1 and A8 smoky charcoal above, contrasting with adjacent segments. Upper half of abdominal segments with smattering of white spots, more numerous rearward of spiracle. A8 slightly humped. Dorsal pinacula black, shiny, pimplelike; those over A1 largest. Secondary setae conspicuous along side of body. Head with orange-red spots over each lobe and numerous black snowflake spots. Larva to 4.5 cm. Middle instars pale green with reddish dorsal patches over T1, A1, A4, A5, and A8. Caterpillar most similar to that of Pleasant Dagger (A. laetifica), which has a continuous orange dorsal stripe, lacks black suffusion over A1, and often bears a pale patch flanking the dorsal pinacula on A1 and A8. J occurrence Woodlands and forests from Manitoba to Maine, south to Florida and Texas. More common in Midwest (Covell 2005). Two generations in Appalachians with mature caterpillars in early summer, then again in fall. Locally common. J common foodplants McCabe (1990) reported elm. Although we have reared larvae to maturity on both elm and apple, basswood was preferred to both by our ex ova larvae. J remarks Young larvae spin down on silk if disturbed. Such belaying behavior is infrequent in middle instars. The last instar’s coloration and its flattened aspect are strongly suggestive that the caterpillar rests on the bark of its host by day—the only mystery is what the hosts are in the wild. It is somewhat remarkable that the species has eluded entomologists and foresters for so long, given its broad distribution throughout the East. In the southern Appalachians this can be a common dagger in mesic mid-elevation forests.

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Bantam Maple Dagger Acronicta retardata J recognition Instantly recognizable black, gray, and orange caterpillar with long white lateral setae. Dorsum crossed by narrow black and gray to lavender lines. Black line passing through D1 setal wart broken into two spots over dorsum of T2–A8. Short, spinelike black setae radiate from raised dorsal warts. Warts above spiracle red-orange, those below smaller, yellow. Sides of body pale, nearly without pigment. Head with conspicuous black spots; those over each lobe often fusing to form bars or lines. Larva to 3 cm. J occurrence Floodplains, swamps, and mesic woodlands from Manitoba to Nova Scotia, south to Florida and Texas. Two generations in Northeast; three or four generations in coastal North Carolina with mature caterpillars from May through October. Common but often overlooked as a caterpillar. J common foodplants Red, silver, sugar, sycamore maples; also boxelder. J remarks The boldly colored late instars perch conspicuously on the upper side of maple leaves, adjacent to their (diagnostic) feeding damage. The caterpillar positions itself out on a leaf lobe and then consumes much of the tissue to either side of the vein. When alarmed, the head is pulled around to one side. Caterpillars are occasionally mistaken for those of foresters, and especially members of the genus Alypia—the resemblance is sufficiently close to reason that at least one of these two moths (and perhaps both) is mimetic of the other. While widely known as the Retarded Dagger Moth, based on a literal translation of the specific epithet, we adopt an alternative that relates to both the insect’s diet and diminutive stature. We have had difficulty getting eggs from captive females held in vials—the figured caterpillar is the offspring of a female sleeved on red maple. Our captive caterpillars sometimes fashioned a gall-like cocoon along a twig.

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Delightful Dagger Acronicta vinnula J recognition Small, pudgy, lime to yellow-green with groups of short, white spiky setae issuing from low warts (above level of spiracles). D1 pinaculum on thoracic segments and D2 pinaculum on A1–A4 often raised and darkened. Dorsum of A8 with pair of closely set, purple-red (D2) warts. White subdorsal stripe extending from T2 to A10, subspiracular stripe running length of abdomen, and broken white middorsal stripe present on abdomen. Yellow connecting dorsal setal clusters along leading edge of prothorax. Upper portion of head with smoky reddish to red-brown patch. Setae along sides white, long, and soft. Larva to 2.5 cm. Chris Schmidt recently drew our attention to the fact that northern populations of what has been called Acronicta vinnula likely represent an undescribed species. Its larva differs from that of nominate A. vinnula in bearing more elevated (conical) warts on A1 and A4 (upper and lower insets). According to Schmidt it occurs from Wisconsin to Nova Scotia south at least into the mountains and Piedmont of North Carolina. J occurrence Fields, woodlands, and bottomland and floodplain forests along Atlantic Coastal Plain from New Jersey, south to Florida and Texas, north to Oklahoma (Chris Schmidt, unpubl. data). Multiple generations with mature caterpillars from May onward. Locally common. J common foodplants American elm (Smith and Dyar 1898); cedar elm (Ulmus crassifolia) in Texas. Winged and American elms widespread over much of its range along Atlantic Coastal Plain. Many literature records for American elm (e.g., Raizenne 1952 and Prentice 1962) refer to undescribed northern segregate. J remarks The head may be withdrawn into the prothorax when the caterpillar is alarmed. The prepupal larva turns yellow-brown to red (lower inset). Presumably the larva pupates in wood as do other daggers that take on such reddish colors prior to pupation. We have no explanation as to why prepupae turn red, or why this color change would be associated with wood tunneling in dagger moths.

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Greater Oak Dagger Acronicta lobeliae [Lobelia Dagger] J recognition Large, hairy, gray to smoky with variously developed pale middorsal and subdorsal stripes; long black dorsal, subdorsal, and lateral setae, and skirt of pale lateral setae. Dorsum of A1 and A8 slightly raised with darkened setal warts. Look for yellow-orange splotch over each spiracle. Body densely peppered with minute black spinules. Actively feeding last instar: gray, primary setal bases black, middorsal and subdorsal stripes vague, and head tan with cherry red dorsal patches and dark snowflake spots over each lobe (figured above). Prepupal (woodtunneling) last instar: charcoal with pale middorsal and subdorsal stripes, head coal black above and white below, and frons gray-black (upper inset). Larva to 5 cm. Middle instars blue with orange and black markings over dorsum (lower inset). J occurrence Woodlands and forests from southern Canada to Florida and Texas. Two generations over much of East with mature caterpillars from June through November; evidently three broods in Missouri. Common. J common foodplants Oak, both reds and whites. J remarks The ultimate instar is well suited for a life on bark—its coloration, corona of lateral hairs, and somewhat flattened aspect enable the larva to “disappear” on oak limbs and trunks. Superficially, the caterpillar resembles a bark-dwelling lasiocampid, e.g., tolypes and the Dot-lined White (Artace cribraria). The color changes that transpire within the last instar are exceptional among lepidopterans—the grayish ground color of the feeding larva gives way to charcoal, and the caterpillar becomes conspicuously striped. Equally remarkable are the changes in the head: the upper portion, white to tannish green with abundant dark snowflake spots and prominent cherry red lobes, becomes progressively black and unmarked in the prepupa. The two caterpillars are so dissimilar that Wagner (2005) initially suspected one of the forms might represent a second species. Prepupal caterpillars tunnel into wood.

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Impressive Dagger complex Acronicta impressa

and others

J recognition  Ground color usually black above and orange below level of spiracles. Spiracles bright white. Setal clusters over A2–A7 white or hoary (usually) to orange, with each made up of more than 20 minutely barbed setae that are bowed or crinkled. Setal cluster over T2, T3, and A1 usually orange or at least contrastingly darker than those from A2 rearward; warts subtending these setal clusters often orange-yellow. Subdorsal and lateral warts on T1, A8, and A9 with long setae that extend beyond ends of body. Head shiny black without markings. Larva to 4 cm. Middle to penultimate instars with orange dorsal tufts on thorax, A1, and A8. Caterpillar sometimes confused with that of Long-winged Dagger (A. longa). J occurrence  Edges of watercourses, fields and meadows, powerline right-of-ways, and forest edges; often open sites with low woody vegetation. Transcontinental; in East extending south to Florida and Texas. Two generations over much of East with mature larvae from June onwards. Common northward. J common foodplants  Alder, apple, beech, birch, blackberry (and other Rubus), blueberry, cherry, currant, dogwood, elm, gale, hazel, larch, oak, plum, poplar, rose, serviceberry, spirea, willow, and witch hazel; occasionally on nonwoody plants, e.g., dogbane. Many reports from willow and poplar. J remarks  It has long been suspected that “Acronicta impressa” might represent a cluster of closely related species. The caterpillars that we have sorted as the Impressive Dagger are highly variable in coloration and degree of hairiness—more so than any other acronictine in this guide, e.g., the caterpillar lower right from Mount Rainier (Washington) strikes us as being distinct. Recent mitochondrial DNA from across the insect’s range is also indicative that more than a single species is going under the name of A. impressa. Our larvae have come from lowgrowing plants, shrubs, and small trees. Impressive Dagger caterpillars are credible mimics of tiger moth caterpillars (Arctiinae). The resemblance is perhaps closest to the larva of the Great Tiger Moth (Arctia caja), a species known for both its beauty and chemical arsenal (Frazer 1965, Rothschild 1985). Prepupal larvae fashion their cocoons in leaves.

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Long-winged Dagger Acronicta longa J recognition Bristly with mostly charcoal ground color, attractively decorated with dark, shiny warts over T2–A1, and contrastingly paler warts rearward. Dorsum of abdomen sometimes with subtle frosty mottling. Sides of abdomen smoky orange. Thorax dark above and laterally. Triangle bounded with white (area between adfrontal sutures and frons). Larva to 4 cm. Middle instars (lower inset) spiky with numerous setae of various lengths. Each setal tuft with one to two dark spines and several finer white setae. Longest setae on T1 about half body length. Caterpillars of the Long-winged Dagger superficially resemble those of the Impressive Dagger (A. impressa) complex (previous page). J occurrence Dry woods, barrens, prairies, and dry fields from southern Canada, south to Florida and Texas. Two generations in Northeast south to Carolinas; flies nearly year-round in southern Florida (Kimball 1965). Locally common, especially southward. J common foodplants Alder, birch, blackberry, cherry, oaks, poplar, roses, willows, and no doubt many other woody plants. J remarks Middle instars preferentially rest on the undersides of leaves. We have occasionally found larvae on low-growing plants such as blackberry and sweet fern. Larvae release a pungent odor, reminiscent of slightly rotten eggs, when molested. The prepupa fashions a cocoon in leaf litter. Larvae that we have seen from the Gulf states are more uniformly orange and lack the dark integument, and may represent distinct taxon (page 259).

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Night-wandering Dagger Acronicta noctivaga J recognition Mostly charcoal ground color with orange subspiracular stripe along abdominal segments. D1 setal warts orange on A3–A7; dorsum of A7 with orange spot that includes D1 verruca. Thoracic segments decidedly blacker than those of abdomen. Setal clusters above spiracular line often with dark plumed (feathery) setae. Spiracles white. Head capsule black, shiny. Larva less than 4 cm. J occurrence Barrens, floodplains, woodlands, and forest edges. Manitoba to Nova Scotia, south to Florida and Texas. One principal generation with mature caterpillars from June through August, and small partial second generation. Common in boreal woodlands and forests, but increasingly local and scarce southward. J common foodplants Poplar commonly listed but also reported from various low plants by Smith and Dyar (1898). Tim McCabe found a wild larva on meadowsweet and has reared ex ova cohorts on cherry and meadowsweet. J remarks Larva in inset bears one white tachinid egg on A1 over the spiracle. Owlet caterpillars are known to chew through tachinid eggs that have been deposited on their bodies. This egg has been placed close enough to the head to be safe from such behavior. There have doubtless been strong selective forces on tachinids to position their eggs well forward on the bodies of their hosts, and equally strong selection of caterpillars to develop behaviors preventing this (such as animated thrashing). Adult Night-wandering Daggers are occasionally common at sugar baits. Like most other daggers, adults commonly come to light.

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Smeared Dagger Acronicta oblinita J recognition  Highly variable in color but usually some combination of black, yellow, red, and white, with black predominating. Consistently with yellow lateral blotches that may coalesce to form broad, interrupted spiracular stripe. Often with raised, reddish, bristly warts over dorsum of thoracic and abdominal segments, and field of white subdorsal spots. Spiracles white. Setae mostly of one length, frequently tinted with orange above. A few longer setae extend from either end of body. Larva to 4 cm. J occurrence  Beaches, marshes, wet meadows, swamps, and other open, usually moist to wet, habitats from southern Canada to Florida and Texas. Two or three generations with mature caterpillars from late May through October over much of East. Sometimes common. J common foodplants  Widely polyphagous with reports from many forbs (such as smartweed), shrubs, and small trees, including some conifers. J remarks  While the Smeared Dagger (or Smartweed Caterpillar) has been reported as a pest of apple and other orchard trees as well as a variety of field crops, it is hard to imagine any dagger being so common as to cause economic injury; more likely, its “pest” status is undeserved and its appearance in economic entomology literature derives more from its beauty and novelty than its monetary importance. The Smeared Dagger often occurs in floodplains and various riparian communities, wet meadows, and various coastal strand communities. Hugo Kons has found caterpillars feeding on emergent vegetation in ponds, lakes, and slow-moving waters in Florida. Unlike many Acronicta, the larva forms a simple cocoon in leaves or debris, perhaps reflecting the fact that the Smeared Dagger lives in early successional habitats, usually devoid of wood into which the prepupal caterpillar might excavate a pupal chamber. It is a beautiful creature—a favorite among caterpillar aficionados and nature photographers. Why the larva is so brightly colored has yet to be explained.

296 Daggers Subfamily Acronictinae

Hackberry Dagger Acronicta rubricoma [Ruddy Dagger] J recognition  Elongate middorsal tufts on A1–A5 and A8 distinguish this hairy, straw-colored dagger. Ground color of body waxy white to somewhat greenish. Middorsal hair pencils—longest on A1–A5 and A8—tan, black at their bases, or entirely black. Black addorsal stripe extends rearward from tuft over A5; in some forms these may fuse to form broad middorsal stripe. Head shiny black; basal segment of antenna and adfrontal sutures white. Spiracles tan to white with narrow black rim. Larva to 4 cm. Most similar to Cottonwood Dagger (A. lepusculina) which lacks a dorsal hair pencil over A2, is vested in setae that are brighter yellow, and as its name suggests, feeds on poplar and willow. J occurrence  Floodplains and other watercourses, woodlands, rocky outcrops, and other plant communities with hackberry. Kansas to southern Ontario, and southeastern New York to Florida and Texas. Two generations over much of East with mature caterpillars from June through October. Common southward. J common foodplants  Hackberry, also elm; report from sumac presumably in error. J remarks  This splendid caterpillar appears to be a mimic of a true tussock moth (Lymantriinae)—the high middorsal tufts are not unlike those of an Orgyia caterpillar. Middle instars of the Hackberry Dagger are especially close in appearance to larvae of the Definite Tussock Moth (Orgyia definita). The caterpillar rests on the underside of leaves with the head drawn against one side of the body. Although the caterpillar is reported to use elm, the range of the species more closely follows that of hackberry, i.e., the moth is absent from many forests and woodlands in the North where elm is common (although evidently in the South the species is occasionally found in forests where hackberry is scarce). Prepupal larvae develop a pale green cast to the integument. They will fashion pupal cells in soft wood or form a cocoon in leaf litter if no wood is provided. F. M. Jones (1928–1932) reported finding cocoons on hackberry bark in winter in Delaware. Adults are seldom encountered in appreciable numbers.

Daggers Subfamily Acronictinae 297

Narrow-winged Dagger Acronicta lanceolaria [Lanceolate Dagger]

J recognition Hairy, pale green with dark middorsal stripe. Middorsal stripe

poorly developed over thorax and often ending over A7. Smaller oblique black spots form variously developed spiracular stripe. Most body setae pale yellow, but with some dark-tipped bristles issuing from dorsal warts. Setal warts of thorax and A8–A10 with one very long, fine seta. Head mostly shiny green-yellow with black to either side of adfrontal suture; triangle also black. Basal segment of antenna bright yellow, contrasting with adjacent black areas of triangle. Spiracles tan with black outer ring. Larva to 3.5 cm. Penultimate instar more yellow in color and lacking black dorsal stripe and lateral spotting (inset). J occurrence Barrens, sand plains, dunes, sand prairies (northward), bogs, and other open, shrubby habitats from Manitoba to Maine, south to Florida and Missouri. One generation with mature caterpillars in June and July over much of range. Uncommon to rare as adult, but see below. J common foodplants Generalist, often on low-growing perennial plants; recorded hosts include alder, blackberry, blueberry, bush clover (Lespedeza) (Tim McCabe, unpubl. data), cherry, swamp doghobble (Eubotrys racemosa), hazel, larch, laurel, (scrub) oak, poplar, red pine, spirea, sweet fern, and willow. J remarks The Narrow-winged Dagger is one of many moths that occur in both bogs and barrens. One can only speculate about the reasons for this; both have acid soils, abundant heaths, and are sunny environs. Adults are rarely encountered at lights and almost never at bait. This is one of three Acronicta (the others being Doll’s, A. dolli, and Hazel Daggers, A. falcula) that we see more often as a larva than as an adult. Many of the records in the literature for the Narrow-winged Dagger, such as those in Farquhar (1934) and Brower (1974), refer to larval collections.

298 Daggers Subfamily Acronictinae

Miller Dagger

Acronicta vulpina (as A. leporina race vulpina in Forbes 1954) Transcontinental in Canada, south in East to Berkshires in Massachusetts, central New York, and Great Lakes states. One generation with mature caterpillars from mid-August to October. Sometimes common northward. Alder, birch, poplar, and willow.

Cottonwood Dagger Acronicta lepusculina

Transcontinental in Canada, south in East to northern Florida and Texas (panhandle). Evidently two generations over southern portion of range with mature caterpillars from July to October. Sometimes common northward. Cottonwood and other poplars; also alder, birch, and willow.

Square-eyed Dagger

Acronicta quadrata [Quadrate Dagger] British Columbia to Quebec; more widely distributed and common in West. A pine barrens species in Wisconsin (Ferge and Balogh 2000). At least partially double brooded over southern portion of Canadian range with mature caterpillars from July to October. Uncommon. Cherry and serviceberry; also willow and poplar.

Lesser Cottonwood Dagger Acronicta sperata [Hopeful Dagger]

Manitoba to Nova Scotia, south to Washington, DC and Missouri. Two generations over much of East with mature caterpillars from July to October. More common northward. Willow and poplar.

Daggers Subfamily Acronictinae 299

Doll’s Dagger Acronicta dolli (= Merolonche dolli) J recognition Attractive jet black, orange-red, and yellow caterpillar with numerous whitish setae. Red-orange subdorsal stripe broken between segments, extending from T2 to A7. Continuous pale yellow spiracular stripe ending on A8. Faint, white middorsal stripe weakly expressed. Some thoracic setal clusters bearing one very long seta whose length sometimes exceeds that of four body segments. Head shiny black. Larva to 4 cm. Early instars marked similarly but with shiny integument. Caterpillar of Doll’s Dagger superficially similar to those of Smeared Dagger (Acronicta oblinita), which is present later in the season, and more likely to occur on forbs. Doll’s Dagger is a species of woodlands and barrens, whereas the Smeared Dagger is a denizen of open marshes, floodplains, and other wetlands. J occurrence Barrens and acid woodlands from Minnesota to Pennsylvania, southeastern New York (Long Island), and New Jersey Pine Barrens south, mostly in mountains, to northern Georgia. One generation with mature caterpillars in late May and June. Rarely collected as adults. J common foodplants Ash, beach plum, blueberry, cranberry, oak, and other woody plants. Heaths and scrubby oaks commonly used in barrens. J remarks Until recently, Doll’s Dagger was classified in the genus Merolonche. Although its caterpillars resemble those of other Acronicta in structure, its larval ecology differs from that of most daggers: adults fly very early in the season, its larvae prefer new foliage, and the insect is strictly univoltine (most eastern Acronicta are bivoltine). (Among our 60 species of daggers, only the univoltine Narrow-winged Dagger (A. lanceolaria) has a similar biology.) We have found early instars perched conspicuously on the upper side of new oak leaves in the New Jersey Pine Barrens. Their shiny integument and red and yellow markings suggest that the caterpillar is unpalatable. Most of our adults collected at light, and several others that we know of, have been taken following unseasonably warm days. The pupa overwinters. For additional information on this elusive animal see Schweitzer et al. (2011).

300 Daggers Subfamily Acronictinae

Marsh Dagger Simyra insularis (= S. henrici) [Henry’s Marsh Moth]

J recognition Beautifully mottled in cream and black with bright

orange setal warts. Dorsum mostly black. Flank between creamy subdorsal and subspiracular stripes mostly yellow with black mottling. Orange setal warts above level of spiracles with mix of both pale and black setae. Spiracles bright white. Larva to 3.5 cm. J occurrence Wetlands and fields from Canada, south to Florida and Texas. Two generations in Connecticut, presumably more southward with mature caterpillars from June through November. Locally common. J common foodplants Broad generalist: mostly grasses, cattails, and sedges, but also forbs and low-growing woody plants. J remarks Until recently, this species was known as Henry’s Marsh Moth (Simyra henrici) or the Cattail Caterpillar. The larva is common in the grassy fields bordering wetlands. Fall semester entomology students in the Northeast routinely turn up individuals in their sweep samples. The two drably colored caterpillars in the upper inset are mummies—each contains more than a dozen Aleiodes stigmator wasp pupae. The adult braconid wasps issued shortly after this image was taken in mid-July. Forbes (1954) made note of the unusual cocooning habit of the Marsh Moth: “The cocoon is made in a distinctive way when a broad grass or sedge blade is available; the blade is sharply folded down near the middle, then after about an inch is sharply folded up again, enclosing a rounded-triangular area in which the fairly strong cocoon is spun.” As in other acronictines, the pupa overwinters. The phylogenetic origins of the genus within the subfamily could prove interesting— superficially the larvae resemble those of a number of our Acronicta, such as the Smeared Dagger (A. oblinita), Impressive Dagger (A. impressa), and others, many of which (also) frequently feed on low-growing vegetation.

Daggers Subfamily Acronictinae 301

Green Marvel Agriopodes fallax J recognition Immediately distinguishable by its teardrop shape. Ground color bright blue to yellow-green with thin, pale subdorsal stripe that extends from T2 to diminutive anal plate. Body densely set with minute rusty setae (spinules); setae longer and pale above prolegs and along caudal end of body. Pinacula reddish or smoky, bearing numerous short setae. Prothorax with small, reddish-brown subdorsal knob. Head roughened, often with dark spots above eyes (right inset), mostly drawn into thorax. Larva to 3 cm. J occurrence Shrub swamps, mesic woodlands, and forests from Manitoba to Nova Scotia, south to Florida and Texas. Two generations in Northeast with mature caterpillars in June and July, then again from August to October. Common. J common foodplants Viburnum; our larvae mostly from wild raisin or possumhaw (Viburnum nudum) and related viburnums; arrowwood viburnum likely in southern New Jersey. J remarks Evidently the distinctive larval shape is largely lost in preserved specimens as Crumb (1956) scarcely made mention of the insect’s body proportions in his description. The larva feeds from a leaf underside. We have found them in July both by turning leaves and beating. Caterpillars are susceptible to dwarfing, so keep them supplied with fresh leaves. Prepupae tunnel into soft wood, if available, to form their pupal chambers. The pupa overwinters. Coloration in our main image is subdued relative to the caterpillar shown in the left inset—perhaps it is approaching pupation.

302 Daggers Subfamily Acronictinae

The Hebrew Polygrammate hebraeicum J recognition Green with pale yellow subdorsal stripe, creamy addorsal spots at midsegment; middorsal spots along caudal end, and often along leading margin of T2–A8. Longest dorsal and subdorsal setae somewhat rusty in color and some longer than segments that bear them. Head smooth, shiny, pale, sometimes with small black spot above and to side of triangle. Larva under 3 cm. Late-stage last instar (a bark-tunneling form) is blue-gray and mottled with black spots (inset). Caterpillars of Cadbury’s Mystique (Comachara cadburyi), also a tupelo feeder, are similar (see below). Its dorsal and subdorsal setae are paler and shorter than the segments that bear them; the addorsal spots are missing. J occurrence Swamps and mesic woodlands from southern Ontario and Maine to Florida and Texas. Evidently two or three generations over much of East with mature caterpillars from June into October. Often locally abundant. J common foodplants Tupelo (Nyssa). J remarks The pale and conspicuously setose early instars “window” leaf blades by removing patches of green tissue from the lower leaf surface (leaving the upper leaf surface intact). The windows that remain are diagnostic. Third instars chew holes in the middle of the lamina, feeding from a leaf underside. Fourth and fifth instars feed inward from a leaf margin, leaving a telltale jagged edge. Caterpillars from the second instar onward move from leaf to leaf to feed, perhaps in an effort to remove themselves from their own feeding damage (and associated volatiles). Three color forms occur in the last instar. The green form described and figured above is the feeding stage; it rests on the lower surface of leaves. It is followed by a specialized transitional form whose charge is to excavate a pupal chamber in bark or rotten wood; its blue-gray coloration (inset) is not unlike that of tupelo bark. The prepupal larva loses much of this pattern and becomes increasingly pink (as is common among wood-boring acronictines); this second color transition usually occurs out of view, within the animal’s pupal chamber. The pupa overwinters.

Daggers Subfamily Acronictinae 303

Cadbury’s Mystique Comachara cadburyi J recognition Pale green with subdorsal line that weakens over thorax and rearward of A7. T2–A8 often with small, middorsal spot at rear of each segment. Wart above spiracle bearing several small setae (visible only with lens); this wart becoming progressively larger rearward. Longest setae shorter or about equal in length to segment that bears them. Head unmarked, shiny, pale tan to yellowgreen. Larva under 2 cm. Middle instar shown (lower right). Caterpillars of The Hebrew (Polygrammate hebraeicum) similar, but longest body setae longer than segment that bears them and addorsal spots present on T2–A8. J occurrence Swamps and mesic woodlands from Missouri to Massachusetts, south to Florida and Texas. One principal generation with mature caterpillars from June to mid-July in much of range; small partial second brood in Carolinas, Texas, and elsewhere in southern portion of range. Locally common. J common foodplants Tupelo (Nyssa). J remarks This moth has had a checkered taxonomic career, having been placed in Arctiinae, Nolidae, and Sarrothripinae (= Chloephorinae) over the past 70 years. Wagner et al. (2006) transferred Cadbury’s Mystique into the Acronictinae, after noting numerous similarities with the larva of The Hebrew (Polygrammate hebraeicum). Eggs of Comachara are laid on leaf undersides, usually tucked against the midrib or a lateral vein. Caterpillars feed on new leaves from the underside. Like Polygrammate, the first two instars “window” leaves by scraping away all the green tissue to the upper leaf surface. The caterpillars have a tenacious grip and need to be handled carefully when on toweling so that one doesn’t accidentally rip away a proleg. Covell’s (2005) common name, Cadbury’s Lichen Moth, was based on a belief that Comachara was a lithosiine tiger moth (and fed on algae and lichens). We offer Cadbury’s Mystique as a substitute, a name that links the Marvel Comics character who could effortlessly change her appearance to the remarkable color transitions that occur in the last instars of Comachara and Polygrammate. The bark-tunneling morph of this “transformer” is shown in the inset. Provide last instars with soft wood into which they can burrow and construct their pupal cell. The pupa overwinters.

304 Daggers Subfamily Acronictinae

Cherry Agate Cerma cerintha J recognition Elongate, green and red, with long dark setae. Cherry to maroon or brick-red dorsum edged by prominent white subdorsal stripe. Sides and subventer lime green or smoky green. Rusty to black setae arise from shiny setal bases. A8 with pair of prominent dark conical warts. Bright, shiny green head with black band extending down from vertex but ending before antenna. Subdorsal stripe broken over A8, then continuing across A9 and A10. Larva to 3 cm. Bright green early and middle instars lack red coloration over the dorsum, and the subdorsal stripe tends to include more yellow; in addition, each lobe of the head is drawn to a point and bears a single, black, bladelike seta (inset). Caterpillar of The Agate (C. cora) mostly black with broad yellow to orange and black spiracular stripe; lobes of head black (page 259). J occurrence Barrens, fields, sandy openings, and woodland edges from Minnesota to Maine, south to Florida and Texas. Two generations over much of East, but second generation increasingly partial northward. Mature caterpillars mostly in July and early August, then again in September. Common. J common foodplants Especially black cherry and hawthorn, but also from apple, cotoneaster, and plum. J remarks Cerma is placed here, next to Harris’s Three-spot (Harrisimemna trisignata) with which it shares many morphological and life history characters (Wagner 2007a, b). For example, caterpillars of Cerma and Harrisimemna (as well as those of Comachara and Polygrammate) tunnel into wood and then seal off their pupal chamber with a sheet of silk and wood chips. The finished closure is virtually undetectable. Upon disturbance, caterpillars of the Cherry Agate and Harris’s Three-spot may shake rapidly from side to side. Cerma caterpillars will wriggle violently if handled and are quick to hurl themselves from their purchase.

Daggers Subfamily Acronictinae 305

Harris’s Three-spot Harrisimemna trisignata J recognition Chimerical caterpillar with grotesquely humped eighth abdominal segment. Integument shiny, reddish brown to nearly black; A3 to A7 with white saddle that “spills” down sides to prolegs. Thoracic legs long with much orange. Often with earlier (shed) head capsules adhering to thoracic setae. Larva less than 3.5 cm. Early instar shown in inset. J occurrence Wetlands and mesic woodlands from southern Canada to Florida and Texas. Evidently at least two generations southward with mature caterpillars from July through October. Widespread but infrequent. J common foodplants Many records from hollies, including inkberry, winterberry, and yaupon (but not American holly); other hosts (all woody) include apple, ash, blueberry, buttonbush, ceanothus, cherry, honeysuckle, lilac, persimmon, seven son flower (Heptacodium miconioides), spirea, viburnum, wiegela, and willow. J remarks By almost any measure, this is an exceptional animal—the caterpillar resembles a bird dropping, a pile of debris, a moldy cadaver, a spider (see below), and who knows what else. While seemingly catholic in diet, Harris’s Three-spot shows a predilection for plants containing iridoid glycosides; these include all the hollies, ash, honeysuckle, lilac, virburnum, and wiegela. To what degree a diet rich in these compounds contributes to the insect’s fitness has yet to be studied. In a most curious fashion, larvae will “retain” the head capsules from previous molts. When alarmed the caterpillar thrashes the head from side to side. Might the retained head capsules function as a mace to knock away parasitoids and other small enemies from the body? In addition, if molested the caterpillar will rock its body with blurring rapidity—the effect is much like that of a spider rapidly tugging on its web (stabilimentum). Prepupal Harrisimemna caterpillars bore into wood and excavate elongate chambers where they will pass the winter as a pupa. The entrance, which is nearly indistinguishable from adjacent wood, is closed with a sheet of silk into which the larva weaves bits of wood that it has gnawed from the walls of the tunnel.

306 Foresters Subfamily Agaristinae

Foresters

Family Noctuidae: Subfamily Agaristinae Our seven eastern species represent four genera. The subfamily is considerably more diverse in the American Southwest, Mexico, and other arid environs. Adults are mostly diurnal, although our two wood-nymphs (Eudryas) are exclusively nocturnal. A number of male agaristines (but no eastern species) make a whistling sound while patrolling over their territories. Eastern North American agaristine caterpillars are boldly ringed in white, black, and orange. Head and body setae are

sometimes long, two SV setae are present on A7, and the spinneret is short and broad with a wide opening at its apex (Crumb 1956). The head is peppered with black spots, and A8 is humped. Five of our eastern species feed on members of the grape family (Vitaceae); the evening primrose (Onagraceae) and loosestrife (Lythraceae) families also host foresters. Eggs are laid singly, usually near new growth. Larvae feed exposed on shoot tips, from the underside of leaves, or in the case of Psychomorpha, from within a shelter. Prepupal larvae require pulpy wood in which to pupate. Winter (or in some cases, multiple winters) is passed as a pupa.

Langton’s Forester Alypia langtoni

Alaska to Newfoundland, northern New England, and Great Lakes states. One principal generation with mature caterpillars mostly in July and August, but evidently at least partial second generation in southern portion of range with larvae from June into September. Local and uncommon. Especially willowherb or fireweed (Epilobium); also evening primrose (Reginald Webster, pers. comm.).

Disparate Forester Androloma disparata

Texas, mostly southward and westward of San Antonio. Presumably multiple generations with mature caterpillars through growing season. Locally common. Peppervine (Ampelopsis arborea) and other Ampelopsis.

MacCulloch’s Forester Androloma maccullochii

Transcontinental in Canada. Generally more common and widespread westward. One principal generation with mature caterpillars mostly in July and August. Willowherb or fireweed (Epilobium).

Foresters Subfamily Agaristinae 307 Paddled Forester Xerociris wilsonii

Central Texas to Corpus Christi, west to Arizona and southward into Mexico. Multiple generations with mature caterpillars through growing season. Locally common. Ivy treebine or sorrelvine (Cissus trifoliata).

Grapevine Epimenis Psychomorpha epimenis ! recognition Body ringed with black and white. Orange prothoracic shield, A8 dorsum, anal plate, and prolegs. Ground color chalky white. T2 and T3 with thickened black rings broken over midline. Head shiny orange with black spots over each lobe. Base of midabdominal prolegs waxy orange with large black spot; distal portion brighter orange with small black spot above crochet. Larva to 2.5 cm. South of Georgia the Grapevine Epimenis is replaced by the Florida Psychomorpha (P. epimenis euryrhoda). We have not seen its larva. ! occurrence Hedgerows, woodlands, and forest edges from Iowa to Massachusetts, south to Florida and Texas. One generation with mature caterpillars in late spring. Common over much of range. ! common foodplants Grape. ! remarks The caterpillar forms a shelter from an entire, mostly fully expanded, new leaf (inset). The leaf edges are pulled upward or downward and tied together with silk. The larva feeds within the shelter, grazing almost to the outer leaf surface, leaving telltale “windows.” We do not know if the caterpillar feeds solely on the tissues within its shelter or also forages outside of it. On fox grape, look for the shelters five to eight leaves back from the shoot apex in late spring. The diurnal adults fly early with Azures (Celastrina species) and the Falcate Orange Tip (Anthocharis midea). In addition to being avid flower visitors, the Grapevine Epimenis is a puddler.

308 Foresters Subfamily Agaristinae

Eight-spotted Forester Alypia octomaculata ! recognition Long, white setae distinguish Alypia from other eastern agaristines. Thoracic shield orange and black. A1–A8 with orange bands; those over A1–A3 broadest. Splotches on A1–A8 below spiracles white to cream; that on A8 enlarged, white. Fleshy, basal portion of prolegs on A3–A6 orange with single black dot. Larva to 3 cm. Wittfeld’s Forester (A. wittfeldii) purportedly replaces the Eight-spotted Forester in parts of Florida. In the inset, we figure a larva reared on grape from Fernadina Beach in Nassau County (Florida) with an enlarged white patch on A7 and A8, but we confess to not knowing of any reliable character that can be used to recognize either the larva or adult of A. wittfeldii. Kimball’s (1965) record from Japanese persimmon (Diospyros kaki) seems exceptional. ! occurrence Woodland and forest edges from South Dakota to extreme southern Quebec and Maine, to northern Florida and Texas. Large generation in spring with mature caterpillars from April to July; partial second and third broods occur widely. Locally common. ! common foodplants Ampelopsis, grape, and Virginia creeper. ! remarks Look for the caterpillars feeding or perched on the underside of leaves near shoot apices, or feeding on tendrils and new stem tissues. On hairyleaved grapes, early instars will feed from upper leaf surfaces and graze patches free of all green tissues. Larvae are quick to vomit an orange, mostly clear fluid when disturbed. Young caterpillars drop from their perch on a belay line. Older larvae simply drop from their perch. When beating, gently position the sheet so as not to forewarn of your presence. Prepupal larvae may perish if not offered pulpy wood, dense peat balls, or other materials into which they can tunnel and fashion their pupal cells. Bryant (1982) had pupae from a single cohort of eggs emerge over a four-year time frame. Holland (1903) credits the house sparrow with helping to control these larvae, which were sometimes “a veritable plague” on ornamental Ampelopsis. The infestation in Harvard Yard noted by Forbes (1960) persisted until at least 1983; we have not seen such densities in natural settings.

Foresters Subfamily Agaristinae 309

Beautiful Wood-nymph Eudryas grata ! recognition Dramatically ringed in black, white, and orange with swollen rump. Bands on A8 and A9 salmon orange. Lacking long setae and prominent white lateral patch on A8 characteristic of Alypia species. Thoracic shield orange. Transverse orange bands extending down sides of T2–A9. Fleshy basal portion of proleg on A3–A6 orange with single black spot. Larva to 4 cm. Caterpillar of Pearly Wood-nymph (E. unio) differentiated on next page. ! occurrence Fields, woodlots, woodland, and forest edges from southern Canada to Florida and Texas. Two generations in Northeast, although second partial, with mature caterpillars from late July to October; two or three generations in Missouri. Often common. ! common foodplants Ampelopsis, grape, and Virginia creeper (all Vitaceae); reports from hops and buttonbush require confirmation. ! remarks The rear of the body forms a crude false head. Agaristines readily regurgitate when disturbed; presumably the false head gives the larva greater opportunity to deliver vomit to its enemies. Wood-nymph adults are birddropping mimics that perch, often conspicuously, on vegetation during the day. Their resting posture is unique: the densely hairy forelegs are held outstretched and the wings are wrapped around the body to make the moth appear somewhat cylindrical. Agaristines have brightly colored caterpillars, and all but the Grapevine Epimenis (Psychomorpha epimenis) feed and rest exposed, on or near new foliage; this suggests that they are chemically protected. The American Southwest has a marvelously rich and varied agaristine fauna—a collection of their larval images would make for a beautiful color publication. Prepupal larvae of the Beautiful Wood-nymph will bore into cedar siding, fence posts, and other wood structures (Richard Heitzman, in litt.).

310 Foresters Subfamily Agaristinae

Pearly Wood-nymph Eudryas unio ! recognition Similar to Beautiful Wood-nymph (E. grata), with white, black, and bright orange rings; white rings predominating in most individuals, black rings in others. Orange rings, and particularly those on A8 and A9, without salmon cast of Beautiful Wood-nymph. Prothoracic shield white and black, lacking appreciable orange pigmentation (although Crumb (1956) indicates that shield may be orange). A8 not as strongly humped as that of Beautiful Wood-nymph. Triangle on face with lower black spots often fusing across midline. Fleshy orange basal portion of prolegs with two offset black spots. Larva to 3.5 cm. In addition to characters given above, black spots on frons usually separated in the Beautiful Wood-nymph. ! occurrence Wetlands, shorelines of lakes and other water bodies, and woodland edges from Manitoba to Maine, south to Florida and Texas. Two generations in Connecticut with mature caterpillars in July, followed by partial second fall brood; two or three generations in Missouri and New Jersey with last instars into October. Present year-round in southern Florida. Locally common. ! common foodplants False loosestrife or primrose-willow (Ludwigia), purple loosestrife (Lythrum), and swamp loosestrife (Decodon) commonly; also reported from evening primrose, gaura, hibiscus, grape, and willowherb (Epilobium). ! remarks Clarification of the brood number is needed. DLW’s larval collections from July and then again in September and October are suggestive of two generations in Connecticut. The most reliable means to locate larvae in the Northeast is to search swamp loosestrife or false loosestrife (primrose-willow) in wetlands from August into October. The caterpillars can sometimes be found on plants growing in standing water—presumably the larvae are able to swim to pupation sites. Rarely, the caterpillar may be common enough to defoliate plants of the pernicious purple loosestrife, a nonnative species. We have only one record of a wild larva on grape; in our experience, Eudryas larvae from grape nearly always prove to be those of the Beautiful Wood-nymph (E. grata). Unlike most members of the subfamily, Eudryas adults are nocturnal and come to light.

Paints or Hooded Owlets Subfamily Cuculliinae 311

Paints or Hooded Owlets Family Noctuidae: Subfamily Cuculliinae

This subfamily recently has undergone taxonomic reorganization—most of its previous members have been shifted into the Noctuinae or Oncocnemidinae. The subfamily’s evolutionary relationships to other lower trifid lineages are unresolved. While many authors have placed the subfamily near the Oncocnemidinae, Mitchell et al. (2006) found little evidence for a close evolutionary association of the two subfamilies. There are nine eastern species, all in the nominate genus. We associate Cucullia with grasslands, meadows, fields, and other open habitats. Paints include some our most spectacularly colored caterpillars, many appearing as if they had been individually painted and glazed. Although adult coloration might be described as mundane and undistinguished, the larvae of paints are considerably more varied in appearance—all nine of the British Cucullia are easily identified to species as caterpillars (see Porter 1997). Eastern North American species are mostly distinct, although members of the C. asteroides group (i.e., C. asteroides, C. postera, C. florea, and C. omissa) are close in appearance and difficult to distinguish. A few are highly polymorphic with green, brown, pink, and lavender forms. The ecological and evolutionary reasons for their outstanding coloration are unstudied, and while one could naturally conclude that the bright orange and black species are distasteful, such colors are also somewhat cryptic when the caterpillars are at rest within the flowers of their host.

The bright coloration and smooth, often shiny integument are diagnostic. The spiracular stripe extends around the anal plate, as it does in the Amphipyrinae and Oncocnemidinae. (It does not bend down and run along the outer face of the anal proleg as in the Condicinae, Heliothinae, and Noctuinae). Setae are short and inconspicuous; SD1 on A9 is unmodified. The crochets are of two lengths and the labial palpus is short. Adults come to light, but some species only in modest numbers; they can be found much more reliably visiting flowers after nightfall—milkweeds are a favorite nectar source. Sweet pepperbush receives many visits on the Atlantic Coastal Plain. Of note is the long proboscis that allows the adults to nectar while hovering in the same fashion as hawkmoths (Sphingidae). Spring generation caterpillars feed on leaf tissue, but late summer and fall broods show a preference for flowers. Females oviposit in captivity if offered flowers of a suitable host. Cucullia overwinter in a tough cocoon fashioned belowground—and at least some European species can overwinter for two or more years. A few paints appear to require specific soil conditions: e.g., we associate the Halloween (C. alfarata) and Speyer’s (C. speyeri) Paints with dry, sandy environs. In Great Britain, cucullias are known as wormwoods and sharks. We offer the common name paints in recognition (and celebration) of their marvelous coloration as larvae. Much additional information on cuculliines, including larval descriptions, can be found in Poole (1995).

Preliminary Key to Larvae of Eastern Members of the Cucullia asteroides Group Jean-Claude Petit and David L. Wagner 1a Spiracles pink, salmon, or red; montane and northern states into Canada . . . . . . . . . . . . . . . . . . . . . . . . Cucullia postera 1b Spiracles white, off-white, cream; variously distributed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2a Middorsal stripe yellow, white and yellow, or entirely white, narrower than that of C. asteroides; spiracles whitish or cream; montane and northern states into Canada . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cucullia florea 2b Middorsal stripe yellow, wider than that of C. florea; variously distributed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3a Thin, black, interrupted line running along top of white spiracles; addorsal stripes without prominent black edging, or black edging broken into series of small spots; widely distributed in East. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cucullia asteroides (first generation) 3b Continuous or almost continuous black line running through white spiracles, broadened to encircle or entirely include each spiracle; addorsal stripe more consistently and prominently edged with black; widely distributed in East . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cucullia asteroides (second generation) 3c Thin, interrupted, black line running through top of spiracles, not broadened (only visible at level of spiracles); addorsal stripes without prominent black edging, or black edging broken into series of small spots; spiracles whitish to cream or tan; montane, northern tier of states, and Canada . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cucullia omissa

312 Paints or Hooded Owlets Subfamily Cuculliinae

Overlooked Paint Cucullia omissa

Transcontinental across lower Canada, south in East through mountainous areas of upstate New York and New England. One generation with mature caterpillars mostly in September. Common northward. Aster, goldenrod, and probably other composites. Broad, lemon middorsal stripe margined with white; addorsal stripes often mottled with black; spiracles white or cream.

Unrusted Paint Cucullia florea

Transcontinental across Canada, south in mountains to North Carolina. Presumably two generations with mature caterpillars mostly in September. More common northward. Aster, goldenrod, horseweed (Conyza canadensis), and likely other composites. Middorsal stripe narrower than that of C. asteroides and C. omissa. Highly variable in coloration.

Rusted Paint Cucullia postera

Transcontinental across Canada, south in East to Massachusetts and Great Lakes states. Presumably two generations with mature caterpillars mostly in September. More common northward. Handfield (1999) gives aster, fleabane (Erigeron), and goldenrod. Spiracles salmon to red; ground color sometimes pink.

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Halloween Paint Cucullia alfarata [Camphorweed Cucullia] ! recognition Boldly marked in orange, black, white, and yellow. Continuous

middorsal stripe yellow, orange, or orange-red. Broad, jet black, subdorsal stripe with spurs “dropping” down to level of spiracle. Thick subspiracular stripe may be orange or yellow. Subventer and venter white. Head mostly black with white cheek patch, clypeus, antenna, and inverted “V” framing triangle. Larva to 4 cm. C. laetifica, which occurs from central Texas westward, is superficially similar, but the black subdorsal stripe is broken and the head is mostly white with black spotting (lower right). Its coloration is highly variable: in some individuals orange is replaced by yellow. Our examples were collected from camphorweed (Heterotheca subaxillaris). Valerie Bugh has reared the species from prairie broomweed (Xanthocephalum dracunculoides), tahoka daisy or tanseyleaf tansyaster (Machaeranthera tanacetifolia), and prairie or plains fleabane (Erigeron modestus). ! occurrence Dry fields, waste areas, pinelands, and other open habitats, with sandy or otherwise coarse soils; from Missouri to Long Island, south to Florida and Arkansas, although apparently absent from many states within this range. One principal brood, with most reports of mature caterpillars in September (New Jersey) and October, although Poole (1995) makes mention of a summer generation. Locally common (as larva), especially along Atlantic Coastal Plain. ! common foodplants Flowers of composites, especially camphorweed and Haplopappus; also some asters, horseweed (Conyza canadensis), slender scratchdaisy (Croptilon divaricatum), narrowleaf silkgrass (Heterotheca graminifolia), and presumably others. ! remarks While C. alfarata has been called the Camphorweed Cucullia, we thought the Halloween Paint a better name, given its coloration and diet. The former name is problematic given the host breadth of C. alfarata and our recent discovery of other Cucullia on camphorweed from Texas westward. And, at least in Georgia, caterpillars of C. alfarata are commonly reported within a week or two of Halloween. While larvae may be locally common, the number of adults in collections is modest. DFS has yet to see an adult in the wild, although he has encountered dozens of caterpillars in southern New Jersey. Most of these bore tachinid eggs. A diurnal ichneumonid wasp also parasitized many of the larvae in New Jersey. Our larvae ignored leaves and fed only on flowers.

314 Paints or Hooded Owlets Subfamily Cuculliinae

Asteroid Paint Cucullia asteroides [Goldenrod Hooded Owlet] ! recognition Usually bright green or brown with yellow, black, and white striping, but exceedingly variable (see Remarks). Integument shiny. Middorsal stripe yellow, often narrowly edged with white, and flanked by variously developed, thin, black stripes. Larva to 4.5 cm. A preliminary key is provided on page 311 that will separate caterpillars of the Asteroid Paint from other members of the C. asteroides species group. ! occurrence Fields, meadows, and woodland and forest edges from Manitoba to Nova Scotia, south to Florida and Arkansas. Two generations in Connecticut with mature caterpillars in July and then again from late August into October. Common at least as far south as Connecticut. ! common foodplants Aster and goldenrod. ! remarks The ground color of the caterpillar varies dramatically, ranging from green to tan and purple-brown. Wild-caught, last-instar green forms sometimes transform into lavender-brown caterpillars over the course of a few days. The first generation is content to eat young foliage, but larvae of the second brood show a decided preference for flowers. Heavy pollen loads on the tongues of adults are a testament to their penchant for flowers (Poole 1995). The genus is in need of life history study and offers much opportunity for someone who has a knack for rearing. The western fauna, which includes many magnificently colored caterpillars, is quite poorly known.

Paints or Hooded Owlets Subfamily Cuculliinae 315

Calico Paint Cucullia convexipennis [Brown-hooded Owlet] ! recognition Stunningly rendered in yellow, red, black, and white. Integument shiny. Middorsal stripe orange-red; subspiracular stripe bright red. Bright yellow lateral patches edged above and below with white, and cut by one thick black line (just behind spiracle). A8 with black hump. Spiracles white. Larva to 4.5 cm. Middle instars yellow, black, and white, without lateral red markings. ! occurrence Fields, meadows, and woodland edges from Saskatchewan to Nova Scotia to Carolinas (mountains) and Missouri. Two generations in Connecticut with mature caterpillars in July and then again from late August into October. Sometimes common. ! common foodplants Our records from aster and goldenrod. ! remarks This is one of our most beautiful and asked about caterpillars. Larvae are conspicuous by day, usually resting on stems in full view. First generation larvae feed on leaves; second generation larvae prefer flowers. Calico Paint caterpillars are seldom common enough to warrant stem by stem searches, but they are certainly numerous enough over most of their range that beating could be expected to yield a larva or two. Cucullia caterpillars are frequently attacked by tachinid flies that lay conspicuous white eggs on the body (see next page). According to Porter (1997), it is important to leave the rather tough cocoons of Cucullia intact to ensure successful eclosion. To see adults, keep an eye on patches of wild bergamot, milkweed, and other flowers in the early evening (Eric Quinter, pers. comm.)—often they are joined by loopers (plusiines) and hawkmoths.

316 Paints or Hooded Owlets Subfamily Cuculliinae

Dashed Paint Cucullia intermedia ! recognition Stout, nearly black caterpillar with prominent yellow to orange lateral spots that get smaller rearward. Integument granular, glossy. Dorsum unmarked, or with middorsal orange spots most evident over thoracic and last two abdominal segments. Lateral spots variously developed: some individuals with spots running length of body; others with spots ending towards front of abdomen. Head black and unmarked. Larva to 4.5 cm. Penultimate instar with broad white and lemon-yellow middorsal and spiracular stripes (lower right). ! occurrence Fields, meadows, roadsides, woodland edges, and other open habitats from Manitoba to Newfoundland, south to Virginia and Great Lakes states. Two generations in southern portion of range with mature caterpillars from late June into October. Common northward. ! common foodplants Flowers of composites. ! remarks Although the larvae readily devour flowers, they can be reared on leaves. The penultimate instar (right) was fed leaves of wild lettuce. The last instar shown above was found crossing a road in mid-August in upstate New York. Like many of the Cucullia that we have found, the larva had been attacked by a tachinid fly—five eggs are visible above the spiracles, all laid in intersegmental folds of the caterpillar’s body. A sixth egg, behind the proleg on A3, is disfigured. Perhaps this egg was crushed by the larva—some caterpillars will chew into and destroy tachinid eggs that they can reach. When disturbed, Dashed Paint caterpillars arch their head backwards over the abdomen and splay their legs. Both their coloration and behavior suggest that the larvae are chemically protected, but we are not aware of studies documenting that members of the genus are distasteful.

Paints or Hooded Owlets Subfamily Cuculliinae 317

Speyer’s Paint Cucullia speyeri ! recognition Frosty white with bold black and yellow patterning (see also Remarks). Yellow middorsal stripe often broken at midsegment and between segments. Subspiracular stripe running length of body. Black spots on thorax ringlike, continuing over dorsum. Head black and white; white framing black frons, white patch extending from first two stemmata (eyes) up to vertex. Larva to 4.5 cm. ! occurrence Sand plains, seaside meadows, pine barrens, oak savannas, waste lots, and other open, xeric, sandy habitats from British Columbia, southern Quebec, and Maine; south to North Carolina (mountains) and Missouri. Widespread in West. Evidently two generations in Ohio and southern New Jersey with mature caterpillars in midsummer, then again in fall. Collection dates for Speyer’s Paint in Missouri suggest that the moth has two or three generations there (Richard Heitzman, pers. comm.). Local and generally rare in East. ! common foodplants Horseweed (Conyza canadensis); also hairy white oldfield aster (Symphyotrichum pilosum), fleabanes (Erigeron), some goldenrods, and undoubtedly additional related composites. ! remarks Forbes (1954) mentions that caterpillars of Speyer’s Paint may also be marked with red, but we wonder if his remark stems from notes he had for one of the nearly indistinguishable western members of the speyeri complex. DFS found first-brood larvae feeding on new leaves of slender fragrant goldenrod in southern New Jersey; his second-brood collections have come from flowers of horseweed. We or our colleagues have found caterpillars in six states and Ontario, sometimes in good numbers, but we have taken adults at light only twice. We suspect that adults are only weakly attracted to light, and are more common than our collection records indicate. To various degrees, such is likely to be true for other members of the genus. We associate Speyer’s Paint with dry, sandy, open habitats, e.g., coastal strand meadows, openings in pine woodlands, as well as disturbed sites with an abundance of horseweed.

318 Sallows, Brocades, and Kin Subfamily Oncocnemidinae

Sallows, Brocades, and Kin Family Noctuidae: Subfamily Oncocnemidinae

Until recently, most of the genera in this subfamily were classified in the Cuculliinae. In Mitchell et al.’s (2006) molecular study of owlets, members of this subfamily grouped with the Agaristinae. The subfamily is represented by about 200 species north of Mexico, with the vast majority of these being western—less than a dozen species occur east of the Great Lakes region and only a few of these are common enough to be treated by Covell (2005). A single genus, Sympistis (Troubridge 2008), with 176 described species, accounts for much of the subfamily’s New World diversity.

The spiracular stripe extends around the anal plate, crochets are of one (most genera) or two lengths, the spinneret is elongate, and the apical seta (of the spinneret) is long relative to that of Cuculliines (Fibiger and Lafontaine 2005). Stemmata 3 and 4 are closely situated, at least in some genera. Oncocnemidines tend to be foodplant specialists on low-growing plants and shrubs. Some of our eastern species are specialists on spring growth; others feed preferentially on flowers or fruits. Such being the case, most are single brooded with seasonal activity timed to the availability of appropriate tissues. Host families containing iridoid glycosides, e.g., Caprifoliaceae, Oleaceae and Scrophulariaceae, are well represented in the diets of the subfamily. Sympistis diapause as eggs or pupae.

Lantana Stick Caterpillar Neogalea sunia [Catabena Moth]

Florida west to California, south to Argentina. Multiple generations with mature caterpillars yearround. Common. Lantana. Introduced as a biological control agent to control lantana in Hawaii, Australia, and elsewhere.

Blueberry Sallow

Sympistis dentata (= Apharetra dentata) Canada to New Jersey, Pennsylvania, and Great Lakes states. One generation with mature caterpillars in June and July. Adults locally abundant northward. Lowbush blueberries (Vaccinium angustifolium and V. pallidum), bog laurel (Kalmia polifolia) (McCabe 1990), and probably other heaths.

Brown-lined Sallow

Sympistis badistriga (=Homohadena badistriga) Southern Canada to Georgia and Texas. One generation with mature caterpillars in May and June. Local and uncommon; now rare or absent across much of Northeast. Honeysuckle, such as Lonicera dioica (but evidently not on many of the common, exotic species).

Sallows, Brocades, and Kin Subfamily Oncocnemidinae 319

Fringe-tree Sallow Sympistis chionanthi (= Adita chionanthi) ! recognition Brown and black with conspicuous lateral striping. Black subdorsal stripe interrupted on T2 and T3, ending on A7 (occasionally absent), thickened over anterior abdominal segments. Middorsal stripe represented by white spots over T2, T3, and line rearward of A7. A1 blackened above, darker than ground color of adjacent segments. Brownish spiracular stripe sandwiched between two white-brown stripes, all three of which continue forward onto head. Head with field of dark pigmentation above (Crumb 1956). Spiracles caramel brown. Larva less than 3.5 cm. An undescribed Sympistis occurs on feverwort (Triosteum perfoliatum) from Iowa to New Jersey (Rummel 1921, Forbes 1954). Its pale, waxy, green caterpillar becomes increasingly infused with red over the dorsum as it matures (inset). Our caterpillar was sent by Mary Jane Hatfield who discovered the moth in Iowa. We know of few recent collections of the Triosteum feeder and suspect the species may be in decline in the eastern part of its range, partly due to overbrowsing of its foodplant by white-tailed deer. ! occurrence Moist bottomlands and other mesic to wet forests from South Dakota to Nova Scotia to Virginia and Kansas, but absent from many areas across this range (see Remarks). One generation with mature caterpillars in late spring. Uncommon. ! common foodplants Ash and fringetree, both in the olive family (Oleaceae). ! remarks The caterpillars favor new leaves. The brown, barklike coloration of the larva suggests that the late instar rests on bark or in leaf litter at the base of the host. We have never found a wild larva, either on or off the host. The longterm survival of this moth and other eastern Fraxinus feeders are threatened by the spread of the Emerald Ash Borer (Agrilus planipennis) (Wagner 2007c). The egg overwinters and the pupa aestivates.

320 Sallows, Brocades, and Kin Subfamily Oncocnemidinae

Saunders’s Sallow Sympistis saundersiana (= Oncocnemis saundersiana) ! recognition Green to pink and perhaps other colors, matching foodplant.

Smooth, elongate, broadest though middle (A1–A6). Ill-defined dorsal and subdorsal stripes. Two supraspiracular stripes, uppermost broken, poorly developed, or absent. Area between supraspiracular and spiracular stripes sometimes filled with white. Prominent white spiracular stripe with upper edge ill-defined; lower edge more sharply defined. Setae short, only twice height of spiracle, inconspicuous. Spiracles tan with black rim. Third stemma (close to antennal base) enlarged. Larva less than 3.5 cm. Prepupal larvae turn pale green prior to pupation. A boreal congener, S. piffardi, feeds on white meadowsweet (Spiraea alba); its caterpillar matures in the spring (McCabe 1985, 1990). The life history of S. riparia, a locally common dune species along the Atlantic coast and Great Lakes, has not been elucidated. ! occurrence Roadsides, rocky outcrops, wet meadows, and other open habitats from British Columbia to southern Ontario, south in East to New Jersey and Texas; however, evidently historic from much of Northeast. One generation with mature caterpillars in early summer. Local and uncommon. ! common foodplants Beardtongue or penstemon, and turtlehead (latter record attributable to Deane Bowers). ! remarks The ground coloration of the caterpillar often is a fair match to the flowers on which its feeds. The resemblance is so close as to suggest the larva is able to transport floral pigments from its gut to the extremity of its body, although we know of no physiological mechanism that would allow this to occur. The larva tunnels underground to pupate and then aestivates through the summer. Adults fly in late August and September, and presumably oviposit on largely senescent plants—the egg overwinters. Saunders’s Sallow has not been seen in Connecticut or New Jersey in decades; presumably it was a victim of reforestation and overbrowsing by deer.

Sallows, Brocades, and Kin Subfamily Oncocnemidinae 321

Scribbled Sallow Sympistis perscripta (= Lepipolys perscripta) ! recognition Bold combination of black, yellow, and white. Mostly black above with irregular, yellow middorsal spots margined by white. Broad white and yellow lateral stripe includes blackened pinacula. Prothoracic shield white over dorsum with four prominent, black spots; lower edges of shield blackened. Head porcelain gray-white with peppering of black spots. Larva to 3 cm. The Toadflax Brocade (Calophasia lunula) has numerous blue-gray lines that connect the yellow middorsal and subdorsal stripes (next page). ! occurrence Sand plains, dunes, rock outcrops, waste lots, and other open, xeric habitats from southern Maine to Florida, west through Minnesota, Colorado, and Arizona to California (but absent from many intervening states, and this range likely conflating other as yet unrecognized, cryptic species). One generation in Connecticut with mature caterpillars mostly in late June. In North Carolina, mature caterpillars occur in April and May. Generally scarce but sometimes locally common (as larva). ! common foodplants Toadflax, especially Canada (or old-field) toadflax (Linaria canadensis); one report from Antirrhinum (Kimball 1965). ! remarks A fugitive species of erratic occurrence, sometimes present in small foodplant patches. The larva, which feeds on buds, flowers, and seeds, is a strikingly handsome insect that perches on the stem of its host. Given the small stature of the foodplant, the caterpillars, especially in the last instar, are conspicuous and easily located. By day, look for them near the base of the host, facing downward; however, others remain high on the stems where their pattern is somewhat cryptic among the developing seeds. Sweeping can be an effective way of securing caterpillars. Westward there are closely similar Sympistis species, misidentified in collections as the Toadflax Brocade. The caterpillar of an ArizonaColorado-Texas sibling species is strikingly distinct in appearance (see bugguide. net), and ranks among our continent’s most beautiful owlet caterpillars. As might be surmised from their coloration, the caterpillars are unpalatable: larvae sequester iridoid glycosides (Boros et al. 1991). Adults occur from April to early June in New Jersey, which might be two broods, with last instars present from late May into early July. Forbes (1954) mentions an August adult from Long Island and there is a single New Jersey specimen from August in the collection at Rutgers University; DFS had one adult emerge in October. Over much of East the larval food is past by midsummer. The pupa overwinters in a substantial cocoon underground.

322 Sallows, Brocades, and Kin Subfamily Oncocnemidinae

Toadflax Brocade Calophasia lunula ! recognition Black, yellow, and waxy blue-gray, with bright lemon-yellow middorsal, subdorsal, and spiracular yellow stripes. Black ground color above subdorsal stripe cut by one to four irregular blue-gray lines; blue-gray ground color above and below spiracular stripe spotted with large and small black spots. One prominent black spot near distal reach of each proleg. Larva to 3 cm. See also Scribbled Sallow (Sympistis perscripta). ! occurrence Fields, roadsides, waste lots, and other open habitats. Eastern Canada south through New England and west to at least Colorado. Two generations in Connecticut with mature caterpillars in July, then again in September and October. Very common. ! common foodplants Butter-and-eggs (Linaria vulgaris), Dalmatian toadflax (L. dalmatica), yellow, and other toadflaxes. ! remarks The Toadflax Brocade or Toadflax Defoliator was introduced from Europe into Ontario in 1968 as a biocontrol agent for Dalmatian toadflax, a naturalized plant that is invasive across large tracts in Canada, the Rockies, and elsewhere. The moth jumped the border and has since spread southward to Ohio and the whole of New England where it feeds principally on another introduced Linaria, butter and eggs. One wonders what the insect’s impact will be on native Linaria. Larvae move up into the flowers to feed, but are more likely to be seen at rest, perched along stems. Deane Bowers has found that the caterpillars sequester antirrhinoside and other iridoid glycosides (unpubl. data), presumably in the same fashion as the Scribbled Sallow (Sympistis perscripta). Up to 15% of the caterpillar’s dry weight may be given over to the iridoid glycosides, bitter compounds that are potentially toxic. The plebian adult, as might be guessed from its coloration, is all but devoid of the toxins. Prepupal caterpillars incorporate soil, litter, chewed leaves, and even cardboard, if such is made available, into their tough-walled cocoons. The pupa overwinters.

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Five-lined Sallow Catabena lineolata ! recognition Elongate, lime-green caterpillar with numerous fine, creamy, wavy stripes; proleg on A3 quarter-sized and that on A4 half-sized. Creamy lateral stripe along lower edge of tan-orange spiracles drops below spiracle on A8, and runs down outer face of anal proleg. Four to five pairs of fine, pale, wavy stripes above spiracles. Anal prolegs elongate, held rearward, often wine colored. Head reddish brown, prominently marked with four white lines above eyes and two along “cheek.” Larva to 2.5 cm. ! occurrence In Northeast, mostly fens, floodplains, and other mesic to wet habitats, but also quarries, prairies, and sandy, open environments. Transcontinental; eastern range extending from North Dakota, southern Ontario, Quebec, and Maine, south to Georgia and Texas. Evidently two generations in Connecticut with mature caterpillars mostly from June into October. Local and uncommon eastward. ! common foodplants Vervain (Verbena); report in Forbes (1954) from goldenrod presumably erroneous. ! remarks Sam Jaffe (in litt.) notes that the caterpillars “have a wonderful dismounting behavior—best brought on by blowing on them—they rear up, curl their heads around and back over their abdomen, forming a loose wheellike bundle or knot, and spring off the plant, bouncing and rolling their way off and away.” In Connecticut, the moth is local in wetlands. The species also occurs widely in drier environs, especially through the Great Plains and in the West, where it utilizes different species of verbena. The reduced prolegs on A3 and A4, and spiracular stripe configuration (not continuing around the anal plate as in other Oncocnemidinae and Cuculliinae) of the Five-lined Sallow are both exceptional for an oncocnemidine. Caterpillars are conspicuous in habit: look for them sitting on upper leaf surfaces or perched near the tips of a given shoot. The pupa overwinters in a slight cocoon on the ground (Forbes 1954).

324 Amphipyrines Subfamily Amphipyrinae

Amphipyrines

Family Noctuidae: Subfamily Amphipyrinae Long have the Amphipyrinae been a repository for waif genera and tribes—its membership ever-changing, waxing and waning with each taxonomic treatment. In 1998, Kitching and Rawlins removed all but the nominate genus (Amphipyra) and a few (mostly Old World) satellite genera, a decision supported by the molecular data of Mitchell et al. (2006). Based on a study of larval morphology, Wagner et al. (2008) argued that the concept of the Amphipyrinae should be expanded to include the lash-eyed sallows (psaphidines) as a tribe. We also add the Phosphilini here because of the many shared larval similarities with the larvae of Cropia and other amphipyrine genera (see page 328). Lafontaine and Schmidt (2010) included the seedcoppers (Stiriinae) as an amphipyrine tribe, a decision that we are reluctant to embrace given the radically specialized larvae and life histories of stiriines. Below, we introduce two amphipyrine tribes separately: Phosphilini and Psaphadini, as well as two subtribes within the latter (Feraliina and Psaphidina). Amphipyrine larvae sometimes have A8 humped and greatly so in some species. The spiracular stripe continues around the anal plate and crochets are of one length. Many

species have small warts or excrescences scattered over the body. Commonly the head is partially retracted into the thorax. Caterpillars of some rest with the head held away from their perch; in the extreme, there are those that crane the head back over the abdomen and splay the legs when threatened (below). The egg overwinters in Amphipyrini, whereas the pupa does so in Psaphidini.

! Threat posture of the Black-eyed Cropia. !

Black-eyed Cropia Cropia templada

South Texas, west to at least Big Bend area of Texas. Mature caterpillars throughout growing season but especially common in wet season (October and November). Locally common. Anacahuita or Texas olive (Cordia boissieri). Larvae from three separate collections from Cordia in Falcon Heights, Texas, made by Berry Nall yielded adults of both C. templada and C. connecta. This leads us to suspect that both entities, and perhaps even C. ruthaea, represent a single valid species.

Mistletoe Marblelet Emarginea percara

Southeastern Virginia to Florida, west to California. Multiple generations with mature caterpillars throughout growing season. Locally common. Mistletoe.

Amphipyrines Subfamily Amphipyrinae 325

Smooth Amphipyra Amphipyra glabella ! recognition Lime green with creamy spiracular stripe extending length of body; stripe noticeably thickened on A7–A10. White, parallel-sided, middorsal stripe weakens on T1 and ends on A9. Subdorsal stripe, composed of a series of white spots, absent rearward of A8. Setae borne from bright white pinacula. Scattered white spots salt body above spiracular stripe. Larva less than 4 cm. Superficially resembling some pinions (Lithophane) and sallows (Psaphida). ! occurrence Early successional forests, watercourses, woodland and forest edges. Mostly western, extending eastward at least to Quebec, Connecticut, and Long Island. One generation with mature caterpillars from June through early July. Uncommon. ! common foodplants Cottonwood; other reported hosts (such as ash and Douglas-fir) require confirmation. ! remarks The caterpillar figured here was found on the University of Connecticut campus near where DLW works. We have not taken the moth in light trap samples from the area, suggesting that adults are only weakly attracted to light. We took a pupa from under a bark flap, more than a meter above the ground, while searching for caterpillars of the Griffin (Ufeus plicatus) (page 474), along the lower boles and exposed roots of large cottonwoods (Populus deltoides) growing along the Connecticut River. DFS regularly observed adults in his yard in Hamden, Connecticut on bait; he also observed them as coinhabitants of Copper Amphipyra (A. pyramidoides) roosts, under shingles, loose roofing, and other crevices (but not at light). Forbes (1954) regarded the species as rare. Females do not live past August.

326 Amphipyrines Subfamily Amphipyrinae

Copper Amphipyra Amphipyra pyramidoides [Copper Underwing]

! recognition Attractive blue-green caterpillar with pale yellow spots and

conspicuously humped eighth abdominal segment. White and yellow spiracular stripe continues around anal plate; often broken across T3 and A1. White middorsal stripe. Subdorsal stripe broken into individual spots anteriorly; rearward, subdorsal stripe nearly continuous, angled sharply upward on A8. Larva less than 4 cm. ! occurrence Woodlands and forests. Transcontinental; Canada to Florida and Texas in East. One generation with mature caterpillars from May through June over much of East. Abundant. ! common foodplants Many woody plants including apple, basswood, blackberry, chokeberry, blueberry, cherry, chestnut, currant, grape, greenbrier, hawthorn, hickory, hydrangea, lilac, maple, oak, poplar, raspberry, redbud, rhododendron, viburnum, Virginia creeper, and walnut. ! remarks This is one of our most familiar and ubiquitous spring caterpillars—expect it on almost any nonconiferous, woody plant. The Copper Amphipyra engages in a “shell game strategy with its would-be predators,” moving away from leaves upon which it has recently fed (Heinrich 1979, 1993). Birds that focus on damaged leaves as a means of locating their quarry are likely to do so fruitlessly with the Copper Amphipyra. Early instars are attacked by a microgastrine braconid wasp that fastens its cocoon (and the host larva) to a leaf underside (inset). The adults gather in cracks, under loose bark, and other tight places; evidently an aggregation pheromone aids assembly. When disturbed, the boldly marked sides of the abdomens are made visible. The effect of several alarmed moths “displaying” suggests a group of agitated wasps, and can be startling when seen for the first time. Each summer DLW gets aggregations of prereproductive adults in the shutters on his front porch—their presence is revealed by a reddish excrement that dribbles down the slats. In late August and September the adults, which eclose mostly in July, become reproductively active and disperse from the aggregations. While adults come to bait during the prereproductive portion of the summer, they are rarely taken at light during this time. Yet, by season’s end the moth can be common at light. Stragglers occur into December in New Jersey.

Amphipyrines Subfamily Amphipyrinae 327

Mouse Amphipyra Amphipyra tragopoginis ! recognition Green with white, parallel-sided middorsal, subdorsal, and

spiracular stripes. Spiracular stripe passing through lower half of spiracles on A1– A7. A8 essentially without hump, although subdorsal stripe on A8 bent downward as if hump were present. Setae from minute white pinacula. Integument with abundant, closely set spinules. Spiracles white with black rim. Head glossy green and unmarked. Larva less than 4 cm. Larva resembles some Psaphida (see pages 331–3). ! occurrence Woodlands and forests. Transcontinental in Canada, south in East to Gulf Coast. One generation with mature caterpillars from May through June. Generally uncommon and local. ! common foodplants Many herbaceous and woody plants including bellflower, burdock, columbine, geranium, hawthorn, penstemon, plantain, Scotch lovage (or Scottish licorice-root), spreading dogbane, stinging nettle, strawberry, and willow. ! remarks The dearth of host records in Prentice’s (1962) survey of the caterpillars of forest trees in Canada, and other compendia for lepidopteran foodplants (e.g., Tietz 1972), suggest that the larva typically feeds on herbaceous plants. The white eggs about the thorax and head of the caterpillar in the inset are those of a tachinid fly. Mounting evidence suggests that the Mouse Amphipyra was introduced from Europe. According to Don Lafontaine, its DNA barcode is identical to that of some European samples; it is associated with urban areas in the West; it commonly enters buildings (which would increase its likelihood for human transport); and an 1870 report in the Canadian Entomologist comments on its increasing prevalence in Ontario. Like other members of the genus, the adults are attracted to fermenting fruits (and only weakly to lights). Amphipyra fashion a cocoon in litter or upper soil layers that houses the pupal stage for six to eight weeks. Female Amphipyra are more apt to lay in captivity if taken late in the season and provided with a roughened surface (Porter 1997). Normally overwintering as an egg, although DFS once collected a much worn A. tragopoginis adult in April in Pennsylvania. Curiously, females will seek out and oviposit clutches of 4–12 eggs into the cases of the bagworm, Apterona helix (Adamski 1984).

328 Greenbriers Subfamily Amphipyrinae: Tribe Phosphilini

Greenbriers

Family Noctuidae: Subfamily Amphipyrinae: Tribe Phosphilini This is a small, mostly Neotropical tribe that is represented in North America by six species in four genera—we illustrate just two members in the nominate genus. Lafontaine and Schmidt (2010) moved the tribe into the Noctuini because the spinneret is dorsally grooved, and they regarded the SD1 seta on A9 to be hairlike. We diverge from their

assessment and follow Poole (1995) who placed Phosphila in the Amphipyrinae based on shared features of the male genitalia. Larval characters that suggest affinity to the Amphipyrinae include the humped eighth abdominal segment and the raised and yellowed pinacula (both in P. miselioides). Phosphila adults resemble those of Cropia (an amphipyrine genus) so closely that we wonder if recognition of a separate tribe is warranted. Phosphila and Acherdoa feed on greenbrier (Smilax).

Spotted Phosphila Phosphila miselioides ! recognition Frosty blue-green to almost luminous, semitranslucent, yellow-green with bright yellow spots (pinacula). Body smooth with short and inconspicuous setae. A8 strongly humped. Intersegmental areas with yellowish hue. Lower and rear portions of body with waxy bloom. Larva less than 3.5 cm. The prepupal larva turns smoky, almost black, although the dorsal pinacula retain their bright yellow color (lower inset). ! occurrence Woodlands, coastal shrublands, scrublands, and powerline right-of-ways from Manitoba to southern Maine, to Florida and Texas. Evidently multiple brooded with mature caterpillars from June through October over most of East. Common. ! common foodplants Greenbrier. ! remarks The solitary caterpillar preferentially consumes new foliage. The larva rests on the underside of leaves with the head curled back and positioned alongside the abdomen. When disturbed the head is withdrawn into the thorax and pushed against the leaf, out of harm’s way. The pupa overwinters in a substantial cocoon, spun in leaf litter, into which the caterpillar incorporates frass, soil, and other matter.

Greenbriers Subfamily Amphipyrinae: Tribe Phosphilini 329

Turbulent Phosphila Phosphila turbulenta ! recognition Black and white pinstriping over dorsum and yellow-orange subspiracular region. Both ends of body swollen, especially so rearward. Caudal end of body with prominent white spotting. Larva less than 3.5 cm. Early instars pale green with shiny black head (lower inset). ! occurrence Open woodlands, scrublands, fields, and powerline rightof-ways from Illinois to southern Maine, to Florida and Texas. At least two generations in Connecticut and North Carolina with mature caterpillars from May to July, then again in the fall (September to November). Common. ! common foodplants Greenbrier. ! remarks Once you develop an eye for bared brier stems, you’ll gain a greater appreciation for the ecological importance of this frequent defoliator. Larvae consume mature and fully hardened leaves, many of which may be discolored and heavily damaged, come late summer. The caterpillars are gregarious, lining up side by side on the underside of leaves (upper inset). By the last instar, caterpillars are solitary—look for larvae on leaf undersides, back from the shoot tip. In some ways the insect is three-headed: the last three abdominal segments form a credible false head in the image above; the black and white prothoracic shield, which conceals the true head, is also “eyed.” The caterpillar’s bold coloration is suggestive that the insect is chemically protected, although it remains to be shown if the Turbulent Phosphila is in fact unpalatable, or if its patterning is largely a ruse. The pupa overwinters in a weak cocoon spun in leaf litter.

330 Lash-eyed Sallows Subfamily Amphipyrinae: Tribe Psaphidini

Lash-eyed Sallows

Family Noctuidae: Subfamily Amphipyrinae: Tribe Psaphidini The taxonomic rank and composition of this small clade has ebbed and flowed markedly over the past couple of decades; until recently, lash-eyes sallows were ranked as their own subfamily. We treat 11 species of this largely western tribe, all of which fall into two subtribes: Feraliina and Psaphidina. In addition to characters given for the subfamily, psaphidines have whitened or yellowed pinacula and the pupae bear dorsal pits anterior to the cremaster. Our species have a single generation

with the adults flying in late winter and early spring. Their nonfeeding adults are short lived. They are best sampled with mercury vapor and other bright lights following warm spring days. The large, handsomely ornamented eggs hatch after two to four weeks (next page). The fast-growing larvae are specialists on young foliage of deciduous trees and shrubs, although our (eastern) species of Feralia—representing a divergent subtribe—eat only conifers. Some Psaphida larvae spin and feed from within leaf shelters. Prepupal caterpillars tunnel deep into soil before forming their cocoons. All overwinter as pupae and some for more than one year.

Boreal Sprawler Brachionycha borealis [Boreal Fan Moth]

Alberta to New Brunswick, south to upper Michigan and Wisconsin; disjunct populations in northwestern New Jersey to central Pennsylvania, and mountains of Virginia and West Virginia. One generation with mature caterpillars in June and July. Locally common in Michigan, Wisconsin, West Virginia, and parts of Canada, but otherwise rare in our area. Alder, birch, blueberry, oak, and likely other woody plants; Les Ferge notes that the larvae may be predatory on other caterpillars. Thoracic legs reddened. Rarely collected eastward; often in sandy barrens northward. See account in Schweitzer et al. (2011).

Gray Sallow Psaphida grandis

Wisconsin to New York, south to Florida and Texas; absent from much of Northeast. One generation with mature caterpillars in late spring. Common southward and westward. Our shelterforming caterpillars were reared on oak.

Lash-eyed Sallows Subfamily Amphipyrinae: Tribe Psaphidini 331

bearded sallows

Family Noctuidae: Subfamily Amphipyrinae: Tribe Psaphidini: Subtribe Psaphidina Eastern representatives include eight species in three genera: Brachionycha, Copivaleria, and Psaphida. Our six Psaphida can be divided into the resumens and styracis groups, each with three species. Members of the first group are green with pale spotting, do not form shelters, and frequently pass more than one winter as a pupa. Members of the second group are more handsomely patterned, form shelters, and do not pass more than one winter as a pupa. The fertile eggs of Psaphida transition from rose pink to reddish gray (right).

Chosen Sallow Psaphida electilis ! recognition Frosty to yellow-green above cream lateral stripe and bluish green below. Conspicuous white setal bases whitened, raised, and pimplelike; these largest over T2 and T3. Lateral stripe beginning on T2; upper portion ill-defined, running just below spiracles; lower edge defined and contrasting with ground color; stripe continuing around anal plate. Middorsal stripe narrow and frequently broken. Subdorsal stripe rudimentary, formed of widely spaced spots. Outer face of each proleg bearing shiny green plate. Pale green head smooth and shiny. Larva to 4 cm. ! occurrence Woodlands and forests from Wisconsin to Quebec south to Florida and Texas, but mostly absent from Atlantic Coastal Plain. One generation with mature caterpillars from middle to late spring. Locally common. ! common foodplants Hickory and walnut. ! remarks Despite the abundance of the adults at light, we have yet to encounter caterpillars of this spring-active moth in the wild. Captive larvae feed and rest on leaf undersides. They do not spin shelters. One of our ex ova cohorts strongly preferred black walnut over pignut hickory (Carya glabra) when both were offered together. Pupae overwinter in the soil, often for more than one year.

332 Lash-eyed Sallows Subfamily Amphipyrinae: Tribe Psaphidini

Figure-eight Sallow Psaphida resumens ! recognition Light green to blue-green caterpillar marked with thin, white, middorsal stripe and yellowish spiracular stripe that is broken anteriorly, but well developed rearward of A6. Abdominal segments (especially on A2–A7) with oblique supraspiracular lines that end below weak, and often broken, subdorsal stripe. Body profile high relative to body width. Head pale blue-green, unmarked, partially withdrawn into prothorax; labrum whitened. Larva to 4 cm. ! occurrence Woodlands and forests from Minnesota and southern Ontario to Maine, south to Florida and Texas. One generation with mature caterpillars from middle to late spring, but always before leaves harden. Common. ! common foodplants Oak; our records from those of white oak group. ! remarks Larvae of oak-feeding Psaphida usually hatch later, but catch up and mature earlier than most other spring Noctuidae. Caterpillars require young leaves in all instars; last instars forced to finish on older leaves may die outright or fail to eclose from the pupae if they mature at all. The late hatching of this species precludes successful maturation on oaks that leaf out early in the season. One cohort reared by DFS refused scrub oak, a favorite among oak-feeding owlets. Old reports of the Figure-eight Sallow from maple are in error and probably refer to misidentifications of Grote’s Pinion (Lithophane grotei) larvae. Larvae do well if sleeved on sunny terminal foliage. Our guess is that the Figure-eight Sallow and other psaphidines are canopy insects, because we encounter few larvae relative to the number of adults seen at lights. No instar of this species makes a shelter, but the caterpillars will hide in the shelters of tortricids and other moth larvae. The lethargic caterpillar usually rests on the main vein of the underside of the leaf, and even mature individuals are difficult to see except when feeding. During an especially warm May, eggs from two females that eclosed on 1 May yielded the first prepupal larvae by 21 May (cohort on white oak), and all larvae had matured by 23 May (cohort on post oak). The pupa overwinters deep in soil. Expect most pupae to overwinter more than one year—DFS has had pupae hold over for as many as seven winters. The Figure-eight Sallow ecloses and flies later than our other four oak feeders, often flying through mid-April in southern New Jersey (matching the late phenology of its white and post oak hosts).

Lash-eyed Sallows Subfamily Amphipyrinae: Tribe Psaphidini 333

Roland’s Sallow Psaphida rolandi ! recognition Bright lime green with white pinacula; somewhat stocky, broadest about A2 and A3; caudal end tapering to anal plate. Middorsal stripe broken over thorax. White spiracular stripe, thickened rearward of A7, continuing around rim of anal plate. Subdorsal stripe composed of scattered white spots and lines. Triangle and front of head blue-green; pale snowflake markings over vertex. Larva to 3.5 cm. ! occurrence Woodlands and forests from southern Ontario to Maine south to central Florida and Texas. One generation with mature caterpillars in spring. Very common. ! common foodplants Oak. ! remarks The caterpillars feed on new, unhardened foliage. They do not make a shelter. Adults are among the most common early spring owlets at light in Connecticut—in late March and early April, we sometimes see dozens of individuals at a mercury vapor light sheet. Yet, like other Psaphida, larvae are seldom encountered. Ninety percent of the Roland’s Sallows reared by DFS have overwintered as a pupa for two to seven years before eclosing (40 from two cohorts). To obtain eggs from Psaphida females, place them in a small jar or vial with a piece of paper towel along one side, one or two twigs with terminal buds oriented upward and not touching lid, and a small piece of dead leaf from an appropriate foodplant. Hold females outdoors, away from direct sun. Females lay about 150 eggs. In captivity, and apparently in nature, up to half are laid the first night, and the rest spread over about a week. Eggs kept outdoors hatch a few days after budbreak of the foodplant.

334 Lash-eyed Sallows Subfamily Amphipyrinae: Tribe Psaphidini

Fawn Sallow Psaphida styracis ! recognition Coloration variable. Commonly waxy blue-green, especially over abdomen, and reddish or greenish along lower half of body. Densely speckled with white spots, especially along sides of body. Prothoracic shield shiny black with central white bar. Head orange and unmarked. Larva less than 3.5 cm. Middle instars more reddish to maroon and lacking prominent, pale dorsal area (but see Remarks). The Gray Sallow (P. grandis) is more evenly mottled and has a black head (page 330). While generally distributed from central Pennsylvania southward and westward, P. grandis is absent from the Northeast. Steve Johnson reports the species as frequent in the oak barrens of Centre County, Pennsylvania. ! occurrence Mostly dry, oak-dominated forests, woodlands, and barrens from Wisconsin to southern Ontario and Massachusetts, to Florida and Texas. One generation with mature caterpillars in late spring. Locally common. ! common foodplants Oak; our records from both red and black oaks. ! remarks Adults emerge early in the spring, beginning in February in the South. Adults in New Jersey occasionally emerge on days when the temperatures remain below 10° C (50° F). They eclose in the morning and crawl up a perch; normally the wings are promptly pumped up, but in cold weather expansion may be delayed for a day or more. Its caterpillars are reluctant to leave their shelters once spun, so it is a good idea to physically transfer larvae to new foliage if you are rearing this species in containers. A better alternative is to sleeve the caterpillars on a shoot with new growth. Evidently, the larva changes color in the last instar, starting off somewhat maroon red, and then becoming increasingly greener with age. If given an opportunity, prepupal larvae will tunnel deep in soil—as much as 15 cm (6 in.) in our rearing containers. The pupa overwinters, but not for more than one year (DFS, unpubl. data).

Lash-eyed Sallows Subfamily Amphipyrinae: Tribe Psaphidini 335

Thaxter’s Sallow Psaphida thaxterianus ! recognition One of North America’s most distinctive and immediately recognizable owlet caterpillars. Reddish to chocolate brown with large, bright, whitish to sea-green patches that reach from dorsum to spiracle. Prothorax with anterior and lateral portions of shield pale green, and rearward areas black. Pinacula behind and below spiracle pale green, contrasting with reddish ground color. Larva to 3.5 cm. ! occurrence Dry open woodlands, barrens, ridge tops, clear cuts, and other scrubby, oak-dominated habitats, usually with acid soil. Southern Manitoba to eastern Massachusetts, south to Georgia (mountains) and eastern Texas, but absent from much of Northeast and other parts of range. One generation with mature caterpillars in late spring. Sometimes common in dry oak woodlands and barrens. ! common foodplants Oak; we have collected larvae from scarlet oak (in Virginia). Our sleeved larvae matured on both white and red oaks. ! remarks Eggs of this species hatch later than those of other psaphidines laid at the same time—at least in frost pockets and some other habitats, the delayed foliation of oaks may be a key requirement for larval survival. This is one of the most handsome eastern owlet caterpillars. Lamentably, like other Psaphidini, caterpillars of Thaxter’s Sallow are rarely encountered. The caterpillar forms a leaf shelter by tying together a cluster of new leaves (inset). We are unable to explain the adaptive significance of the larval coloration given that the caterpillars are normally concealed within shelters. Thaxter’s Sallow is among the earliest springemerging noctuids, usually flying with Roland’s Sallow (P. rolandi) and the Fawn Sallow (P. styracis), ending its flight by the middle of April, even in Wisconsin. The sex ratio of adults at light is much more strongly male-biased than in other Psaphida—expect to find dozens of males for every female.

336 Lash-eyed Sallows Subfamily Amphipyrinae: Tribe Psaphidini

Grote’s Sallow Copivaleria grotei ! recognition Pale or frosty green, marked with white middorsal, broken yellow subdorsal, and solid yellow-green and white spiracular stripes. Ground color of venter and subventer waxy bluish green. Middorsal and subdorsal stripes reduced or absent on A8–A10. Spiracular stripe lime green to yellow-green above and white below; white portion thickened on A8 and A9. Dorsal (and some lateral) setae borne from round, white to yellowish bases. Middorsal stripe absent on T1 and broken into spots and lines on T2 and T3. Prothoracic shield whitened along leading edge. Straw-colored spiracles with darkened rim. Head pale green and unmarked. Larva to 4 cm. ! occurrence Woodlands and forests from Manitoba to Nova Scotia, south to Florida and Texas. One generation with mature caterpillars in late spring. Locally common. ! common foodplants Ash. ! remarks Grote’s Sallow is a common moth in most forested regions of eastern North America, yet the caterpillars are rarely encountered. Perhaps many of the larvae are active in canopy foliage. This owlet and other ash feeders may be lost if the Emerald Ash Borer’s (Agrilus planipennis) (Buprestidae) decimation of Fraxinus cannot be brought under control (Wagner 2007c).

Lash-eyed Sallows Subfamily Amphipyrinae: Tribe Psaphidini 337

mossy sallows

Family Noctuidae: Subfamily Amphipyrinae: Tribe Psaphidini: Subtribe Feraliina Body shiny, emerald green with prominent white to cream stripes, and slightly humped eighth abdominal segment. In

our three eastern species, the spiracular stripe is edged above with red. Young larvae feed on new spring growth, whereas older larvae also eat mature needles. Pupation occurs in the soil; pupae overwinter, often for more than one year. Eastern species feed on conifers.

Pine Sallow Feralia major [Major Sallow] ! recognition Deep green with strong, white middorsal and addorsal stripes; cream spiracular stripe edged with burgundy above. Subspiracular stripe made up of elongate yellow spots on thoracic segments and A1–A8; those on A1–A6 nearly connected. Thoracic legs wine red; midabdominal prolegs with wine-red spot at base. Larva to 3.5 cm. In both the Comstock’s Sallow (F. comstocki) and Jocose Sallow (F. jocosa), the subdorsal and spiracular stripes are broken into lens-shaped spots and the subspiracular spots are reduced or absent. Superficially similar to Pine Pinion (Lithophane lepida) (page 439) and some pine-feeding Xestia (e.g., page 543), all of which lack the burgundy edging above the subspiracular stripe. ! occurrence Pine barrens, woodlands, and forests from southern Canada to northern Florida and eastern Texas. One generation with mature caterpillars from late April through July, with most encountered from late May through June over much of East. Locally common. ! common foodplants Principally pines, both hard pines and white pine. Reports from spruce and other conifers may be in error. ! remarks The Pine Sallow flies early in the spring northward; adults are seen southward beginning in late December and January. It is most apt to be encountered in middle to low elevation and Coastal Plain pine forests. Feralia adults rest on tree trunks where their coloration blends with that of foliose lichens. Adult color and pattern varies markedly in all our Feralia and especially so in the Pine Sallow—its forewings range from mostly green-white to sea green, gray-green to nearly black in melanic individuals. In southern New Jersey, an area of relatively low melanism in other moths, all adults are of the blackish form.

338 Lash-eyed Sallows Subfamily Amphipyrinae: Tribe Psaphidini

Jocose Sallow Feralia jocosa [The Joker] ! recognition Emerald green, with broken spiracular stripe made up of maroon upper portion and white to yellow lower portion. Subdorsal stripe continuous along thorax but broken or at least constricted between abdominal segments. Abdominal prolegs with yellow spot at base. Larva to 3.5 cm. In Comstock’s Sallow (Feralia comstocki) (inset) spiracular spots nearly fully connected along abdomen; spiracular spots often have prominent yellow lower component; upper portion of spiracular spots frequently tinted with rose or red; and spots above prolegs yellow and ovoid. The Jocose Sallow usually with spiracular spots often fully separated between segments; little to no yellow in spiracular spots; red portion of spiracular spots often burgundy; and spots above prolegs on A3– A6 frequently elongate, but they may also be small or absent. Comstock’s Sallow occurs in northern and boreal forests across Canada, south in East to Connecticut, Georgia (mountains) and Great Lakes states. Several larval images of both F. comstocki and F. jocosa are given in Duncan (2006); all three eastern Feralia are figured by Maier et al. (2004). ! occurrence Woodlands and forests. Transcontinental in Canada, south in East to northern New Jersey, Georgia (mountains), and Great Lakes states. One generation with mature caterpillars mostly in June and early July. Common. ! common foodplants Balsam fir, hemlock, spruce, and other conifers (Maier et al. 2004). ! remarks Through much of the East, the Jocose Sallow is the most commonly encountered member of the genus, especially in low to midelevation hemlock forests. It flies early with the Pine Sallow (F. major) and its season ends about the time Comstock’s Sallow (F. comstocki) takes wing. In the Appalachians, where all three Feralia fly together, the Jocose Sallow has the greatest altitudinal span of the three, ranging from low-elevation hemlock stands upwards into high-elevation Fraser fir forests.

Seedcoppers Subfamily Stiriinae 339

Seedcoppers

Family Noctuidae: Subfamily Stiriinae Seedcoppers are most diverse in arid areas: Texas is home to more than three dozen species while the whole of the Northeast can claim only two. They share important morphological similarities with the Psaphidini (page 330), and may actually be best treated as a tribe within the Amphipyrinae, as proposed by Lafontaine and Schmidt (2010). Stiriine caterpillars are among the most morphologically and behaviorally specialized owlets, and include among their ranks many of the most amazing flower mimics in the animal kingdom. They would make a wonderful group to study how Darwinian

Malvastrum Seedcopper Chamaeclea basiochrea

Extreme southern Texas. One principal generation with mature caterpillars mostly from October to November. Common. False mallow (Malvastrum).

Beggarticks Seedcopper Cirrhophanus triangulifer [Goldenrod Stowaway]

Kansas to Ohio and southern New York, to northern Florida and Texas. One principal generation with mature caterpillars mostly in September and October. Locally common. Spanish needles or beggarticks (Bidens), with most records from bearded beggarticks (B. aristosa). Jones (1937) provides details on this insect’s life history in Delaware.

Ragweed Seedcopper Plagiomimicus pityochromus [Black-barred Brown]

Minnesota, southern Ontario, and Long Island, New York, south to Florida and Texas. Becoming more generally distributed and common southward. One principal generation with mature caterpillars mostly from September to November. Locally common. Ragweed; great ragweed (Ambrosia trifida) over much of East. Figured individual from Cuman ragweed (A. psilostachya) in south Texas.

natural selection molds both form and behavior. The caterpillars have a short, scalelike spinneret. SD1 on A9 is hairlike. North American species are flower and seed feeders, often on composites. Life histories and brief larval descriptions for several North American stiriines appear in Poole (1995), although as a group their life histories are poorly known and much in need of study. Stiriines can be challenging to rear as many are seed predators that require flowers of the proper age to complete development. To make matters more difficult, prepupal larvae are finicky about their choice of soil for cocooning— when collecting caterpillars for rearing, also gather ample duff and soil from below the foodplants. Pupae overwinter.

340 Seedcoppers Subfamily Stiriinae

Crownbeard Seedcopper Basilodes pepita [Gold Moth] ! recognition Distinctly patterned orange, black, and white caterpillar that cannot be confused with any other in our fauna. Dorsum crossed by alternating smoky black and pale rings. T2–A8 with orange supraspiracular and white subspiracular spots. Prothoracic shield and A10 mostly orange with black spots. Head pale orange and unmarked. Larva less than 4 cm. ! occurrence Fields, meadows, glades, and edges of woodlands from North Dakota, southern Ontario, and western New York, south to Florida and Texas. A single generation with mature caterpillars in September and October. Locally common. ! common foodplants Crownbeard (Verbesina); commonly wingstem (V. alternifolia). ! remarks Both the coloration and behavior are suggestive of a sawfly. Like a sawfly, the larva is rather slow moving and seemingly thick skinned. When disturbed, it often stops, rolls the head under, and elevates the anterior abdominal segments. Although the species is not known to be toxic, its coloration, sluggish behavior, and foodplant suggest that the caterpillar enjoys some level of chemical protection. Middle and early instars eat mostly at night, consuming both ray and disk flowers. In the last instar, larvae feed at all hours and eat both flowers and seeds (Laura Miller, pers. comm.).

Gilded Seedcopper Basilodes chrysopis

Kansas to central Texas, west through Arizona. One principal generation with mature caterpillars mostly September to November (following monsoons in Arizona and elsewhere). Locally common. Cowpen daisy or golden crownbeard (Verbesina encelioides). See Poole (1995).

Seedcoppers Subfamily Stiriinae 341

Sunflower Seedcopper Stiria rugifrons [Yellow Sunflower Moth]

! recognition Unmistakable in shape, coloration, behavior, and association with sunflowers. Dorsal setae borne on high, yellow-orange (D1) and smoky red to black (D2) warts on T2–A8. Larva less than 3.5 cm. Penultimate instars bright yellow with incomplete dark ring passing through the D1 warts. Other members of the genus occur in Texas and in the West. ! occurrence Prairies, grasslands, outcrops, woodland clearings, and other sunny habitats from Canadian Great Plains to Ohio and Virginia, south through Gulf states. Common through Great Plains, Ohio, and Southeast, but rare and local east of Appalachians. One generation with mature caterpillars from August through October; later southward. Local and uncommon; absent from many seemingly suitable sites. ! common foodplants Sunflower; small woodland sunflower (Helianthus microcephalus), confederate daisy or Porter’s sunflower (H. porteri), and no doubt others (but see below). ! remarks Eloise Carter and Steve Baker of Oxford College, Emory University, generously provided the following unpublished life history information on this marvelous animal. The caterpillars are dependent on reproductive (floral) tissues. Young larvae consume pollen; middle instars eat flowers, especially the ray flowers or petals; and last instars perch below the flower and tunnel through the receptacle, gaining access to and feeding on the developing ovules. Sometimes it is possible to locate last instars by looking for the accumulation of discarded tissues below the feeding site (mostly the feathery pappi that transport mature seeds). The superbly camouflaged larvae perch on the disk of open sunflowers. Middle instars, which rest curled about the disk, have been fine tuned by natural selection; their coloration (particularly their alternating yellow and black pattern), texture, and behavior match their sunflower hosts to perfection (above right). As if that were not enough, the caterpillar’s coloration and resting posture change in the last instar, and are reminiscent of a past flower that is on its way to setting seed (right). Adult coloration is suggestive of a fallen, yellowed leaf. Both sexes come to light but seldom in number. Pupae may hold over for more than one winter.

342 Sun Moths Subfamily Heliothinae

Sun Moths

Family Noctuidae: Subfamily Heliothinae This subfamily contains more than 150 North American species. Sun moths are especially well represented in the American Southwest and other arid environs: e.g., prairies, sand plains, and coastal strand communities. A single genus (Schinia) accounts for much of this diversity: more than 120 species are known from north of Mexico and several new species await description. Two genera (Heliothis and Helicoverpa) include important agricultural pests. Approximately 40 species of heliothines occur in our area, although many of these are restricted to portions of Florida and Texas. The integument bears numerous, fine, spine-tipped granules (visible with a strong lens), and often raised or pimplelike setal bases. On the prothorax, the two setae immediately forward of the spiracle (L setae) are usually aligned parallel to the body axis (in most other noctuoids the L setae are positioned vertically); in a few the two L setae are offset by as much as 45 degrees. The crochets are of two lengths (biordinal) in most. Heliothine caterpillars are commonly flower or seed feeders. The foodplants are typically low-growing dicots, including many annuals. Pyrrhia are our only heliothines commonly found feeding on trees. In a pinch, many can be reared on green beans. Several are known to be cannibalistic (above right),

especially those that feed on plants of small stature. The caterpillars frequently perch with the thorax and anterior abdominal segments arched upward, and roll the head under when alarmed. Pupae overwinter in underground cells, sometimes for more than one year.

Adults are highly dependent on nectar and Schinia species may feed only on the flowers of the larval foodplant. Some even oviposit while nectaring. The floral lure phenylacetaldehyde is attractive to Helicoverpa (Meagher 2002) and likely its relatives. Sugar bait is visited by some, especially those with migratory tendencies. Many are rare, localized, or otherwise make worthy conservation targets. Larval surveys are often the most time efficient and reliable means by which to census heliothine populations.

False Foxglove Sun Moth

Pyrrhia aurantiago (= Rhodoecia aurantiago) [Orange Sallow] Wisconsin to Maine, south to Florida and Texas, although uncommon to rare over much of range. One generation with mature caterpillars from late August through October or later. Local and uncommon. False foxglove (Aureolaria flava, A. grandiflora, A. pedicularia, but not A. virginica) and perhaps mullein foxglove (Dasistoma macrophylla) (all Scrophulariaceae). Look for holes in ripening seed pods. Middle instars pale to yellow tan. See Wyatt (1938) and account in Schweitzer et al. (2011).

Horse-balm Sun Moth

Psectrotarsia hebardi (= Erythroecia hebardi) Appalachian endemic with narrow range from southern Ohio to northwestern New Jersey, south into Virginia. One generation with mature caterpillars in September. Rare. Horse-balm or richweed (Collinsonia canadensis). See account in Schweitzer et al. (2011).

Sun Moths Subfamily Heliothinae 343

Noble Sun Moth Pyrrhia cilisca (Pyrrhia adela) [American Bordered Sallow] ! recognition Variable in coloration—ground color ranging from green to pink, olive, brown, gray, and black, with broad, lemon spiracular stripe. If striped (inset), look for five black stripes separated by four paler stripes above spiracular stripe. Setal bases shiny black; those on A1 and A2 and over dorsum of A8 largest. Often with triangular black plate on outer face of each proleg, dark spiracles, and orange head. Larva to 3.5 cm. Co-occurring with Purple-lined Sun Moth (P. exprimens). Because of the extreme variability of the caterpillars of Pyrrhia, we are uncertain how to reliably separate our two eastern species. Don Lafontaine (pers. comm.) suggests that some P. exprimens caterpillars may be distinguished by having the dorsal stripes broken up into splotches, and by their pale yellow spiracles. Identifications are best assured by rearing to the adult stage. ! occurrence Fields and other open habitats from Manitoba to Newfoundland, south to South Carolina and Texas. Apparently two or more generations with mature caterpillars through summer and fall months. Uncommon to locally common. ! common foodplants General feeder on leaves, buds, and reproductive tissues. Many records from alder and sumac, but also balsam fir, cabbage, monkshood, mustard, penstemon, poplar, rose, ticktrefoil, and walnut (see Remarks). ! remarks The Bordered Sallow (P. umbra) is a European species with which P. cilisca (and its synonym, P. adela) were previously confused. Presumably the Noble Sun Moth and its Old World counterpart have similar biologies. Caterpillars of P. umbra show a predilection for flowers and seeds of low-growing plants. Like many flower-feeding Lepidoptera, its caterpillars are given to cannibalism and will consume smaller larvae that they encounter. Adults of the Bordered Sallow come to light and bait, and oviposit in captivity if held with a suitable host (Porter 1997). Our species, P. cilisca, frequently includes small trees in its diet—both of our larval collections came from sumacs growing in open areas. The pupa overwinters.

344 Sun Moths Subfamily Heliothinae

Purple-lined Sun Moth Pyrrhia exprimens ! recognition Exceptionally variable in color, ranging from almost white, yellow, or green to black, with striping variously expressed. Spiracular stripe ranging from narrow, creamy line to broad, orange to yellow belt. Subdorsal stripe present, although vaguely expressed in many forms. Two sets of faint addorsal stripes variously developed (one along inner side of D1 and other inside of D2). Integument with short macrospinules (visible at 12× magnification); microspinules absent (visible at 50× magnification) (Hardwick 1996). Larva to 3.5 cm. Cooccurring with Noble Sun Moth (P. cilisca). Caterpillars also approach some Schinia in appearance, e.g., False Boneset Flower Moth (S. grandimedia). Because of the extreme variability of Pyrrhia caterpillars, identifications are best based on (bred) adults. ! occurrence Fields, gardens, prairies, and mostly open habitats from British Columbia to Newfoundland, south to Georgia and Texas, but mostly in mountains southward. Two generations north to southern Quebec with mature caterpillars from June onward. Uncommon to locally common. ! common foodplants General feeder on both forbs and woody plants; many records from beggarticks and other legumes; other hosts include chicory, columbine, dogbane, monkshood, pea, penstemon, poplar, rose, strawberry, sweet fern, and willow. ! remarks The spectacular coloration of the caterpillar invariably draws attention. Three images are not enough to capture the species’ variability—likely your caterpillar will not match any shown here. Not infrequently, the Purple-lined Sallow turns up in flower beds and gardens, especially westward through the Great Plains. The caterpillars consume both leaves and reproductive tissue, but prefer buds, flowers, and fruit; when opportunity allows, the larvae will sometimes tunnel into these. The pupa overwinters.

Sun Moths Subfamily Heliothinae 345 The Corn Earworm (Helicoverpa zea) and Tobacco Budworm (Heliothis virescens) are among the most economically significant lepidopterans in North America. Below we provide a larval key (based largely on that of Godfrey 1987) to five eastern Helicoverpa and Heliothis (only the latter genus is abbreviated in the text below). Heliothis ononis and H. acesias appear in the key but are not figured. A sixth Canadian species, H. borealis, occurs from northern Ontario westward (like H. ononis the dorsal pinacula on A1 and A2 are not warted). Hardwick (1996) figures all but H. ononis and H. acesias. Larval identifications are difficult even for experts, and require microscopic examination of minute setae, integumental spines, and dissection of the mandibles.

Key to Larvae of Common Eastern Helicoverpa and Heliothis 1a Dorsal pinacula on A1, A2, A8 (and others) mostly smooth, without spinules reaching up pinaculum towards setal insertion (see figure below); dorsal pinacula not connected by diffuse, darkened bar (patch); inner surface of mandible with or without enlarged basal tooth . . . . . . . . . . . . . . . . . . . . . . . . . 2 1b Dorsal pinacula on A1, A2, A8 (and others) with spinules reaching up pinaculum towards setal insertion (see figure below); dorsal pinacula on A1 and A2 often connected by diffuse, darkened bar; inner surface of mandible with enlarged basal tooth (although this sometimes missing, broken free at its base) (see figures below) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Heliothis virescens 2a Dorsal pinacula on A1 and A2 scarcely differentiated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Heliothis ononis 2b Dorsal pinacula on A1 and A2 raised and sometimes darkly pigmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3a Height and diameter of D1 pinaculum on A1 nearly equal to height of spiracle on same segment; no minute SD2 seta visible anterior to spiracle; mandible without or with only slightly enlarged basal tooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Helicoverpa zea 3b Height of D1 pinaculum on A1 nearly equal to height of spiracle on same segment, but diameter at base often exceeding height; minute SD2 setae anterior to spiracle often borne from brown spot; mandible with enlarged basal tooth; often feeding on groundcherry (Physalis) and nightshade (Solanum) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Heliothis subflexa 3c Height and diameter of D1 pinaculum on A1 nearly twice height of spiracle on same segment; apex of dorsal pinacula black; minute SD2 setae anterior to spiracle present but scarcely differentiated; mandible without enlarged basal tooth . . . . . . . . . . . . . . . . . . . . . . Heliothis phloxiphaga and H. acesias 1 A fifth eastern Heliothis, H. acesias, was long confused with H. phloxiphaga. It occurs across southern Canada, and in our region, south through the Great Lakes states to Texas. Its larva and life history remain unstudied—presumably it would key out here.

1

phloxiphaga

ononis inner tooth virescens zea subflexa

1st abdominal segment

zea

! First abdominal segment of Heliothis and Helicoverpa showing condition of the d2 pinaculum. !

virescens

broken inner tooth virescens

! Inner surface of mandible. Redrawn from Godfrey (1987). !

346 Sun Moths Subfamily Heliothinae

Tobacco Budworm Heliothis virescens ! recognition Extremely variable in coloration: green, yellow, pink, red, maroon, and other colors. Dorsum of A1 and A2 frequently with diffuse, darkened, transverse bar or patch connecting enlarged, wartlike D pinacula. Dorsal pinacula on A1, A2, A8, and many others with integumental microspinules. SD1 seta (that above spiracle) enlarged, especially on anterior abdominal segments. Often with blackened subdorsal patches. Inner surface of mandible with central tooth (or scar where tooth had been). Integument with rows of minute, whitened spinules (visible under magnification). Larva to 4 cm. Groundcherry Sun Moth (H. subflexa) (next page) caterpillar similar, but with small SD2 seta on abdominal segments proportionately longer; often with four pronounced black setal bases about abdominal spiracles (typically only three present on H. virescens); closely associated with groundcherry and nightshade. If our reference collections of Heliothis caterpillars are identified properly, then the integument of H. subflexa is granulose and without distinct lines of spinules. ! occurrence Fields, gardens, and waste lots from Nebraska to southern Ontario and Maine, southward through Gulf states. One principal generation in North, two or three in New Jersey, and breeding continuously in southern Florida. Migratory, becoming increasingly common in late summer at least as far north as southern Great Lakes states and New England. ! common foodplants Many reported hosts: abutilon, cotton, geranium, groundcherry, tobacco, zinnia, and plants with glandular hairs among preferred hosts (see below). Numerous records from various Malvaceae and soybean. Larvae have been found in New Jersey and Virginia eating seeds of the federally endangered Virginia jointvetch (Aeschynomene virginica). ! remarks As with other heliothines, the caterpillars have a penchant for buds, flowers, fruits, and seeds. Somehow the larval coloration frequently ends up matching that of the foodplant: on cotton the caterpillars are green, while on geranium flowers maroon colors prevail. Our colleague Scott Smedley and his students recently discovered that Tobacco Budworm caterpillars are able to transfer

Sun Moths Subfamily Heliothinae 347 the glandular defensive secretions of their foodplants onto their own setae, and in so doing accrue chemical protection from ants and other natural enemies. The pupa overwinters, at least north to coastal New Jersey. The Tobacco Budworm is a migrant to New England, with most larval records falling in the months of September and October.

Spotted Straw Sun Moth Heliothis phloxiphaga

Mostly western; eastward to Ontario, Ohio, and Arkansas, occasionally with strays to East Coast. Continuously brooded through growing season southward; northward most likely to be encountered in fall (as offspring from migrants). Common westward. Generalized feeder on reproductive tissues of low-growing plants: e.g., aster, delphinium, dustymaiden or pincushion (Chaenactis), gilia, gladiola, gumweed (Grindelia), phlox, strawberry, sweet pea (Lathyrus), tarweed (Madia), and others (Crumb 1956, Covell 2005).

Groundcherry Sun Moth Heliothis subflexa

Upper Midwest to New England (where it is rarely encountered), south to Florida and Texas, and westward to Pacific; northern records represent seasonal migrants. Common from Ohio southward and westward. One principal generation in North with mature caterpillars from September to November on black nightshade (Solanum) and groundcherry (Physalis). Common southward. SD2 pinaculum larger than that of Tobacco Budworm (H. virescens). Note: Figured (yellow) larva did not produce an adult, and pinacula about spiracle smaller than those described by Forbes (1954). See account for Tobacco Budworm.

348 Sun Moths Subfamily Heliothinae

Corn Earworm Helicoverpa zea ! recognition Highly variable in color, ranging from pale tan to yellow, orange, red, maroon, green, dark brown, or nearly black. Stripes variously developed. Setal bases often black and shiny, typically not as enlarged over A1 as in Tobacco Budworm (Heliothis virescens). Raised or warted portion of dorsal setal bases on A1, A2, and A8 nearly smooth, without microspinules. Mandible without a central tooth or tooth scar. Larva to 4 cm. Many individuals may be distinguished from those of the Tobacco Budworm by examining microscopic details of dorsal pinacula: in the Tobacco Budworm the integumental spinules often continue onto these setal bases (Godfrey 1987), while the dorsal pinacula are nearly smooth in the Corn Earworm (see figure on page 345). The mandible is the most reliable means for separating the two. ! occurrence Agricultural fields, gardens, waste lots, and other open habitats, from Canada south through Florida and Texas. One principal (fall) generation in North. Two or three generations over most of our area with mature larvae typically from late July into October. Multiple generations with mature caterpillars through growing season in parts of Florida and Texas. Common pest through late summer and fall months. ! common foodplants Widely polyphagous, especially on herbaceous plants, consuming mostly buds, flowers, and fruits. Especially fond of corn, peppers, and tomato. Other economically important hosts include artichoke, asparagus, beans, cotton, cucumber, okra, soybean, spinach, squash, sunflower, and tobacco. Native foodplants include groundcherry, various composites, a range of low perennials, and perhaps some larger grasses. In addition, frequently cannibalistic.

Sun Moths Subfamily Heliothinae 349 ! remarks The Corn Earworm’s many aliases attest to its polyphagous nature and its considerable economic importance: Cotton Bollworm, Soybean Podworm, Tomato Fruitworm, Sorghum Headworm, and Vetchworm. It is among the most economically injurious North American crop pests. Its penchant for young tissue, and especially those of reproductive organs, makes its presence especially problematic because harvested parts of the plant are likely to be damaged. On corn, larvae tunnel into the kernels towards the apical portion of the ear and render it unsalable. Individuals that bore into host tissue tend to have more subdued coloration (as in image below) than caterpillars feeding externally on seeds and flowers. Typically, larvae pass through six instars. The entire life cycle may be completed in as little as four weeks during warm summer months when mean daily temperatures exceed 30° C (86° F) (Butler 1976). Larvae are both cannibalistic and predatory—two of those figured here killed smaller caterpillars of the Olive-shaded Bird-dropping Moth (Ponometia candefacta) and Pinkbarred Marvel (Pseudeustrotia carneola) before we had a chance to isolate them. Although pupae overwinter to about latitude 40° north, roughly Kansas, Ohio, and southern New Jersey (Schweitzer 2006, Capinera 2007), populations in most of our region are augmented annually by late-spring and summer migrants from the Gulf states and Mexico. In Arkansas, and probably New Jersey, migrants arrive before overwintered pupae eclose. The occasional appearance of large numbers in the South, coinciding with northerly cold fronts, suggests that there may be a southern migration in the fall (Johnson 1995 and references therein). Larvae are attacked by a multitude of tachinid flies, wasps, and predators. In our larval collections from noncrop hosts, tachinids (possibly an Archytas) have taken a heavy toll. Three tachinid fly eggs are visible on the individual figured in the lower left panel on previous page.

350 Sun Moths Subfamily Heliothinae

Purple-top Sun Moth Heliocheilus lupatus ! recognition Slender, purplish-black and yellow-striped caterpillar with tapered abdomen. Integument minutely spinulose. Thin, black middorsal stripe sandwiched between broader yellow addorsal stripes. Thin, white subdorsal stripe thickened and accentuated through shiny black prothoracic shield; often with obscure white to reddish stripe running between dorsal setae. Broad, lemon-yellow spiracular stripe edged with white above and below. Head reddish brown with diffuse dark patch over antenna. Larva to 3.5 cm. Penultimate instar more yellow with black and yellow markings about evenly divided. ! occurrence Fields, roadsides, and waste lots from Missouri, Kentucky, and New Jersey, south to Florida and Texas. One generation with mature caterpillars from late August to early October in New Jersey, later southward. Locally common. ! common foodplants Flowers and seeds of purpletop tridens (Tridens flavus) and perhaps other grasses. ! remarks This moth has been known as Heliothis turbatus, a name Hardwick (1996) restricts to a rare Floridian species. The Purple-top Sun Moth is expanding its range. At the time Forbes (1954) wrote his book, the moth occurred north only to the Carolinas. It appeared in southern New Jersey in the 1970s, where it is now common and widespread. Smaller larvae are nearly impossible to locate, but caterpillars become conspicuous once their girth exceeds that of the stems upon which they are perched. Larvae seldom rest on the main stem, but rather on the seed-bearing branchlets below the flowers or seeds. Caterpillars can be extremely common, especially relative to the numbers of adults found in major collections. There are colonies in southern New Jersey where DFS has found hundreds of larvae over the course of a few hours. Weeping lovegrass (Eragrostis curvula) is suspected as a secondary foodplant for both this species and H. paradoxus in New Jersey. The latter species, when reared on corn kernels, completes its larval development in as few as 12 days (Hardwick 1996).

Sun Moths Subfamily Heliothinae 351

flower moths Schinia

With more than 120 North American species, Schinia ranks as one of the largest and most evolutionarily successful genera of Lepidoptera in the New World. The genus is most diverse in semiarid and arid environments; sand plains, savannas, scrublands, grasslands, prairies, dune, and coastal strand communities harbor comparatively rich faunas in our region. Connecticut has but nine species, Missouri has 42, and Texas has 66. We list 70 species as residents of our region in the table on page 361. The brightly colored adults may be diurnal, nocturnal, or both. Hardwick’s (1996) monograph is a must for anyone interested in the immature stages of the genus. His detailed accounts of egg placement and larval development reveal his passion for these fascinating moths. High species diversity, narrow geographic ranges, and specialized habitats suggest the genus is speciating rapidly. Schinia caterpillars can usually be recognized by their minutely spined integument, and their association with plant reproductive tissues. The prothoracic plate is often shiny and more pigmented than adjacent portions of the thorax. Seedfeeding Schinia are macrocephalic, i.e., their head—being specialized for crushing developing seeds—appears so large as to be out of proportion with the rest of the body. Many are broadest through the thoracic segments and taper rearward. A fair proportion, especially among the composite feeders that feed within flowers, bear thin dark stripes. The relative sizes of the pinacula and (integumental) spinules provide important clues to a caterpillar’s identity. Neither Crumb (1956) nor Hardwick (1996) offered characters to distinguish members of the genus from other heliothine genera. Most of our eastern species are cryptically colored, although the Clouded Crimson (Schinia gaurae) is boldly marked and seemingly aposematic; elsewhere the genus includes several gaudy species. Coloration may be highly variable—be cautious when making larval identifications. Larvae of more than 65 North American species are figured in Hardwick (1996). We illustrate just 17 eastern species, although the brief notes on hosts and life histories given in the table on pages 361–3 will aid considerably in the identification of all eastern species. Most members of the genus are specialized in diet and some extremely so. Their evolutionary radiation has penetrated into a diverse array of plant taxa that includes members of more than a dozen plant families. Composites host some 80% of the North American fauna; other important families include the evening primrose (Onagraceae) and legume (Fabaceae) families. Schinia frequently exhibit a tight association with their foodplant; not only is the host used by the caterpillars, but for many Flower Moths the host is the only plant at which the adults are likely to nectar. Adults of many Schinia preferentially roost on or in the flowers of their host. Some have a bladelike ovipositor that they use to insert their eggs into flowers and developing fruits. Early instars are usually concealed feeders, feeding within a flower or fruit. In many, only the penultimate

or ultimate instar will be found exposed, perched on the inflorescence or nearby stem. A notable exception is S. snowi, an onion-feeding species whose larvae tunnel into, and feed on, leaves and bulbs. Schinia caterpillars are often cryptic in habit, coring out nests within the host flower, tunneling into fruits, crawling downward to hide under leaves and other debris below the host, burrowing underground by day, or in other ways finding means to conceal their whereabouts. Development is fast: the egg stage is as short as two to three days in desert species, while the larval stage rarely lasts more than three weeks in warm weather. In nature, pupation occurs belowground. Held females oviposit readily if housed with flowers of an appropriate host. While many are easy to rear, others seem to languish in captivity—keep the caterpillars supplied with fresh flowers or seeds of various ages and use ventilated rearing containers. Hardwick (1996) found that most could be lab-reared on green beans. A reliable means by which to locate Schinia caterpillars is to revisit sites (and hosts) where adults were seen nectaring two to three weeks earlier. Late instars can be collected by carefully inspecting inflorescences. Watch for abdomens protruding from an inflorescence, feculae, or evidence of bored seeds or flowers; the last includes bulging disk flowers, ray flowers silked over the disk, or the fluff of damaged seeds working free from an “occupied” flower. Damage is characteristic and conspicuous once recognized. Nighttime searches may be necessary to locate caterpillars of those species that seek shelter under leaves or debris of the host, or that burrow underground by day. If the host is tall enough to allow it, beating works for collecting Schinia that perch on stems or feed from the outside of the host fruit or flower, but is largely ineffectual for “internally feeding” early instars and those that form nests within the flower. One option, especially where the hostplant is abundant, is to pick inflorescences two to four weeks after adults were active, hold the flowers in plastic bags, and then examine the bags daily for caterpillar feculae and damaged flower heads—we have collected Northern Flower Moth (Schinia septentrionalis) caterpillars in this way.1 Be forewarned that there are many lepidopteran flower and seed predators. Two microlepidopteran families, in particular, share Schinia’s predilection for reproductive tissues: the Pyralidae (especially members of the subfamily Phycitinae) and the Tortricidae (especially members of the tribe Cochylini). From our perspective, these other taxa will enrich your efforts—rear and photograph these too. Some flower moths are cannibalistic, even in early instars—important collections should be reared individually. Hold Schinia pupae over a lightly moistened mixture of sand and peat, varying the proportion of sand and moisture to reflect the substrate where the caterpillars were found. Expect some to overwinter more than a single winter as a pupa. Consult Hardwick (1996) for additional rearing suggestions. 1 If either the host or moth is deemed to be rare or even uncommon locally, we strongly discourage any collecting that could prove detrimental to the plant or insect.

352 Sun Moths Subfamily Heliothinae

Impatient Flower Moth Schinia petulans

Georgia south to central Florida west to Oklahoma and Texas. One generation with mature caterpillars from October to November. Very local. Scrubland goldenaster (Chrysopsis subulata) (Jeff Slotten, pers. comm.).

Bleeding Flower Moth Schinia sanguinea

Core range North Dakota, Minnesota, and Ontario south to Florida and Texas. One generation with mature caterpillars in September and October over much of range, but into November and December southward. Locally common. Blazing star (Liatris); our records from L. cylindracea, L. punctata, L. scariosa, L. spicata (Hardwick 1996; Chuck Harp and Ron Panzer, pers. comm.).

Thoreau’s Flower Moth Schinia thoreaui

Southern Canada to Massachusetts, south to Florida and Texas, but scarce and local in Northeast. One generation with mature caterpillars from August and September (northward) into October and November (southward). Common through Midwest. Giant Ragweed (Ambrosia trifida). See also page 357.

Sun Moths Subfamily Heliothinae 353

Arcigera Flower Moth Schinia arcigera ! recognition Brown with numerous subtle stripes and abundant integumental hairs and spinules. According to Hardwick (1996), these can be “so long and dense in some that larva” appears to be “covered with fur.” Ground color varies from brown to almost pinkish green (Fig. X9 in Hardwick 1996). Shiny prothoracic shield with narrow white middorsal stripe and prominent subdorsal stripe bounded on both sides by black patch. Broad white spiracular stripe divided by reddish-brown stripe. Middorsum with dark stripe that may bear trace of fine white line. Pinacula shiny, black, small, and somewhat raised. Head large with dark bar to either side of midline. Larva to 2 cm. ! occurrence Fields, waste lots, prairies, woodland edges from Manitoba to Nova Scotia, south to central Florida and Texas. Evidently one protracted generation with mature caterpillars in late summer and autumn. Common over much of East. ! common foodplants Aster, camphorweed, horseweed (Conyza canadensis), sea lavender or lavender thrift (Limonium carolinianum), and spotted knapweed. ! remarks Adults are on the wing from July into September in the Northeast, and early August into October in southern New Jersey. The flight is so drawn out that Forbes (1954) suggested there might be two annual generations. The Arcigera Flower Moth is the only Schinia known to us that feeds on members of two plant families. In Cumberland County, New Jersey, females have been observed ovipositing in horseweed, two asters (A. linariifolius, A. lateriflorus), and spotted knapweed (Centaurea maculosa); however, no attempt was made to rear larvae from the last host. DFS also reared adults from two last instars collected from flowers of sea lavender (Limonium carolinianum). Additionally, we have collected Arcigera Flower Moth caterpillars from camphorweed (Heterotheca subaxillaris). The caterpillar feeds on developing seeds, ignoring the pappi (seed plumes) which may expand and collect over the larva, concealing exposed portions of its body. About a third of reared pupae of the Arcigera Flower Moth overwinter two or three times before eclosing.

354 Sun Moths Subfamily Heliothinae

Evening Primrose Flower Moth Schinia florida

[Primrose Moth]

! recognition Pale green, elongate caterpillar, tapering rearward with numerous, minute spine-bearing spots (visible with lens); we have not encountered brownish-red forms as mentioned by Crumb (1956) and Hardwick (1996). Deep green middorsal stripe. Upper portion of subdorsal stripe creamy; lower portion wine to maroon, with red more pronounced anteriorly. Broad, creamy spiracular stripe contains tan-white spiracles along its upper edge; spiracles rimmed with brown. Stripes indistinct rearward, especially over A8–A10. Prothoracic shield mostly wine colored. Head pale brown, green, or yellow, and unmarked. Crochets more or less of two lengths (Crumb 1956). Larva to 3.5 cm. ! occurrence Fields, roadsides, waste places, coastal communities, grasslands, prairies, and other open habitats from Alberta to Nova Scotia, south in East to northern Florida and Texas. One generation with mature caterpillars when evening primrose is in fruit, mostly August to October in North. Locally common. ! common foodplants Biennial gaura and evening primrose. ! remarks While adults come to light, the most reliable means to locate the exquisitely rendered adults is to search evening primrose blooms by day. The adult roosts with the head pushed into a partially closed flower. The larvae feed on flowers (inset) and especially green seed pods. The feeding damage on pods is distinctive—the larva often begins tunneling at the end of the fruit and bores its way toward the peduncle, hollowing out the pod (of its seeds) as it goes. The caterpillar is surprisingly well camouflaged: its shape, coloration, and behavior all help it escape attention of the casual observer.

Sun Moths Subfamily Heliothinae 355

Clouded Crimson Schinia gaurae ! recognition Attractive yellow to orange, white, and black-ringed caterpillar with orange head. Bright lemon-yellow to orange coloration over dorsum and along sides of thorax and abdomen. Each segment with short black bar toward front and larger black ring that extends below spiracle at midsegment. Larva to 3 cm. ! occurrence Fields, roadsides, prairies, and other open habitats from Nebraska to Ohio, south to Florida and Texas. One generation, at least in northern portion of range, with mature caterpillars when gaura is in fruit (mostly September and October in North). Apparently two generations in Missouri, with second being much more numerous. Locally common. ! common foodplants Gaura. ! remarks In contrast to the Clouded Crimson, most of our Schinia caterpillars are cryptically colored, rendered in greens and browns. Hardwick (1996) pointed out that the larva is patterned much like the caterpillar of a Monarch (Danaus plexippus), and wondered if it might be a Batesian mimic of the milkweed-feeding butterfly. Alternatively, the caterpillar’s mimicry might be of a general nature, i.e., not linked to any single (model) species. Similar bright orange, red, black, and white ringing occurs in forester moths, the Convict Caterpillar (Xanthopastis regnatrix), and in dozens of unrelated lineages in the Neotropics. Miller et al. (2006) argue that such bold patterning taps into the innate fear that many birds have of brightly ringed cylinders (Smith 1977)—in essence, protection occurs because of an innate fear of coral and other poisonous snakes common to many avian predators. To find caterpillars of the Clouded Crimson, search inflorescences that are going to seed.

356 Sun Moths Subfamily Heliothinae

Goldenrod Flower Moth Schinia nundina ! recognition Highly variable in coloration, ranging from bright green to lemon yellow with dark subdorsal patches that form incomplete chevrons over dorsum. Green forms commonly with pale, broken subdorsal stripe (see Hardwick 1996). Middorsal line faint. Integument with minute black plates (spinules absent); these slightly raised over dorsum of thorax. Prothoracic shield with yellow to white middorsal and subdorsal lines; latter capping irregular black spot. Anterior margin of shield blackened between middorsal and subdorsal stripes. Pinacula smoky to black, shiny. Spiracles tan with dark brown rim. Larva under 2 cm. ! occurrence Fields, meadows, woodland edges, and other open habitats from Minnesota to southern Ontario and Nova Scotia, south to central Florida and southern Texas. One generation with mature caterpillars in late summer and fall. Common northward. ! common foodplants Goldenrod. ! remarks The caterpillar’s bright yellow ground color would be a dead giveaway on most plants, but is perfectly suited for a life among goldenrod flowers. Look for partly curled larvae with their heads buried deep into the flowers. Not surprisingly, a beating sheet can be used to great advantage. Feeding larvae leave piles of petals and silky pappi over the bottom of their rearing containers—like other Schinia, seeds and lower portions of the receptacle have primacy in the diet. Feeding occurs mostly at night.

Sun Moths Subfamily Heliothinae 357

Ragweed Flower Moth Schinia rivulosa ! recognition Usually green but occasionally yellow, brown, gray, or nearly

black, with dark subdorsal spots and white to yellow-green spiracular stripe. Conspicuous dark patches run obliquely back from addorsal area to spiracular stripe. Head and T1 broadest, tapering to rear. Addorsal and subdorsal stripes present as faint traces, often at anterior and posterior ends of each segment, if present at all. Thoracic shield, sometimes with black or cherry addorsal patch and pale middorsal line, dotted with minute pale spots. Setae white, stout, and conspicuous. Integument salted with numerous, minute white spines. Head green with numerous snowflake spots above and mostly unmarked below. Spiracles tan to white with dark rim. Larva to 2.5 cm. Caterpillar of Thoreau’s Flower Moth (S. thoreaui) similar (page 352). It is associated with great ragweed (Ambrosia trifida). It has a long flight season with adult records spanning the period from July through October. According to Crumb (1956), the white dorsal setae of S. thoreaui are sharper at their apex and the integumental granules are often embedded in pale spots; it also averages a bit larger—mature larvae may reach lengths of 3.5 cm. ! occurrence Fields, waste lots, sand plains, coastal communities, and other open mostly dry habitats from Nebraska, Quebec, and central New England, south to central Florida and southern Texas. One generation with mature caterpillars from August to November. Common. ! common foodplants Ragweeds (Ambrosia), especially annual ragweed (A. artemisiifolia). ! remarks We find larvae of the Ragweed Flower Moth in nearly every patch of ragweed that we search in late August and September. The larva will often be found feeding on staminate (male) flowers. In the last instar, the macrocephalic (big-headed) caterpillars consume many seeds (Hardwick 1996). The caterpillars readily tumble onto a beating sheet where they will at first curl themselves up into a ball.

358 Sun Moths Subfamily Heliothinae

Three-lined Flower Moth Schinia trifascia ! recognition Smooth, pale, green to (more commonly) tan, with numerous stripes; body narrowing rearward from thorax. Darkened green or brown middorsal stripe, and broad, smoky-green to brown supraspiracular stripe separated by cream to tan subdorsal stripe. Spiracular stripe pale green to nearly white. Pinacula often blackened, especially those in supraspiracular stripe. Integument with abundant fine brown spinules (visible with lens). Head with faint spotting and bar over each vertex to either side of midline. Larva to 2.5 cm. ! occurrence Wetlands, grasslands, and prairies, wet meadows to dry fields, edges of watercourses, openings in woodlands and forests from southern Ontario to Massachusetts southward into central Florida and Texas. One generation with mature caterpillars from August to November. Relatively common (seemingly more so as a larva). ! common foodplants Thoroughwort (Eupatorium); e.g., joe pye weeds, or trumpetweed (E. fistulosum) in many mesic habitats and northern locales. We found it abundant on tall thoroughwort (E. altissimum) in Illinois; Wyatt (1938) listed several other white-flowered bonesets; yankeeweed (E. compositifolium) reported in central Florida (Hardwick 1996). ! remarks The caterpillar intercalates itself among flowers. Given the density of the floral heads, the caterpillar can be completely hidden from view. It is quick to drop to the ground if disturbed; beating while joe pye weeds or bonesets are in flower will often yield larvae. Despite their stocky appearance, they are rapid climbers. Prepupal larvae take on a slight pinkish cast. Ben Williams has found the caterpillars to be common on his property in Pomfret, Connecticut, during the latter half of September, yet adults of the Three-lined Flower Moth are, at best, infrequent at his light. Occasionally, adults will be seen nectaring at flowers during the day.

Sun Moths Subfamily Heliothinae 359 Fuller’s Flower Moth Schinia fulleri

Central Florida. One generation with mature caterpillars in October and November. Local and uncommon. Yellow buttons or coastal plain honeycombhead (Balduina angustifolia).

Iva Flower Moth

Schinia gracilenta [Slender Flower Moth] Nebraska to southeastern Massachusetts, south to Florida and Texas. This range may include two species. If so, we illustrate the one commonly feeding on Iva frutescens in salt marshes from southeastern Massachusetts to at least North Carolina. One generation with mature caterpillars from September to November (southward). Locally common. Sumpweed or marsh-elder (Iva).

False Boneset Flower Moth Schinia grandimedia

Nebraska to Ohio, south to Alabama and central Texas, and westward. One generation with mature caterpillars in September and October. Locally common. False boneset (Brickellia eupatorioides; formerly Kuhnia eupatorioides).

Amorpha Flower Moth

Schinia lucens [Leadplant Moth and False Indigo Flower Moth] Great Plains from southern Canada to Texas, and along Atlantic Coastal Plain from North Carolina, south and west. One generation with mature caterpillars in July in North Carolina. Locally common. Principally desert false indigo (Amorpha fruticosa), leadplant (A. canescens), but also Psoralea and possibly clover and other legumes.

360 Sun Moths Subfamily Heliothinae

Lynx Flower Moth Schinia lynx

Wisconsin, southern Ontario and Quebec, and Maine, south to Florida and Texas. One generation with mature caterpillars mostly in August to September northward. Locally common. Camphorweed (Heterotheca subaxillaris) in Texas (Hardwick 1996); our wild larvae from horseweed (Conyza canadensis).

Matutinal Flower Moth Schinia mitis

Oklahoma to North Carolina, south to central Florida and eastern Texas. One principal generation with mature caterpillars usually in spring; mostly April and May in Florida, June in North Carolina. Locally common (diurnally). Desert-chicory (Pyrrhopappus).

Camphorweed Flower Moth Schinia nubila

Oklahoma to Long Island, south to Florida and Texas; range expanding in Northeast (first record for New Jersey in 1992, now common). One generation with mature caterpillars from September to November (southward). Locally common. Camphorweed (Heterotheca subaxillaris). Caterpillar figured here did not yield an adult.

Brown Flower Moth Schinia saturata

South Dakota to at least Long Island, south to Florida and Texas. One generation with mature caterpillars from September to November (southward). Locally common. Camphorweed (Heterotheca subaxillaris) and silkgrass (Pityopsis).

Sun Moths Subfamily Heliothinae 361

S CHINIA life histories Adapted largely from Hardwick (1996). Metzler et al. (2005) was also consulted. In addition to cited references, the table includes unpublished information of our own, as well as that provided by Chuck Harp (CH), John Richard Heitzman (JRH), Mark Mello (MM), Ron Panzer (RP), Michael Pogue

(MP), and Jeff Slotten (JS). Chuck Harp and Michael Pogue added several species based on their unpublished distributional records for the genus. Ranges and biologies for seven rare or localized members of the genus (S. arefacta, S. carolinensis, S. indiana, S. petulans, S. rufipenna, S. scissa, and S. septentrionalis) are summarized in Schweitzer et al. (2011).

SPECIES

FOODPLANTS

accessa alencis

Unknown Suspected to be Heterotheca canescens (Pogue and Harp 2005) Aphanostephus ramosissimus Aster: A. laevis, A. multiflorus, A. puniceus, etc.; Centaurea maculosa, Conyza canadensis, Heterotheca subaxillaris, Limonium carolinianum, and Machaeranthera tenuis Chrysopsis subulata (JS)

antonio arcigera

arefacta

avemensis bicuspida

Helianthus petiolaris Isocoma drummondii and Machaeranthera annua

bimatris biforma

Unknown Amblyolepis setigera

bina

Amblyolepis setigera, Crepis pulchra, Gaillardia pulchella, Hieracium gronovii (JS), Verbesina encelioides, and likely other composites Carphephorus corymbosus, Garberia fruticosa, and G. heterophylla Unknown Heterotheca villosa var. nana (Pogue and Harp 2005) Amphiachyris dracunculoides Gutierrezia sarothrae Croton californicus, C. dioicus, and C. texensis (CH) Grindelia nuda var. aphanactis (CH) Unknown Artemisia frigida Calylophus berlandieri and C. hartwegii (Pogue and Harp 2003b) Rayjacksonia annua

carmosina carolinensis chryselloides chrysellus ciliata citrinellus coercita crenilinea cumatilis cupes diffusa

errans florida

Machaeranthera tanacetifolia (CH) Oenothera, especially O. biennis; also O. nuttallii, O. parviflora, and O. rhombipetala (CH); occasionally Gaura according to Wyatt (1938)

COMMENTS

Perches on stems below flowers; tunnels down into disk flowers to feed.

Perches on stems below flowers and tunnels down into disk flowers. Occurs alongside Schinia petulans, often on the same plant (JS). Bores in heads. Early instars feed in heads; third to last instar seed predators, tunneling into heads through receptacle; last instar feeds on browned flowers. Forms nests in flowers; these may include ray flowers in last two instars. Perches on stems or disk; tunnels down from above to feed on florets (mostly corollas).

Bores into receptacle through fourth instar; last instar shelters belowground by day. Females have been observed ovipositing in buds and open flowers (Pogue and Harp 2005).

Feeds on buds and blossoms, but often resting among flowers; also consumes foliage (Hardwick 1996).

Feeds in flowers, fashioning a loose silken nest; heads with larvae have severed flowers bulging upward from disk; last instars in browned flowers. Seed predator; tunnels deep into young fruits.

362 Sun Moths Subfamily Heliothinae SPECIES

FOODPLANTS

COMMENTS

fulleri

Balduina angustifolia

gaurae

Gaura biennis, G. coccinea, G. filipes, G. mollis, and others; Stenosiphon linifolius Iva annua and I. frutescens Brickellia (Kuhnia) eupatorioides

Larva cores out disk florets; concealed feeder, occupied flowers with little evidence of larval presence. Feeds on developing seeds; rests on or among flowers.

gracilenta grandimedia hanga hulstia indiana jaguarina lucens

lynx

maculata meadi meskeana mitis

Heterotheca subaxillaris Artemisia ludoviciana (CH) Phlox pilosa Psoralidium tenuiflorum and Pediomelum rhombifolium and possibly other Fabaceae Amorpha, especially A. canescens; Dalea canescens (JRH); possibly also Psoralea, Trifolium, and other legumes Conyza canadensis (DFS) and Heterotheca subaxillaris; Erigeron (Forbes 1954) (although this record may refer to obscurata) Unknown

oleagina

Psoralidium lanceolatum (CH) Unknown Pyrrhopappus carolinianus, P. pauciflorus (= P. geiseri and P. multicaulis), and possibly P. grandiflorus Grindelia lanceolata (and others) Heterotheca subaxillaris Solidago canadensis, S. rigida (CH), S. stricta, and others Erigeron philadelphicus, E. annuus, and likely other fleabanes Unknown

olivacea

Hermannia texana

parmeliana

Adults perch on Pityopsis graminifolia (JS), which by fiat becomes the suspected host Chrysopsis subulata (JS)

mortua nubila nundina obscurata

petulans psamathea regia regina

reniformis rivulosa roseitincta rufipenna

Unknown; possibly woody goldenrod (Chrysoma pauciflosculosa) (Pogue 2010) Vernonia texana Palafoxia sphacelata

Artemisia filifolia (CH) Ambrosia, including A. trifida and A. artemisiifolia (CH) unknown Pityopsis graminifolia

Perches and feeds externally on flower heads. Seed predator; perches and rests in flower heads; cryptic and easily overlooked.

Feeds on developing seeds. Perches on stem and floral rachis; cores out contents of pods. Adults sit on Amorpha and other host flowers.

External flower feeder, tunneling into heads.

S. maculata = S. blanca of Hardwick (1996) (see Pogue 2006).

Seed predator; tunnels into head from side (through receptacle) or from above; often on browned, dry heads. Adult flies in early morning (CH). Tunnels into flower heads. Perches and rests in flower heads; cryptic and easily overlooked. Tunnels into flower heads (DLW) early in season. Life history information in Hardwick (1996) applies to Schinia sara (MP). Feeds on flowers and developing seeds, other parts of fruit (seed pod), and even leaves (captive larvae).

Seed predator: tunnels into seed heads, including ones that have started to brown.

Feeds deep in flowers, boring down to feed on achenes. Presence of early instars in flowers evident because severed florets push outward from receptacle; last instars feed externally (at night) (Pogue and Harp 2003a). Perches and feeds on flower heads; seed predator in last instars. Initially concealed within flowers; last instar rests along peduncles/pedicels/stems; bores into flowers to feed.

Sun Moths Subfamily Heliothinae 363 SPECIES

FOODPLANTS

COMMENTS

sanguinea

Liatris, including L. cylindracea, L. punctata, L. scariosa, and L. spicata (CH, RP) Heterotheca subaxillaris and Pityopsis graminifolia Polygonella is suspected (JRH) Polygonella gracilis (JS)

Early instars among florets; last instar shelters in cell belowground by day.

saturata scissa scissoides septentrionalis

simplex siren snowi sordida spinosae

subspinosae tertia sensu Forbes (1954) thoreaui

Aster; mostly A. linariifolius and A. spectabilis in Northeast; A. oolentangiense (= A. azureus) in Midwestern prairies; also A. puniceus (CH), A. concolor (MM), and others Ipomoea leptophylla (CH) Heterotheca subaxillaris Allium, including A. canadense var. mobilense Evidently tunnels into leaf and from there bores into bulb (in late spring and early summer). Haplopappus divaricatus and Pityopsis pinifolia External flower feeder. Polygonella (P. articulata over much of Most colonies along coast, especially beaches, sand plains, northern range) powerlines, and gravel pits; larvae possibly belowground during day. Polygonella gracilis and other Polygonella (JS) Unknown Ambrosia trifida

trifascia

Eupatorium: many species—see page 358

tuberculum ultima

Pityopsis falcata and P. graminifolia Mainly Chrysopsis pilosa in TX, OK, KS, and southeast CO; may use Heterotheca subaxillaris where both hosts grow together (CH) Unknown Gaillardia pulchella

varix volupia

Inland sand areas; larvae possibly belowground during day. Perches on stems and flowers; seed predator; prefers mature (browned) flowers; damaged heads conspicuous. A detailed account for this species appears in Schweitzer et al. (2011).

Lantana Groundling

Diastema tigris [Lantana Moth] (Subfamily Condicinae) Mostly coastal and southern Florida and central and southern Texas; one historic record from Louisiana (Vernon Brou, in litt.). Mature caterpillars occur through growing season. Locally common. Lantana. The caterpillar is used as a biological control agent to suppress Lantana camara in Africa, Australia, and on islands worldwide, including Hawaii in the United States.

Feeds externally on flower heads; some larvae hide by day in dead leaves at base of plant. Perches and rests in flower heads; cryptic and easily overlooked. Rests along stems and peduncles; feeds on flowers.

Tunnels in floral heads; last two instars form nest in heads, sometimes folding over ray flowers and silking these over feeding shelter.

364 Groundlings and Kin Subfamily Condicinae

Groundlings and Kin

Family Noctuidae: Subfamily Condicinae The 49 described North American species fall into two tribes, the Condicini and Leuconyctini, each with five genera. More than half the species belong to the nominate genus, Condica (including many formerly classified in the genus Platysenta). The monophyly of the subfamily as proposed by Poole (1995) was not supported by Mitchell et al.’s study (2006). Poole’s (2009) Nearctica website (http://www.nearctica.com/) has accounts for all eastern species. In many condicines, the prolegs on both A3 and A4 are equally reduced but large enough to retain function; the dorsal setae are borne from white pinacula; the spinneret is reduced and scalelike (Crumb 1956); SD1 on A9 is hairlike; and the SV group on A1 is trisetose (Condica) or bisetose (Crambodes, Ogdoconta, and Leuconycta) (Lafontaine in litt.). In addition, the crochets are uniordinal. A8 is often humped, and the lateral stripe drops down along the anal proleg. Many can be recognized by their distinctive head markings (several have

the triangle outlined with white lines, and Condica may bear distinctive black bars across the head). At rest, their bodies are often flattened against the substrate, especially the thoracic segments, and the head is extended forward. Prepupae take on a reddish cast. Most feed on composites, including 10 of the 12 illustrated in this guide (all but the two Homophoberia). According to Holloway (1989), hosts with pyrrolizidine alkaloids are especially likely to serve as foodplants. Many, and Condica in particular, are migratory and regularly show up north of where their pupae are capable of overwintering successfully. The species epithet given to the Mobile Groundling (C. mobilis) is indicative of its distribution, which ranges south to Argentina and west to Arizona, and includes many small, remote islands in the Caribbean. Members of the genus Condica are frequently attacked by braconid wasps—parasitism rates exceeding 50% are common in collections of the Boneset Groundling (C. albigera). Plan for such when your intent is to rear out adults from larval collections.

Mobile Groundling Condica mobilis

Resident of Neotropics and Gulf states, often moving north as far as Missouri and New Jersey during summer and fall months. Mature caterpillars occur through most of growing season and year-round in southern Florida and Texas. Increasingly common southward. Spanish needles (Bidens).

Confederate Groundling

Condica confederata [The Confederate] Breeding range unclarified, perhaps Arkansas to Virginia southward into Neotropics, autumn migrants frequent north to Long Island. Mature caterpillars occur through growing season in Deep South. Common. Climbing hempvine (Mikania). Co-occurring with Splotched Groundling (C. cupentia), which has similar caterpillar. In our limited experience, its caterpillar tends to have less pronounced (often smoky) cranial band and setal bases in the triangle are frequently blackened.

Groundlings and Kin Subfamily Condicinae 365

Common Pinkband Groundling Ogdoconta cinereola

[Common Pinkband]

! recognition Pale green caterpillar with well-defined middorsal and subdorsal white stripes that continue onto anal plate. Upper portion of body with abundant creamy to frosty white suffusion, especially between subdorsal stripes. Setae arise from conspicuous white bases (although these sometimes obscured by white “frosting” over dorsum). Yellowish spiracles bounded by thin white stripe, both above and below, that continues down anal proleg. Head shiny with middorsal white stripe that splits and runs down either side of triangle and thick subdorsal line that ends above eyes; spiracular stripes fusing, continuing through eyes, nearly reaching antenna. Labrum white. Proleg on A3 reduced, that on A4 somewhat smaller than those that follow; anal prolegs comparatively long and directed behind body. Larva to 2.5 cm. Middle instars shiny emerald green with conspicuous white stripes; thin spiracular stripes fused into single stripe, somewhat flattened; and reduction of first two pairs of prolegs especially evident (inset). Middle instars easily confused with caterpillars of Cobbler Groundling (Condica sutor) (page 367). ! occurrence Fields, meadows, roadsides, weedy lots, and other open sites from southern Canada to Florida and Texas. Multiple broods with mature caterpillars from June until first frosts. Common. ! common foodplants Our wild larvae from burdock, crownbeard or wingstem (Verbesina), parthenium, various ragweeds, sunflower, and Viguiera (all composites); also reported from asparagus and beans, which we regard as erroneous. ! remarks We have found caterpillars of the Common Pinkband Groundling on numerous occasions while beating ragweed. The insect often occurs in early successional (e.g., beaches and sandbars) and disturbed areas (e.g., about roadsides, suburban parking lots, and abandoned building lots). Caterpillars feed from the underside of leaves. The pupa overwinters.

366 Groundlings and Kin Subfamily Condicinae

Boneset Groundling Condica albigera ! recognition Green or brown with faint dorsal chevrons and abruptly humped A8, that drops steeply to anal proleg. T3 and especially A1 greatly swollen, at least in resting larvae. Middorsal stripe often broken over A1–A3, best developed over anterior half of these segments, ending over hump on A8. Many forms with dark addorsal patch on A1 and A2 (inset). Body densely speckled with minute pale spots, especially laterally and below. Obscure, ill-defined, oblique subdorsal patches form herringbone pattern over dorsum. White spiracular stripe thickens rearward, forking on A8 with upper portion running up over hump and lower portion continuing down anal proleg. Head green or brown, variously mottled, but always with diffuse whitish edging along sides of triangle. Mature larva to 3 cm. Early instars smoky to gray-green in appearance (not green), and T3–A2 conspicuously swollen. ! occurrence Meadows, fields, roadsides, and other open habitats across southern Texas. Multiple generations with mature caterpillars year-round in Rio Grande Valley. Very common to abundant through south Texas. ! common foodplants Blue boneset (Tamaulipa azurea) and related eupatoriums. ! remarks The false head is the most credible of the Condica species that we treat: the posterior face of the hump is drawn into the abdomen proper, and the subdorsal cavities that form under the abdominal ridge presumably serve as the ruse’s “eyes.” Our larvae ignored flowers, preferring mature leaves. Late instars rest along upper stems with the head and thorax flattened against their perch—they are easily spotted once a search image is formed. Other individuals will sit on or wedge themselves among clusters of florets. Early instars spin down on belay lines but older instars stay attached to the host when disturbed. The Boneset Groundling is heavily parasitized by a gregarious microgastrine braconid wasp that spins chocolate- to gray-colored cocoons in the vicinity of the host larva, often in two discrete piles to either side of the victim (see page 29, row four). Prepupal larvae turn maroon.

Groundlings and Kin Subfamily Condicinae 367

Cobbler Groundling Condica sutor [The Cobbler] ! recognition Bright green to reddish green and maroon, speckled with minute creamy spots. A8 humped with posterior face branded with circular to teardrop-shaped, middorsal white spot. Broken, white middorsal stripe enlarged towards front of each abdominal segment. White, conspicuous pinacula. Thin, crooked subdorsal stripe, often dipping downward toward rear of given segment. Darkened prothoracic shield cut by white middorsal and subdorsal stripes. Ground color usually appreciably darkened above thin white spiracular stripe; in crowded, captive-bred specimens, upper half of body mottled with maroon patches. Spiracles white with black rim, often capped by wine-red patch. Head with thick brown line over each lobe that narrows abruptly above antenna (below right). In well-pigmented forms middle segments of thoracic legs blackened. Larva to 3.5 cm. Middle and penultimate instars smoky green above and pale green below the spiracular stripe, striping more pronounced, and anal plate cut by three white stripes (above lower right) (sometimes confused with caterpillars of the Common Pinkband Groundling, Ogdoconta cinereola, page 365). ! occurrence Fields and other early successional habitats. Resident in South; regular fall migrant northward at least to Ohio and Long Island, less commonly straying into southern Canada. Single fall generation in Northeast with caterpillars from late August to October, but usually arriving too late to complete development; active throughout growing season in South. Very common southward, and occasionally so northward in autumn. ! common foodplants Reported as pest on celery in Florida (Crumb 1956). Marigolds and wedelia (Covell 2005). We have found larvae on false ragweed or parthenium, ragweed, sunflower; our ex ova larvae accepted aster, goldenrod, and sneezeweed (all composites). ! remarks We do not know the degree to which Condica can endure prolonged freezing temperatures. In most of our area, the Cobbler Groundling must reestablish its populations each summer—the first individuals usually appear across the Northeast and upper Midwest sometime in September or October, but occasionally some adults arrive as early as July. Although larvae attain normal size if fed solely on leaves, they will avidly consume aster petals if offered. At rest, the head is extended forward and pressed downward; the caudal segments are drawn inward, enhancing the degree to which the swollen eighth abdominal segment resembles a head.

368 Groundlings and Kin Subfamily Condicinae

Dusky Groundling Condica vecors ! recognition Yellow-green with white to yellow spiracular stripe edged with red above. A8 distinctly raised and angulate with triangular, white middorsal patch, and roughly oval subdorsal rump patch; dorsum of A9 and A10 often reddened (lower inset). Hair-thin, white middorsal stripe edged with red, especially over thorax and anterior abdominal segments; vague, incomplete subdorsal stripe evident over thoracic segments. Scattered, minute, white and yellow spots over trunk. Setae borne from minute white bases; D1 pinacula on A1–A8 often with adjacent maroon spot. Spiracles small, black. Head green with broad black line that thickens rearward. Larva to 3.5 cm. Crumb (1956) provides a key to six members of the genus. The three most common eastern Condica may be distinguished by examining the markings on the head. In the Dusky Groundling the broad black stripes converge rearward; in the Cobbler Groundling (C. sutor) the stripes do not converge; and in the White-dotted Groundling (C. videns) there are three pairs of thin stripes, with no red along the upper margin of the spiracular stripe as in the Dusky Groundling. ! occurrence Meadows, fields, roadsides, yards, and other open habitats from southern Canada to Florida and Texas; more common southward. At least two generations over much of East, with mature caterpillars from June onward; more generations in South. Moderately common. ! common foodplants Ours from sweetscented joe pye weed (Eupatorium purpureum) and white snakeroot (Ageratina altissima); dogfennel (E. capillifolium) (Heppner 2007); lettuce in captivity; perhaps eating a variety of composites. ! remarks At rest, the eighth abdominal segment is bent sharply downward. Markings on the rump form a faux head, or at least a passable facsimile. The real head is only half its width and more modestly rendered. The caterpillars perch on the underside of low-growing vegetation late in the summer and fall. Unlike the previous species, the Dusky Groundling appears to be a resident species over much of our area.

Groundlings and Kin Subfamily Condicinae 369

White-dotted Groundling Condica videns ! recognition Yellow-green or reddish brown with strong lateral stripe edged above and below with white. A8 with oval, white middorsal spot. Thin, vague, white middorsal, subdorsal, and supraspiracular stripes. Pinacula small, whitish, conspicuous, especially over dorsum. White spiracular stripe infused with pink, extending from anal proleg to antenna, passing through lower half of spiracles; upper edge of stripe sometimes edged with wine red through thoracic segments and around spiracles. Creamy speckles, visible with lens, especially obvious below spiracular stripe. Head with prominent black line over each lobe, sandwiched between two whitish lines; black line not appreciably thicker rearward (as in related species), running more or less parallel to line over other lobe (below right). Larva to 3.5 cm. Middle instars with thin, white middorsal, subdorsal, and supraspiracular stripes bounded by dark green; thickened spiracular stripe creamy to white; and two dark green lines over top of head (above lower right). ! occurrence Fields, salt marshes, moist pinelands, powerline right-ofways, vacant lots, and other open, usually moist, habitats from southern Canada to Florida and Texas. Two or more generations over much of East with mature caterpillars from May onward; three or four generations in southern New Jersey and more in Deep South. ! common foodplants Aster, goldenrod, sneezeweed, tatalencho or gumhead (Gymnosperma glutinosum), and other composites. ! remarks Early instars loop when moving forward, reflecting the fact that the first two pairs of prolegs are reduced. Crumb (1956) notes that flowers are preferred. Our captive caterpillars supplied with aster flowers fed on petals (as well as leaves) but avoided disk flowers and seeds; those offered sneezeweed consumed leaf tissue and ignored flowers. Our larvae rejected clover, sorrel, and lettuce. The caterpillar rests extended along a leaf or the axis of a stem. The posture and shape are distinctive: the body is somewhat flattened and spindlelike, widest through the thoracic segments, then narrowing to the rear of the body, and the short anal prolegs are directed backward and held alongside one another. The pupa of the White-dotted Groundling overwinters at least as far north as New Jersey (where adults are common in April).

370 Groundlings and Kin Subfamily Condicinae

Smartweed Groundling Homophoberia apicosa [Black Wedge-spot]

! recognition Pale yellow-green, smooth, salted with minute yellowish spots.

Proleg on A3 reduced, about half size of those that follow. Middorsal, subdorsal, two wavy supraspiracular, and subspiracular stripes all faint. Development of spiracular stripe variable: sometimes represented by thin faint stripe, while in other individuals area between spiracular and subspiracular stripes filled with bright pink to wine red (upper inset). Spiracles white with dark outer rim, and often embedded in reddish spot, giving spiracular stripe what Forbes (1954) called a “beaded” appearance. Dorsal pinacula whitish or yellowish. Setae short, rusty (under magnification), especially those on head, T1, and A10. Thoracic and abdominal legs with pink to maroon cast. Head flattened, shiny green with black cheek stripe that reaches to long, red antenna; often with curving row of darkened spots over each lobe that drops down and joins cheek stripe and second row to either side of midline. Larva under 2.5 cm. ! occurrence Marshes, wetlands, fields, waste lots, and other open habitats from Minnesota to Quebec and Nova Scotia, south to Florida and Texas. At least two generations in Northeast with mature caterpillars from June onwards. Locally common. ! common foodplants Smartweed or knotweed (Polygonum). ! remarks Larvae of the Smartweed Groundling feed from the lower side of the blade, typically carving out holes from the center of the lamina. At rest, the thoracic segments are somewhat flattened and the head is extended forward. Adults come to both light and bait. Its congener, the Yellow Water-lily Groundling (H. cristata) [Waterlily Moth] is a specialist on pond-lily (Nuphar) (lower inset). It ranges from southern Canada to Massachusetts, south to Florida and Arkansas (Covell 2005). Prepupae of both of our Homophoberia redden somewhat prior to pupation. The winter is passed as a pupa.

Groundlings and Kin Subfamily Condicinae 371

Verbena Groundling Crambodes talidiformis [Verbena Moth]

! recognition Lime to smoky green with prominent spiracular stripe

infused with pinkish red (usually more so than in example shown). Upper edge of spiracular stripe running through middle of spiracle on T1, but extended upward to envelop spiracles on segments A1–A8; upper edge of stripe wavy on A7 and A8; rearward extending down outer face of anal proleg; lower portion of stripe straight, pure white. White (and sometimes red) middorsal stripe, weak, thin, broken; similarly, subdorsal and supraspiracular stripes faint, thin, often broken, composed of minute spots and lines. Ground color sometimes darkened between supraspiracular and spiracular stripes. Dorsal and lateral setae from contrasting white pinacula. Prolegs on A3 and A4 smaller than those that follow. Head with tracing of spiracular stripe reaching to antenna and vague white snowflake spots over each lobe (lower inset). Spiracles black. Larva to 3 cm. The caterpillars of the Nutmeg (Anarta trifolii) and leuconyctas are superficially similar. ! occurrence Fields, beaches, waste lots, sand plains, wetlands, and other open areas from Minnesota, southern Quebec, and Maine, to Georgia and Texas (and westward). Two generations over much of East with mature caterpillars in June and July, then again from August to October. Locally common. ! common foodplants Vervains (Verbena), but shunning some introduced species. ! remarks The larvae rest on the underside of leaves, extending along the midrib or lateral vein. The posture is distinctive: at rest, the rear of the caterpillar slopes back to the outstretched anal prolegs. Reds in the lateral stripe match those that develop along the petiole and stem of many vervains. Ours preferred leaf tissue over that of flowers. Caterpillars are easily sampled with a beating sheet. Like other members of the subfamily, prepupal larvae take on a reddish cast. The pupa overwinters.

372 Groundlings and Kin Subfamily Condicinae

Green Leuconycta Leuconycta diphteroides ! recognition Elongate, pale green to blue-green, with pale spiracular stripe whose lower boundary is blurred. Well-differentiated, white middorsal and subdorsal stripes. Thorax somewhat flattened (at rest). Spiracular stripe edged with rosy red above, especially on T1–T3 and A8, sometimes fused with thin subspiracular stripe that runs down anal prolegs. Dorsal setae from prominent white setal bases. Yellow to tan spiracles with narrow, outer black ring. Prolegs on A3 reduced in girth. Head also appearing somewhat flattened, shiny pale green and unmarked. Larva to 4 cm. ! occurrence Fields and meadows from Manitoba to Nova Scotia, south to Florida and northern Arkansas. Evidently with two rather poorly differentiated generations in Connecticut and over much of East with mature caterpillars from June onward, but with majority of caterpillars showing up in late summer. Common. ! common foodplants Fleabane, goldenrod, aster, and likely other composites. ! remarks A detailed larval description was published by Neil (1988). At rest, the thoracic segments, and especially the prothorax, are depressed; likewise, the rump is flattened, partly because the anal prolegs are extended backward from the rear of the body. Watch for larvae stretched out along a leaf or stem. Unlike many summer-active goldenrod feeders, the larvae prefer leaves over flowers, as might be surmised from their green coloration. Disturbed larvae drop to the ground and may thrash violently. Leuconycta caterpillars are common in sweep and beating samples from fields and other open habitats from July onward. The pupa overwinters.

Groundlings and Kin Subfamily Condicinae 373

Marbled-green Leuconycta Leuconycta lepidula ! recognition Yellow-green with prominent white, yellow, or occasionally orange spiracular stripe edged above with red. Midabdominal prolegs increasing slightly in size rearward. Spiracular stripe with red to maroon edging most conspicuous over thoracic segments and rearward, extending down anal proleg. Middorsal and subdorsal stripes thin, pale, fading over A9. Dorsal setae from small, pale pinacula. Thoracic legs sometimes blackened apically. Head green, shiny, unmarked. Larva under 3.5 cm. Ground color of Green Leuconycta (L. diphteroides) more blue-green or with bluish-white bloom. Dorsal setal bases of Marbled-green Leuconycta smaller and less conspicuous than those of Green Leuconycta. ! occurrence Fields, wet meadows, marshes, and mesic woodlands from Manitoba, Great Lakes states, and Nova Scotia, south to Georgia (mountains) and Missouri. One generation with mature caterpillars in July and August in Connecticut. Common over parts of Midwest, but often more local than Green Leuconycta. ! common foodplants Dandelion as laboratory host; Handfield (1999) also lists dock. Wild hosts not known. ! remarks Neil (1988) provides a detailed larval description. Rings et al. (1992) regarded the moth as abundant in Ohio. The Marbled-green Leuconycta is curiously scarce and local in Connecticut, and apparently absent in most of New Jersey. Adults come to both light and bait. The image in the inset is of an ex ova caterpillar reared in rather humid, cramped quarters, and thus may not reflect coloration of wild larvae. According to Tim McCabe, there may be additional, unrecognized Leuconycta species in the East. Until the taxonomy of the genus is clarified, we encourage those so inclined to photograph larvae and rear adults, save and voucher specimens, and take notes on larval biology.

374 Fern Moths Subfamily Eriopinae

Fern Moths

Family Noctuidae: Subfamily Eriopinae This small subfamily is represented in the New World by a single genus: Callopistria. All five Nearctic species occur in the East. The subventral group is bisetose on A1; SD1 on A9 is hairlike and issues from a hardened pocket; and paired flaps cover the apex of the dorsally grooved spinneret (Beck 1999, 2000). Callopistria caterpillars often have green, red, and cinnamon forms. All of our species are believed to feed on ferns. The pupa overwinters.

Florida Fern Moth Callopistria floridensis

Virginia south to Florida and Texas. In greenhouses north to Canada. Continuous generations in Florida. Very common. Ferns. Ground color green, brown, black, or reddish with thin, jagged, creamy addorsal and subdorsal stripes and prominent white to yellowish spiracular stripes. Occasionally with black, bat-shaped spots over dorsum and small, black, elongate spots subtending spiracles. Middle instar shown (above lower right).

Fern Moths Subfamily Eriopinae 375

Silver-spotted Fern Moth Callopistria cordata ! recognition Red or yellow-green larva with white stripes and oblique lines. Fine white addorsal stripe extends rearward from T2. Continuous, white subdorsal stripe of thoracic segments broken into shallowly oblique white lines on abdomen that get wider (and sometimes more lens-shaped) rearward; spot on A8 continued through A9 onto anal plate. Thin, white, broken spiracular line on T2 and T3, and set of oblique supraspiracular lines nearly fusing with corresponding subdorsal series. Several setae on T1 from shiny black pinacula. Head with two prominent black lines extending across lobe to antenna. Spiracles black. Larva under 2.5 cm. Middle instars lime green, shiny, with reduced markings on head (right inset). Caterpillar of Florida Fern Moth (C. floridensis) highly variable in coloration (previous page). We have images of one larva of C. granitosa found on bracken; the larva is light green with conspicuous longitudinal striping and black spots over the head. It flies in open coastal plain pinelands from New Jersey southward. ! occurrence Barrens, woodlands, forests from Wisconsin to Nova Scotia, south to Florida and Texas. Two generations with mature caterpillars from July to October, from Quebec south through New England. Locally common. ! common foodplants Ferns; all our larval records from bracken. Adult collections also consistent with bracken as principal host. ! remarks One wonders if the prominent silvery spots on the adult forewings serve as imitation water droplets, breaking up the outline of the moth and making it less recognizable as prey. While not restricted to them, this species seems more common in dry habitats than the Pink-shaded Fern Moth (C. mollissima). The cocoon is spun in leaf litter.

376 Fern Moths Subfamily Eriopinae

Pink-shaded Fern Moth Callopistria mollissima ! recognition Emerald green or reddish with whitish, oblique, lens-shaped

spots above each abdominal spiracle; these capped above by darkened, mirror image spot. Finer zigzagging (lightning bolt) lines above each spiracle. Pale halo ringing each black spiracle. Head with two pairs of black lines, one pair reaching to antennae and other to eyes. Larva less than 2.5 cm. ! occurrence Swamps, wetlands, mesic woodlands, and forests from Michigan to Nova Scotia, south to central Florida and Texas. Evidently two generations with mature caterpillars from June to October in Northeast; two or three generations in Missouri. Very common. ! common foodplants Ferns, such as hayscented and New York fern. ! remarks In captivity we commonly see cinnamon-red forms, but it is unclear if these represent an inducible phenotype caused by crowding, high humidity, or both. Also we do not know if such forms occur in nature. The degree to which the diets of Callopistria are specialized on different fern genera is unexplored: many ferns are unsuitable for larval development. Fern Moth caterpillars rest on the upper surface of fronds by day. They are easily collected with a beating sheet or by sweeping, and where common, by visual inspection.

Fern Moths Subfamily Eriopinae 377

Marsh Fern Moth Fagitana littera ! recognition Handsome, emerald to yellow-green to cinnamon, with thick subdorsal stripe and narrow, parallel-sided, white middorsal, supraspiracular, and spiracular stripes. Weak stripes, composed of minute spots, intercalated between these. Body mottled with minute white spots that gather into lines. Head with dark line to either side of triangle. Anal prolegs short. Larva to 3 cm. Middle instars (inset) bright green with stripes more prominent. Head yellow-green, mostly unmarked or with faint line above eyes, and second line to either side of triangle. ! occurrence Mostly acidic shrub swamps, fens, bogs, wet pine barrens, and other wetlands from Wisconsin to Nova Scotia, south to North Carolina—see Remarks. One generation in Ohio and Connecticut with mature caterpillars in summer; two generations in North Carolina. Local and uncommon. ! common foodplants Ferns. Marsh fern widely reported in literature (Forbes 1954, Rockburne and Lafontaine 1976), apparently based on single collection on Long Island by Roy Latham. Franclemont reared larvae on brackenfern. Our ex ova larvae accepted Virginia chainfern (Woodwardia virginica) and royal fern (Osmunda regalis). ! remarks This is a curiously scarce moth over much of its range, but it is locally common along the Atlantic Coastal Plain and through parts of the Great Lakes region. Caterpillars often rest on leaf undersides directly over the midrib with the stubby anal prolegs extended rearward and held together. Caterpillars from a single female were comprised of bright green and cinnamon-colored larvae with the proportion of the latter increasing in later instars. Perhaps in nature the green form is the norm and the reddish and cinnamon forms occur when larval densities are high, foliage quality is poor, or the caterpillars are experiencing other kinds of stress—analogous darker (crowding) forms occur in various other Lepidoptera. The phylogenetic origin of Fagitana is unclarified—we are even uncertain of its subfamily placement. We place it here with Callopistria, because it is a fern feeder, shares a similar green to red color polymorphism, and because both genera are small headed with a strong black line along each lobe. Lafontaine and Schmidt (2010) tentatively placed Fagitana in the Xylenini.

378 Marvels Subfamily Noctuinae: Tribe Pseudeustrotiini

Cutworms, Lash-eyed Sallows, Sallows, and Kin Family Noctuidae: Subfamily Noctuinae

This is the largest subfamily treated in this work with more than 1400 species occurring north of Mexico. The scope of the subfamily was greatly enlarged by Mitchell et al. (2006) whose molecular phylogeny suggested that a number of widely recognized subfamilies, e.g., the Hadeninae and Xyleninae, were phylogenetically subordinate within the Noctuinae. Lafontaine and Schmidt (2010) recognize 17 tribes for the North American fauna. The spinneret is dorsally grooved (right and page 14, Fig. 8); in those with a short spinneret, e.g., Euxoa, the spinneret is flattened with its upper side open and troughlike. A2 always bears two SV setae. The SD1 seta on A9 is hairlike (except in a number of borers). The larvae are extremely diverse in coloration, form, and habit; such being the case, we introduce each tribe separately.

Miranda Moth

Proxenus miranda (Tribe Caradrinini) Manitoba to Nova Scotia south through Carolinas and Texas. Two broods over much of East with mature caterpillars in June and July, then again in late summer. Locally common. Presumably polyphagous on forbs; Godfrey (1987) lists cantaloupe, strawberry, sugarbeet, and sweet potatoes. Rough skinned; crochets biordinal.

Marvels

Family Noctuidae: Subfamily Noctuinae: Tribe Pseudeustrotiini Only two species occur in our region; we include figures of both. Prolegs on A3 reduced in both species; sometimes those on A4 are also reduced in Pseudeustrotia.

Gray Marvel

Anterastria teratophora Manitoba to Quebec, south to Georgia and Arkansas, absent from much of Atlantic Coastal Plain. One generation with mature caterpillars from early to late summer. Locally common eastward, but widespread and sometimes abundant in Midwest. Mint and horsemint or bee balm (Monarda).

Marvels Subfamily Noctuinae: Tribe Pseudeustrotiini 379

Pink-barred Marvel Pseudeustrotia carneola [Pink-barred Midget]

! recognition Dusky maroon to pale green and brown-green with somewhat reduced prolegs on A3 and A4. Supraspiracular area and sometimes entire dorsum may be wine red. Weak middorsal line consisting of minute, scattered, white spots embedded in darker heart line. Pure white to mostly red spiracular stripe usually present (but sometimes green and obscure), running down outer side of anal proleg. Spiracles white with dark outer rim; midabdominal spiracles minute. Prothoracic shield sometimes with dark pigment to either side of midline and dark edging; more rarely, entire shield lightly darkened (melanized) (bottom right). Dorsal pinacula conspicuous, contrasting with ground color; other pinacula small, whitened. Head smooth, shiny yellow-green to suffused with red, especially above; some individuals bearing dark “freckles” over each lobe. Frons usually clear and unmarked. Larva to 2 cm. ! occurrence Fields, croplands, waste lots, wet meadows, and other open habitats from Manitoba to Newfoundland, south to Florida and Texas. Very common. ! common foodplants Dock and smartweed (Polygonum). ! remarks The thoracic segments are dorsoventrally compressed when the caterpillar is resting (as in Leuconycta and related genera). Initially, we had difficulty reconciling our caterpillars with the descriptions of Forbes (1954) and Crumb (1956), but now believe theirs and ours represent a single variable species. Of some 30 caterpillars collected by DLW in an abandoned sand quarry only one (mature larva) was appreciably reddened at the time of its collection; but over 80% of the larvae took on reddish coloration within a week. Part of this shift may be a natural consequence of maturation, although many of our larvae turned red well before they had finished feeding. Another possibility is that the red forms were induced, and that the transition had been triggered by crowding, high humidity, or other stimuli not present in the quarry. In addition to leaf tissues, our captive Pink-barred Marvel caterpillars fed on the bracts enclosing the seeds of smartweed.

380 Midgets Subfamily Noctuinae: Tribe Elaphriini

Midgets

Family Noctuidae: Subfamily Noctuinae: Tribe Elaphriini This small tribe includes four genera and 20 North American species (Lafontaine and Schmidt 2010). The head is small relative to the swollen thoracic segments with the disparity being especially marked in early instars. A8 is humped and in some resembling a (false) head. The head often bears white adfrontal lines. Many Elaphria, Gonodes, and perhaps others can be reared on the green algae (Protococcus) that grows on tree bark. Curiously, some Elaphria can be reared on the leaves of boxelder (Acer negundo) (Tim McCabe, unpubl. data; Wagner et al. 2008). Given their coloration and scarcity in our beating samples (except for Variegated Midget, Elaphria versicolor), we suspect that Elaphria caterpillars feed mainly at night, and along tree trunks, low vegetation, or the ground. The Grateful Midget (E. grata) and Sugarcane Midget (E. nucicolora) become increasingly common through the autumn months, which is indicative that they are migratory.

Northern Festive Midget

Elaphria alapallida [Pale-winged Midget] Southern Canada to mountains of North Carolina. One principal brood with mature caterpillars mostly in June and July. Two closely related species (E. cornutinus and E. festivoides) at least double brooded in Piedmont and southern Atlantic Coastal Plain, respectively. Common. Foodplants unclarified: Rockburne and Lafontaine (1976) list boxelder, but dearth of subsequent records from this host suggests other substrates normally used (see above). The name E. festivoides was, until recently, misapplied to this species (Pogue and Sullivan 2003).

Festive Midget Elaphria festivoides

Southern New Jersey to Florida and Texas. Two generations over much of range with mature caterpillars in summer and again in the fall. Common. Tim McCabe reared three cohorts on green algae and boxelder including the individual figured here. A southern species with a penchant for xeric sandy habitats.

Midgets Subfamily Noctuinae: Tribe Elaphriini 381 George’s Midget Elaphria georgei

Michigan to Nova Scotia, south to Florida and Texas. One generation with mature caterpillars in summer. Formerly scarce, but increasingly common northward to Pennsylvania and New Jersey. Tim McCabe reared cohort on green algae and fungi (Wagner et al. 2008).

Grateful Midget Elaphria grata

Wisconsin, southern Quebec, and Maine, south to Florida and Texas. Multiple generations with mature caterpillars from late May into October. Common. Foodplant unclarified; Tim McCabe reared cohort on green algae and fungi (Wagner et al. 2008).

Sugarcane Midget Elaphria nucicolora

Arkansas to Virginia (and New Jersey once), south through Gulf states. Multiple generations with mature caterpillars through growing season. Locally common southward. Pest of sugarcane; also bahia, rye, and other grasses (Kimball 1965, Lyle Buss, pers. comm.). The second larva is fully mature and nearing pupation.

382 Midgets Subfamily Noctuinae: Tribe Elaphriini

Variegated Midget Elaphria versicolor ! recognition Brown to charcoal, variable in pattern; body swollen about thorax, narrowing to A6, thickening again rearward. A8 swollen: raised and ridged rearward, with white supraspiracular patch. Often with irregular pale saddle that ends above spiracle on A4. Pale middorsal stripe sometimes with orange, especially over abdominal segments. Larva less than 2 cm. ! occurrence Woodlands and forests from Canada south to Florida and Texas. Two or three generations in Missouri with mature caterpillars from June to October. Common. ! common foodplants Many literature records for conifers: fir, hemlock, larch, pine, red cedar, spruce, white cedar, but also birch and other broadleaf plants; yet larvae often fail if fed only leaves from these plants (see Remarks). Other hosts given below. ! remarks Caterpillars rest on stems, away from foliage. We have beaten caterpillars from a variety of deciduous and coniferous trees and shrubs. Enigmatically, larvae almost always fail when offered foliage from the plants on which they were collected. In captivity we had a caterpillar graze extensively on the green bark of new black cherry shoots, yet the same larva showed no interest in cherry leaves. A second larva beaten from hemlock ate a pupa of the False Hemlock Looper (Nepytia canosaria) en route to DLW’s lab, before the two could be separated. Three of our caterpillars beaten from a hawthorn ignored foliage but eagerly devoured green (recently formed) stem galls. Later, the same larvae cored green grapes of Vitis labrusca that had been added to their rearing vial. Tim McCabe has reared several midgets (Elaphria) from egg to adult on Protococcus, a widespread and abundant epiphytic green alga; Protococcus may have primacy in the diet of this and related genera, especially in early instars (Wagner et al. 2008). McCabe has also reared several midgets, again from ova, on the foliage of boxelder (Acer negundo). The Variegated Midget overwinters as a pupa in a shallow chamber carved into tree bark—frass from the excavation is incorporated into the wall of the cocoon.

Midgets Subfamily Noctuinae: Tribe Elaphriini 383

The Wedgeling Galgula partita ! recognition Shape (and host) immediately distinguish this caterpillar. T3– A2 greatly swollen and A8 strongly angulate. Deep brown with white middorsal stripe especially well developed over A8–A10. White spiracular stripe includes small, dark spiracles. Head coloration diagnostic: central, inverted, white “V” bounded by shiny black lobes, which are divided by white stripe. Larva to 2.5 cm. ! occurrence Meadows and woodlands from Canada, south in East to Florida and Texas. Three or more generations with mature caterpillars from May onward over much of East. Common. ! common foodplants Wood sorrel, including introduced species. ! remarks Godfrey (1981b) provides a detailed description of the larva. The shape and posture of The Wedgeling are quite remarkable. The adaptive significance of its form, assuming there is one, eludes us, particularly because the larva feeds close to the ground where it would be inconspicuous and overlooked by many birds, the most abundant and chronic visual predators of temperate owlet caterpillars. The early instars of the Rustic Quaker (Orthodes majuscula) also have a swollen anterior, and crudely resemble The Wedgeling. Presumably pupae of Galgula overwinter but we are unsure how far north they are able to do so perennially. Our collection records suggest that the species is migratory with relatively few pupae surviving southern New Jersey winters.

384 Midgets Subfamily Noctuinae: Tribe Elaphriini

Cloaked Marvel Chytonix palliatricula ! recognition Body chunky, smooth, intricately mottled with pale snowflake markings. Pale subdorsal spot on A1. Broken, white middorsal stripe, strongest over A3–A8, poorly developed over thorax. Lab-reared individual figured here (lower left) with yellow subventral spot on T1, T2, and A3–A7. Shallow, transverse creases extend across abdominal segments. Head small, shiny, with two broad, parallel transverse bands and long, white antennae. Larva to 3 cm. The Barrens Marvel (C. sensilis) is similarly marked, but darker in the images that we have for study (bottom right)—see below. ! occurrence Barrens, woodlands, and forests from southern Canada, south in East to Florida and Texas. At least one extended and possibly two generations in Connecticut with mature caterpillars from midsummer onward; presumably with more generations southward. Common as an adult. ! common foodplants Larvae feed on a variety of gilled and pore-bearing mushrooms. Tim McCabe has reared ex ova cohorts on smut fungi. ! remarks The small head and chunky thorax suggest that the larva draws its head into the thorax when alarmed. Don Lafontaine and William Forrest (pers. comm.), who reared both C. palliatricula and the Barrens Marvel (C. sensilis) on store-bought Agaricus, were struck by the “amazingly” rapid larval development on mushrooms; they concluded that the caterpillars were well adapted to exploit ephemeral fungal hosts. Natural host ranges of our Marvels are undocumented. The pupa overwinters in a cocoon spun in litter. The Barrens Marvel is a habitat specialist associated with pitch and longleaf pine communities from southern Michigan to Quebec and Maine, south to Florida. It is most apt to turn up in barrens, dry oak woodlands, flatwoods, and savannas that have burned with some regularity. The moth has one brood late in the season; mostly August in Connecticut, September in North Carolina, and October in Florida when numbers of the Cloaked Marvel adults are falling off for the year. The taxonomic position of the genus is unresolved, but genitalic features suggest that Chytonix is related to elaphriines (Don Lafontaine, unpubl. data). The caterpillar’s small head, interrupted middorsal stripe, and enlarged rump are features shared with midgets (Elaphria).

Armyworms Subfamily Noctuinae: Tribe Prodeniini 385

Armyworms

Family Noctuidae: Subfamily Noctuinae: Tribe Prodeniini This tribe is represented by the genus Spodoptera in North America. Several members of the genus are important agricultural pests of field crops and cereals, especially in the tropics and subtropics. Spodoptera is a large, taxonomically difficult genus, although much of the species diversity is found in the Old World. There are nine established species in our coverage area. Pogue’s (2002) comprehensive monograph of the world fauna includes a wealth of information on the taxonomy and bionomics of the genus, keys to both larvae and adults, and numerous illustrations. Steven Passoa (USDA-APHIS-PPQ) authored the key below, supplied literature, and confirmed the identities of most of the larvae used to illustrate this work. Spodoptera (and a few related genera) have a minute (hardened) plate below the SD1 seta on T2 and T3. Most color forms have a dark lateral spot on A1 or T2 (the latter only in the Beet Armyworm, S. exigua, in the East). The frontal triangle extends four-fifths of the distance to the top of the head, and is commonly edged with white. While most late instars can be identified with this text, some color forms cannot—coloration is remarkably variable within species and only a fraction of the forms are shown here. Moreover, coloration and patterning of early and middle instars differs decidedly from that of the last instar: e.g., see the Southern Armyworm (S. eridania). Crumb (1956), Levy and Habeck (1976), and Pogue (2002) have written larval keys to the species in our fauna. The small eggs are laid in clusters of 50 to 100 or more, usually on the underside of a leaf, and then covered with a mass of deciduous scales from the female’s abdomen (photograph above right shows egg clutch of S. exigua). Early instars are scrapers, feeding only part way through the leaf from the underside, avoiding veins. Larvae pass through five or six instars. Spodoptera are widely polyphagous, consuming a sweeping variety of grasses, herbs, and less frequently, woody plants.

Egyptian Cotton Leafworm Spodoptera littoralis [African Cotton Leafworm]

Nonresident: Old World species taken once in Ohio greenhouse. Widely polyphagous, mostly on herbaceous plants and by extension numerous plants of economic significance: e.g., alfalfa, clover, cotton, crucifers, cucurbits, lettuce, potato, soybean, sunflower, and tomato, but also grains (wheat and corn) and fruit trees. Dorsum contrastingly paler than sides. Typically with large triangular spots on A1 and A8.

Armyworms get their name from the mass marches made by the caterpillars when abandoning fields that they have ravaged. Heppner (2007) provides extensive hostplant data for our species. Organically grown (i.e., pesticide free and not genetically modified) lettuce and spinach can be used to rear armyworms year-round; during the summer months, we rear our collections on dandelion, plantain, clover, and any number of available weeds. Larvae are parasitized commonly by braconid wasps (Chelonus, Cotesia, and Meteorus) and gregarious, externally-feeding eulophid wasps (Euplectrus and kin) (see page 29) (Ashley et al. 1980; DLW, unpubl. data). The pupa overwinters in a cell belowground, free of silk. None are known to survive prolonged periods of freezing. Many, and perhaps all, are migratory, moving northward over the summer and fall months. In the Northeast, the Yellow-striped Armyworm (S. ornithogalli) is the first to arrive, often in spring; the Fall Armyworm (S. frugiperda) is the next most commonly encountered eastern species and seldom appears in appreciable numbers before mid-August or September. Any others that arrive in New Jersey and northward do so from September to November.

386 Armyworms Subfamily Noctuinae: Tribe Prodeniini ! A RMYWORM HEADS. Coloration and patterning varies within species, e.g., the melanized (dark) forms in Spodoptera exigua and S. ornithogalli shown below represent crowding or outbreak forms. In S. androgea, S. eridania, and S. ornithogalli, the first image represents either a middle or penultimate instar and the second image represents a last instar. !

Field Key to Late Instars of Spodoptera of America North of Mexico Steven Passoa The following key, based on Levy and Habeck (1976), Passoa (1991), Godfrey (1987), and Pogue (2002), will facilitate identification of the ten Spodoptera species known to breed in the continental United States and Canada, as well as two exotic species, the Rice Armyworm (S. litura) and Egyptian Cotton Leafworm (S. littoralis). The key was written to enable extension entomologists, regulatory officials, and others to make preliminary larval identifications and increase the chance that nonnative species will be more quickly recognized. To simplify the key, emphasis is placed on the most frequently encountered color forms. Keep in mind that the markings in Spodoptera are often highly variable, e.g., exceptions and intergrades in shapes of the dorsal markings are common. Try to secure and examine more than a single individual whenever possible—a reasonable sample should include typical forms that will key correctly. The key is for living late instars; consult the text and photographs in Pogue (2002) for the identification of preserved larvae. We also caution that the color variation in the Androgea (S. androgea) and Caribbean Armyworm (S. pulchella) is unstudied. Interested people may also want to consult Heppner’s (1998) key to the seven widely established Spodoptera in Florida (www.doacs.state.fl.us/pi/enpp/ento/entcirc/ent390.pdf). 1a Dorsal abdominal pinacula larger than diameter of spiracles on A1–A7; these pinacula either conspicuous, especially on A8 and A9 (brown or black color forms), or pale (green form) . . . . . . . . . . . . . . . . . . . . frugiperda 1b Dorsal abdominal pinacula inconspicuous and smaller than the diameter of spiracles on A1–A7 . . . . . . . . . . . . . . . . . . . . . . 2 2a Lateral dark spot on T2 normally present; large lateral spot on A1 absent; dorsum with an irregular series of white dots and broken lines, or more rarely a series of paired thin black dashes . . . . . . . . . . . . . . . . . . . . . . . . .exigua 2b Lateral dark spot on T2 almost always absent (present in praefica from western US); large lateral spot on A1 commonly present; abdominal dorsum with either triangular markings or no markings . . . . . . . . . . . . . . . . . . . . . . . 3

Armyworms Subfamily Noctuinae: Tribe Prodeniini 387 3a Dorsal triangle on A1 much larger than dorsal triangle of A6, or triangles present only on A1 and A8 . . . . . . . . . . . . . . . . . . 4 3b Dorsal triangle on A1 absent, equal to, or smaller than dorsal triangle of A6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 4a Spiracular stripe less intense in front of lateral dark spot on A1 than behind it; if lateral dark spot or spiracular stripe on A1 absent, then dorsal triangles lack white spots at their apex; head usually light orange brown; native to US . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . eridania 4b Spiracular stripe equal in intensity on thorax to abdomen; dorsal triangles, especially those over T2 and T3, with dorsal white spots at their apex; head usually dark; Old World species, most likely to be found on or in areas near imported produce and nursery stock . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5a Ground color usually brown or gray, rarely green; subdorsal area strongly contrasting with paler dorsum; middorsal line usually absent; spiracular stripe not interrupted by black band or spot; dorsal triangular markings usually present only on A1 and A8, or just A8; if abdominal segments have black dorsal triangular markings, then these lack white spots at their apex; nonresident, potential pest . . . . . . . . . . . . . . . . littoralis 5b Ground color varying from green to brown; subdorsal area not strongly contrasting with paler dorsum; middorsal line often present; spiracular stripe often interrupted on A1 by black band or spot; at least some dorsal triangular markings with white spot at their apex, these triangles either found on all abdominal segments, or just on A1 and A8; nonresident, potential pest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . litura 6a T2 marking trapezoidal and equal in size to dorsal triangle on A8 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 6b T2 marking usually triangular or hemispherical, rarely reduced to white spot or small trapezoid; these markings smaller than dorsal triangles on A8 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 7a Larva from southern Florida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . androgea or dolichos (see text) 7b Larva from central Florida northward and westward, but infrequent north of Gulf states . . . . . . . . . . . . . . . . . . . . . . dolichos 8a Dorsal abdominal triangles with single white spot at middle or close to apex; if dorsal abdominal triangles absent, then at least some white spots bordered with black semicircles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . albula 8b Dorsal abdominal triangles solid or with lines, many fine spots, or dashes passing through them; if triangles include single white spot, then spot smaller than half diameter of spiracles of A1–7; rarely, white dashes may be present at base of dorsal triangles; if dorsal abdominal triangles absent then white spots never bordered with black semicircles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 9a Dorsal abdominal triangles absent on A1, thin or absent on A2, and with several thin narrow lines passing through them on other segments; southern Florida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .pulchella 9b Dorsal abdominal triangles either present on A1–A2, or if absent, then any remaining dorsal triangles lack thin narrow lines passing through them; widely distributed throughout US . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 10a T2 with small, dark, lateral rectangular spot; restricted to western US (southern Canada south into Mexico, west to Pacific coast) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .praefica 10b T2 lacks small dark lateral rectangular spot; widely distributed throughout US. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11a Dorsal abdominal triangles solid, without lines or dots passing through them . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 11b Dorsal abdominal triangles not solid, with lines and/or dots passing through them (faint or absent in dark forms). . . . . 14 12a Dorsal abdominal triangles on A1 to A8 equal in size . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ornithogalli 12b Dorsal abdominal triangles on A7 and A8 larger than others . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 13a Larva found in Gulf states. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ornithogalli or latifascia 13b Larva found northward of Gulf states . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ornithogalli 14a Pale markings through dorsal abdominal triangles form solid, or nearly solid, white or yellow line; subdorsal line white or yellow; T2 spot triangular, elongate-oval, or semicircular; widely distributed north of Mexico . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ornithogalli 14b Pale markings through dorsal triangles composed of a series of large dots; subdorsal line white or orange; T2 spot semicircular or faintly trapezoidal, never triangular or elongate-oval; mostly from Gulf states . . .latifascia

388 Armyworms Subfamily Noctuinae: Tribe Prodeniini

Androgea Armyworm Spodoptera androgea

Southern Florida to Brazil; possibly only temporary or occasional resident in our area. Polyphagous; cacao, Piper, and Xanthosoma confirmed as foodplants in Central America. Lab hosts include amaranth and lettuce. According to Pogue (2002), caterpillar similar to S. dolichos except dorsal triangles over A6, A7, and A8 gradually increase in size and never touch at midline on A8. Also, dorsal markings on metathorax absent in S. androgea but present or absent in S. dolichos. Broad, pale spiracular stripe.

Rice Armyworm

Spodoptera litura [Cluster Caterpillar, Tobacco Cutworm, Oriental Leafworm, and Tropical Armyworm] East Asia; adults collected twice from southern Florida but not established. Widely polyphagous: alfalfa, amaranth, apple, beans, cabbage, corn, cotton, sorghum, tobacco, various orchids, and many other forbs and grasses. Similar to Southern Armyworm (S. eridania) but spiracular stripe usually not obscured forward of supraspiracular black spot on A1, dorsal triangles often with small white spot at apex, and head capsule smoky brown with pale edging to triangles (not orange-brown) (left and below left).

Caribbean Armyworm Spodoptera pulchella [Not illustrated]

Southern Florida. Multiple generations throughout the growing season. Increasingly common and widespread across southern Florida. Cotton and orchids; presumably a general feeder on nonwoody plants. Dorsal triangles absent on A1; narrow lines passing through triangles on other segments.

Armyworms Subfamily Noctuinae: Tribe Prodeniini 389

White-spotted Armyworm Spodoptera albula (= S. sunia)

[Unbarred Armyworm]

! recognition Exceptionally variable in color, ranging from brightly variegated in yellow, orange, red, and black (lower left), to greenish, gray, or tan, with subdued patterning. Dark dorsal triangular markings when present with white spot at apices (to inside of D1 seta); in poorly patterned individuals look for evidence of faint dorsal patches over T2 and A7. Middorsal and subdorsal stripes range from brightly colored to being wholly undifferentiated from ground color; subdorsal stripe, when present, often edged with white above and below. Body below often tinted orange, with subspiracular stripe generally unpatterned. Prothoracic shield brown above with broad subdorsal creamy area. Head mottled with dark brown; orange-brown triangle bounded with cream lines. Larva to 3.5 cm. Middle instars often handsomely rendered in yellows, reds, and other bright colors; middorsal stripe often yellow to orange-red. ! occurrence Waste lots, agricultural fields, and other open sites. Mostly southern Florida and Texas, southward through Caribbean and Central America to Argentina. Multiple generations with mature caterpillars throughout the growing season. Common. ! common foodplants Broadly polyphagous on low-growing plants, including grasses. Economically important hosts include cabbage, corn, cotton, flax, pea, peanuts, rice, snapdragons, sorghum, soybean, and tomato (Passoa 1983; Pogue 2002; Lyle Buss, unpubl. data). ! remarks The White-spotted Armyworm was previously known as S. sunia. While the species is occasionally destructive to crops in Central America, it is not a pest in the United States. In addition to leaves, caterpillars sometimes damage flowers (cotton) and fruits (tomatoes). Early and middle instars are often beautifully rendered in a checkerboard of oranges, yellows, and other colors. Mature caterpillars are more subdued, typically brownish with a wood grainlike aspect.

390 Armyworms Subfamily Noctuinae: Tribe Prodeniini

Sweetpotato Armyworm Spodoptera dolichos [Large Cotton or Dolichos Armyworm]

! recognition Like other armyworms, color highly variable, ranging from ochre to pale brown or gray with paired, black dorsal spots over thorax and abdomen. Dorsal spots on T2 large, trapezoidal, and approximately same size as those on A8. Head with brown coronal bars; triangle edged with white (adfrontal area). Larva to 5 cm. Black spots over T2 and A8 not triangular, elongate-oval, or round as in Orange-striped Armyworm (S. latifascia) and Yellow-striped Armyworm (S. ornithogalli). In the Orange-striped Armyworm, the species most likely to be confused with S. dolichos, the dorsal black spots on T2 are distinctly smaller than those over A8. See also Androgea Armyworm (S. androgea) (page 388). ! occurrence Agricultural fields, waste places, and other open habitats. Resident in Gulf Coast states, straying northward to Wisconsin and Maine in fall. Continuously brooded in southern Florida and Texas. Common. ! common foodplants Polyphagous on grasses and herbaceous plants including asparagus, beans, clover, corn, cotton, sweet potato, strawberry, and turnip; many records from plants in nightshade family, e.g., potato, tobacco, and tomato. ! remarks When alarmed, the larva rolls the head under, which serves to fully expose its “eyes” (the dark trapezoidal spots over T2) (lower inset). Adults come to light, flowers, and baits; migrating adults are avid visitors to sugary solutions. Predictably, floral scents (e.g., phenylacetaldehyde and acetic acid + 3-methyl1-butanol) can be used to collect members of the genus (e.g., see Meagher and Mislevy 2005). The sex ratios of Spodoptera attracted to floral baits are nearly equal, whereas those from light trap samples are invariably male biased.

Armyworms Subfamily Noctuinae: Tribe Prodeniini 391

Southern Armyworm Spodoptera eridania ! recognition Variable, ranging from mundane gray-green to smoky grayblack, with patterning obscure to beautifully festooned with orange, yellow, and/ or violet markings (especially through middle instars). Anterior of body often more darkly pigmented. Lateral spot over spiracle on A1 enlarged. White to yellow middorsal, subdorsal, and subspiracular stripes. Broad subspiracular stripe, frequently cream to yellow with flush of orange across abdominal segments, less intense or sometimes obscured (by black suffusion) anterior to spot on A1. Dorsum with large, black triangular spot on A1, and shallowly hemispherical or linear black spots along inner side of subdorsal stripe on T2–A9. Head usually orange-red to red-brown, with or without coronal bars. Larva under 4 cm. Early instars more smoky green, thickened through T2, T3, and A1, with pair of diffuse black spots over A1 and A8 (lower middle right), lacking the bright coloration characteristic of middle instars (lower right). ! occurrence Agricultural fields, waste places, and other open habitats. Gulf Coast states (south to Argentina); adults stray northward to Missouri and New Hampshire (rarely) during summer and fall. Continuously brooded in southern Florida and Texas with five or more annual generations. Very common. ! common foodplants Broadly polyphagous on grasses and herbaceous plants including celery, corn, cotton, crucifers, cucurbits, legumes, potato, sugarbeet, sweet potato, tobacco, and tomato; many records from amaranth, nightshade, pigweed, and pokeweed. ! remarks The Southern Armyworm is a frequent agricultural pest in Florida and Texas, and south of our border through Mexico and portions of Central and South America. Typically the larvae are leaf feeders, although they sometimes damage buds, flowers, or fruits. On tomato plants the larvae sometimes tunnel into fruits. The caterpillars are somewhat secretive and tend to feed near the ground and from leaf undersides, at least during the day. The larval stage lasts 17–21 days in warm weather. Pupation occurs in a cell belowground.

392 Armyworms Subfamily Noctuinae: Tribe Prodeniini

Beet Armyworm Spodoptera exigua ! recognition Various shades of green, brown, or gray with broad lateral (subspiracular) stripe. Dorsum flecked with pale spots or dashes, usually lacking prominent, paired, black, triangular spots of other Spodoptera. Thin middorsal stripe usually broken and/or obscured by other patterning. Prothoracic shield cut by pale middorsal stripe. Ground color often darkened below white to yellow subdorsal stripe (stripe sometimes obscure except through shield). Pale lateral stripe often edged above with black. Many forms with black supraspiracular spot (above lateral stripe) on T2 (an analogous black spot occurs on A1 in other Spodoptera); dark subdorsal spot sometimes present on A8. Infrequently with series of dorsal dashes (as in individual in lower right of larval images). Coronal bar and dark reticulations sometimes present; occasionally head mostly blackened as in Armyworm Head Plate. Larva under 3.5 cm. Closest in appearance to Fall Armyworm (S. frugiperda)—see next page. ! occurrence Agricultural fields, waste places, and other open habitats. In East, from Canada southward through Florida, Texas, and into northern South America; however, an erratic migrant and for many years not present over much of East or north of Georgia. Common resident in Florida, Texas, and Southwest. Multiple generations with mature caterpillars throughout the growing season in Gulf states. Common. ! common foodplants General feeder on low-growing plants, especially grasses and grains: e.g., corn, rice, sorghum, sugarcane, and timothy. Other commonly reported hosts include asparagus, bean, cabbage, carrot, cotton, lettuce, pea, pepper, pigweed, potato, soybean, spinach, sugarbeet, and tomato. ! remarks The Beet Armyworm was introduced to the United States from Europe. It established in Oregon by 1876 and from there spread eastward to Florida by the 1920s. Infestations are commonly reported by farmers in the South and West. The moth is infrequent to rare in Ohio (Rings et al. 1992) and Connecticut. Older larvae can be aggressive towards one another, and more rarely cannibalistic; this is somewhat surprising given that the eggs are deposited in clutches of 50 or more. Development is rapid with generations cycling through in a month during periods of warm weather. Pupation occurs belowground, often at depths of 2–8 cm. Pupal survivorship over the winter months drops off quickly with freezing conditions; few pupae are thought to successfully overwinter north of the Gulf states. Adults come to light and are avid flower visitors; we have not taken adults at sugar baits.

Armyworms Subfamily Noctuinae: Tribe Prodeniini 393

Fall Armyworm Spodoptera frugiperda ! recognition Usually shades of brown but gray, green, yellow-green, and blackish forms also occur. Abdominal girth increasing slightly rearward. Setal bases large, shiny black, raised, especially over dorsum and above spiracles. Dorsal pinacula on A8 and A9 especially prominent. Supraspiracular area darkened below subdorsal stripe. Broad, pale lateral stripe often infused with brownish mottling. Prothoracic shield brown, well differentiated, cut by middorsal and subdorsal stripes. Dorsal integument densely set with small, rounded granules (cobblestonelike) visible at magnifications of 25×. Larva to 3.5 cm. Middle instars often more yellow-green in color. ! occurrence Fields, waste places, and agricultural lands from southern Canada to Florida and Texas (south to Argentina), becoming increasingly common southward. Multiple generations with caterpillars only in late summer and fall northward; active year-round in extreme south. Abundant southward. ! common foodplants General feeder on grasses, forbs, and some woody plants; crop hosts include alfalfa, corn, clover, cotton, strawberry, tobacco, and wheat. ! remarks The Fall Armyworm is a chronic pest of corn in the South where its characteristic damage has earned it the common name whorlworm (or cogollero in Spanish). The caterpillars tunnel through the husks to get to the developing seeds (kernels), whereas larvae of the Corn Earworm (Helicoverpa zea) tunnel downward from the tassel. The Fall Armyworm is also a frequent pest of turf grass. Development is rapid with generations cycling through in as few as 30 days in warm weather. The rapid development time reflects, at least in part, the willingness of caterpillars to feed at any time of day. Recent DNA data and sex pheromone trapping studies suggest that the Fall Armyworm is a complex of two sibling species (Pashley 1988; Pashley et al. 1985, 2004). One of these is associated with rice, low grasses, and occasionally corn. The other has been reared most frequently from corn, sorghum, and cotton. Morphological characters that would allow separation of the two entities, either as larvae or adults, have not been identified. Like most members of the genus, it is susceptible to winter freezes: Johnson (1995) reported that cold weather in January 1977 virtually eliminated populations north of Homestead, Florida. After the Yellow-striped Armyworm (S. ornithogalli), this is the Spodoptera most likely to be seen in the North. In New Jersey, at least in recent years, adults arrive mostly in September, probably in time to breed successfully.

394 Armyworms Subfamily Noctuinae: Tribe Prodeniini

Orange-striped Armyworm Spodoptera latifascia ! recognition Highly variable, ranging from rather undistinguished to

brightly colored. Dorsal black hemispheres or triangles (if present) increasing in size rearward. Dorsal spot on T2 consistently smaller (about half the size) than that on A8; also dorsal spot on A7 tends to be smaller than that on A8. Dorsal spot on T2 often hemispherical (not trapezoidal). Subdorsal stripe white, or more commonly, infused with orange. Head reddish orange with sides of triangle edged with white. Larva to 5 cm. Well-marked individuals can be confused with those of the Yellow-striped Armyworm (S. ornithogalli), which has a (more) prominent yellow subdorsal stripe (although this may be white in some forms); also, the subdorsal stripe tends to have less orange than that of the Orange-striped Armyworm. The area below the subdorsal stripe in the Yellow-striped Armyworm bears a series of fine longitudinal stripes that are rarely expressed in the Orangestriped Armyworm. Caterpillars of the Sweetpotato Armyworm (S. dolichos) are somewhat similar to S. latifascia, but the dorsal spots over T2 are trapezoidal and nearly equal in size to spots over A8. Additional color forms of the Orange-striped Armyworm are posted on Dan Janzen’s Costa Rican caterpillar website (http:// janzen.sas.upenn.edu/). ! occurrence Agricultural fields, waste places, and other open habitats. Gulf states south through Central America, sometimes straying northward. Continuously brooded in southern Florida and Texas. Very common in Deep South. ! common foodplants Broadly polyphagous on grasses, forbs, and, less commonly, woody plants. Crop hosts include beans, corn, cotton, and tomato; Heppner (1998) lists 80 plant genera as hosts, most of which are of economic importance. ! remarks The Orange-striped Armyworm is variously known as the Black, Garden, Lateral-lined, or Velvet Armyworm. Significant outbreaks have been reported from a variety of field crops, e.g., in Honduras it is a pest of young sorghum and maize (Portillo et al. 1996). Armyworms are often attacked by gregarious eulophid wasps in the genus Euplectrus (upper right pane). The wasp larvae mature quickly—once the maggots take on a greenish color the caterpillar’s demise is imminent. After they are fully fed, the diminutive wasp larvae abandon the corpse and crawl beneath their victim to spin wispy, funguslike, cocoons (page 29). The Orange-striped Armyworm is becoming more common and widespread—DFS took the moth for the first time in New Jersey in 2006.

Armyworms Subfamily Noctuinae: Tribe Prodeniini 395

Yellow-striped Armyworm Spodoptera ornithogalli ! recognition Patterning variable: ground color tan to purple-brown, marked

with colorful stripes. Most forms with shallowly hemispherical to triangular black spots that extend inward from white, yellow (usually), or brown subdorsal stripe; dorsal spots, when present, commonly cut by fine line near their base (lower right). Dorsal spot on A7 approximately same size as that on A8. Abdominal segments frequently with set of four, thin, white stripes beneath subdorsal line; these sometimes confluent and appearing as single, broad, lavender stripe. Frequently with rusty dash beneath each spiracle, which may join to form thin subspiracular stripe. On head, sides of triangle outlined with yellow or white. Larva to 4.5 cm. Caterpillars of Sweetpotato Armyworm (S. dolichos) may be differentiated by the size and shape of the black spots on T2 and A8 (see page 390). In many forms of the Orange-striped Armyworm (S. latifascia), the dorsal spots are interrupted by minute white spots (not a line as in the Yellow-striped Armyworm); additionally, the subdorsal stripe is often orange rather than yellow. Other characters are given in the account for the Orange-striped Armyworm. The Western Yellow-striped Armyworm (S. praefica), present in the Rockies and westward, has more contrasting mottling above the antenna—see treatment in Pogue (2002). ! occurrence Fields and other open habitats from southern Canada to Florida and Texas (south to Costa Rica). Multiple generations with mature caterpillars through summer and fall months. Very common. ! common foodplants General feeder, usually on low-growing forbs and woody plants, including many field crops and garden plants: e.g., beans, corn, cotton, crucifers, sugarbeet, and sweet potato; occasionally reported as pest of fruit trees (Rings 1977). ! remarks The Yellow-striped Armyworm is the most commonly encountered member of the genus northward. Adults migrate north each spring with storm fronts that blow up from the Gulf of Mexico. Caterpillars are occasionally found feeding well above the ground on shrubs and trees. For a concise summary of the Yellow-striped Armyworm’s life cycle, foodplants, and natural enemy complex, visit the University of Florida’s Featured Creatures website (http://entnemdept.ufl. edu/creatures/veg/leaf/yellowstriped_armyworm.htm), written by John Capinera (2001, 2005).

396 Caradrines Subfamily Noctuinae: Tribe Caradrinini

Caradrines

Family Noctuidae: Subfamily Noctuinae: Tribe Caradrinini This small, mostly western tribe includes five genera (AnorthAnorthodes, Athetis, Caradrina, Protoperigea, and Proxenus)) and 22 species. Immature stages and biologies are poorly known, perhaps because the larvae feed on or near the ground, and are generalized in diet.

The Slowpoke Athetis tarda ! recognition Dark brown, mottled cutworm, appearing somewhat roughened and corrugated with enlarged and strongly humped eighth abdominal segment. Tan middorsal stripe runs from head to middle of A8. Posterior edge of A8 beige; A9 with dark subdorsal spot. A2–A8 with pale, oblique subdorsal lines; these frequently with dark pigment about their lower end and enclosing vague dorsal chevrons. Shiny dark head capsule that may be partially withdrawn into thorax. Each proleg bears black stripe. Integument peppered with minute dark granules. Larva to 3 cm. Superficially similar to Rustic Groundcat (Orthodes majuscula), page 517. ! occurrence Woodlands, forests, and barrens from lower Great Lakes states to New Hampshire, south to Florida and Texas. Two generations in Connecticut with mature caterpillars through winter and early spring, then presumably again from late June through early August. Very common in oak woodlands. ! common foodplants Dead oak leaves. ! remarks The Slowpoke is an apt name for this species. The caterpillars are decidedly lethargic and feign death when disturbed, and their coloration resembles the litter and detritus upon which they feed. When exposed, the caterpillar hunkers down against its perch, which is often a dead leaf. The swollen A8 and dark subdorsal spots on A9 make for a credible false head. If the caterpillar is threatened, The Slowpoke will draw further attention to the rear of its body by releasing the anal prolegs and elevating the terminal abdominal segments. In addition to dead leaves, our lab-reared larvae occasionally consumed siblings (inset). Larval collections made by Crumb (1956) in February, and in March by us, indicate that the larva overwinters. In Mitchell et al.’s (2006) molecular study, Athetis and Caradrina sit near the base of the Noctuinae, sister to the core group of Noctuinae tribes. The spring and fall generations get further separated in time as one moves southward—in coastal North Carolina adults fly in April and then again mostly in September.

Dipterygiines Subfamily Noctuinae: Tribe Dypterygiini 397

Dipterygiines

Family Noctuidae: Subfamily Noctuinae: Tribe Dypterygiini As circumscribed by Lafontaine and Schmidt (2010), the North American fauna consists of 11 species distributed among five genera, two of which are represented here: Dypterygia and Magusa. A8 is strongly humped in both.

American Bird’s-wing Moth Dypterygia rozmani

Manitoba to Nova Scotia, south to Florida and Arkansas. Probably two broods over much of East with mature caterpillars in July, and then again in fall. Locally common. Dock and climbing false buckwheat (Polygonum scandens). Our captive larvae refused aster and other forbs.

Divaricate Narrow-wing Magusa divaricata ! recognition Aposematically colored with black, white, yellow, and orange. Head, prothoracic shield, legs, and anal plate bright orange. A8 swollen, raised, often yellow to red-orange. Dorsum with three, thin, white to yellow stripes. Bold white to yellow supraspiracular stripe capping prominent black lateral stripe. Orange head with black setal insertions. Venter yellow-green and unpatterned. Larva to 4 cm. The larva of the Orbed Narrow-wing (M. orbifera) was described by Forbes (1954) as “pale green with about six fine white or yellowish lines on each side, some faint white mottling.” It occurs in southern Florida, from about the latitude of Miami southward, and south through the Caribbean (Lafontaine et al. 2009). DLW reared a few from black ironwood (Krugiodendron) in Homestead, Florida. ! occurrence Thickets, scrublands, and coastal strand communities along

continued overleaf

398 Dipterygiines Subfamily Noctuinae: Tribe Dypterygiini

Gulf coast, south through Neotropics to Argentina, but curiously scarce from southern Florida; in some years, migrating north to southern Canada. Presumably resident and continuously brooded in northern Mexico and southern Texas, at least in mild years, with mature larvae year-round. Likely breeding northward on introduced Rhamnus species to Great Lakes states during summer months. Common southward except in Florida. ! common foodplants Buckthorn or coffeeberry (Rhamnus), coyotillo (Karwinskia), and presumably other Rhamnaceae. ! remarks While the adults of our two narrow-wings (Magusa) are similar enough in appearance to have been confused as a single species until recently (Lafontaine et al. 2009), their caterpillars are notably distinct: that of the Orbed Narrow-wing (M. orbifera) is yellow-green and cryptic, while the Divaricate Narrow-wing is boldly colored and gregarious. The latter is two headed: the red-orange eighth abdominal segment is swollen and marked with the same black spotting as the insect’s true head. Larvae occur in clusters, and sometimes dozens can be seen on a single shrub or tree; large aggregations, sometimes with more than a hundred individuals, are less commonly encountered, but often defoliate their host. Larval clusters are reminiscent of those formed by Datana (Notodontidae)—in the same fashion, the caterpillars lift both ends of the body and splay their thoracic legs. In other instances, the caterpillars scatter themselves, and feed solitarily. Alarmed individuals are quick to release their grip and drop to the ground. The taxonomic position of Magusa long puzzled lepidopterists. Crumb (1956) placed the genus at the end of his work, unsure of its subfamilial placement. Lafontaine et al. (2009) place the genus with Trachea and Dipterygia based on similarities in the mitochondrial sequence data of the three genera. Magusa is a tropical lineage and cannot tolerate prolonged freezing conditions in any stage.

St. Johnswort Moths Subfamily Noctuinae: Tribe Actinotiini 399

St. Johnswort Moths

Family Noctuidae: Subfamily Noctuinae: Tribe Actinotiini A small tribe with two widespread eastern species. The hypopharynx bears a lateral row of 30–40 peglike spines and the apical margin of the spinneret is entire, not fringed as in many related groups. In our species, the spiracles on A5, A6, and A7 are often reduced in size and the outer face of prolegs bears a dark T-shaped plate. Larvae feed on St. Johnswort (Hypericum).

Gray St. Johnswort Moth Nedra ramosula

[Gray Half-spot]

! recognition Sienna to red-brown with prominent white to yellow spiracular stripe (composed of whitish dots). Thin and incomplete middorsal, subdorsal, and supraspiracular stripes. Ground color often darkened between dorsal and supraspiracular stripes. Spiracular stripe extends forward to antenna and back down outer face of proleg; its upper edge touches the spiracles on T1–A7. Spiracles black. Antenna and sometimes entire head red. Anal plate with cream subdorsal stripe. Middle instars yellow-green with thin middorsal and subdorsal stripes; spiracular stripe delimits deeper green upper half of body from pale subventer and venter. Larva to 3.5 cm. ! occurrence Fields, meadows, bogs, moist pinelands, and edges of woodlands from Canada, south to Florida and Texas. Two or more generations throughout much of range with mature caterpillars from May through October. Locally common. ! common foodplants St. Johnswort (Hypericum). ! remarks On several occasions while beating bushy St. Johnswort (Hypericum densiflorum) in the Appalachian foothills for larvae of Nemoria tuscarora (Geometridae), we have serendipitously found Nedra caterpillars on our sheet. The early and middle instars feed from the lower side of leaf blades; older larvae tend to position themselves along a stem or among the flowers of a browning inflorescence. The moth is also frequent in wet meadows and marshes where other species of St. Johnswort grow. In all instars the anterior end of the body is flattened and appressed to the caterpillar’s perch. Adults will visit bait in the fall. Presumably, the pupa overwinters.

400 St. Johnswort Moths Subfamily Noctuinae: Tribe Actinotiini

Burgundy St. Johnswort Moth Iodopepla u-album ! recognition Tan or yellow-brown to warm red-brown with broad, creamy spiracular stripe. Hair-thin, white middorsal, subdorsal, and supraspiracular stripes; both middorsal and subdorsal stripes sometimes obliterated by brown shading. Small, white dorsal pinacula, contrasting with ground coloration. Broad lateral stripe running from antenna to anal proleg, along or just below lower edge of spiracles, distinctly narrowed forward and rearward; central portion of stripe sometimes infused with vague vertical reddish lines. Spiracles tan to nearly black with dark outer rim. Head as in upper inset. Larva to 3.5 cm. ! occurrence Marshes, wet meadows, bogs, and wet, coastal-plain pinelands from Great Lakes states, southern Ontario, and Massachusetts, south to Georgia. At least double brooded—see below. Local northward, but at least locally common along much of coastal plain from New Jersey southward. ! common foodplants Captive larvae accepted St. Johnswort (Hypericum). ! remarks Our efforts to rear this moth were made possible by Jim Troubridge who noted that the male genitalia are remarkably similar to those of the Gray St. Johnswort Moth (Nedra ramosula), a known Hypericum specialist. Armed with this clue, we obtained eggs of Iodopepla and reared caterpillars on Hypericum. Larvae of the Burgundy St. Johnswort Moth and Gray St. Johnswort Moth, two very different looking moths, are remarkably similar in appearance. In addition, both share a similar resting posture: the thoracic and terminal abdominal segments are somewhat flattened. The number of annual generations is in need of study. Flight records in Rings et al. (1992) suggest two generations in Ohio. In southern New Jersey, adults appear from April into October, in three or four peaks. Adults occur throughout the growing season in coastal North Carolina. The prepupal larva is shown in the lower inset. The pupa overwinters.

Angle Shades Subfamily Noctuinae: Tribe Phlogophorini 401

Angle Shades

Family Noctuidae: Subfamily Noctuinae: Tribe Phlogophorini This small tribe is represented in our area by three North American genera and five species. Adults fold the wings about the body in a characteristic fashion. Diagnostic larval characters, based on details of the mouthparts, are given by Ahola and Silvonen (2008). Phlogophorines often have both green and brown forms. While all are regarded to be polyphagous, ferns account for a disproportionate share of the hostplants in our wild larval collections.

American Angle Shades Euplexia benesimilis ! recognition Yellow to emerald green or cinnamon brown with distinctive pair of white “rump” spots over A8. Dark spots form oblique lines over subdorsal area of abdominal segments. Prothorax lightly pigmented anteriorly. Sides mostly waxy green or pale brown with conspicuous white mottling. Spiracles black with minute brown centers. Distal portion of prolegs sometimes with pinkish cast. Head pale green, shiny, unmarked, or with dark reticulations over each lobe. Larva to 3.5 cm. ! occurrence Woodlands and forests from southern Canada to Florida and northern Arkansas. Two or three generations over most of East with caterpillars from June to October. Common. ! common foodplants Ferns principally; also reported from alder, aster, huckleberry, sunflower, trillium, and willow. ! remarks We have found caterpillars on several occasions while sweeping or beating ferns. Certainly the larva’s coloration is a good match for ferns. Its British counterpart, the Small Angle Shades (E. lucipara), is commonly associated with ferns but also feeds on a variety of other woody, and especially herbaceous, plants. The caterpillar’s modestly swollen rump marked with white “eyespots” yields a somewhat credible false head. The pupa overwinters in a cocoon belowground.

402 Angle Shades Subfamily Noctuinae: Tribe Phlogophorini

Olive Angle Shades Phlogophora iris ! recognition Light green, olive, or reddish-brown, cutwormlike caterpillar with rounded and slightly swollen A7 and A8 (at least broader than anterior of body). Anal prolegs stubby, sometimes held under body and together (with crochets intermeshed). Most forms with vague, oblique, subdorsal lines that meet over dorsum to form backward directed “V”s over abdominal segments. These oblique marks may be replaced by weakly defined subdorsal lines over thoracic segments. Middorsal stripe thin and inconspicuous, often interrupted by black spot between adjacent segments, at least over abdomen. Spiracular stripe poorly expressed or absent, running under spiracles on A7 and A8, continuing down anal proleg. Spiracles with dark rim and narrow, tan inner area. Head with mottling over upper half with pigment coalescing to form coronal bars. Larva to 4.5 cm. ! occurrence Woodlands and forests. Saskatchewan to Newfoundland, south to central New Jersey, Georgia (mountains), and Great Lakes states. One generation with mature caterpillars from fall through following spring. Common. ! common foodplants Widely polyphagous. Literature records include dandelion, dock, and thistle (Crumb 1956), and sensitive fern (Rockburne and Lafontaine 1976). We have found larvae on alder, blackberry, black birch, St. Johnswort (Hypericum prolificum), and Vernonia. Our captive larvae were reared on apple. ! remarks The caterpillars feed on forbs, shrubs, and small trees. The nearly full-grown caterpillar overwinters, presumably nibbling on forbs and other lowgrowing plants, and possibly dead leaves, through the winter months.

Angle Shades Subfamily Noctuinae: Tribe Phlogophorini 403

Brown Angle Shades Phlogophora periculosa ! recognition Pinkish-tan to brown, mottled cutworm with smooth integument and thin middorsal stripe. A8 somewhat swollen, rounded. Middorsal stripe broken, weakest between dorsal setae. Setae on head arising from minute, darkened bases (visible with lens). Spiracular stripe faint, dissipating rearward of A8, often broken between segments. Sometimes with pink cast about spiracles. Anal proleg with white line down outer face. Larva to 5 cm. Middle instars green. Larva most cutwormlike in last two instars. Superficially resembling larva of Sloping Sallow (Epiglaea decliva) (page 461), but middorsal stripe more continuous and not broken up into well-separated spots. Duncan (2006) figures middle, penultimate, and last instars. ! occurrence Woodlands and forests. Transcontinental in Canada, south in East to Georgia, Mississippi, and Arkansas. One generation with mature caterpillars from May into July. Common. ! common foodplants Widely polyphagous: alder, blueberry, cedar, cherry, ferns, fetter-bush or swamp doghobble (Eubotrys racemosa), fir, hemlock, maple, meadowsweet, plum, sheep laurel, spruce, wild raisin, and many others. ! remarks The caterpillars are extreme generalists, accepting ferns, forbs, and many woody plants, including a wide range of conifers. Caterpillars may be reared on romaine lettuce. Adults come to bait. Both green- and brown-form larvae overwinter in leaf litter, consuming herbs and even dead leaves. Come late winter or early spring, the caterpillars move into shrubs and small trees, to eat buds and new growth.

404 Borers Subfamily Noctuinae: Tribe Apameini

Borers

Family Noctuidae: Subfamily Noctuinae: Tribe Apameini This tribe has long been in taxonomic flux, with its membership ever changing. For many years the tribe was placed in the Amphipyrinae, but more recently has been subsumed under the Hadeninae (Kitching and Rawlins 1998), Xyleninae (Fibiger and Lafontaine 2005), and now Noctuinae (Lafontaine and Schmidt 2010). Apameines have garnered much interest from lepidopterists because of their rich species diversity, frequent rarity, and often specialized habits; the last have led to numerous species-level radiations over evolutionary time (e.g., Apamea, Oligia, and Papaipema). Forbes (1954) had accounts for about 120 eastern species—the largest number for any tribe in his noctuid volume. Lafontaine and Schmidt’s (2010) North American checklist recognizes 219 species with numerous others awaiting description. Because most apameine caterpillars are internal or concealed feeders (consequently often lacking appreciable color and pattern) and because they are seldom encountered, they are underrepresented in this treatment. Few caterpillars are described in Mikkola et al.’s (2009) apameine monograph. Eric Quinter, the North American authority on apameine immature stages and their biology, helped considerably with this section of our guide. Apameine caterpillars tend to be mostly unpigmented, as is the norm for concealed caterpillars. Those that bore are usually whitish (fatty), smooth, and unpatterned, although Papaipema and related genera provide a handsome exception, especially in their middle instars. There is a field of tonofibrillae (internal muscle attachments) between XD1 and D2 setae on the prothoracic shield that may close to form a circle (Matti Ahola in Fibiger and Lafontaine 2005). The pigmentation pattern of the prothoracic shield and armoring of the anal plate are diagnostic for many. Morphological differences that can be used to distinguish species tend to be subtle, and often require access to a reference collection and/or microscopic inspection of the mouthparts and chaetotaxy. Because apameines are often dietary specialists, foodplant associations are helpful in making identifications.

All are univoltine. The eggs, usually laid in rows on senescing or dead foodplants, are covered with a varnishlike secretion and sealed within leaf sheaths or crevices in dried foliage.

The larvae are borers, subterranean cutworms, or root feeders. Monocots host a large fraction of our fauna (especially if Papaipema is excluded from consideration). Little or no silk is employed in the formation of the pupal chamber. North American species generally overwinter as eggs or early to middle instars, although at least Apamea, and perhaps some others, are sometimes well grown by the onset of winter. Most of the borers can be reared on carrot, potato, or sweet potatoes; artificial diets are also accepted. Corn roots or stalks are often acceptable to the grass feeders. Eggs may be obtained by providing gravid females with tight spaces into which they can insert their ova. Eric Quinter supplies strips of tightly pleated paper, stapled at both ends. Dead leaves of the foodplant will sometimes stimulate egg-laying behavior.

Double Lobed Borer Lateroligia ophiogramma (= Apamea ophiogramma)

Palaearctic species established throughout Great Lakes region and Northeast, and expanding southward and westward. One generation with mature caterpillars in late spring and early summer. Locally common in wetlands. Feeds internally on large grasses, such as reed canarygrass (Phalaris), common reed (Phragmites), and mannagrass (Glyceria) (Mikkola et al. 2009).

Borers Subfamily Noctuinae: Tribe Apameini 405 Iris Borer

Macronoctua onusta Minnesota to Nova Scotia, to South Carolina and Kansas. One generation with mature caterpillars in midsummer. Common. Destructive to iris corms and roots; also reported from gladiolas. Early instars mine leaves. Larva often with pinkish cast.

Broken-lined Borer Mesapamea fractilinea [Broken-lined Brocade]

Minnesota to New Brunswick, south to Florida and Mississippi. One generation with mature caterpillars in late spring and early summer. Common pest of timothy, less often of corn; native hosts unrecorded.

Green-washed Culm Borer Oligia chlorostigma

At least Wisconsin to southern Quebec, south to Georgia and Texas. One generation with mature caterpillars in late spring and early summer. Locally common. Grasses. Our example is a penultimate instar nearing molt.

Marbled Minor Culm Borer Oligia strigilis

Palaearctic species established in southern Ontario, Quebec, and eastern Massachusetts. One generation with mature caterpillars in late spring and early summer. Mostly local and uncommon. Grasses.

406 Borers Subfamily Noctuinae: Tribe Apameini

Veiled Grass Moth

Loscopia velata (= Amphipoea velata) [Veiled Ear Moth] ! recognition Lime green with broad, white spiracular stripe and characteristic black line (veil) on head that reaches to eyes. Thin, white middorsal and subdorsal stripes beginning on T2 and extending back to anal plate. Broad spiracular stripe including straw-colored spiracles narrowly rimmed with black. Prothoracic shield with weakly developed black addorsal and supraspiracular spots. Side of head with black commalike line; thickest portion over each lobe and tail passing through eyes. Larva to 3.5 cm. ! occurrence Lawns, fields, meadows, woodland, and forest edges from Manitoba to Newfoundland, south to Georgia and eastern Kansas. One generation with mature caterpillars from late April through June. Very common. ! common foodplants Grasses. ! remarks The early instars bore in stalks while the later instars are external leaf feeders, as their coloration would suggest. We do not know which instar diapauses and whether or not the larva does so within a tunnel, or if the caterpillar moves externally to feed on leaves by the onset of winter. In the late spring, this is among the most commonly encountered grass-feeding owlets—caterpillars perch conspicuously by day on leaves or seed heads. They can be readily sampled by sweeping, especially at night. It is curious that Crumb (1956) did not include the Veiled Grass Moth in his work on noctuoid larvae. Pupation occurs in a cell near the soil surface—supply prepupal larvae with soil or an equivalent.

Borers Subfamily Noctuinae: Tribe Apameini 407

Glassy Borer Apamea devastator (= Crymodes devastator)

[Glassy Cutworm]

! recognition Glossy, transparent, smooth integument; pale green to greasy white or gray with shiny, yellowish to reddish-brown head. Prominent, shiny orange-brown to light brown prothoracic and anal shields, each edged with black. Pinacula lightly pigmented, inconspicuous. Spiracles twice as high as broad. Head with vague coronal bars sometimes present; vertex deeply lobed. Three bladelike setae borne from basal segment of the maxillary palpus (inset). Prothoracic and anal plates darkened. Coxae on T2 separated by diameter of femur. Larva to 4 cm. Other Apamea caterpillars, of which there are many, are similar in appearance— Crumb (1956) provides a key to common Apamea species. ! occurrence Fields, meadows, and other grassy communities. Transcontinental in Canada, south in East to North Carolina (mountains) and Missouri; common northward. One generation with mature caterpillars in late spring and early summer. Common. ! common foodplants Grasses; pest of numerous cereal crops, including corn and wheat, as well as turf grass. Also feeds on roots of herbaceous plants (including alfalfa) growing with grasses. ! remarks We associate Apamea with wetlands, sand plains, and other grassy habitats. McCabe (1990) reported finding eggs of the Glassy Borer in the seed heads of rattlesnake grass (Glyceria canadensis). As might be guessed from its coloration, the larva is subterranean. Infestations are most commonly reported when corn or other row crops are planted in fields that were previously given over to sod, pasture, and other grass-dominated communities (Rings 1977). The partially grown larva overwinters. Additional details on its larval biology and pest status are reviewed in Mikkola et al. (2009).

408 Borers Subfamily Noctuinae: Tribe Apameini Apameine Borers Specimens and much of the biological information for this page and the next were provided by Eric Quinter. Most images are of preserved specimens (photographed in a dish of alcohol). Characters mentioned below may apply to all members of a given genus or tribe.

American Borer

Amphipoea americana [American Ear Moth] Manitoba to Newfoundland, south to Georgia and Arizona. One generation with mature caterpillars in late spring and early summer. Common. Often grasses and sedges, including corn, although figured individual (collected by Eric Quinter) was boring in iris. Setal plates enlarged, darkened, especially that which follows spiracle. Larva of Strawberry Root Borer (Amphipoea interoceanica) [Strawberry Cutworm] an increasingly significant pest of strawberries, especially throughout the Canadian prairie provinces; presumably close in appearance.

Yellow Sedge Borer

Capsula subflava (= Archanara subflava) Western Canada to Nova Scotia, south to at least southeastern Virginia and Great Lakes states. One generation with mature caterpillars from late spring through midsummer. Locally common. Bulrushes (Scirpus), including softstem bulrush (Scirpus tabernaemontani), and rushes (Juncus) (Eric Quinter, unpubl. data). Among most slender and elongate noctuid borers.

American Crescented Borer Helotropha reniformis

Transcontinental in Canada, south in East to Virginia and Great Lakes states. One generation with mature caterpillars from late spring into August. Local and uncommon. Grasses and sedges, including corn. Trailing edge of anal plate upcurved.

Borers Subfamily Noctuinae: Tribe Apameini 409 Hop Vine Borer Hydraecia immanis

Manitoba to New Brunswick, south to Virginia and Great Lakes states. One generation with mature caterpillars from late spring through midsummer. Local; declining eastward. Hop, grasses including corn, Silphium, and others. Violet bands ringing body, these interrupted by pale middorsal and subdorsal stripes (Godfrey 1987).

Newman’s Borer

Meropleon ambifusca [Newman’s Brocade] North Dakota to southern Ontario, south to coastal Georgia and Texas. One generation with mature caterpillars from midsummer to late summer. Local, generally uncommon. Paul Johnson and Eric Quinter have found larvae feeding within big bluestem (Andropogon gerardii). Head small, blackened spiracle on A8 elevated above others, transverse dorsal plate over A9.

Phragmites Borer

Rhizedra lutosa [Large Wainscot] Ontario and New Brunswick, south through Chesapeake region and Ohio; native to Europe. One generation with mature caterpillars in midsummer. Locally common. Bores in lower stems and rhizomes of common reed (Phragmites australis).

Reed Canarygrass Borer Xylomoia chagnoni

Saskatchewan to Nova Scotia, south into North Carolina and Colorado. One generation with mature caterpillars in late spring. Generally scarce, but often locally common. Reed canarygrass (Phalaris arundinacea); native foodplant is unknown. Note subdorsal plate on A9.

410 Borers Subfamily Noctuinae: Tribe Apameini

stalk borers Papaipema

This handsome genus contains more than 50 described and at least a half dozen undescribed species in our region. While the genus is often, and not incorrectly, thought of as characteristic of prairies, its global center of diversity apparently was in the Northeast, along the New Jersey-New York-Connecticut corridor. The genus appears to be speciating rapidly as evidenced by the number of species that are known to be geographically localized and those that are scarcely differentiated either phenotypically or genitalically. As might be expected of a large genus with specialized habits, a number of species appear to be slipping toward extinction. Close to a third of Connecticut’s 30 species have not been seen in more than three decades; several of these are thought to be extirpated and six receive legal protection (CT DEP 2010). The situation in northern New Jersey mirrors that of Connecticut with more than a third of the state’s historical fauna believed to be extirpated. We provide images for nine species below and additional eastern species are figured on the website associated with this guide (see the link at http://press.princeton.edu/titles/9420.html). Crumb’s (1956) larval key treats nearly two dozen species, including all common eastern members of the genus. The caterpillars are distinctively pigmented with broad, whitish middorsal, subdorsal, and subventral stripes that cut through their maroon, pink, purple, or chocolate-brown ground color; dark pigments are heavily expressed on the first four abdominal segments where the patterning may obliterate the upper stripes. A few exceptional taxa (Meadow Rue Borer, P. unimoda, and P. beeriana) are ringed rather than striped. Patterning across the genus is most evident in early to middle instars; penultimate and ultimate instars become increasingly “fatty” white and the markings obsolescent (compare images for Northern Burdock Borer, P. arctivorens, next page). Many characters given below were extracted from Crumb’s (1956) larval key. Identities often can be deduced from foodplant associations (see pages 414–16), characteristics of the tunnel, and by making comparisons to images in this guide. However, final determinations are best based on reared adults; this is because the genus contains numerous, closely allied taxa, with the caterpillars somewhat undifferentiated in character, and because a few members of the genus are generalized in diet (and could turn up in any host). Decker’s (1931) detailed life history for the Stalk Borer (P. nebris), an occasional agricultural pest, applies generally across the genus. Adults lay eggs on and in the vicinity of the larval host. First instars bore into a side branch or the central stalk. A few excavate new, aboveground shoots (e.g., P. araliae and P. furcata), but most tunnel downward toward or into the roots. Frass is pushed out of the tunnel and accumulates about and below the entrance. A few species induce the formation of galls. The majority make their tunnels just above the soil surface; but many tunnel into roots, rhizomes, rhizoids, and other belowground tissues. Prepupae aestivate for one to two

months, especially southward where maturation is delayed into the fall months. Pupation occurs within the tunnel or in nearby soil, and lasts about four weeks. Adults emerge late in the season with most species flying from the second half of August into early October, but into November and December in the Deep South. Larvae and/or larval damages are usually more reliably located than are the adults, especially once one learns each hostplant species’ symptomatology: wilted or flagged shoots, necrotic stems and leaves, weakened stems, galled tissue, etc. Frass accumulation below or about the tunnel entrance can be used to pinpoint those stems or roots with actively feeding larvae. While the adults come to light, and less commonly to bait, many collecting efforts will be thwarted by the cool nighttime temperatures that are the norm during the fall when adults are on the wing. Furthermore, some species fly late, after midnight. Papaipema are thought to be parochial in nature, seldom straying far from colonies of their preferred host. Not surprisingly, light trapping is most effective when traps are deployed in the immediate vicinity of the foodplant. Some success can be had by searching foodplant patches at night with a headlamp (or a small flashlight held at eye level) and watching for the eye shines of the adult moths. A few members of the genus are generalized in diet: the Stalk Borer (P. nebris) feeds on more than 100 different plant species. However, most are foodplant specialists that feed in only one or a few closely related foodplants in nature. In the laboratory, generalist and most specialist Papaipema can be reared on carrots, potatoes, and sweet potatoes. Acceptance of such artificial hosts can be accelerated by introducing the caterpillars into holes bored in the root tissue (e.g., with a drill bit or Phillips screw driver) of a diameter roughly equal to that of the larva. Artificial diets, available from BioServe® and other biological supply companies will work for most members of the genus. The interested reader is encouraged to consult Jones (1928–1932), Kwiat (1916), Bird (1944), Hessel (1954), and Schweitzer et al. (2011) for more details on this fascinating and attractive genus of moths.

! Pitcherplant Borer Moth (Papaipema appassionata). !

Borers Subfamily Noctuinae: Tribe Apameini 411

Northern Burdock Borer Papaipema arctivorens ! recognition Ground color mostly smoky brown forward of A5. Subdorsal stripes obliterated by brown to purplish pigment on A1–A4. Pale middorsal stripe running length of body, constricted through prothoracic shield. Setae borne from shiny brown plates, some of which are greatly enlarged. Head with black line running back from eyes that more or less joins darkened lateral margins of prothoracic shield. Dorsal pinacula conspicuously enlarged on A8. Anal plate caramel-colored, blackened laterally. Spiracles not twice as high as broad (compare with Burdock Borer, P. cataphracta). Anterior prolegs with 11–16 crochets. Larva to 4 cm. The Burdock Borer co-occurs with the Northern Burdock Borer over much of the East and shares some of the same hosts. Its pale middorsal and subdorsal stripes run uninterrupted along the length of the body. ! occurrence Agricultural fields, beaches, farms, and other open, weedy habitats from Minnesota to Quebec, south to Georgia and Missouri. One generation with mature caterpillars from June into early August. Common. ! common foodplants Burdock, teasel, thistle, and presumably other members of aster family. ! remarks The Northern Burdock Borer is one of few Papaipema that initially tunnels into an upper shoot tip. The larva bores down into the stem, but does not tunnel into the root as is typical for Burdock Borer (P. cataphracta) (next page). External signs of the larva can be inconspicuous because only occasionally will bored shoots flag or discolor. Thistle tends to show damage more than burdock. The entrance hole, often as high as half meter above the ground, is left open. Like other Papaipema, larvae can be found by first locating frass deposits that collect on the ground below the tunnel entrance. Papaipema larvae lose much of their coloration over the course of the last instar, in part because their bodies become densely packed with fat (as in the fully mature larva shown in the inset). The fatty nature of Papaipema is a chronic problem for collectors who sometimes go to lengths to extract fat (e.g., in acetone, xylene, or white gas) to keep their adult specimens from “greasing up.” While many Papaipema have become rare, P. arctivorens has fared well, benefitting substantially from introduced burdock, thistles, and teasel that now are major foodplants eastward.

412 Borers Subfamily Noctuinae: Tribe Apameini

Burdock Borer Papaipema cataphracta ! recognition Unusual among our common Papaipema in having pale middorsal and subdorsal stripes continuous and running uninterrupted along length of body (in most Papaipema subdorsal and sometimes middorsal stripes get obliterated by reddish-purple to brown pigmentation on anterior abdominal segments). Broad brown addorsal stripe including both dorsal pinacula. Pinacula flat and well differentiated. D1 pinacula on A1–A3 and L1 pinacula on A1–A6 enlarged, with diameters of D1 pinacula exceeding those of L1 (those immediately following spiracles). Prothoracic shield conspicuously blackened laterally and along leading edge, but the blackened areas not merging in our examples. Spiracles roughly twice as high as broad. Larva to 4 cm. Middle instars are strongly striped (inset). According to Crumb (1956), the larva of the Seaside Goldenrod Borer (P. duovata), a species of East Coast salt marshes and bay shores, is patterned similarly. ! occurrence Fields, waste lots, wetlands, grasslands, glades, and woodlands from Minnesota to Maine, south to Florida and Louisiana. One generation with mature caterpillars from late June into early August. Common. ! common foodplants Aster, burdock, corn, Culver’s root, dahlia, joe pye weed and other Eupatorium, iris, lily, lizard tail, mayapple, sunflower, swamp loosestrife or water-willow (Decodon), thistle, tomato, wild indigo (Baptisia), and many other plants—along with young shoots of trees such as tuliptree. ! remarks The larva enters the main stem well above the ground and then tunnels upward typically; but in smaller hosts the larva may reverse its direction and bore downward, occasionally even tunneling into the root crown. The overall tunnel length can exceed 0.5 m. Caterpillars can sometimes be identified even before they are seen: relative to other Papaipema, the larva extrudes greater volumes of sawdustlike frass from its gallery. Pupation occurs either in the tunnel or in nearby soil.

Borers Subfamily Noctuinae: Tribe Apameini 413 Wild Indigo Borer

Papaipema baptisiae [Indigo Stem Borer] Illinois and southern Canada to North Carolina. One generation with mature caterpillars in July. Very common. Wild indigo (Baptisia) and dogbane; also recorded from dock, cacalia, ironweed, mayapple, and rosinweed (Silphium integrifolium). The closely related P. rutila bores in mayapple.

Sensitive Fern Borer Papaipema inquaesita

Great Lakes states and southern Ontario to Nova Scotia, south to Georgia and Louisiana. One generation with mature caterpillars in July. Among our most common members of genus. Sensitive fern (Onoclea sensibilis), less commonly other ferns.

Columbine Borer Papaipema leucostigma

Rocky Mountains to New Brunswick, south at least into West Virginia and Kentucky. One generation with mature caterpillars in July. Locally common northward and westward; rare southward. Columbine; an occasional pest in gardens (Forbes 1954).

Stalk Borer

Papaipema nebris Southern Canada to Georgia and Texas. One generation with mature caterpillars from July to September. Very common. Recorded from more than 100 forbs and grasses including corn; commonly ragweeds. Higher in stem than many Papaipema. A1– A4 tending to be more brown (rather than maroon). Decker (1931) provides a detailed life history.

Meadow Rue Borer Papaipema unimoda

Alberta to Newfoundland, south in mountains to Virginia and Colorado. One generation with mature caterpillars from June into August. Common. Meadow-rue.

414 Borers Subfamily Noctuinae: Tribe Apameini

life history summaries for eastern PAPAIPEMA Adapted from Hessel (1954) with added annotations from Eric Quinter (in Winter 2000). New hosts and observations have been provided by Tony McBride (TM), Mark Mello (MM), Ron Panzer (RP), Eric Quinter (EQ), David Wagner

(DW), and Jim Wiker (JW). Schweitzer et al. (2011) provide life history data for five rare or declining species: P. araliae, P. astuta, P. cerina, P. duplicatus, and an undescribed species that feeds in fly poison (Amianthium muscaetoxicum).

SPECIES

FOODPLANT

SYMPTOMATOLOGY

POSITION OF POSITION OF LARVA PUPA

aerata

burdock (Arctium), native hosts unrecorded pitcherplant (Sarracenia spp.)

wilted branch

lower stem

tunnel or soil

orange frass

crown, moves to new crown in July in bulb 5–8 cm belowground (TM)

among roots or in sphagnum near plant tunnel

stem tip, new growth

soil

stem

soil

stem; later cell in root high in stem to upper root

soil

lower stem to root crown

tunnel or near root

stem, up to 1 m from ground down into roots

soil, 10–25 cm from plant, 2–5 cm deep

stem; at maturity near ground

stem, sometimes root

stem; perhaps starting in bottlebrush grass and moving to other plants stem (early) to root; usually above break in stem (JW)

tunnel

stem

soil

stem (early) to root

tunnel

appassionata

new species

araliae

arctivorens

astuta baptisiae

beeriana

birdi

cataphracta

cerina

cerussata

circumlucens duovata

fly poison (Amianthium muscaetoxicum)

nonblooming plants; 2–5 discolored central leaves; reddishorange frass plug to one side of leaf whorl (TM) Hercules club (Aralia spinosa) bored new growth; withered leaves; dead, blackened terminal shoot (JW) burdock (Arctium lappa), thistle thistle: branching below crown (Cirsium), etc. or blackened crown; frass below entrance hole horse-balm, stoneroot, or richweed dry stem; white frass (Collinsonia canadensis) wild indigo (Baptisia tinctoria and discolored foliage; frass; holes in stem; often fallen and/or dead B. alba), Indian plantain (Cacalia tuberosa), dogbane (Apocynum stem cannabinum), etc. blazing star (Liatris) wilted tip; 1–3 brown and dry leaves; plants not producing inflorescences (JW) water hemlock (Cicuta maculata), bending or fallen top or branch water parsnip (Sium suave), and at point of entry; multiple sites other umbels. (Warning: these (holes) along stem where frass has plants toxic to handle.) been extruded general feeder, especially burdock plants stunted, drooping, (Arctium), thistles (Cirsium), and discolored; stem swelling; copious crownbeard (Verbesina) frass poorly understood: at least eastern small hole; drying or dry stem; bottlebrush grass (Elymus hystrix) little frass in early instars; turk’s-cap lily (Lilium superbum), mayapple, and other plants later New York ironweed (Vernonia plant stunted; often broken noveboracensis) or bent at larval entrance; top may be much branched (witch’s broom); bore hole often 15–20 cm aboveground hop (Humulus lupulus) elongate or spindle-shaped stem gall seaside (Solidago sempervirens) and several openings in stem; often tan one report from giant goldenrod frass on soil (S. gigantea)

soil

soil, 5–30 cm from root

Borers Subfamily Noctuinae: Tribe Apameini 415 SPECIES

FOODPLANT

SYMPTOMATOLOGY

duplicatus

horse-balm, stoneroot, or richweed (Collinsonia canadensis) button eryngo (Eryngium yuccifolium) joe pye weed (Eupatorium purpureum and likely other purple-flowered species)

wilted leaves on dry stem; sometimes white frass little or none; yellow or dead leaf on shoot without inflorescence leaning stem usually still living; slightly wilted crown leaves on hot days (EQ); sometimes frass

green, white, and black ash (Fraxinus pennsylvanica, F. americana, and F. nigra), and probably others cow parsnip (Heracleum maximum and H. lanatum) and angelica (Angelica atropurpurea)

dry branch or blackened tip of shoot; hole to burrow in older growth; most feeding believed to be in canopy various: yellow or wilted leaf; bore holes along stalk; drooping umbels; chlorotic or browning plants, or sometimes leaning stems (TM) large opening; frass accumulation can be modest

eryngii eupatorii

furcata

harrisii

impecuniosa

inquaesita

leucostigma

aster (Aster laevis (DW), A. puniceus, A. umbellatus, A. novae-angliae) and sneezeweed (Helenium autumnale) sensitive fern (Onoclea sensibilis); older larvae at least occasional in ostrich fern columbine (Aquilegia canadensis and others)

limpida

Missouri ironweed (Vernonia missurica) (EQ)

lysimachiae

loosestrife (Lysimachia quadrifolia and L. terrestris)

marginidens maritima

biology unknown giant sunflower (Helianthus giganteus); other species in prairies general feeder, especially ragweed (Ambrosia artemisiifolia) and burdock (Arctium)

nebris

necopina

nelita

nepheleptena

sunflower (Helianthus decapetalus, H. strumosus, H. tuberosus (TM), and probably others) and Indian plantain (Cacalia tuberosa) cutleaf coneflower (Rudbeckia laciniata), burdock (Arctium), and probably other composites turtlehead (Chelone glabra) and blue vervain (Verbena hastata) (JW)

POSITION OF POSITION OF LARVA PUPA stem (early) to root leaf or stem (early) to root stem to root; older larvae at or belowground level first in new growth, later in shoot near wood

root

leaf petiole or flower rachis to root crown

soil

lower stem to root

base of stem or root

yellow, brown, to dry stem; hole at entry; orange frass

stem (early) to root

root

wilted to entirely withered and dead foliage; inconspicuous frass about crown (DW) very small bore hole near base of plant; base of plant may be slightly swollen or galled (JW) yellow, brown or dry stem; brown withered leaves, and later tilted or fallen stems (DW)

rhizome, often well away from crown base of stalk, tunneling into root crown stem (early) to root

soil

frass and gall-like swelling; stem may be broken at top of tunnel swellings or galls; holes in stem

stem gall

base of stalk (in gall) tunnel or soil

elongate enlargement or cigarshaped gall at base of stem

lower stem; often tunneling upward from entry hole stem to root

tunnel in root tunnel at base

soil

tunnel or soil nearby (JW) soil: 1–7 cm deep, 5–38 cm from plant

root tunnel or soil

gall

base or root

soil

white frass; on vervain, plant somewhat wilted; exit hole with frass below

stem to root

plant or soil

416 Borers Subfamily Noctuinae: Tribe Apameini SPECIES

FOODPLANT

polymniae

leafcup (Polymnia uvedalia)

pterisii new species

rigida

rutila

sciata silphii speciosissima

stenocelis sulphurata

unimoda

SYMPTOMATOLOGY

irregular swelling; boring length of stem; bore hole at 15–20 cm (JW); small frass plug brackenfern (Pteridium aquilinum) yellow or brown frond; orange frass ostrich fern (Matteuccia slightly necrotic to dead frond; struthiopteris) sometimes frass, but difficult to locate; bored stipes easily break free (DW) oxeye (Heliopsis helianthoides), holes in stem near base; slight sunflower (Helianthus spp.), swelling; frass golden alexanders (Zizia aurea), and likely others mayapple (Podophyllum peltatum) yellow leaf (early in season) and much frass; bore hole high on stem but below leaves (JW) Culver’s root (Veronicastrum dry, black, dead, sometimes broken virginicum) stems; frass prairie dock, rosinweed, and cup brown leaf or two; frass plant (all Silphium) yellow (June) to dry (Aug.); frond cinnamon fern (Osmunda bends as larva matures; frass cinnamomea), interrupted fern (O. claytoniana), but especially royal fern (O. regalis) chain fern (Woodwardia virginica) browning frond; pile of orange frass at base of frond swamp loosestrife (Decodon exit hole; often broken stem; verticillatus) and dock (Rumex) scattered (wet) frass on ground (MM) (MM, DW) Meadow-rue (Thalictrum) slightly dwarfed plant; rarely yellow or bending stem; hollow blackened stem breaking easily when pulled

Pitcherplant Borer Papaipema appassionata

Southern Canada and Great Lakes states south into the Carolinas along Coastal Plain. One generation with mature caterpillars in July. Generally rare, but locally common in many bogs. Pitcherplants (Sarracenia). Infected plants are usually killed and pitcher plant mortality can be high in some populations, to the extent that the insect seems to be among few in the temperate zone that play a major role in regulating their own population numbers. Adult figured on page 410.

POSITION OF POSITION OF LARVA PUPA midstem to base

soil

stem to upper root base of frond and “crown”

soil near root

stem (early) to root

soil

root

soil

root

soil

root

soil

stem (near tip of frond) to root

soil

frond into rhizome stem, mostly 15–45 cm above soil tip of stem (early) to root

soil

soil near root

in stem

soil, 1–8 cm deep, 2–38 cm from plant

Borers Subfamily Noctuinae: Tribe Apameini 417

Elder Shoot Borer Achatodes zeae ! recognition Mostly unpigmented with large, round, shallowly convex, shiny brown-black pinacula. A10 strongly sclerotized and coarsely roughened; posterior margin armed with three pairs of rearward-projecting teeth (outer pair apparently reduced in caterpillar figured here). Head and prothoracic shield shiny black. Unpigmented portions of integument smooth, shiny, whitish, perhaps due to subcuticular fat deposition. Spiracles with thick anterior rim. Larva to 3.5 cm. ! occurrence Fields, riparian, and other wetland habitats from southern Canada to Florida and Texas. One generation with mature caterpillars in late spring and early summer. Generally uncommon. ! common foodplants Elderberry commonly; Godfrey (1972) and older literature also report corn, dahlia, and grasses. ! remarks Elderberry is the primary host but like many apameines, larvae occasionally end up in other plants. The caterpillar, also known as the spindle worm, tunnels into new shoots of elderberry. In early June search near the bases of the year’s new growth, and look for a small hole through which the frass and feculae are ejected and particles of frass collecting on leaves and ground. Older larvae bore downward into the previous year’s growth. In Connecticut, caterpillars are mature in June and early July; the adult flies principally in July and August. The extreme sclerotization (hardening) and melanization (blackening) of the terminal abdominal segment is convergent with numerous other, taxonomically unrelated, insects that bore in stems. The urogomphus (sclerotized projection or horn on the terminal abdominal segment) of many beetle larvae is similar in form and presumably function. Perhaps the urogomphus allows the larva to hold its position within the stem when it is boring—according to Eric Quinter, apameines that tunnel in woody hosts have more developed urogomphi. No doubt, the sclerotization also serves to protect the rear of the body: the hardened plate plugs the tunnel, protecting the caterpillar’s rear end from attack by a wide range of tunneling predators, parasitoids, and other enemies. Such may be especially important during defecation because many apameines back out of their tunnels to push away frass and release feculae. Silver (1933) published on the biology of the Elder Shoot Borer. The egg overwinters.

418 Divers Subfamily Noctuinae: Tribe Arzamini

Divers

Family Noctuidae: Subfamily Noctuinae: Tribe Arzamini This taxonomically isolated and distinctive tribe is represented in North America by a single, semiaquatic genus, Bellura. Eric Quinter, an authority on the genus, recognizes six eastern species: B. obliqua and B. brehmei (sensu Forbes) feed on cattail (Typha); B. densa feeds on pickerelweed (Pontederia) and the three introduced water hyacinths (Eichhornia); B. anoa feeds on arrowhead (Sagittaria); B. gortynoides feeds on yellow pond-lily (Nuphar); and B. vulnifica (= B. melanopyga = B. diffusa) feeds on white waterlily (Nymphaea) (Forbes also lists yellow pond-lily for this species). Our application of the names anoa and brehmei is tentative. Likewise, ranges given below are incomplete, based principally on the collection and

experiences of Quinter with adjustments by Don Lafontaine. Eggs are laid on above-water plant tissue in all species; upon hatching the caterpillars tunnel into submerged leaf or stem tissues. Bellura gortynoides mines leaves into the third instar (Levine and Chandler 1976). At least some Bellura are gregarious when young. Once fully fed, Bellura tunnel out of their host, float to the surface, and then swim to shore. Swimming caterpillars whip the abdomen from side to side in much the same way as does a water snake. Upon reaching the shore, the caterpillar seeks shelter beneath refuse, logs, and other objects, typically carving out a small cell in soil or downed, rotting wood, where it will wait out the winter. As necessary, such overwintering caterpillars can move to higher ground to escape flood waters. In May the larva pupates in the overwintering cell.

Pickerelweed Diver Bellura densa [Pickerelweed Borer] ! recognition Elongate, greasy brown caterpillar with smooth, shiny integument; brown band forming where adjacent abdominal segments telescope into one another. Prolegs reduced. A9 much smaller than A8. Spiracles raised and facing rearwards. Ground color lighter below. Head orange-brown, darkened along ecdysial lines. Prothoracic plate somewhat thickened and darkened. Each body segment with transverse dorsal crease rearward of midsegment. Larva to 4 cm. ! occurrence Ponds, ditches, and other bodies of water from at least southern Ohio to Florida and Texas; records from farther north may refer to a second entity (see below). One generation with mature caterpillars from October through May. Common. ! common foodplants Pickerelweed and water hyacinth. ! remarks According to Don Lafontaine (in litt.), as presently sorted in collections, B. densa has two DNA barcode types; haplotypes of individuals from north of Ohio sort with those of Cattail Diver (B. obliqua). Until a revisionary study of the genus is completed, identifications, ranges, and hostplant records for Bellura should be used with caution.

Divers Subfamily Noctuinae: Tribe Arzamini 419 White-tailed Diver Bellura gortynoides

Wisconsin, southern Ontario, and Maine south to Florida; distribution uncertain due to confusion with similar species. One generation northward with mature caterpillars from late fall into spring (fully fed larvae overwinter on shore). Possibility of second generation southward, as suggested by Forbes (1954), needs confirmation. Common. Yellow pond-lily (Nuphar).

White Waterlily Diver Bellura vulnifica (= B. melanopyga = B. diffusa)

At least Manitoba to Newfoundland, south to Florida (Eric Quinter, unpubl. data). One generation with mature caterpillars from September through May. Common. Our image of initial larval mine in host, white waterlily (Nymphaea odorata).

Cattail Diver

Bellura obliqua [Cattail Borer Moth] Transcontinental in Canada, south in East to Florida and Texas (although species limits, and thus ranges, should be regarded as tentative until the genus has been revised). One generation with mature caterpillars from October through May. Common. Cattail. Our identification is tentative—this caterpillar did not produce an adult.

420 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

Pinions, Sallows, and Kin Family Noctuidae: Subfamily Noctuinae: Tribe Xylenini

There are about 194 species of Xylenini found north of Mexico, about half of which occur in our region. The tribe is diverse in the Northeast and southern Canada. Forests, barrens, bogs, heathlands, and other acid-soil habitats are rich in pinions and sallows. The caterpillars often have round, clearly delineated, white dorsal pinacula (especially in the pinions, Lithophane). Most have a broad spiracular stripe. Early instars tend to be green with middorsal, subdorsal, and broad spiracular stripes. Later instars are commonly green or rendered in grays, brick reds, browns, black, and other earth tones. Many pinions (Lithophane) and some others may exhibit a marked change in color between the penultimate (fifth) and final (sixth) instars. Green instars rest on leaves; the last instars, rendered in grays, browns, and other earth tones, rest on tree trunks or beneath litter, and move back onto foliage under cover of darkness. The caterpillars are new leaf specialists and rank among the most ubiquitous and ecologically significant elements of the spring caterpillar fauna in eastern woodlands and forests. Caterpillars of several pinions (Lithophane) are facultatively predaceous on other insects, especially other caterpillars and pupae (Sanders and Dustan 1919, Schweitzer 1979a). Pupation occurs belowground (except in a few Lithophane) in a cell lined with a film of silk. All are univoltine. Xylenines include the bulk of our winter moths. Among the 71 species illustrated here, all but the Red-winged Sallow (Xystopeplus rufago) emerge in the fall.1 Many mate and lay eggs in the fall and early winter and then die; but others, e.g., all pinions (Lithophane), wait until the first warm nights of late winter or early spring to engage in reproductive activities. The spermatophore is substantial—one can often feel if a female has mated simply by squeezing down gently on the abdomen, and in some cases even determine if she has mated more than once. Pinions and sallows “shiver” prior to taking wing on cold evenings, a behavior that warms the flight musculature of the thorax; the flight muscles are rapidly stretched and contracted but in such a way that no lift is provided by the wings and the moth is able to remain in place. “Shivering” pinions are capable of raising their internal thoracic temperatures as much as 15–20° C (~25°–35° F) above ambient temperatures! Once sufficiently warmed, the moths lift off and begin their searches for mates, food, or suitable oviposition sites. Xylenine adults are attracted to sugar and wine baits (see introductory sections). Baiting is particularly effective in late winter before red maples have bloomed. On a good night in the Northeast, one might see several Eupsilia and Lithophane (especially, L. hemina, L. bethunei, L. grotei, and L. patefacta); 1 The Red-winged Sallow is unusual among temperate moths in that two life stages are known to overwinter; some individuals pass the winter as a recently eclosed adult (only a few), but most diapause as a pupa.

one to three species of Pyreferra; and in some areas, Xystopeplus, Sericaglaea, and Xylena. Under natural conditions in late winter and early spring, these same moths would be feeding at wounds on maples and birches, e.g., where there has been winter breakage of limbs or at sites where squirrels have gnawed through the bark to get a taste of sap. Red maple and willow blossoms provide much of the diet later in the spring. (The common name “sallow” derives from the association of xylenines with willow flowers—in Europe willows are widely known as sallows.) Autumn food sources are poorly documented but include asters, goldenrods, and other late-blooming flowers, as well as damp, senescing leaves. Gravid females readily release eggs if confined, especially if they have been fed. On the whole, xylenine caterpillars are decidedly nocturnal—nighttime efforts may be the only means of securing last instars of many “glaeas” (page 456). Larvae must be supplied with new foliage or they will starve. Sleeving is often the most efficient option for rearing. Nearly all of the general feeders can be raised on black cherry. Provide mature larvae with about 4–5 cm of moist peat for pupation. We use cups as large as 8 to 16 ounces for single larvae. Mortality will be high if the peat dries out. Keep aestivating larvae and especially pupae below 85° F (30° C). A basement is an ideal location. Reared larvae sometimes produce adults in July or August, perhaps due to malnutrition. Over much of our area, pupation is in August or September and adult emergence follows about a month later. Members of the subtribe Antitypina aestivate as pupae. Eggs kept indoors, even in an unheated garage, may hatch too early to be reared successfully—hold them outdoors in a shaded location so that their hatch will be synchronized with leaf out.

! The Bethune’s Pinion (Lithophane bethunei) larva shown here is consuming a scale cluster on walnut. !

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 421

Acadian Swordgrass Moth Xylena thoracica ! recognition Elongate orange-brown with spiracular stripe and faint chevrons over dorsum. Thin, pale middorsal and subdorsal stripes. Prothoracic shield with darkened anterior edge. The dorsal setae (D1 and D2) arise from off-white bases. Narrow spiracular stripe runs through spiracles and continues down anal proleg. Body paler below spiracular stripe, and particularly so on thoracic segments. Spiracles tan with narrow black rim. Head orange-brown, smooth, shiny, and mostly unmarked except for mottling over each lobe. Larva to 6 cm. Penultimate instar waxy blue-green with well-defined white middorsal, subdorsal, and spiracular stripes and considerable white speckling (inset). ! occurrence Barrens, bogs, and open grassy areas. Transcontinental in Canada, south in East to Maine, New Hampshire, and northernmost New York. One generation with mature caterpillars from June through at least July. Locally common. ! common foodplants Alder, birch, blueberry, cherry, currant, dogwood, oak, poplar, rose, spruce, and willow, presumably many other woody plants; heaths often favored. Ours reared on young cherry and oak leaves. ! remarks Swordgrass moths are largely northern in distribution—evidently none occur south of Pennsylvania in the Appalachians. Eggs are laid in large rafts. Initially the larvae, like many trifid Noctuidae, are loopers—the first two pairs of prolegs are not used in walking. Prepupal swordgrass moth larvae form cells belowground and aestivate for a month or more before pupating. Adults of the genus can be abundant at bait, especially northward, and usually much more so in the spring than the fall.

422 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

Dot-and-dash Swordgrass Moth Xylena curvimacula ! recognition Smooth, large, yellow-green to rusty brown cutworm with faint to strong dorsal chevrons. Pale middorsal line edged with smoky green to black, absent over prothorax and A8; obscure subdorsal stripe composed of disconnected minute spots. Oblique, dorsal lines meeting over midline at rear of abdominal segments A1–A8, sometimes forming weak spot at rear of segment. Ground color significantly paler below thin, green and white, or brown and white spiracular stripe. Venter sometimes tinged with pink. Spiracles orange-brown with black outer rim in our material; Forbes (1954) examined larvae with white spiracles. Prothorax somewhat swollen with modest, reticulate brown patterning. Pale subdorsal area on A8. Head pale orange-brown with dark mottling over each lobe. Larva to 5 cm. Caterpillars of Gray Sword Moth (X. cineritia) similar, but lacking dark (green or brown) edging above spiracular stripe that is present in most forms of Dot-and-dash Swordgrass Moth (Crumb 1956). Waxy blue-green penultimate instar (lower right) resembling Speckled Green Fruitworm (Orthosia hibisci). ! occurrence Almost any kind of forested or brushy habitat. Transcontinental in Canada, south in East to New Jersey (rare) and Great Lakes states. One generation with mature caterpillars in June and July. Most common Xylena across most of our region. ! common foodplants Alder, aspen, birch, cherry, meadowsweet, oak, poplar, and willow. Dandelion listed by Crumb (1956). Our captive larvae fed on young American hornbeam and black cherry leaves. ! remarks Caterpillars of this genus apparently forage primarily on low shrubs and small trees. The caterpillars are more cutwormlike in habit than those of the American Swordgrass Moth (X. nupera), which are more likely to remain on vegetation. Caterpillars of Xylena have a more pronounced blue-green aspect and often bear a hint of dark edging above the spiracular stripe.

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 423

American Swordgrass Moth Xylena nupera ! recognition Last instars range from green to brown or black. Proleg on A3 somewhat reduced in size; that on A4 slightly smaller than those that follow. White to pale yellow middorsal and subdorsal stripes well defined from T2 to A9. Three prominent white spots (setal bases) over dorsum of T2–A8; A9 with only two white dorsal spots. White to yellow spiracular stripe edged with black above, running along lower edge of spiracles, broadened on A7–A9. In green forms, ground color darker above spiracular stripe; in brown forms, ground color between subdorsal stripes sometimes darker than that below. Spiracles often orange. Head green to orange-brown, smooth, shiny, without brown reticulation and spots. Crumb’s (1956) account suggests that markings on head capsule are more weakly expressed than those of other swordgrass moths. Larva to 5.5 cm. Middle and penultimate instars green. ! occurrence Open grassy, sedgy, or shrubby habitats such as bogs, fens, heathlands, pine barrens, floodplains, and, less commonly, mesic woodlands and forests. Transcontinental in Canada, south in East to southern New Jersey (where it is the only resident Xylena), Pennsylvania (mountains), Illinois, and other Great Lakes states. One generation with mature caterpillars mostly from June through middle July. Common northward. ! common foodplants Generalized in diet, eating grasses, forbs, and woody plants. Forbes (1954) mentions young cherry as a favorite, but notes that larvae may drop from host and mature on grasses and other plants. Other hosts include apple, blueberry, cranberry, iris, poplar, rush, spirea, sweet gale, and willow. ! remarks While the American Swordgrass Moth is generally considered to be uncommon, it can be locally abundant in bogs and northern barrens, and can be an occasional pest of cranberry (Landry et al. 2002). Adults emerge in late August and September northward and October southward, fly through the fall, and then reappear from February to May. While swordgrass moths are commonly seen at lights (especially mercury vapor lights), their numbers are better assessed with baits. A European counterpart (X. vetusta) visits willow blossoms in the spring (Porter 1997). Females deposit their eggs in masses of a hundred or more. Larvae are active early in the year and can do great damage to unopened buds. Cranberry growers refer to the species as the False Cutworm. Caterpillars do well when offered a variety of “forage.” Our lab-reared larvae of X. nupera remained as prepupae well into August.

424 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

American Brindle Lithomoia germana ! recognition Last instars mostly brown to nearly black with broad white spiracular stripe and conspicuous white dorsal setal bases. Dorsum frequently with oblique dark patches that form chevrons over abdomen; dark patches best developed on A1, A2, A7, and A8. Middorsal, subdorsal, and supraspiracular stripes, progressively weakened. Posterior portion of A8 somewhat raised; A9 also somewhat swollen. Subdorsal stripe continued through prothoracic shield. Spiracular stripe passing along lower edge of black spiracles and then dropping down outer face of anal proleg. Prolegs on A3 and A4 somewhat reduced. Head shiny orange-brown with pale snowflake spots over each lobe and black patch to either side of triangle. Larva to 5 cm. ! occurrence Fields, meadows, bogs, heathlands, woodland and forest openings of Canadian and Hudsonian life zones. Transcontinental in Canada, south in East to northern New Jersey, Pennsylvania, and Great Lakes states. One generation with mature caterpillars mostly from June through mid-July. Common northward. ! common foodplants Alder, basswood, birch, blueberry, Labrador tea, poplar, spirea, willow, and presumably other shrubs and small trees (see also below); ours reared on black cherry. European host records for its congener and our collection records suggest members of Ericaceae (heaths) are favored. ! remarks The American Brindle was formerly thought to be conspecific with the European Goldenrod Brindle (L. solidaginis). American Brindle larvae may be common in sweep samples from heathlands and fields with low shrubbery in late May and early June. In New England, we associate Lithomoia with open (northern) habitats with an abundance of spirea and lowbush blueberry. The moth is among the earliest-flying members of the tribe and even southward few adults persist past early October. Look for adults nectaring on goldenrod and other flowers in late August and September. They also come to bait and light. Similarities in larval characters indicate that swordgrass moths (Xylena) and brindles (Lithomoia) are related. Captive females oviposit freely if provided with dried grass stems but lay few eggs otherwise. Our females inserted their eggs in grass sheaths. The egg overwinters.

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 425

Goat Sallow Homoglaea hircina ! recognition Chocolate, pinkish, or gray-brown with sprinkling of minute pale spots; dark prothoracic shield cut by white subdorsal lines. Middorsal stripe nearly obliterated; A5–A8 often with dark middorsal spot toward anterior end of each segment. Faint, yellowish subdorsal stripe broken through abdominal segments; subdorsal stripe conspicuous, nearly pure white where it cuts through prothoracic shield and anal plate. Spiracular stripe cream to yellow or pinkish orange, running along lower edge of black-rimmed spiracles, although dipping well below spiracle on A8 and occasionally also on A7. Lower part of abdominal prolegs frosty. Head orange with brown patterning and brown patch to either side of midline that drops down and runs along either side of triangle; Crumb (1956) mentions dark area above each antenna. Larva to 3.5 cm. Middle and penultimate instars apple green with white middorsal, subdorsal, and spiracular stripes; the last twice width of other two (inset). Mature larva superficially similar to the Lost Sallow (Eupsilia devia) and Red-winged Sallow (Xystopeplus rufago). ! occurrence Woodlands and forests. Transcontinental in Canada, south in East to New Jersey, Pennsylvania, and Great Lakes states. One generation with mature caterpillars in late spring. Common northward. ! common foodplants Aspen and poplar; hosts outside of Salicaceae listed by Prentice (1962) are exceptional and likely incidental or in error. ! remarks The caterpillars spin a crude shelter and feed within, up until the last instar. The brown color of the last instar suggests that it shelters on bark or below leaf litter by day. The Goat Sallow emerges in the fall but hibernates soon after eclosion. Adults begin flying as early as February in Connecticut. Although the moths are often attracted to lights, they tend to seek shelter in folds and other nearby dark areas. They also come to bait. Like other spring-active noctuids, adults shiver to generate heat within their thorax prior to flight on cool evenings.

426 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

pinions Lithophane

This is a large and successful group with more than 30 eastern species. Identification of the adults can be challenging, although male genitalia are reliable for most species (see Forbes 1954). The rate of phenotypic (observable) evolution differs between life stages: adult wing patterns (and habits) are largely similar across the genus, while larval coloration and behavior may differ substantially between related species. Lithophane caterpillars are a remarkable case study in background matching and natural selection; within a single genus, there are species that mimic deciduous foliage (mostly green with subtle markings), pine foliage (longitudinally striped with white), and cedar foliage (white-spotted). Members of the Nameless Pinion group (L. innominata group = L. semiusta through L. oriunda in Forbes 1954), match leaves as middle instars but the last instars (variegated in browns and grays) resemble bark. A divergent scheme has been adopted by the Shivering Pinion (Lithophane querquera); its white- and yellow-banded body appears to mimic a sawfly, or perhaps the Mustard Sallow (Pyreferra hesperidago) and its congeners. Many of the observations and foodplant records in this section are those of DFS, who worked on the genus as part of his doctoral studies. Pinion caterpillars may be recognized by the round, clearly delineated, and often whitened dorsal spots (pinacula); a third, smaller spot is sometimes evident anteriorly, above the subdorsal stripe, although it is commonly hidden in an intersegmental fold. Many have a broad, pale, lateral stripe; this is referred to as a spiracular stripe here but some call it a subspiracular or subventral stripe because its upper edge may lie just beneath the level of the spiracles. Although many larvae can be identified to species with this work, especially if the host is known, identifications are best based on reared adults. With the exception of Lemmer’s Pinion (L. lemmeri) and a few relatives, all pinions have six instars. Coloration may differ markedly between instars. As noted above, middle to penultimate instars of the Nameless Pinion group are pale green with white spots and stripes, whereas their corresponding last instar tends to be rendered in grays and browns with black spots and chevrons. Middle to penultimate instars of this lineage sit on the underside of leaves by day, whereas the last instars rest on bark or under litter until nightfall. While some species are polyphagous on woody plants, most are somewhat specialized, so be sure to make an effort to identify and record hostplant information for the caterpillars that you find. And be forewarned that bark resters sometimes wander onto nonhosts. Caterpillars of several pinions are facultatively predaceous on other insects, especially the caterpillars and pupae of Lepidoptera. The larva shown to the right is consuming a geometrid caterpillar (see also page 420). Additional life history notes appear above in the introduction to the tribe (Xylenini), and below in the individual species accounts.

The most reliable means to acquire pinion livestock is to bait for mated females in late winter to early spring and then hold these for eggs. Gravid females of a few Lithophane, e.g., Grote’s Pinion, will be encountered as early as January in New Jersey. The best nights are mild evenings (above 10 C° or 50° F at dusk), before red maple has come into flower. If it is early in the season and there is a possibility that females are unmated, secure and feed a couple of male companions too. Most will mate readily if placed in a screen-covered box or cage placed outdoors in a shady place. Keep mated females fed, cool, and provide paper towels or other roughened surfaces on which they can lay their eggs. It is often necessary to hold (and feed) females for a few nights, before they will begin laying. Lithophane lay as many as a thousand eggs. In nature, the eggs are laid singly or in small clusters of two or three; only the Hoary Pinion (L. fagina) lays in rafts. If kept outside, eggs should hatch soon after the foodplant buds open. While young and middle instars can be found by inspecting the lower surfaces of leaves, beating will yield larvae in number. Last instars of the bark resters can be found by inspecting crevices and looking under strips of bark at and below eye level; on smooth-barked trees search under leaf litter in the vicinity of the trunk. Burlap banding works well for bark-resting pinions.

! This Shivering Pinion (L. querquera) has consumed the anterior half of a Fall Cankerworm (Alsophila pometaria). !

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 427

Ashen Pinion Lithophane antennata ! recognition Pale green with broad, yellowish spiracular stripe that runs length of body, then drops down along anal proleg. Integument smooth, shiny. Dorsal white spots circular and well defined. Middorsal stripe thin, creamy, continuous from T2 rearward. Subdorsal stripe white, broken into spots; supraspiracular area also with spots that more or less form a weak stripe. Head pale green and mostly unmarked, shiny; labrum whitened; mandibles black (bottom right). Thoracic legs frequently blackened. Larva to 4 cm. Similar to Russet-spotted Pinion (L. laticinerea) (next page); often confused with middle instars of many other Xylenini and even Hadenini (e.g., Orthosia). ! occurrence Woodlands and forests, especially common in dry oakhickory woodlands. Minnesota to southern Ontario, Quebec, and New Brunswick, south through Carolinas, northern Georgia, and Mississippi. Due to old misidentifications, range limits uncertain; e.g., many southeastern records may refer to L. laceyi. One generation with mature caterpillars in late spring, mostly June in Connecticut. Often common as a larva. ! common foodplants Oak and hickory account for many of our records. Also apple, ash, basswood, blackberry, buckeye, cherry, desert false indigo, elm, hawthorn, leadplant, peach, plum, walnut, willow, and many other woody plants; occasionally nonwoody plants (e.g., feverwort). In addition, other caterpillars (and pupae) are consumed. ! remarks The larva is known to agricultural entomologists as the Green Fruitworm (it is one of several moths to which this name has been applied), because the caterpillar consumes young apples and related fruits (illustration in Holland 1903). This is a poor common name for the animal given that fruits are not often part of its diet and its coloration (“green”) applies across nearly the whole of the Xylenini and many Orthosiini, at least in middle instars. The caterpillar can be aggressive, reeling around and biting at objects that contact its body. It will also hold a threat posture with the head and thorax raised, black mandibles flared, and the legs splayed (right). It is a predatory species, with a proclivity to attack and consume the larvae and pupae of other noctuoids, geometrids, and sawflies that are confined with it. All instars can be found resting on the undersides of leaves; late instars often rest over the midrib.

428 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

Russet-spotted Pinion Lithophane laticinerea ! recognition Waxy, yellow-green caterpillar with prominent striping and spotting, and sometimes shiny integument. Body deeply creased and more roughened in appearance than that of other pinions. Creamy middorsal stripe. White subdorsal stripe of irregular width and expression. Three conspicuous, slightly elevated, white dorsal pinacula over abdominal segments. Thick, poorly delineated, creamy spiracular stripe with spiracles embedded along upper edge. Supraspiracular stripe represented by white spots that become less numerous rearward. Head pale green, shiny, unmarked, modestly roughened, with whitened labrum. Thoracic legs not blackened in our specimens. Straw-colored spiracles with dark rim. Larva to 4.5 cm. Similar to Ashen Pinion (L. antennata) but more strongly marked, and with spiracles located centrally in spiracular stripe. In the Ashen Pinion, the spiracles are situated along the dorsal margin of the spiracular stripe. ! occurrence Woodlands and forests from southern Canada, south at least to Maryland, Missouri, and Nebraska. One generation with mature caterpillars mostly in late May and June. Usually uncommon. Range and abundance unclarified because of moth’s elusive habits and difficulty of adult identifications. ! common foodplants Red and silver maple. Other confirmed hosts include basswood (DLW), crab apple (Rings et al. 1992), and black oak (Schweitzer 1974). Some older reports from ash, aspen, birch, cherry, elm, hawthorn, poplar, and willow may be based on misidentifications. ! remarks Although we find this species to be relatively uncommon, L. laticinerea is one of few owlets known to occasionally defoliate forest trees— specifically red (and silver) maples (Rings 1969, Schweitzer 1974). Drooz (1989) mentions the defoliation of maples on islands of the Penobscot River in Maine. Like many other insect pests, it fluctuates in abundance—in most years we see none or just a few individuals. Like its close relative Grote’s Pinion (L. grotei), which also commonly feeds on red maple, this species mates and finishes flying early in spring. While adults are similar to those of Grote’s Pinion (to the extent that the two are often confused in collections), caterpillars of the two are immediately separable, and thus provide another example of larval (phenotypic) evolution outpacing that of the adults.

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 429

Grote’s Pinion Lithophane grotei ! recognition Large, pale, sea green, with numerous minute white lines and spots; trunk appearing leathery. Stouter than other pinions; broadest through A3–A7, body constricted between segments. Most individuals with faint, white middorsal and subdorsal stripes; broad, yellowish spiracular stripe weakly expressed; yellow tinting sometimes evident where segments overlap. Other individuals more uniformly green with only trace striping (as in that illustrated here). Dorsal setae borne from circular, pale green bases. Prothoracic shield bright green and smoother than adjacent integument. Head sea green, slightly roughened, with or without pale snowflake spots. Thoracic legs green. Spiracles straw-colored with dark rim. Larva to 4.5 cm. Middle and penultimate instars with striping more evident, resembling those of Russet-spotted (L. laticinerea) and Ashen (L. antennata) Pinions. ! occurrence Mostly forests from Wisconsin to Nova Scotia, south to northern Georgia (mountains) and Missouri. One generation with mature caterpillars from late May to early July; among first pinions to mature in spring. Common. ! common foodplants Reported from apple, birch, cherry, maple, oak, and other woody plants, but see below. We have found caterpillars on aspen, red maple, yellow buckeye, and various species of oak. ! remarks If misidentifications and unvouchered records are disregarded, it appears that red maple is the primary foodplant—the mature larva is cryptic on the undersides of its leaves. Slingerland and Crosby (1915) reported Grote’s Pinion as an occasional pest of green apples. Roy Rings, despite years of work on Ohio fruitworms, failed to record this species from fruit trees (see Rings et al. 1992). This distinctive pinion can be among the more predatory (and cannibalistic) members of the genus, although some cohorts, at least when sleeved, appear to be entirely phytophagous. Grote’s and Ashen (L. antennata) Pinions remain on foliage when not feeding, and sometimes feed during the day (on foliage or caterpillars). This may explain why they tend to mature more rapidly than the brown pinions that seek shelter along the trunk, remaining hidden and inactive by day. In southern New Jersey, females of Grote’s Pinion mate in January and February; in some years few or no adults are found after the middle of March.

430 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

George’s Pinion Lithophane georgii ! recognition Waxy to pale blue-green above with well-developed middorsal and spiracular stripes. Thick middorsal stripe white, parallel-sided, constricted over prothorax. Lemon-yellow spiracular stripe thickest rearward. Subdorsal stripe vague, most evident as run of broken spots over A1–A8. Dorsum with abundance of minute white spots, giving upper portion of body frosted aspect. Dorsal setae from porcelain white setal bases. Head green and unmarked or with pale snowflake spots. Spiracles straw-colored with dark rim. Larva under 4.5 cm. Penultimate instar yellow-green, with middorsal, broken subdorsal, and weak supraspiracular stripes (larva shown bottom right is preparing to molt). Amanda’s Pinion (L. amanda) (page 441) is similar but ground color more uniformly blue-green; middorsal stripe thicker through prothoracic shield; spiracular stripe on T1 and T2 often infused with red; and head with frosty white middorsal line splitting and running along sides of triangle. Amanda’s Pinion is also a species of the North. Willow is favored; other recorded hosts include alder, birch, cherry, oak, poplar, rose, and serviceberry (Handfield et al. 1999). ! occurrence Canadian and boreal zone forests. Transcontinental in Canada, south in East to northern New England, upstate New York, and northern Great Lakes states; rarely, and probably temporarily, south to coastal Massachusetts and northern Pennsylvania. Disjunct populations in North Carolina mountains mostly above 1600 meters. One generation with mature caterpillars found in June and early July. Generally scarce, becoming more common northward and westward. ! common foodplants Alder, apple, blackberry, maple, and sweet fern listed as principal hosts by Handfield et al. (1999); with ash, birch, boxelder, buffaloberry (Shepherdia), hazel, and rose added as occasional foodplants. We have found caterpillars on blueberry, blackberry, currant, and pin cherry in Smokies. ! remarks George’s Pinion represents one of many disjunct species in the southern Appalachians. The core of its eastern range is in Canada and the moth is otherwise absent south of northern New York and New England. In June 2001 on Balsam Mountain, North Carolina, George’s Pinion was the most frequent owlet in beating samples from woody, broad-leaved plants. In the Rockies and westward, it can be among the more common Lithophane.

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 431

Dowdy Pinion Lithophane unimoda ! recognition Waxy blue-green over dorsum, giving way to green along sides, with prominent yellow spiracular stripe. Robust and chunky for a pinion. Spiracular stripe, running just under black spiracles, thickening rearward and continuing down anal proleg. Middorsal stripe white and thin. Dorsum with somewhat corrugated or leathery texture (visible with magnification). Dorsal pinacula poorly differentiated relative to those of other pinions. Head sea green, smooth, shiny, unmarked except for white clypeus, labrum, and mandibles. Spiracles black. Larva to 4.5 cm. Penultimate instar with less white speckling or “frosting” (inset); spiracular stripe strong with yellow cast, resembling Ashen Pinion (L. antennata). ! occurrence Fields, orchards, yards, woodlands, and forests. Transcontinental in southern Canada, south in East to Georgia (mountains), Missouri, and Nebraska. One generation with mature caterpillars in June in Connecticut; mid-May to mid-June in New Jersey. Often common. ! common foodplants Cherry, especially wild or black, and pin cherry, but also crabapples, plum, and tupelo; other recorded hosts (perhaps in error) include aspen, maple, oak, and willow (Rings et al. 1992). ! remarks The common name, appropriate for the adult, is an injustice to the caterpillar, which we regard as handsome. Over much of the East, it is among the most common pinions at bait and light, especially in late fall and early winter. We have collected many caterpillars inadvertently while gathering cherry foliage for other caterpillars. Check for larvae by examining leaf undersides of new growth— the caterpillar often will be found extended along a leaf midrib. Gravid females readily oviposit in captivity, especially once fed. To the oviposition container, add a sprig of black cherry and a piece of paper toweling. The caterpillars of most pinions can be reared on cherry. All pinions that feed on deciduous trees require new foliage and will straggle and starve if offered only older leaves. Although this species mates and oviposits relatively late for a pinion, its larvae grow rapidly and are among the first to mature each spring. We once had a Dowdy Pinion caterpillar consume a prepupal Spring Azure (Celastrina ladon) before we could separate the two, but in our experience this species is not usually predaceous.

432 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

Pale Green Pinion Lithophane viridipallens ! recognition Waxy or whitish green with middorsal, subdorsal, supraspiracular, and spiracular “stripes” of small white spots. First eight abdominal segments with three small white dorsal spots. Prothoracic shield waxy blue-green, mostly unmarked. Larva to 4 cm. Caterpillars of Bailey’s Pinion (L. baileyi) (page 441) and White-eyed Pinion (L. tepida) (page 442) similarly marked and perhaps indistinguishable. Both of these, however, are essentially northern in distribution (see Remarks below). Middle instars easily confused with those of Red-winged Sallow (Xystopeplus rufago) and myriad other green xylenines. ! occurrence Shrub swamps, woodlands, and forests from southeastern Massachusetts to Georgia, west at least to Louisiana (and into Texas if L. pruena proves conspecific). Northward it is limited largely to the coastal plain. One generation with mature caterpillars from April (southward) through June. Common. ! common foodplants Lab-reared caterpillars accept blueberry, cherry, fetter-bush or swamp doghobble (Eubotrys racemosa), and holly; our two wild caterpillars from greenbrier and hophornbeam. Presumably a generalist on shrubs and trees. ! remarks The caterpillar rests on the underside of new leaves with its head curled to one side. It is often predatory—rearing sleeves typically produce one or a few large larvae. Even though caterpillars of Pale Green, Bailey’s, and White-eyed Pinions are similar, those of the first can usually be identified by their geographic location. The Pale Green Pinion is a species of the Atlantic Coastal Plain, southern Piedmont, and Gulf Coast; it is common from New Jersey southward. In the Northeast it is local and rare, being confined mostly to shrubby swamps and pine barrens where it occurs with the much more numerous White-eyed Pinion. Bailey’s and White-eyed Pinions are northern species that are confined to mountains south of Connecticut and New York. Bailey’s Pinion is a denizen of northern hardwood forests. The White-eyed Pinion is associated with barrens, woodlands, forests, and wetlands on acidic sandy or granitic soils where heaths are abundant.

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 433

Bethune’s Pinion Lithophane bethunei ! recognition Usually brown to gray with weak to strong charcoal patterning. Dorsum on A1–A7 with backward projecting chevrons as in other members of innominata group; dark patches enlarged, connecting over dorsum on A8 (but not A7). Thin, broken middorsal stripe often infused with red, rust, peach, or yellow; white subdorsal stripe often broken into dashes and spots. Thin, black, wavy, often broken stripe running along upper edge of broad, white to yellowish subspiracular stripe. Body mostly free of pigmentation below. Dorsal setae from small, white pinacula; diameter of D2 pinaculum about a third distance of that between D1 and D2. Thoracic plate with dark quadrangular patch to either side of midline; decidedly paler below subdorsal stripe. Head mahogany with black reticulations and coronal bars. Larva to 4 cm. Penultimate instar green with nearly solid middorsal, dotted subdorsal, and broad creamy lateral stripes (inset). ! occurrence Woodlands and forests from Saskatchewan to Nova Scotia, south to northern Georgia and Missouri. Rare on coastal plain south of Long Island. One generation with mature caterpillars in June and July; among latest Lithophane. Common. ! common foodplants Apple, ash, basswood, beech, birch, cherry, crabapple, cranberry, elm, hawthorn, hickory, hophornbeam, maple, oak, poplar, willow, witch hazel, and other woody plants. Often predatory. ! remarks Adults show up at light and bait early in the spring, and persist until the end of April. The species mates relatively late, usually after the early spring bait season. Caterpillars of Bethune’s Pinion (as well as those of related species) hide in crevices and under loose bark on tree trunks, or they move into leaf litter during daylight hours if the bole of their host lacks good hiding places. The larvae are somewhat predatory, at least in the last instar. DFS observed larvae of Bethune’s Pinion eating tent caterpillar (Malacosoma) pupae in Connecticut— the caterpillar may even rest partially within the cocoon after feeding upon the contents. Sanders and Dustan (1919) observed this same habit and even suggested this species might be more beneficial as a predator than destructive as a fruitworm in Nova Scotia orchards, under some conditions.

434 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

Brown Pinion Lithophane hemina ! recognition Usually gray-blue ground color with strong charcoal markings above level of spiracular stripe. Mottling coalescing into dorsal chevrons that become more apparent rearward to A8. Often with faint, thin, tan to orange-yellow middorsal and subdorsal stripes. Numerous dark spots and striations below level of white to cream spiracular stripe, which includes tan spiracles along its upper edge. Prothoracic shield, or at least that part between subdorsal stripes, black and well differentiated. Head with considerable mottling. Larva to 4 cm. Penultimate instar waxy, green-white, with broken middorsal and subdorsal stripes, and prominent white dorsal pinacula; broad white spiracular stripe (inset of premolt larva). Larva may be inseparable from those of related species, especially the Nameless Pinion (L. innominata) and the recently described L. lanei. The latter species, described from specimens collected in the Ottawa area by Troubridge (2006), presumably occurs more widely across the Northeast. Identifications are best based on reared adults. ! occurrence Woodlands and forests from Manitoba to Nova Scotia, south to New Jersey, Missouri, and in mountains to Georgia. One generation with mature caterpillars from late April through June. Very common northward. ! common foodplants Many records from black birch; also reported from aspen, other birches, boxelder, cherry, crabapple, hickory, oak, willow, and many other woody plants. Our wild caterpillars from big-toothed aspen, black birch, black cherry, hazel, hickory, red maple, red oak, and witch hazel. ! remarks Pinions usually feed from the lower surfaces of leaves, at or very close to the growing shoots. Caterpillars are quick to release their grip and drop if perturbed. Often just blowing lightly on a late instar will send it tumbling to the forest floor. Like the Nameless Pinion (L. innominata), the last instar of the Brown Pinion rests on bark (and probably also in debris at the base of smooth-barked trees). Look for larvae under exfoliating pieces of bark or in furrows. Alternatively, burlap banding can be used to sample larvae of this species and other pinions.

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 435

Nameless Pinion Lithophane innominata ! recognition Light or dark brown to gray or steel blue, and often with middorsal, subdorsal, and spiracular stripes. Middorsal stripe often reddish brown to rusty. Dorsum usually marked with vague, dark chevrons; these darkest rearward. Head brown with darker mottling and coronal bars. Larva to 4 cm. Penultimate instar with broad, creamy spiracular stripe. Broken middorsal stripe thickest between segments, and subdorsal stripe composed of whitish spots (below right). The caterpillars of several other eastern Lithophane, and in particular the Brown Pinion (L. hemina), are easily confused with those of the Nameless Pinion. Until a reliable key to Lithophane larvae is authored, it is best to base identifications on the adults, which can be difficult enough. ! occurrence Woodlands and forests across Canada, south in East to Georgia (mountains) and Great Lakes states. One generation with mature caterpillars mostly in June and early July. Very common northward and in mountains. ! common foodplants Alder, apple, basswood, birch, cherry, fir, hawthorn, hemlock, hickory, hornbeam, maple, oak, pine, spruce, willow, and many other woody plants. ! remarks Last instars descend along the trunk and hide in crevices or under flaps of bark by day. The Nameless Pinion is one of the most polyphagous eastern Lithophane; larvae have even been reared from conifers. The species becomes increasingly common northward: in New England, the Nameless and Brown Pinions (L. hemina) are often the most commonly encountered Lithophane at bait. Adults are only weakly attracted to light, or at least bait collections can usually be expected to yield more than ten times the number of individuals found at blacklights or mercury vapor lights in the same location.

436 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

Buckeye Pinion Lithophane joannis ! recognition Strongly mottled, often with straw-colored to yellow tint where adjacent segments overlap, and/or tan to straw-colored flushes to middorsal and lateral stripes. White, dorsal pinacula edged with black; both dorsal pinacula often embedded in diffuse, dark patch that is best developed on A8. Head heavily marked: usually with coronal bar, black spot within triangle, and black bar above antenna. Larva to 4 cm. Middle instars easily confused with those of the Bailey’s Pinion (L. baileyi) and kin. Penultimate instar waxy to lime green with strong, creamy spiracular stripe; broken, white middorsal stripe; and generous salting of white spots (above lower right). Last instar may be inseparable from that of Brown Pinion (L. hemina), Nameless Pinion (L. innominata), and kin (although these, presumably, would not be found in leaf shelters of buckeye). ! occurrence Woodlands and forests from southeastern Michigan to North Carolina (mountains) and northern Alabama, and likely elsewhere within range of foodplant. One generation with mature caterpillars mostly in May and June. Rarely collected, but sometimes locally abundant. ! common foodplants Buckeye; especially yellow buckeye in Smokies. ! remarks We include this species because of the larva’s unusual shelter-residing habit, and as a lesson about rarity. The Buckeye Pinion went unnamed until 1992 and was not represented in major United States museums prior to its description. Yet, at the type locality in Ohio and elsewhere in the Appalachians, this Lithophane may outnumber all other members of the genus at bait (adults of the Buckeye Pinion are rarely if ever taken at light—see Covell and Metzler 1992). In May 2005, near the Chimneys picnic area of Great Smoky Mountains National Park, we discovered some two dozen last instar Buckeye Pinion caterpillars in less than ten minutes by opening leaf shelters made by the Brokenbanded Leafroller (Choristoneura fractivittana) and American Ermine Moth (Yponomeuta multipunctella) on a single buckeye tree. Last instars also can be found by looking in or adjacent to moss clumps growing along the trunks of buckeye. Early and middle instars are easily located by simply turning leaves of yellow buckeye in May and looking for larvae perched on the undersides, often over the midrib. See also Wagner (2006).

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 437

Branded Pinion Lithophane patefacta ! recognition Variegated in steel blue, gray, and white with thin yellow to orange middorsal and subdorsal stripes; some individuals pale while others quite dark (especially southward). Dorsal pinacula white, prominent. Pale spiracular (lateral) stripe may be edged with black above. Head variable, pale and unmarked to extensively mottled with black. Larva to 4 cm. Penultimate instar shiny pale green with prominent white striping and spotting (lower right). The caterpillars of the Wanton Pinion (L. petulca), Brown Pinion (L. hemina), and other members of this group are closely similar. ! occurrence Woodlands and forests from Wisconsin, east across southern Canada to northern Florida and Kentucky. One generation with mature caterpillars from late May through early July. Common. ! common foodplants Especially sugar maple northward; also blueberry, cherry, hickory, oak, red maple, and others. ! remarks Until recently, this was a rather poorly known moth. Adults of the southern form are sometimes misidentified in collections as the Nameless Pinion (L. innominata). It is among the most widespread and abundant pinions in the Southeast, especially in the Piedmont. Its propensity to eat other caterpillars (including its own siblings) is curiously variable. In some of our larval cohorts (especially those from the South) larvae are both cannibalistic and predatory, while others show no interest in carnivory even when reared in crowded conditions. It would be interesting to know if the predatory behavior of pinions is influenced by environmental factors, and if so, what factors are involved. Pale greenish forms are more common northward. Last instars, regardless of their color, rest on tree trunks and in the litter; larvae with greenish elements are especially cryptic when resting on algae and lichens. The first record of the Branded Pinion in Connecticut was of an adult collected at bait in March 1978. Within a year or two, the species was common in the state, with as many as 100 seen along a single bait trail. Soon thereafter, the moth was collected in western Massachusetts and the Albany, New York area for the first time. Even where the species is common, adults can be scarce at light; for example, none at all were found over 25 seasons in southern New Jersey where this is usually the most common pinion at bait. Adults have been observed nectaring at Chickasaw plum.

438 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

Shivering Pinion Lithophane querquera ! recognition Waxy white with bright yellow markings. Yellow spiracular

stripe broad, sometimes broken below spiracle. Dorsal yellow patches, at anterior and posterior ends of T2 and T3 and caudal end of A1–A7, may extend laterally to join spiracular stripe. Prothoracic shield and head pale and unmarked. Larva to 4.5 cm. Caterpillars of Mustard Sallow (Pyreferra hesperidago) and other Pyreferra species are also white and yellow (pages 444–6), but feed only on witch hazel, American hornbeam, hazel, and hophornbeam. In Pyreferra, the yellow markings over the dorsum are located centrally or anteriorly on each segment and extend down to the prolegs on the abdominal segments. The immature stages of the recently published and closely related Connecticut Sallow (Lithophane scottae) have not been described, but potentially they are close in appearance to the caterpillars of the Shivering Pinion; L. scottae occurs from Wisconsin, southern Ontario, and (formerly) the Boston area, south to northern Georgia (Schweitzer et al. 2011). The Connecticut Sallow is rare in collections, although Herman Wilhelm vouchered more than 80 adults in the vicinity of his home in Willimantic, Connecticut. ! occurrence Woodlands and forests from Ohio to New Hampshire, south to northern Georgia and Mississippi; possibly also Texas. One generation with mature caterpillars in late May and June from New Jersey northward. Often common; evidently scarcer in historical times. ! common foodplants We have larvae from apple, ash, black birch, cherry, hickory, hophornbeam (Ostrya) (Tim McCabe), and oak; presumably a generalized feeder on woody plants. ! remarks This is a consistently predatory pinion, consuming other caterpillars and pupae, especially in later instars (Schweitzer 1979a). It is perplexing why the caterpillars of Pyreferra and the Shivering Pinion should be similar in appearance. Before 1980 or so, the Shivering Pinion was uncommon to decidedly rare across much of the northeast (Schweitzer 1974). In the 1970s, the moth became common in southern New England and expanded into new areas, including the Albany, New York region and southern New Jersey. In the latter area in October 1989, DFS encountered 76 adults on the first three trees of a 35-tree bait line; he saw more than 500 in that one hour. In recent years, finding fewer than five would be more typical for these same areas.

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 439

Pine Pinion Lithophane lepida ! recognition Deep green with conspicuous, white to creamy middorsal, subdorsal, and subspiracular stripes. Dorsal setae not borne from white pinacula as in other pinions. Prothoracic shield usually maroon-brown along leading edge; thoracic legs also wine colored. Larva to 4 cm. Larva of Pine Sallow (Feralia major) and several conifer-feeding darts (Xestia) are similar in appearance. ! occurrence Pine barrens and woodlands from Alberta to Nova Scotia, south in mountains to Georgia and on coastal plain to South Carolina—see Remarks. One generation with mature caterpillars from June into early July northward; May in coastal Carolinas. Uncommon to rare. ! common foodplants Pines, at least Jack, pitch, red, Virginia, and probably other hard pines. ! remarks Traditionally, northern populations have been referred to as L. lepida lepida and southern ones as L. l. adipel. Troubridge and Lafontaine (2003) suggested that the two are specifically distinct and that the true L. lepida should be restricted to mostly historic populations from northern New York and eastern Ontario to Maine and Nova Scotia. Others they consider L. adipel; its adults tend to be paler, with a marked pinkish flush over the hindwings, and a washed out appearance to the forewing. Both entities fly at Clintonville, New York, and in central New Hampshire’s Ossipee Pine Barrens. Adults of the Pine Pinion are notoriously difficult to collect even where and when they are present. Many of the (sparse) northern records in collections are of fall-flying individuals captured at lights; apparently this is one of few Xylenini that does not often come to bait. Beating for larvae might well prove to be the most effective method for finding this elusive insect. Like other conifer-feeding owlets, the eggs are large and fecundity is low; females produce around 200 eggs, far fewer than most pinions. See account in Schweitzer et al. (2011). The caterpillar in the main image is L. lepida (Tennessee) while that to the right is L. adipel (New York).

440 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

Alder Pinion Lithophane pexata ! recognition Deep reddish to violet-brown with broken, tan middorsal and subdorsal stripes. Dorsal pinacula small, tan to white, and inconspicuous, sometimes surrounded by black; dark triangular patches meeting over dorsum on A8. Middorsal stripe nearly absent over thoracic segments, best developed over posterior half of A1–A8. Narrow subdorsal stripe nearly continuous, well developed through prothoracic and anal shields. Supraspiracular and spiracular stripes nearly obsolete. Spiracles white and sharply contrasting with ground color. Lateral areas frequently with faint reddish flush. Head with pale snowflake spots. Larva to 4.5 cm. Penultimate instar with reddish-brown cast; dorsum of each abdominal segment with three pairs of white spots (pinacula); continuous white middorsal and subdorsal stripes; supraspiracular stripe broken into minute spots; and spiracular stripe also consisting of closely approximate spots (inset). ! occurrence Swamps, riparian corridors, and northern woodlands. Transcontinental in Canada, south in East through New Jersey, Washington, DC, and upper Great Lakes states. One generation with mature caterpillars from June into early July northward. Uncommon. ! common foodplants Alder. ! remarks Mature caterpillars are darker than any other pinions with which we are familiar. The reddish to purplish cast is well suited for an alder feeder. Forbes’s (1954) description implies that in some forms the caterpillar stays green into the last instar. All of ours from Quebec and Connecticut turned purple-brown in the last instar. In captivity, the dark brown last instars are reclusive by day, hiding under leaves. The moth can be common at bait in the fall and very early spring—expect any female taken from late winter onward to have paired.

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 441 Amanda’s Pinion Lithophane amanda

Transcontinental across Canada, south in East to Connecticut and northern Great Lakes states. One generation with mature caterpillars in June and July. Uncommon. Willow. The related European Pale Pinion (L. hepatica) feeds on birch, willow, and other woody and herbaceous plants (Porter 1997).

Bailey’s Pinion Lithophane baileyi

Transcontinental; eastern portion of range includes Ontario and Quebec, New England, south in mountains to northern Georgia, Kentucky, and Great Lakes states. One generation with mature caterpillars in June and early July. Uncommon. Our larvae from alder, buckeye, currant, and hickory; Handfield et al. (1999) list birch, jack pine, and poplar.

Dashed Gray Pinion Lithophane disposita

Alberta to Quebec, south to New Jersey, Pennsylvania, and Great Lakes states; an unnamed sibling species occurs from Connecticut southward (DFS). One generation with mature caterpillars in June and early July. Uncommon. Especially willow and aspen, but also birch, elm, and likely others.

Hoary Pinion Lithophane fagina

Transcontinental in Canada, south in East to Pennsylvania and New Jersey Pine Barrens, but local southward. One generation with mature caterpillars in June and July. Locally common. Paper and gray birch; one report from pin cherry (Raizenne 1952).

442 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

Wanton Pinion Lithophane petulca

Transcontinental in Canada, south in East to Kentucky and North Carolina (mountains). One generation with mature caterpillars in June and July over most of range. Locally common. Alder and birch commonly; also ash, blueberry, cherry, elm, hickory, maple, oak, poplar, and willow. Our wild larvae mostly from black birch.

Sycamore Pinion Lithophane signosa

Lower Great Lakes states to Vermont, south to Georgia and Mississippi. One generation with mature caterpillars mostly in May and June. Locally common. Sycamore, both native and cultivated. Caterpillars readily found by searching under flaps of bark or in litter at base of trees. Both middle instar (green) and last instar shown.

White-eyed Pinion Lithophane tepida

Manitoba to Nova Scotia, south to Pennsylvania and in mountains to Georgia. One generation with mature caterpillars in June and July. Uncommon to locally common. Basswood, birch, blueberry, currant, oak, sweet fern, and presumably other woody plants; also recorded from spruce.

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 443 Thaxter’s Pinion Lithophane thaxteri

Transcontinental in Canada, south in East to northern New York, Maine, and eastern Massachusetts; disjunct populations in New Jersey Pine Barrens. One generation with mature caterpillars in early summer. Local, uncommon, mostly in bogs and pine barrens. Andromeda, Douglas-fir, larch, leatherleaf, New Jersey tea, sweet fern, sweet gale, willow, and presumably other woody plants; accepts many heaths in captivity.

Lemmer’s Pinion Lithophane lemmeri

Coastal plain and Piedmont from southeastern Massachusetts to Georgia, Florida panhandle, and Alabama. One generation with mature caterpillars mostly in June in New Jersey. Local and uncommon. Mainly Atlantic white cedar where it occurs, but red cedar over much of range. See account in Schweitzer et al. (2011).

Canadian Cedar Pinion Lithophane thujae

Ontario to New Brunswick, south to Maine and northern parts of Michigan and Wisconsin. One generation with mature caterpillars mostly in June and July. Local and uncommon. Northern White Cedar. Bald Cypress Pinion (L. abita) not illustrated. Coastal plain from Maryland to Texas. Associated with bald cypress throughout its range.

Citrine Sallow Pyreferra citrombra

Ontario to New Hampshire, south to Georgia (mountains), but range confused with that of P. moffatiana. One generation with mature caterpillars in May and June. Common. At least hazel (Corylus) and likely related genera such as American hornbeam and hophornbeam. Our images likely refer to P. moffatiana and not true P. citrombra—see Remarks in main account, next page.

444 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

Citrine Sallow Pyreferra citrombra ! recognition Waxy white with bands of lemon yellow. Yellow bands absent

on T1, A9–A10, and often above spiracle on A8. Amount of yellow on intervening segments highly variable, usually best developed along subdorsum and at level of spiracles. Larva to 4 cm. Two or three other Pyreferra feed on witch hazel in the East: the Mustard Sallow (P. hesperidago) and Anointed Sallow (P. ceromatica) illustrated on pages 445, 446, respectively, A third species, P. moffatiana is discussed below. Until we have reared more examples of each of these, and the group’s taxonomy is revised, we are reluctant to suggest diagnostic characters for any (see Remarks below and Mustard Sallow account). Caterpillars of the Shivering Pinion (Lithophane querquera) share a similar coloration scheme, but its yellow rings are confined to the posterior portion of the abdominal segments (page 438). ! occurrence Woodlands and forests from Ontario to New Hampshire, south to Georgia (mountains). One generation with mature caterpillars in May and June. Common. ! common foodplants At least hazel (Corylus)—see below. ! remarks According to Chris Schmidt, P. citrombra is a complex of two species. One is pale and smaller (typical) (our live adult), and the other warmer (due to greater numbers of orange scales) and somewhat larger (about the size of P. hesperidago). He is tentatively calling the second P. moffatiana. Franclemont’s type series of the Citrine Sallow’s were reared from hazel. We have raised what we believed were Citrine Sallow on witch hazel (Hamamelis) American hophornbeam (Carpinus), and hophornbeam (Ostrya). The first was a lab host that our ex ova larvae accepted; the latter two hosts resulted from collections of larvae in the wild. At least two larval forms are included in the present concept of P. citrombra— some have almost complete yellow banding (main image above), and thus closely resemble P. hesperidago, while other individuals are yellow-green with the yellow band broken into large subdorsal and spiracular spots (previous page and right). The three caterpillars shown in this work yielded P. moffatiana-type adults. We have not knowingly collected or imaged caterpillars of true P. citrombra. Until the group is revised, the description, range, and foodplants given above should be regarded as tentative. We encourage others to help solve this conundrum. Adults reared from wild larvae are especially needed. Alternatively, cohorts of larvae obtained from gravid females might be divided and reared on the hosts listed above. The Citrine Sallow’s foodplants may include members of both the birch and witch hazel families—an odd couple of sorts, although the caterpillars of Zale phaeocapna (page 184) also specialize on these same hostplants, and no others.

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 445

Mustard Sallow Pyreferra hesperidago ! recognition Waxy white with bands of lemon yellow. Yellow bands almost fully occupying T2, T3, A1, and A2, then narrowing rearward where they extend around middle or anterior of each segment. Bands on A3–A6 continuing down outer face of prolegs. Larva to 4 cm. Perhaps indistinguishable as a larva from Citrine Sallow (P. citrombra), P. moffatiana, and the very rare Anointed Sallow (P. ceromatica), all of which may feed on witch hazel (see previous account). Our examples of the Anointed Sallow (next page) had brighter yellow markings than any other Pyreferra, although we have seen only a few examples of its caterpillar. ! occurrence Woodlands and forests from Wisconsin to Nova Scotia, south to Florida and southern Louisiana. One generation with mature caterpillars in May and June in most of range. Often very common northward. ! common foodplants Witch hazel. ! remarks Pyreferra emerge in the fall and overwinter as unmated adults. The pale, hemispherical eggs, visible to the eye, are laid near a leaf scar. Pyreferra caterpillars may be collected by crouching beneath a host and scanning upward. The yellow and white caterpillars, perched on leaf undersides, are readily apparent, especially if the sun is high enough to backlight the leaves. Beating works at any time of day. When alarmed, the caterpillars may release the abdominal legs and curl the abdomen about the head, in sawfly fashion. Prepupal caterpillars of the witch hazel-feeding Pyreferra turn an even brighter yellow—the waxy white ground gives way to lemon yellow, especially over the dorsum. The Mustard Sallow sometimes shows up in numbers far from known populations of its foodplant. While none are seen in most years in southern New Jersey, in other years adults appear in November and will be seen at bait throughout the winter and then persist into March. On occasion, as many as two dozen will be encountered along a single bait line. Apparently these individuals leave in the spring, returning to breeding areas to the north and west. Adults of all Pyreferra come to light and especially to bait.

446 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

Anointed Sallow Pyreferra ceromatica

Presently Gulf states, north along Atlantic coast through the Carolinas; prior to about 1935, north to southern Canada and west to Chicago area. One generation with mature caterpillars in late spring. Rare and local. Witch hazel. See account in Schweitzer et al. (2011).

Pettit’s Sallow Pyreferra pettiti ! recognition Body translucent green with yellow tints to thorax. Anterior third of prothorax waxy white (as in other Pyreferra). Spiracular stripe formed from closely set creamy white spots. Most setae borne from bright white pinacula; those over dorsum enlarged. Traces of middorsal stripe may be present as spots, especially rearward on each segment. Subdorsal stripe composed of widely spaced spots. Each proleg with prominent white pinacula above base. Head pale sea green to pale yellow-orange, smooth, shiny. Larva less than 4 cm. ! occurrence Woodlands and forests from Ontario and Quebec to New England, south to Florida and Texas. One generation with mature caterpillars mostly in June. Locally common. ! common foodplants Hophornbeam; old reports from black birch and yellow birch (e.g., Forbes 1954) may trace back to hostplant misidentifications. Tim McCabe and DFS have had four ex ova cohorts refuse various birches. ! remarks The largely green larva immediately distinguishes Pettit’s Sallow from its yellow, sawfly-mimicking congeners. Its coloration, which more closely resembles that of a pinion (Lithophane) or middle instar quaker (Orthosia), may represent the ancestral phenotype for the genus. Conspicuous tracheal branching, extending outward from each of the abdominal spiracles, is visible through the body wall (at least if a flash is used to image a caterpillar). The caterpillar can be taken reliably in the Northeast by beating hophornbeam plants in late spring and early summer. Alternatively, one can examine leaf undersides: the larva often perches over the midrib with the head curled to one side.

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 447

eupsilia sallows Eupsilia

Seven species occur in the East; one is well known to collectors but remains undescribed. While Lost (E. devia), Morrison’s (E. morrisoni), Three-spotted (E. tristigmata), and Straighttoothed (Eupsilia vinulenta) Sallows are readily identified as adults in the field, the other three are a truly challenging lot. Definitive identifications are best based on the examination of genitalic characters (see Forbes 1954). The larvae, too, are exceedingly similar. Two or three Eupsilia would have sufficed for representation in this work, but because they are among the most abundant forest owlets, we illustrate all the eastern species. Use characters provided here cautiously: final determinations should be based on reared adults or genetic evidence. Keep in mind that coloration changes both between instars (see Franclemont’s Sallow, E. cirripalea, page 450) as well as within the last instar (see Three-spotted Sallow, E. tristigmata, page 453). In general, fourth, fifth, and early sixth instars are the most colorful. The sixth (final) instar loses color as it matures, becoming dull brown and nondescript as a prepupa. While larvae can be reared in captivity or sleeved on cherry and oak, little is known about their host preferences in na-

PUDDLING

Many species of Lepidoptera “puddle,” a behavior where adults congregate on moist soil to collect sodium. This is an essential element, but one that is in short supply in plants. To circumvent their deficiency, Lepidoptera gather sodium through puddling, i.e., by imbibing salts in soils, or else in carrion, dung and urine, eye secretions, and sweat. Such indelicate behaviors are usually the purview of males. Females of puddling species get their sodium principally during copulation (Smedley and Eisner 1995)—in addition to sperm, male Lepidoptera commonly transfer amino acids, salts and minerals, carbohydrates, defensive compounds, and other substances during the process of mating (Scoble 1992). In moths, these substances are frequently bundled into a spermatophore, which in the extreme may be so large as to preclude additional matings! Xylenines, for example, produce a substantial spermatophore—one can often feel which females have

ture, especially those of the last two instars. Early and middle instars are new-leaf specialists that fashion crude leaf shelters in soft spring foliage or take refuge in the abandoned leaf shelters fashioned by other caterpillars. The darkly rendered last instars are more apt to be found on the ground, and it is our guess that most mature on fallen plant matter or low-growing plants. Wagner et al. (1995) found 33 early and middle instars on oak foliage and 13 on blueberry foliage, but no mature caterpillars were recovered from leaves and only one was found under burlap. Late instars to which we offered only fallen hickory catkins grew and matured. Prepupal larvae tunnel into the soil and form a cell in which they aestivate for three or more months before pupating. As prepupae they are susceptible to desiccation—their containers should be kept lightly moistened. Eupsilia emerge in the fall, but they are seldom seen for a month or so; it is not known if they enter a short, second diapause. Adults start visiting bait in number beginning in late October in the Northeast. Mating occurs from midwinter to early spring, with Franclemont’s, Morrison’s, and the Straight-toothed Sallow initiating activity before the others. These three also tend to be more active through midwinter, at least in temperate areas such as southern New Jersey and southern Connecticut. mated recently with a gentle squeeze of the abdomen. Many of the minerals and nutrients contributed by a male will find their way into the eggs … his paternal investment. In Gluphisia septentrionis (Notodontidae), the eggs sired by “puddled” males contain two to four times more sodium than those from males denied access to sodium as an adult (Smedley and Eisner 1996). Gluphisia is the reigning champion among puddlers: males can drink and process more than 600 times their own body weight in a single evening (Smedley and Eisner 1995). In Smedley’s image below, a male is forcibly expelling excess water, “scrubbed” of its sodium. From “Sodium uptake by puddling in a moth, Science (1995): Volume 270, Issue 5243.” Reprinted with permission from AAAS.

448 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

Lost Sallow Eupsilia devia ! recognition Last instar with thin, mostly broken, white middorsal and

subdorsal stripes. Broad, immaculate, white lateral stripe runs below spiracles, bordered above by black spots that may coalesce into stripe. Dorsum mottled with tan and darker brown; dorsal setal bases white. Subventer yellow-green and mottled with gray-brown spots, but without pink or maroon flush below. Head shiny, orange-brown above, black below. Thoracic legs shiny black. Larva to 3.5 cm. Antepenultimate and penultimate instars with more distinct patterning (inset). ! occurrence Fields, grasslands, wet meadows, marshes, and open woodlands. Transcontinental in Canada, south in East to northern Georgia and Missouri, although increasingly uncommon southward. One generation with mature caterpillars in late spring. In Connecticut last instars occur throughout June. Common northward. ! common foodplants Mostly aster and goldenrod in the wild. Covell (2005) also lists aspen, oak, and cherry, but these represent lab hosts used by John Franclemont and us. ! remarks In contrast to other Eupsilia, the Lost Sallow is a species of fields and open habitats. The caterpillar forms a leaf shelter by silking together leaves near a shoot apex. Two or more shelters are constructed by each larva. Adults emerge in late September and October and fly a month or two before hibernating. Like other Eupsilia, they are strongly attracted to bait. The caterpillar in the inset is a penultimate instar approaching its last larval molt—the head capsule of the sixth instar appears as a swelling beneath the prothoracic plate.

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 449

Barrens Sallow Eupsilia sidus ! recognition Last instar dirty purple-brown to reddish brown, markings obscure, only modestly paler below spiracular stripe than above. Supraspiracular and subspiracular regions more or less same color (contrast between these two portions of body greater in other eastern Eupsilia). Venter of thorax smoky purple (not reddish or pinkish as in congeners), sometimes with an orange cast; contrast in coloration between thoracic and abdominal segments more subdued than in some other Eupsilia. Dorsal and lateral pinacula only slightly whitened. Spiracular stripe well developed rearward, but seldom pure white. Subdorsal line on prothoracic shield prominent. Only upper portion of head orange-brown. Larva to 4.5 cm. Middle instars also darker than those of congeners; ground color purplebrown. Individual in lower right is 11 mm long. ! occurrence Dry oak forests, savannas, woodlands, ridgetops, and barrens from Wisconsin, southern Ontario, and central New Hampshire, south to northern Georgia and Texas. One generation with mature caterpillars from June through early July (northward). Local over much of range; becoming very common in barrens and dry oak woodlands northward and eastward. ! common foodplants Wild larvae have been collected from lowbush blueberry, sand hickory, and oak. Our ex ova cohorts were reared on cherry, hickory, oak, and walnut; oaks and heaths probably the usual hosts. Records in Prentice (1962) apply to other species. ! remarks Little published information on the Barrens Sallow actually refers to this species (most applies to the Straight-lined Sallow, E. vinulenta and the undescribed species treated on the next page). The Barrens Sallow’s phenology is geared toward the late-flushing oaks of barrens and other xeric habitats. Adults mate weeks later than other Eupsilia and fly deeper into spring, often well into May in southern New Jersey. Young larvae do well on cherry or oak, but last instars may have some specialized dietary needs—few of our captive larvae attained normal size and hatch rates were poor. It is associated with sandy areas in the Great Lakes region, along the coastal plain in southeastern Massachusetts, and on Long Island. In Connecticut, Pennsylvania, New York, and elsewhere it occurs in acid rocky sites. Over much of the Northeast, but not in southern New Jersey, the Barrens Sallow is associated with scrub oak. While the Satellite (E. transversa), a European moth, is notoriously predatory, we have not noted cannibalistic behavior in any North American Eupsilia; however, captive larvae bite when handled roughly and we have observed them occasionally feeding on larvae that have died during a molt or for other reasons.

450 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

Franclemont’s Sallow Eupsilia cirripalea ! recognition Last instar dark with white subspiracular stripe weakly developed across T2 and T3. Thoracic venter red-purple; abdominal venter usually faint waxy green. Red-purple flush to supraspiracular and subventral regions of abdomen, especially above prolegs. Thoracic shield with prominent white subdorsal lines. Dorsum between subdorsal stripes sometimes dark greenish brown with deeper brown mottling, although much of this patterning obscured in late instars and deeply pigmented individuals. Lower three-quarters of head black, including frons; upper portion orange-brown. Larva to 4.5 cm. Earlier instars shown on facing page. Subspiracular stripe often more prominent than those of related Eupsilia (although appreciably thinner than that of Lost Sallow, E. devia). Caterpillar similar to, and perhaps inseparable from, the Straight-toothed Sallow (E. vinulenta) and an as yet undescribed Eupsilia species (see below). Generally both of these caterpillars have less white in the subspiracular stripe, the subdorsal white lines in the prothoracic shield are not as prominent, and they are less likely to have the red-purple flush carried back to the supraspiracular and subspiracular regions of the abdomen. Larvae of the unnamed species are sometimes identifiable by the absence of reddish purple ventrally on the abdomen, and more prominent dark patches on the head. ! occurrence Woodlands, especially those with abundance of oak and hickory, from Missouri, Ohio, and Massachusetts, south to northern Georgia and northern Mississippi. One generation with mature caterpillars from May through mid-June (northward). Common southward. ! common foodplants Forbes (1954) lists cherry, but presumably polyphagous on woody plants. Captive larvae accept birch, cherry, hickory, maple, oak, persimmon, and walnut. We have collected larvae on blackberry and oak and know of a collection from red maple. ! remarks Franclemont’s Sallow is often the most abundant Eupsilia in the southern parts of its range, but it becomes scarce and local north of southern New Jersey, central Ohio, and Missouri. It was rare or absent in Connecticut before 1980. A very common but unnamed species replaces Franclemont’s Sallow in abundance north of Missouri and southern New Jersey; it ranges to Wisconsin and southern Ontario, and tends to be the more common of the two in cooler mountain areas. Both species are widely sympatric from southern Connecticut to Georgia. Adults of Franclemont’s Sallow mate in February or early March in New Jersey (the new species usually mates in March). Females “call” again after laying their first few hundred eggs; older females may contain multiple spermatophores. Captive females of Franclemont’s Sallow will lay 1000–2000 eggs if well fed and supplied with males. On the facing page, images to the left are those of E. cirripalea and those to the right illustrate the new species. The series also illustrates how coloration changes through development. Adults of the two moths have reliable genitalic characters in both sexes.

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 451

Eupsilia cirripalea 4th instar (15 mm)

Eupsilia n. sp middle instar

Eupsilia cirripalea 5th instar

Eupsilia n. sp 5th instar (early)

Eupsilia cirripalea 6th instar (early)

Eupsilia n. sp 6th instar

Eupsilia cirripalea 6th instar (prepupal larva)

Eupsilia n. sp 6th instar

452 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

Morrison’s Sallow Eupsilia morrisoni ! recognition Last instar dark purple-brown with normal complement of thin middorsal, subdorsal, and spiracular stripes; often with faint supraspiracular stripe represented by series of scattered spots. Pattern elements more obscure than most other Eupsilia. Venter mostly greenish with less red to purple flush below thorax (relative to congeners); little red along subventer of abdomen. Subdorsal lines in prothoracic shield commonly with orange hue. Spiracular stripe often weakened anteriorly, sometimes reduced to trace on T2–A2. Subspiracular and subventral regions of abdomen with brownish patterning, and little red to purple tinting. Larva to 4.5 cm. Antepenultimate instar dark above a thin, white spiracular stripe and greenish below. Dorsum with faint middorsal, weak subdorsal, and fine supraspiracular stripes. Subventer of thorax dull wine red; subventer of abdomen with red-brown spots. We are unaware of features that reliably distinguish larvae from those of the Straight-toothed Sallow (E. vinulenta) and others. ! occurrence Woodlands and forests from Ontario to Nova Scotia, south to northern Georgia and Missouri; often uncommon to rare on coastal plain. One generation with mature caterpillars from May through early June. Abundant, except on coastal plain. ! common foodplants Widely polyphagous on woody plants. Literature records from cherry, elm, maple, and oak. Our records include scrub oak, sugar maple, sweet gale, viburnum, and wild strawberry. ! remarks While we commonly find young Eupsilia caterpillars on foliage, we have yet to find a mature caterpillar on aboveground foliage. Moreover, their dark red-purple-brown coloration seems ill suited for a “green-leaf feeder” or “bark rester.” We initially suspected that Eupsilia caterpillars behaved as climbing cutworms, migrating between foliage (at night) and the ground each day. But after finding numerous late instars under objects far removed from woody vegetation, we now suspect that many if not most Eupsilia caterpillars mature on low-growing plants, fallen catkins, and various other organic substrates without ascending into woody vegetation to feed. Morrison’s and the Straight-toothed Sallows (E. vinulenta) are among the most abundant moths in northeastern forests—in most years they account for nearly all the owlet moths in your headlights between November and early March. DFS recalls evenings in Connecticut where his bait trail yielded over 1000 Eupsilia adults, among which were all seven eastern members of the genus. Morrison’s and the Straight-toothed Sallows accounted for most, and sometimes over 90%, of the individuals seen on such occasions. Not surprisingly, Eupsilia are common but unwelcome visitors in sugaring buckets used to collect maple sap (particularly those that manage to drown themselves in the buckets).

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 453

Three-spotted Sallow Eupsilia tristigmata ! recognition Rather undistinguished member of genus and difficult to identify as a larva. In our examples, white spiracular stripe pure white and thickened on A3–A8, but weakly expressed on anterior segments. Subdorsal line on prothoracic shield commonly orange. With exception of burgundy flush, subventer and venter of abdomen pale and unpigmented. Larva to 3.5 cm. Middle instars handsomely patterned with prominent white striping and bright pink-red venter below T1–A2. Often with bright burgundy flush above each proleg, rest of venter approaching white, and sharply contrasting with dorsum (upper inset). ! occurrence Woodlands and forests, barrens and heathlands, as well as swamps and bogs from southern Canada to Georgia (mountains) and Missouri. One generation with mature caterpillars in late spring. Common, at least northeastward. ! common foodplants Often feeding on apple, blackberry, blueberry, cherry, mountain ash, rose, serviceberry, strawberry, and other members of cherry and heath families. Also recorded from alder, aspen, birch, butternut, oak, and willow; but some of these host records may be unreliable. ! remarks Flowers and young fruits are preferred to new leaves, especially by early instars. The middle instars are beautiful with considerable pink to red tinting along the venter (upper inset). As in other Eupsilia, the attractive red colors are lost in mature larvae (lower inset).

454 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

Straight-toothed Sallow Eupsilia vinulenta ! recognition Velvety purple-brown with weak stripes. Spiracular stripe somewhat reduced relative to other members of genus: anterior portion over T1–A2 frequently consisting of series of white spots (as in Morrison’s Sallow, E. morrisoni). Subdorsal line in prothoracic shield sometimes incomplete. Pink to wine-red flush of venter subdued in last instar, somewhat intermediate between conditions seen in Franclemont’s (E. cirripalea) and Morrison’s Sallows. Larva to 3.5 cm. A fifth instar is shown in the inset. As is true across the genus, positive identification is best based on the adults. ! occurrence Yards, parks, barrens, woodlands, and forests from southern Canada to coastal North Carolina, Georgia (mountains), and Missouri; old reports from Texas. One generation with mature caterpillars in May and June over much of East. Abundant. ! common foodplants Widely polyphagous on woody plants. Cherry, maple, and oak are commonly listed foodplants; also some forbs such as woodnettle and violets. We have records from hackberry (including psyllid galls), hawthorn, oak, red maple, sycamore, walnut, and many from black cherry. ! remarks This is sometimes the most common owlet of eastern forests, especially from late autumn to March after numbers of the Bicolored Sallow (Sunira bicolorago) have fallen off for the season. We routinely see hundreds, and occasionally more, along a bait line. The main image (above) is a sixth instar; note especially the red-purple aspect of the venter and velvety brown-purple dorsum. As is common for the genus, coloration of the prepupal caterpillar is more uniformly brown (not purple-brown) and all striping is more subdued. Mating occurs in late winter and early spring. Whether a female has mated, or even the number of times she has paired, often can be ascertained by gently squeezing her abdomen between thumb and forefinger; hardened lumps, representing male spermatophores, persist within the female’s abdomen for weeks. Females of the Straight-toothed Sallow apparently spend a great deal of time nectaring at red maple flowers, and we suspect that they oviposit among the flowers as well.

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 455

Red-winged Sallow Xystopeplus rufago ! recognition Speckled, red-brown with dark, shiny prothoracic shield and anal plate. Subdorsal stripe prominent through prothoracic shield, but otherwise weakly expressed. Anal plate black, edged with white, contrasting sharply with rest of rump. Spiracular stripe continuing to anal proleg, although somewhat weakened rearward. Larva to 3 cm. Antepenultimate and penultimate instars reddish brown (lower middle right); neither prothoracic shield nor anal plate darkened. Early instars green, striped, with prominent dorsal pinacula (lower right), and easily confused with those of many pinions (Lithophane). Mature larva superficially similar to that of the Goat Sallow (Homoglaea hircina). ! occurrence Woodlands, forests, and barrens from Minnesota to Maine, south to Florida and Texas; increasingly associated with scrub oak and barrens northward. One generation with mature caterpillars from May to July. Common in most of range. ! common foodplants Many records from oak and blueberry; also beach plum, hazel, poplar, (sand) hickory, sourwood, witch hazel, and other woody plants. DFS has observed captive late instars avidly feeding on young (green) blueberry fruits. ! remarks The Red-winged Sallow is most common in dry, scrubby oak woods and barrens. Our captive-bred pupae have yielded adults both in the fall (few) and March (most), but even in the latter instances, the adults were fully developed within the pupal shell by late fall. Although seemingly anomalous, because almost all temperate Lepidoptera have a single stage that overwinters, these observations are consistent with museum records—i.e., there are modest numbers of records for fall adult collections relative to those for the spring. Mating is in midspring, later than in most Xylenini. The dark color of the late instar suggests that it feeds at night, and moves from foliage into the litter during the day or feeds near the ground. On several occasions we have found Red-winged Sallow caterpillars as “contaminants” in our rearing sleeves deployed to field-rear other owlet species on oaks, but also on blueberry and sand hickory. Larvae tolerate older foliage than most xylenines, and occur somewhat later into the spring than other sallows. DFS once observed a Red-winged Sallow caterpillar eating the near-dead body of a parasitized larva of the same species; the parasitoid larva that had killed the Xystopeplus caterpillar was consumed as well.

456 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

glaeas

various xylenine genera We use this informal grouping for Xylenini that mostly fly, mate, and lay their eggs in the fall. Ova are dropped among low plants or laid singly on, or adjacent to, unopened buds of woody plants. Larvae develop early in the spring on young leaves and then tunnel underground to pass the summer as prepupae in silken cocoons. Caterpillars are nocturnal and may rest in litter around the base of their host, at least in the late instars. The feculae are large and irregular in form. Adults of most are strongly attracted to sugar baits, and come well to lights; they also can be seen feeding at fermenting apples and berries. Many are abundant in barrens.

Scalloped Sallow Eucirroedia pampina

Transcontinental in Canada, south in East to Georgia and Texas. One generation with mature caterpillars in late spring. Common. Cherry, lowbush blueberry, oak, poplar, red maple flowers, and many other woody plants in captivity; lowbush blueberry and wild strawberry documented in nature. DFS twice watched ovipositing females place eggs singly at base of dry grass stems in sandy woodland openings at dusk.

Crimson Sallow Psectraglaea carnosa

Southern Ontario, Quebec, and Maine, south to Maryland and Wisconsin in barrens and heathlands. One generation with mature caterpillars in late spring. Sometimes locally common. We have reared captive larvae on blueberry, chokeberry, and cherry. See account in Schweitzer et al. (2011). Caterpillars superficially similar to those of Scalloped Sallow (Eucirroedia pampina).

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 457

Variable Sallow Sericaglaea signata ! recognition Smooth, tan brown with abundant speckling above spiracular stripe and pale, unmarked venter. Body widest through A7 and A8. Spiracular stripe frequently infused with subtle yellow, salmon, orange, or brown splotches, especially below spiracles. Pale middorsal stripe thin and poorly expressed. Dorsum with vague, darkened, V-shaped spots at either end of segment; these sometimes forming darkened Xs between adjacent segments. Subdorsal stripe obliterated except where it cuts through prothoracic shield. Front of head mostly brown with paler spotting over each lobe. Spiracles caramel colored with black rim. Larva to 5 cm. Middle instars paler with somewhat more defined striping. Similar to Silky Sallow (Chaetaglaea sericea), but ground color deeper brown; bottom edge of spiracular stripe more defined and more apt to be infused with yellow, salmon, or orange. In addition, the Variable Sallow’s coloration below the spiracular stripe mostly lacks pigment and is paler than over dorsum. ! occurrence Woodlands and forests from Missouri to Connecticut (since late 1970s), south to Florida and Texas (see also below). One generation with mature caterpillars from April through June. Common. ! common foodplants Ash, cherry, chokeberry, hackberry, hickory, linden, oak, plum; undoubtedly other woody plants. Common on oak and cherry. ! remarks The Variable Sallow can be an abundant moth in southern woodlands and forests. It is one of many moths whose range has been expanding northward in recent decades: adults were uncommon to rare in New Jersey as late as the mid-1970s, but now hundreds sometimes can be seen across much of the lower half of the state, particularly on damp evenings. In some larval resting positions segments A7 and A8 appear thickened. We suppose, especially with extended anal prolegs, that this end of the body could be mistaken for the head end by a would-be predator. Larvae can be found during the day by searching beneath leaves and duff at the bases of shrubs and trees. The adult is most common in the fall and late winter, but flies well into May in the Northeast. Unlike other “glaeas,” this and the Goat Sallow (Homoglaea hircina) overwinter unmated. In New Jersey, mating takes place in late February or March.

458 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

Silky Sallow Chaetaglaea sericea ! recognition Pinkish tan, pale brown, or beige with mottling of short white lines and spots. Bold white spiracular stripe, sharply defined above; lower side diffuse; upper margin running along lower edge of black spiracles on A1–A7, weakening rearward of spiracle on A8. Middorsal, subdorsal, and supraspiracular stripes, obscure, broken into pale spots, often absent. Dorsal pinacula white, about twice diameter of small subdorsal spot near leading edge of abdominal segments A1–A8. Ground color below spiracular stripe only slightly paler than that above. Prolegs pale, without markings. Prothoracic shield somewhat darker than ground color with thin middorsal stripe and pale setal bases. Head with or without coronal bars, numerous snowflake spots over each lobe, and pair of short, dark brown lines behind eyes. Larva to 4 cm. We have seen too few Chaetaglaea caterpillars to offer characters that will distinguish caterpillars of the Silky Sallow from those of other Chaetaglaea. Larvae of Variable Sallow (Sericaglaea signata) are similar—features that will distinguish the two are given in that account (previous page). ! occurrence Barrens, woodlands, and forests from Manitoba to Nova Scotia, south to Florida and Texas. One generation with mature caterpillars from late spring and early summer. Common in oak forests and barrens eastward. ! common foodplants Blueberry and oak commonly. Aspen, cherry, chokeberry, and oak verified as wild hosts by us. Ex ova larvae rejected hickory, maple, and walnut. ! remarks The Silky Sallow is the mostly widely distributed and frequently encountered Chaetaglaea. Caterpillars were common in the foothills of Virginia (Wagner et al. 1995). Adults can be locally abundant in oak barrens—we have encountered dozens along bait lines on warm nights in October and November in New Jersey and southern New England. Yet over much of the inland portion of its range, the Silky Sallow is uncommon to rare (Rings et al. 1992, Covell 2005). Adults fly in the fall, beginning in September and continuing into early winter; in the South the flight extends from mid-October into February. The eggs overwinter; if kept outdoors in the shade, expect them to hatch shortly after red oaks break bud.

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 459

Trembling Sallow Chaetaglaea tremula ! recognition Brown to pinkish, smooth, with broad white spiracular

stripe that may include yellowish spots below the abdominal spiracles. Obscure middorsal, subdorsal, and supraspiracular stripes; these somewhat more obvious through prothoracic plate. Dorsal setae borne from small, white pinacula. Spiracles black. Setae short and inconspicuous, scarcely longer than height of midabdominal spiracles. Head with numerous snowflake spots and weakly developed lateral stripe (as extension of subspiracular stripe). Larva to 4.5 cm. Both Silky Sallow (C. sericea) and C. cerata are similar. At least in our examples of the Trembling Sallow, the spiracular stripe appears to be a bit broader and better defined along its lower margin than that of Silky Sallow. Identifications should be based on reared adults. ! occurrence Heathlands, barrens, scrublands, pinelands, and open, dry woodlands from Ohio (once) to extreme southern Ontario (rare); otherwise southern Maine to southern New Jersey with isolated colonies in the Poconos, and from coastal North Carolina south to Florida and Texas. One generation with mature caterpillars in late spring and early summer. Locally abundant along Atlantic coast from Maine to Florida, but local and sporadic inland. ! common foodplants Lowbush blueberries and small oaks in many habitats; also huckleberry, sweet fern, and undoubtedly others. Ours reared mostly on cherry. ! remarks In New England, this is a species of barrens, heathlands, and open oak woodlands on sandy or thin rocky soils. The larva figured here was among several swept by our colleague Mark Mello from huckleberry (Gaylussacia baccata) and lowbush blueberry (Vaccinium angustifolium) in early June on Nantucket Island, Massachusetts. In New Jersey, adults are among the first “glaeas” to appear in September and are often done flying by November; they start later and persist into the winter months from the Carolinas southward. Adults can be reliably sampled with bait or lights.

460 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

Cranberry Sallow Epiglaea apiata ! recognition Reddish to pinkish brown with broad, pale spiracular stripe. Weak, thin middorsal, subdorsal, and supraspiracular stripes often broken. Spiracular stripe passing along lower edge of spiracles on T1 and A1–A7. Body pale pinkish brown to ochre below spiracular stripe, and mostly without pigment ventrally. Dorsal setae from whitish bases. Prothoracic plate more caramel brown and shiny than adjacent areas. Head shiny with brown reticulations. Spiracles dark; spiracle on A8 twice height of that on A7. Larva to 4 cm. Middle instars more striped and with green flush to dorsum (inset). ! occurrence Cranberry bogs, barrens, heathlands, powerline right-ofways, moist southern pinelands, and other open wooded habitats with heaths. Transcontinental in Canada, south along coastal plain to northern Florida and southern Louisiana, and into Great Lakes states. One generation with mature caterpillars from June through early July in New England. Locally common. ! common foodplants Cranberry and some lowbush blueberries, e.g., it is often abundant in bogs, northern barrens, and heathlands dominated by Vaccinium angustifolium or V. myrtilloides; also black huckleberry, leatherleaf, and likely other heaths. ! remarks Eggs hatch in May northward. The larva, known in the applied literature as the Cranberry Blossomworm, feeds on young leaves and flowers, mostly at night. The Cranberry Sallow is a good example of how a species’ demographics can change across its range. Cranberry growers in New Jersey, Massachusetts, and Quebec regard the caterpillar to be an occasional pest (Landry et al. 2002), yet in Ohio the moth is listed as an endangered species (Rings et al. 1992). Prepupae tunnel into the ground and spin a weak cocoon where they hold for several weeks to months (longer southward) before pupating. Adults come to light and bait beginning in late August in Canada; from New England to New Jersey the species flies mostly from late September through October; and is on the wing through November in the South. The eggs, which are laid loose over plant debris around the base of the foodplant, overwinter (Landry et al. 2002).

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 461

Sloping Sallow Epiglaea decliva ! recognition Brown to red-brown, sometimes with pinkish cast; dorsal stripe composed of elongate, pale middorsal spots at back end of T3–A7. Weak middorsal stripe on first two thoracic segments. Subdorsal stripe nearly obliterated. Indistinct white to yellow spiracular line; body decidedly paler below stripe. Prothoracic plate well differentiated, divided over midline. Head mottled with black and prominent coronal bars. Spiracles cream or brown with black rim, or wholly darkened. Larva to 5 cm. Last instar of the Brown Angle Shades (Phlogophora periculosa) somewhat similar in appearance (see page 403). ! occurrence Barrens, thickets, woodlands, and forests from southern Ontario and Quebec to Maine, to Georgia (Piedmont) and Missouri; apparently absent from coastal plain south of New Jersey. One generation with mature caterpillars from late May (New Jersey) through June. Somewhat local southward. ! common foodplants Apple, buckeye, cherry, maple, oak, viburnum, and likely other woody plants. We suspect late instars feed generally on shrubs and small trees. ! remarks The caterpillars abhor light; even the first instars will leave their feeding sites and seek shelter when exposed to bright lights. Middle and late instars apparently leave foliage to seek out hiding places in bark crevices and litter, then move up on to the host to feed at dusk. The Sloping Sallow can be abundant in forests of the Appalachian and Piedmont regions, yet it is curiously absent from many forest types. We often see it in woodlands with rich soils, and unlike many other “glaeas,” this moth is not common in barrens, nor is it associated with heaths. Adults emerge and fly in the fall and come to bait mostly from October to early December; stragglers are present into January in southern portions of range. Winter is passed as an egg. Expect hatching about the time wild cherry leafs out in the spring.

462 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

Roadside Sallow Metaxaglaea viatica ! recognition Stout, variegated brown cutworm with zigzagging, thin, black spiracular stripe. Body brown and heavily mottled above spiracular stripe; tan, mostly unmarked, and frequently tinted with red below stripe. Prothoracic shield darkened with pale subdorsal line. Vague middorsal line, mostly broken. Spiracles black, small. Head as in inset. Larva to 4.5 cm. The Footpath Sallow (M. semitaria)—thought to be primarily a blueberry feeder—with spiracular stripe also zigzagging but white; middorsal stripe cream and more conspicuous, and less mottled in aspect (page 464). The Viburnum Sallow (M. inulta) occurs widely northward, but is mostly restricted to Piedmont and mountains south of Pennsylvania. Its patterning is much subdued and the pinkish-gray to brown ground color is densely salted with whitish spots below the spiracular stripe (see page 464); a thin middorsal stripe is often present. Little is known about the life history of the Southern Sallow (M. australis), a species of the Atlantic Coastal Plain from Maryland southward (see page 464). Additional information on the genus can be found in Schweitzer (1979b). ! occurrence Woodlands and forests from Missouri to Ohio; extreme southern Maine to northern Florida and northeast Texas. One generation with mature caterpillars from April through June. Common. ! common foodplants Natural hosts include apple, chokeberry, crabapple, and mountain ash (all Rosaceae). Our captive larvae accepted black cherry, blueberry, red maple flowers and seeds, and oak, although some larvae refused the last. ! remarks Metaxaglaea emerge and lay their eggs in the fall or winter on and around buds. Eggs either have no diapause or it is terminated after just a few warm days. Eggs hatch asynchronously and may start eclosing in late January in New Jersey (Schweitzer 1979b). In the Roadside Sallow, most eggs eclose so early in the spring that we surmise red maple flowers or buds—the most obvious plant tissue—serves as a staple for nascent larvae. As their drab coloration would suggest, late instar Metaxaglaea and most other glaeas are climbing cutworms that spend the day in leaf litter and ascend the stems of hosts after nightfall to feed.

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 463

Holly Sallow Metaxaglaea violacea ! recognition Tan to flesh ground color with gray mottling and black spotting; some individuals predominantly gray, even waxy bluish. A1–A8 with black middorsal spot towards front edge of each segment. Setal bases smoky gray to black, contrasting with pale ground; setae inconspicuous. Wavy, ill-defined, yellow-orange to peach middorsal and spiracular stripes most pronounced across thoracic segments. Short, peach subdorsal stripe through prothoracic shield. Head with pale snowflake markings over each lobe and directly above triangle; sometimes partially withdrawn into T1. Spiracles black. Elongate, black spot (L1 pinaculum) posterior to each spiracle on A1–A6. Antennae and true legs translucent. Larva to 5 cm. Middle instars more boldly marked (lower right). ! occurrence Woodlands, forests, swamps, yards, and other habitats from Kentucky, southeastern Pennsylvania, and Cape Cod region of Massachusetts, south to northern Florida and Texas. One generation with mature caterpillars by mid-April in Florida; mostly late May into June in New Jersey. Locally common. ! common foodplants Holly; American holly (Ilex opaca) confirmed in nature, but some other evergreen Ilex species, notably inkberry (I. glabra), documented in captivity. ! remarks The gray to bluish body with yellow stripes and black pinacula distinguish the caterpillar from at least the third instar onward—it is the only caterpillar with this coloration on American holly. The first instars hatch in early spring before holly buds open. In captivity, recently eclosed larvae can subsist on the cuticle of buds and old leaves until budbreak, when they switch to new leaves and flowers. The boldly marked early and middle instars often rest on undersides of old holly leaves where they do not seem cryptic. Last instars can be found by searching litter and surface soil at the base of the host in late spring. In most years, the caterpillars mature about the time that hollies start dropping their blossoms. The significance of the larval coloration, which borders on aposematic, is unknown and departs radically from that of other glaeas. Adults can be abundant in southern New Jersey during October and November; DFS has encountered over a thousand adults along a single bait trail. Despite these numbers, the moth is seldom so common as to seriously injure holly. Eggs overwinter, but many adults survive the winter and continue ovipositing into March, especially when persistent cold weather curtails activity from November through February, the normal oviposition time.

464 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

Southern Sallow Metaxaglaea australis

Maryland south along coastal plain to northern Florida, and west to Louisiana. One generation with mature caterpillars probably from April (southward) to May. Local and uncommon. Natural hosts unknown; our caterpillars reared on young cherry and oak.

Viburnum Sallow Metaxaglaea inulta

Manitoba to Nova Scotia, south to Georgia (mountains) and Missouri. One generation with mature caterpillars mostly in May and June. Local southward; more common and generally distributed northward. Viburnums, especially possumhaw (Viburnum nudum) and related species; rejects mapleleaf viburnum.

Footpath Sallow Metaxaglaea semitaria

Eastern Kentucky, Ohio, and coastal New Hampshire, south to northern Florida and Louisiana. One generation with mature caterpillars from April (southward) to June. Locally common. Blueberry, chokeberry, and oak; DFS has reared cohorts on apple, chokeberry, crabapple, and mountain ash.

Poplar Catkin Sallow Anathix puta

Transcontinental in Canada, south in East to Connecticut, Pennsylvania, and Great Lakes states. One generation with mature caterpillars in spring. More common northward. Quaking aspen; coloration suggests late instars feed on ground, perhaps on fallen leaves and fruit (compare with Bicolored Sallow, Sunira bicolorago, opposite).

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 465

Bicolored Sallow Sunira bicolorago ! recognition Variegated, brown cutworm with dark brown, dorsal patches (“shields”) that intensify rearward. Body thickening rearward to A7–A8. Faint middorsal and subdorsal stripes. Ground color abruptly paler below level of spiracles. Prothoracic shield darker than adjacent areas with middorsal and subdorsal lines. Somewhat thickened setae from pale pinacula, longer anteriorly. A10 with small dark plate. Head small, shiny brown with diffuse, dark patch over each lobe. Larva under 3 cm. Caterpillars of Anathix similar in appearance. ! occurrence Parks, woodlands, and forests from Canada, south to central Florida and Texas. One generation with mature caterpillars in late spring. Abundant. ! common foodplants Initially buds, flowers, young seeds, and less often leaves of blueberry, cherry, elm, maple, oak, peach, plum, poplar, willow, and presumably many other trees (Forbes 1954, Rings et al. 1992, and our records). Crumb (1956) reports cabbage, dock, maple, and tobacco (often in vicinity of maples) for older larvae. Additional hosts listed below. ! remarks We use only the common name of the adult here because the caterpillar, known as the Shield-marked Sallow, is seldom seen. The Bicolored Sallow is among the most abundant eastern forest owlets. This is the strawcolored, fall-flying owlet seen darting around woodlands just after sunset, with its numbers peaking through the time of fall color change. DFS once worked a bait trail in Connecticut that he guessed was serving 10,000 adults. We have found young, wild larvae on numerous occasions in red maple flowers picked to feed other caterpillars. Larvae also have turned up in our sleeves on flowering highbush blueberry, red and sugar maple, and scrub oak. DLW collected two late instars on the ground, beneath leaf litter, one of which was reared to maturity on dead blueberry leaves. Hajek et al. (2000) collected 47 larvae from leaf litter. While larvae will eat dead leaves, they prefer new growth of almost any tree or shrub. Related European species shelter in leaf litter by day and ascend plants by night to feed on new foliage and especially flowers (Porter 1997). Crumb (1929) made the curious observation that freshly collected larvae give off a strong odor of heliotrope. Captive females lay the eggs in masses, commonly on red maple buds and flowers.

466 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

Common Hyppa Hyppa xylinoides ! recognition Brown, mottled, rather variable in color, with characteristically humped A8. Pronounced white spiracular line on T1 weakens on T2 then enlarges into vague, broad stripe whose upper edge includes spiracle on A8; rearward stripe becomes constricted and extends as well-defined white line along outer face of anal proleg. Poorly developed middorsal and subdorsal stripes. Larva to 3.5 cm. The closely related Summer Hyppa (H. contrasta) flies in July between the two broods of the Common Hyppa. Its caterpillars presumably mature in August and September. Characters have not been identified that allow for the separation of its caterpillar from that of the Common Hyppa. ! occurrence Woodlands and forests from southern Canada to northern Georgia (mountains). Two generations with mature caterpillars in June and July, then again from September to November in Missouri and Connecticut. Common. ! common foodplants Broadly polyphagous on forbs and low woody plants, especially low forest herbs. ! remarks The Common Hyppa may be found at night by searching low vegetation, such as asters, clover, and willowherb (Epilobium). Roadsides running through woodlands and forests can be productive. Curiously, the integument collects moisture; on cool wet nights the caterpillar may accumulate a hoary glaze of condensation over its thorax and abdomen (upper image). Adults come to both light and bait. The last instar or prepupa overwinters. Our figured individuals were not reared to maturity, so it is possible that both species are represented among the three images.

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 467

American Dun-bar Cosmia calami ! recognition Shape diagnostic: short, stocky, thickened through middle of abdomen, posterior end rounded downward, appearing somewhat abbreviated. Yellow, lime, sea, or waxy green, densely mottled with irregular pale spots. Welldeveloped middorsal stripe; sometimes with thin, faint subdorsal and spiracular stripes. Head pale green. Larva less than 3 cm. ! occurrence Barrens, woodlands, and forests from southern Canada to Florida and Texas, but local across parts of coastal plain. One generation with mature caterpillars from April to June. Very common to abundant over much of range. ! common foodplants Oak. Also consuming other caterpillars—see Remarks. ! remarks Over many areas of the East, the American Dun-bar is among the most common moths seen at light in early summer. The caterpillar is active in the spring, feeding on young oak foliage. Like some of its Old World congeners, early instars will spin and reside within a crude shelter. Mike Singer has encountered wild larvae on four occasions. Twice, the caterpillars were concealed; one appeared to have spun its own shelter and one was found in a curled, dry leaf. The two others were found resting on the underside of a leaf, over the midrib. According to Forbes (1954), a principal component of the diet of the American Dun-bar is other caterpillars. We have reared several caterpillars to maturity on a diet of foliage only. Its European counterpart, the Dun-bar (C. trapezina), is also omnivorous, feeding on a variety of woody trees and shrubs as well as other caterpillars. Inchworms, such as the Winter Moth (Operophtera brumata, Geometridae), are a favorite prey of the Dun-bar (Carter and Hargreaves 1986). The egg overwinters.

468 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

tentmakers Enargia

The taxonomy of the genus was recently revised by Schmidt (2010b). McGuffin (1958) provides notes on two of our species. Life histories of our three Enargia are similar. There is one generation per year; the caterpillars hatch in early spring and mature by early July. Most foodplant records are from poplar (especially trembling or quaking aspen), but willow, alder, and birch are also listed; white birch is a favored host of the Tawny

Tentmaker, E. fausta (Schmidt 2010b). Larvae are infrequently encountered in our experience—the degree to which hosts other than willows, aspens, and poplars are used warrants further study. Larvae spin purselike shelters, usually between two overlapping leaves, lining the inside with silk. The caterpillars are reported to be “notorious cannibals” (Forbes 1954). Adults are attracted to light, but come more readily to bait. Eggs overwinter. Larval characters that would allow for species-level identification of Enargia caterpillars are unknown to us.

Straw-eyed Tentmaker Enargia decolor

Woodlands and forests. Transcontinental in Canada, south in East to northwestern New Jersey and Great Lakes states. Common northward; most frequently encountered member of genus. Poplar, with quaking aspen a favored host; also recorded from alder, birch, and willow. See also Wong and Melvin (1976).

Tawny Tentmaker

Enargia fausta (= E. infumata of previous authors) Woodlands and forests. Central Alberta to Nova Scotia, probably southward into northern border regions of United States. Uncommon. White birch and quaking aspen; also balsam poplar (see Schmidt 2010b). Due to taxonomic confusion concerning previous application of names, we are uncertain if our image, in fact, represents E. fausta.

Smoky Tentmaker

Enargia infumata (= E. mephisto of previous authors) Woodlands and forests. Alaska to New Brunswick, south to New Jersey and Great Lakes states. Locally common. Poplar, especially quaking aspen; fewer records for alder, birch, and willow (but see Schmidt 2010b). Our image appears to be of a molting or prepupal larva that has lost its patterning.

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 469

Blackcheeked Tentmaker Ipimorpha pleonectusa

[Even-lined Sallow]

! recognition Stout, shiny, translucent, green with prominent white middorsal stripe that begins along trailing margin of T1. Body surface leathery, corrugations more evident over dorsum. Dorsal setae from small pale bases; those bearing D1 setae most evident. Weak subdorsal stripe sometimes present. White to yellow subspiracular stripe fading rearward, best defined along thoracic segments. Head tricolored: pale sea green behind black bar and porcelain white across front. Spiracles straw colored to brown, with dark rim. Larva to 4 cm. Middle instars pale green, translucent with white middorsal, subdorsal, and spiracular stripes. ! occurrence Woodlands and forests from Alberta to Nova Scotia, south in East to North Carolina. Pennsylvania, and Great Lake states. One generation with mature caterpillars in June in Connecticut. Common. ! common foodplants Aspen and poplar. ! remarks As suggested by Forbes (1954), the Blackcheeked Tentmaker shares affinities with Cosmia. Both are stout, leathery, and bear a broad dorsal stripe and distinctively rounded rump. The caterpillar forms a shelter, often between two overlapping leaves. A generous sheet of silk is spun within. Forbes (1954) wrote: “When disturbed the larva will stick its head out of the leaf [shelter], and then it looks like a lady-bird.” Many shelter-feeding Lepidoptera are brightly or otherwise strikingly colored (e.g., some skipper butterflies, Thaxter’s Sallow (Psaphida thaxterianus), and The Laugher (Charadra deridens)); or they have a boldly marked head much like the Blackcheeked Tentmaker. Presumably, such coloration serves to startle the many birds that forage by tearing open leaf shelters (Janzen et al. 2010). Its British congener, the Olive (I. subtusa), pupates in a belowground cell, and emerges after about two months, in July and August (Porter 1997). Adults are attracted to sugar baits. Eggs overwinter on bark.

470 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

antitypine sallows

Family Noctuidae: Subfamily Noctuinae: Tribe Xylenini: Subtribe Antitypina About 29 species occur in North America, although most are western. Principal genera include Andropolia, Fishia, Mniotype, Platypolia, Sutyna, and Xylotype. In most, the larva is green through the penultimate instar, but then may become

pink, tan, or brown in the last instar. Coloration within a species can be highly variable. As in other xylenines, the dorsal pinacula are often white. Antitypines aestivate as pupae, unlike other Xylenini which pass the summer as prepupae. Foodplants are poorly known; we suspect most are generalists on shrubs or low plants. Our species overwinter as diapausing eggs laid in the fall.

Wandering Sallow

Fishia illocata (= Oligia illocata) [Wandering Brocade] Transcontinental across Canada, south in East to Georgia (mountains) and Missouri. One generation with mature caterpillars in late spring. Locally common, especially on acid soils. Alder, birch, poplar, buffaloberry (Shepherdia), willow, and likely many other woody plants. First image is a penultimate instar.

Three-horned Sallow Pachypolia atricornis

Northern tier of states, south in Appalachians to North Carolina, and in Midwest to Kansas. One generation with caterpillars maturing in midspring. Uncommon. Natural hosts unrecorded; our captive larvae were started on red and silver maple flowers and then switched to cherry after budbreak.

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 471 Gray-dusted Sallow Platypolia anceps

Canada and northern tier of states, south in high Appalachians to North Carolina. One generation with mature caterpillars in late spring. Uncommon. Aster macrophylla (A. E. Brower in Proctor 1946); ex ova cohorts have been reared on blueberry and cherry by DFS and Tim McCabe.

Checkered Sallow Sutyna privata

At least Wisconsin to Nova Scotia, south to Florida and Kansas, assuming all eastern populations represent a single entity. One generation with mature caterpillars in late spring and early summer. Usually uncommon. Presumably a generalist: one wild larva from Vaccinium. Captive larvae accept aster, blackberry, and lettuce. The first image is a penultimate instar preparing to molt.

472 Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini

Adorable Sallow “Platypolia” mactata (= Oligia mactata)

[Adorable Brocade]

! recognition Plump, pale, greenish, orangish, pinkish to tan caterpillar with dense, pale speckling; ground color paler below spiracular stripe. Thin, broken, white middorsal and subdorsal stripes. Spiracular stripe edged above with brown, fading rearward and dropping down outer face of proleg; pale spotting below forming ill-defined subspiracular stripe. Dorsal setal bases white. Rump broadly rounded. Spiracles pale tan with dark outer rim. Head with reticulate markings; triangle pale tan without mottling. Larva to 4.5 cm. Penultimate instar waxy green with more well-defined stripes, bright white dorsal pinacula, and abundant white speckling (inset). Wandering Sallow (Fishia illocata) (= Oligia illocata) similar (page 470); in our examples speckling more subdued, especially below spiracular stripe. Middorsal, subdorsal, supraspiracular (weak and broken), spiracular, and subspiracular stripes more prominent than those of Adorable Sallow. The caterpillar also resembles that of Checkered Sallow (Sutyna privata) (previous page). ! occurrence Woodlands and forests, mostly on rich soils. Transcontinental in Canada, south in East to Georgia. One generation with mature caterpillars in spring and early summer. Common. ! common foodplants Presumably many woody plants; one wild caterpillar from Carolina silverbell. Our ex ova caterpillars were reared on cherry and gray birch. ! remarks The absence of published larval foodplants and the coloration of the last instar of this common sallow suggests that the caterpillar may behave as a climbing cutworm, i.e., feeding at night and then descending into leaf litter during the day. We follow Lafontaine and Schmidt (2010) in placing the Adorable and Wandering Sallows in Antitypina, near Sutyna and Platypolia, which their larvae and life histories closely resemble. Until recently, both were mistakenly classified in the apameine genus Oligia. The egg overwinters.

Pinions, Sallows, and Kin Subfamily Noctuinae: Tribe Xylenini 473

Broad Sallow Xylotype capax ! recognition Smoky black to slaty gray or purplish with pale spots and broad spiracular stripe that narrows appreciably along thorax. Subtle, orangeyellow supraspiracular spots on A1–A8. Middorsal stripe thin, mostly obliterated. Dark pigment sometimes forming diffuse spot between dorsal setae on A1 and A2; A8 with dark subdorsal spot that may fuse over midline. Spiracular stripe often with dark patch below each abdominal spiracle. Pinacula white. Black spiracles run along upper edge of lateral stripe. Head shiny orange with brown reticulations. Larva to 4.5 cm. We do not know of characters that will separate its caterpillars from those of the closely related Arcadian Sallow (X. arcadia) (next page). ! occurrence Pine barrens (especially with wetlands), balds, forested and heath swamps, and in mountains, dry oak forests and other shrubby, acid soil communities. Northeastern New York to southern Maine, south along coastal plain to southern New Jersey, and from Pennsylvania to northern Georgia in mountains. One generation with mature caterpillars in late spring. Locally common. ! common foodplants Wild larvae from cherry and highbush blueberry in New Jersey; captive larvae have been reared on apple, cherry, and red oak. ! remarks Eggs are laid in autumn and then overwinter; they hatch in spring before most buds open, usually in March in New Jersey. (This suggests that buds, such as those of blueberry, serve as an initial food.) Second and third instars of the closely related Arcadian Sallow (X. arcadia) have been swept in number from blueberry in Canada. There is some question as to whether our two Xylotype are distinct—adults of X. capax from northern parts of New England tend to be smaller in size and thus approach X. arcadia in stature. The Broad Sallow is a lateseason moth; adults appear in the last days of September northward and are on the wing even later southward. Adults are attracted to both light and bait. Its numbers fluctuate markedly from year to year.

474 Griffins and Satyrs Subfamily Noctuinae: Tribe Ufeini

Arcadian Sallow

Xylotype arcadia (Tribe Xylenini) Transcontinental in Canada, south in East to at least Maine. One generation with mature caterpillars in late spring. Locally common. Alder, blueberry, bog laurel (Kalmia polifolia), cherry, larch, and spruce (Robinson et al. 2002, Duncan 2006); we suspect that blueberry and bog heaths serve as its primary hosts.

Hops Angleshade

Niphonyx segregata (Tribe Xylenini) Asian; recently established along Eastern Seaboard from Connecticut south to at least Delaware. Two generations with mature caterpillars in July and then again in fall. Locally common. Hops. The taxonomic position of this genus within the Noctuinae is unresolved.

Griffins and Satyrs

Family Noctuidae: Subfamily Noctuinae: Tribe Ufeini We diverge from the Lafontaine and Schmidt (2010) checklist by recognizing a separate tribe for the genus Ufeus based on its exceptional larval characters and life history. The prolegs bear more than 50 crochets and there are two L group setae on A9. Both our eastern species feed on poplars. In contrast to other Xylenini, the adults hatch in early summer and then aestivate until the fall. Adults overwinter and then mate in the spring.

Griffin Ufeus plicatus ! recognition Stocky, smooth, brown, thickened through anterior abdominal segments; dorsum mostly unpigmented between D1 pinacula. Pale, broken, middorsal and subdorsal stripes, with latter being nearly obliterated in some individuals. Dorsal setae from nearly white pinacula; D2 pinacula touching subdorsal stripe. Broad, dirty salmon, spiracular stripe. Prothoracic and anal plates darkened, shiny, cut by middorsal and subdorsal stripes. Prolegs with more than 50 crochets. Larva to 4.5 cm. We do not know of characters that will separate its caterpillars from those of the Satyr (U. satyricus) (see below). ! occurrence Poorly delineated; at least river courses with mature cottonwoods from southern Quebec, central Connecticut, and southeastern Pennsylvania westward to Iowa. One generation with mature caterpillars from late spring into midsummer (northward). Rarely encountered. ! common foodplants Ours from cottonwood (Populus deltoides). Western species of Ufeus also associated with Populus. Handfield et al. (1999) lists alder, but this should be confirmed given the wandering capacities of larvae. ! remarks Ufeus caterpillars can be distinguished from those of all other North American owlets by examining the prolegs, which bear more than 50 needle-thin crochets (other noctuoids tend to have less than 35). Because of their exceptional crochets and the presence of an extra L seta on A9, Crumb (1956) recognized the

Griffins and Satyrs Subfamily Noctuinae: Tribe Ufeini 475

Ufeinae as a distinct subfamily within the Noctuidae. Our images derive from an early June collection of seven caterpillars recovered from a mature cottonwood (Populus deltoides) that was growing along a highly developed stretch of the Connecticut River in East Hartford. The larvae were secreted beneath loose bark and in recesses of the tree’s exposed roots. Ufeus caterpillars are muscular and capable crawlers, as would be suspected of a species that shelters on the trunks of canopy trees by day. The cottonwood on which we found our caterpillars was massive; at night, larvae were moving no less than 10 to 20 meters to feed, and perhaps twice that distance. Both eastern Ufeus are listed as endangered in Ohio. Despite years of moth collecting in southern New England, ours was the first record of the species from Connecticut. This suggests that Ufeus plicatus adults are only weakly attracted to light and that the rarity of U. plicatus, and perhaps other members of the genus, is more apparent than real. Unlike other xylenines, which generally enter the soil to pupate, ours fashioned a slight cocoon in paper toweling above the floor of the container—pupation followed two to three days later. Adults emerged after three weeks and then aestivated until the fall. The Griffin is a long-lived moth—adults will fly into March or April of the year following their emergence. One wonders if the species might be reliant on the availability of loose bark beneath which the larvae (and adults) can shelter. And it is even conceivable that females seek out such crevices to lay their eggs. Younger trees rarely have bark with subcortical spaces where the larvae could shelter. While our larvae were solitary, at least in the last instar, our image for the Satyr (U. satyricus) (below) indicates that this species is gregarious. One wonders if pheromones left by the larvae have any role in coordinating larval movements back to the diurnal sheltering sites. Neither species comes to baits.

Satyr

Ufeus satyricus Alberta to Nova Scotia, south in East to Pennsylvania (mountains) and Great Lakes states. One generation with mature caterpillars in late spring. Locally common northward. Alder and poplar (Handfield et al. 1999).

476 Quakers, Woodgrains, and Kin Subfamily Noctuinae: Tribe Orthosiini

Quakers, Woodgrains, and Kin Family Noctuidae: Subfamily Noctuinae: Tribe Orthosiini

The orthosiines are a bridge group of sorts, sharing characteristics with both Xylenini and Hadenini. The 14 widespread eastern species, representing five genera, are all figured. The hypopharynx is divided by a groove on its dorsal surface. Orthosiines overwinter as pupae, but remarkably, the adult moths are fully formed within the pupal casing by midsummer, save for their abbreviated wings—eight or nine months in advance of their eventual emergence. Not surprisingly, they are among the first moths to eclose in the spring. Females lay their eggs

in rafts. Last instars of the Confused Woodgrain (Morrisonia confusa) spin a dense shelter. The Distinct Quaker (Achatia distincta) and a few Orthosia form flimsy shelters, commonly prior to a molt. Like the xylenines, the caterpillars feed on trees and other woody plants well above the ground. Because all are broadly polyphagous, we make no attempt to provide comprehensive foodplant listings; see Rings et al. (1992) and Robinson et al. (2002, 2011) for more complete summaries of foodplant records. Caterpillars of Orthosia require new foliage. Collectively, orthosiines are among the most abundant spring caterpillars in forested ecosystems, and by extrapolation are quintessentially important in the diet of songbirds and other insectivorous animals. There are six larval instars.

Gray Woodgrain Morrisonia mucens

Mostly coastal plain from southeastern Massachusetts to Florida and west to Texas. One generation with mature caterpillars from May at least into July. Uncommon. Oak; associated with scrub or blackjack oak (habitats) northward; wild larvae from farkleberry (Vaccinium arboreum) and hawthorn in Florida (DLW). Actively feeding larvae are green (upper image). The pinkish form (lower image) sent to us by Lyle Buss may represent a (long-lived) prepupal form.

MOTHS AND EARS AND BATS By the time most moths crawl from their roosts and take wing, the night sky has already filled with hungry bats using high-pitched cries to echolocate prey (Fullard 1987, Ratcliffe 2009). This takes place in the same fashion that a trooper employs radar or laser pulses to identify speeding vehicles. Under this chronic and highstakes pressure, moths have repeatedly and independently evolved organs—on their abdomen, wings, thorax, and even mouthparts—that detect the ultrasonic frequencies emitted by hunting bats. The ear of all owlets is located below the hindwing; a pale membrane, the tympanum (Tym), stretches across the cavity with two or three attached nerves. The tympanum is sensitive to ultrasound and serves as the animal’s early-warning system. An owlet’s first response will be to fly away from the ultrasound source, but if the bat continues to close in, the auditory nerves will trigger a series of additional evasive behaviors (Fullard et al. 2003). Some moths immediately initiate a rapid, erratic path of flight that diminishes the likelihood of their capture; others power dive to the ground, and another subset draws the wings upward over their thorax and drops from the sky (Ratcliffe 2009). This last, evidently, can be duped simply by the jingling of keys—some summer night when ! Left ear of a wainscot (Leucania). Note the hood or you are passing a street light with a bevy of whirling moths, try taking out a set of flap to the right of the tympanum (Tym) that serves keys and giving them a good shake. the same role as our auricle or pinna, i.e., to direct sound toward the tympanic membrane. !

Quakers, Woodgrains, and Kin Subfamily Noctuinae: Tribe Orthosiini 477

Speckled Green Quaker Orthosia hibisci ! recognition Green or blue-green with continuous, white middorsal and spiracular stripes, and broken, subdorsal stripe; creamy speckles abundant. Spiracular stripe running through or above spiracles on A1–A7 and below spiracles on T1 and A8. Stripe thickened across A7–A10, sometimes with incomplete dark edging above, particularly in blue-green forms (upper inset). Spiracles pale with dark rim. No transverse line across dorsum of A8 (as in Gray Quaker, O. alurina). Dark-form caterpillar smoky green, especially dark between spiracular and subdorsal stripes (lower inset). Larva to 4 cm. Earlier instars of many Orthosia and some Xylenini resemble caterpillars of the Speckled Green Quaker. ! occurrence Woodlands and forests from Canada, south to Florida and Texas. One generation with mature caterpillars from late April to end of June. Very common to abundant. ! common foodplants Many shrubs and trees including both broadleaf and coniferous species: our records from apple, ash, aspen, autumn olive, birch, cherry, Douglas-fir, elm, gooseberry, larch, maple, oak, poplar, spruce, and willow. ! remarks This is one of the most ubiquitous forest insects of spring and early summer. In boom years its caterpillars can be found on almost any tree or shrub. While the adults of Orthosia were characterized by Forbes (1954) as “fairly homogeneous in appearance and vestiture,” the caterpillars are a diverse lot; some species are green and cryptic, others brown and barklike, a few are brightly colored, etc. Orthosia species with caterpillars colored in brown and other earth tones rest in bark crevices or even descend into litter by day. Some members of the genus are shelter forming (at least in early and middle instars). Old World species, of which there are many, are sometimes cannibalistic (Sugi 1987, Porter 1997). Pupation in all Orthosia occurs belowground in a slight cocoon. Adults of the Speckled Green Quaker and other Orthosia come to light, bait, maple and black birch sap, and later in the spring, to red maple and willow blossoms.

478 Quakers, Woodgrains, and Kin Subfamily Noctuinae: Tribe Orthosiini

Gray Quaker Orthosia alurina ! recognition Ground color ranges from yellow-green to waxy green, to nearly black; coloration often darker above spiracular stripe than below. Densely speckled, more so than other Orthosia. White middorsal stripe parallel-sided. Subdorsal stripe often broken into spots, weakly developed across thoracic segments and rearward of A8. White spiracular stripe either thin or thickened. When thin, running through spiracles on A1, A2, and A7, passing along upper edge of those on A3–A6, and then dropping below spiracle on A8. When stripe thickened, running through all spiracles. Spiracles tan, rimmed with black. Dorsal setae from bright white bases. A8 with transverse white line extending across hump. Head pale green, shiny black, or mixture of both. Forms with black head often have black plates on outer face of prolegs on A3–A6, and black anal plate. Even in black forms, prothoracic shield may remain green (lower left). Larva to 4 cm. ! occurrence Woodlands and forests from Great Lakes region, southern Ontario and Quebec, to Florida and Texas. One generation with mature caterpillars in late spring, before foliage hardens. More common and general southward. ! common foodplants Cherry commonly; also basswood, crabapple, holly, serviceberry, and presumably other woody plants. ! remarks The moth was regarded as rare by Forbes (1954), especially eastward. Beginning in the 1950s eastern collectors started seeing the species for the first time: DFS (Pennsylvania), Joe Muller (New Jersey), and Sydney Hessel (Connecticut). In recent years, the moth has become increasingly common and widespread across the Northeast. Population numbers fluctuate markedly among seasons. All Gray Quaker caterpillars start out green but may darken appreciably if crowded. Three-quarters of our larvae reared in sleeves at higher densities were dark, while all individuals reared in sleeves at lower densities on the same plants remained green throughout their development (upper left). Interestingly, when a cohort of larvae was resleeved at lower density, half of the dark-form larvae reverted back to their previous green coloration. Analogous melanistic forms, induced by high larval densities, occur in many geometrid defoliators: the Cabbage Moth (Mamestra brassicae) (Goulson 1994), several armyworms (Spodoptera) (e.g., Wilson et al. 2001), and others. Adults of this and most quakers feed for weeks before producing eggs, visiting red maple, willow, plum and other flowers, tree wounds, and other sugary substrates.

Quakers, Woodgrains, and Kin Subfamily Noctuinae: Tribe Orthosiini 479

Garman’s Quaker Orthosia garmani ! recognition Drab, smoky green-brown to frosty gray, red, or tan brown with broken, white middorsal, subdorsal, and spiracular stripes. Ground color often with pinkish cast, especially where segments overlap. Caramel-colored prothoracic shield, divided by pale middorsal stripe, darkened along its leading margin. Pale, caramel-colored anal plate. Body decidedly paler below spiracular stripe. Spiracles black or tan with black rim. Faint bars (coronal stripes) to either side of midline. Larva to 4 cm. Middle and penultimate instars green (inset). ! occurrence Woodlands and forests from Great Lakes region, extreme southern Canada, and Massachusetts, south to Georgia and Texas, but absent from many areas within this range. One generation with mature caterpillars in late spring before foliage hardens. Locally common. ! common foodplants Beach plum, blueberry, cherry, dogwood, elm, hawthorn, maple, walnut, and a variety of other woody plants (McFarland 1967, Godfrey 1972)—presumably a generalist on woody plants. ! remarks The green instars can be found by beating, but last instars probably spend the day in the litter. While generally considered scarce, this is not the case in Connecticut oak woodlands in recent decades where the adults fly in the early spring (mid-April). The first captures for Garman’s Quaker in Connecticut date to the late 1960s. Forbes (1954) did not have records of the moth from New England; see Alternate Woodling (Egira alternans) and Gray Quaker (O. alurina) for related discussions. When sheet collecting, watch for adults hiding under folds or seeking shelter underneath your drop cloth. The larva in the inset is a penultimate, readying for its final larval molt—the head of the new instar is visible as a swelling within the prothorax of the figured individual. Quakers are among the first moths to issue from their pupae in a given year. Males of Garman’s Quaker will fly on very cool evenings, even when temperatures drop to 3° or 4° C. Flight activity occurs earlier on cold evenings, but not until later, if at all, on warm evenings. Unlike most members of the tribe, the adult does not feed.

480 Quakers, Woodgrains, and Kin Subfamily Noctuinae: Tribe Orthosiini

Ruby Quaker Orthosia rubescens ! recognition Coloration of last instar usually diagnostic: dark above broad white spiracular stripe and pale below (upper row images); frequently with black patches about white spiracle on A3–A7. Eighth segment swollen. Thin, poorly developed, white middorsal and broken subdorsal stripes. Trunk with abundant white flecking. Green and blue-green forms difficult to characterize (lower row images); usually with ground smokier above spiracular stripe and more white flecks over dorsum. In darker forms, head pale brown with snowflake spots and coronal bar to either side of midline; broad, dark, transverse bar extends between eyes, passing through triangle. Larva to 4 cm. Middle instars resemble those of Speckled Green Quaker (O. hibisci) and some pinions (Lithophane). ! occurrence Woodlands and forests from Wisconsin to Nova Scotia, south to Georgia and northern Texas. One generation with mature caterpillars from late April into early July. Very common to abundant. ! common foodplants Many shrubs and trees including both broadleaf and coniferous species: e.g., beech, birch, blueberry, cherry, chestnut, elderberry, gooseberry, greenbrier, hemlock, holly, hophornbeam, maple, oak, and viburnum; less commonly on forbs and other low-growing plants. ! remarks A common caterpillar in most years and most places, with occasional banner years when larvae will be found on almost any woody plant. The green early instars through penultimates rest on the underside of new foliage. They are easily mistaken for young pinions (Lithophane), but quakers have one white spot to either side of the midline (compared with two or three in pinions). The brown and white last instar rests on bark by day, often well away from foliage. We have found dozens of last instars under burlap bands while censusing Gypsy Moth (Lymantria dispar) larval populations in the foothills of the Appalachians. Adults of most quakers come to bait in the spring, at least until red maple and other nectar becomes available.

Quakers, Woodgrains, and Kin Subfamily Noctuinae: Tribe Orthosiini 481

Subdued Quaker Orthosia revicta ! recognition Body brown with broad, pale spiracular stripe; A8 somewhat humped and swollen. Middorsal, subdorsal, and supraspiracular stripes obscure and broken. Dorsal setae from inconspicuous white pinacula. Spiracular stripe narrowed at each end of body, width on T1 and A9 about half that of A3; usually with spiracle on A1 and A2 positioned close to upper edge of lateral stripe (spiracles more distant from upper edge in Alternate Woodling, Egira alternans). Orange-brown head usually with darker reticulations and coronal bars. Larva to 4 cm. Penultimate instar with striping more evident (lower inset). Similar in appearance to Alternate Woodling—see next page. ! occurrence Woodlands and forests from British Columbia to Nova Scotia, south in East to Georgia (mountains) and Kentucky. One generation with mature caterpillars in late spring into early summer, before foliage hardens. Common northward. ! common foodplants Apple, ash, basswood, beech, birch, cherry, elm, hickory, maple, oak, poplar, spirea, walnut, willow, and presumably other woody plants. ! remarks According to notes in the Canadian National Collection, there are six instars. The generic assignments of species in the Orthosiini (especially Egira, Orthosia, and Morrisonia) are in much need of phylogenetic study and taxonomic reshuffling. The appearance of the larva of the Subdued Quaker, has much in common with that of the Alternate Woodling (Egira alternans). Phenologically, the Subdued Quaker differs from other eastern Orthosia in having a more extended flight season (it flies from late March to early June in New Jersey). All other eastern Orthosia species emerge and finish earlier in the spring. Adults are common at light and occasional at bait.

482 Quakers, Woodgrains, and Kin Subfamily Noctuinae: Tribe Orthosiini

Alternate Woodling Egira alternans ! recognition Brown with broad white spiracular stripe. Hair-thin, white middorsal, subdorsal, and supraspiracular stripes obscure and broken. A8 somewhat humped. Dorsal pinacula white. Spiracles black. Head orange-red and unmarked or with bar over each lobe and additional mottling. Larva to 4 cm. Middle and penultimate instars with white dorsal pinacula, continuous white middorsal and subdorsal white stripes, and darkened supraspiracular area. Similar to Subdued Quaker (Orthosia revicta) (see Remarks), but dorsal pinacula more conspicuously whitened and lateral stripe tending to be broader (fully enveloping spiracles on A1–A6); also less often infuscated with the browns and reds that are common in many forms of the Subdued Quaker. ! occurrence Pinelands, flat woods, and woodlands; often on acid soils from southern Canada to Florida and Texas. One generation with mature caterpillars from late April into early July. Locally common; abundant over much of coastal plain. ! common foodplants Many woody plants including blueberry, buckeye, cherry, honeysuckle, leatherleaf, mountain holly, oak, rhododendron, silverbell, sourwood, walnut, and willow. ! remarks The moth’s common and scientific names derive from the light and dark bands that encircle the adult’s abdomen. Caterpillars of the Subdued Quaker (Orthosia revicta) are surprisingly similar in appearance. One wonders if the similarity of the two is due to shared ancestry or evolutionary convergence, and if the latter, why such a pattern might be adaptive. The Alternate Woodling abounds in the coastal woodlands of North Carolina and New Jersey, especially those with an abundance of heaths. This moth was called “rare” by Forbes (1954). Curiously, the first records of this common moth in New Jersey and Pennsylvania date from the 1960s, and the first Connecticut records from the 1970s. Adults of the Alternate Woodling visit flowers—chickasaw plum is a favorite in southern New Jersey; they do not come to bait. The flight season is brief, lasting just three weeks in most years. The pupa overwinters belowground.

Quakers, Woodgrains, and Kin Subfamily Noctuinae: Tribe Orthosiini 483

Cunning Woodling Egira dolosa ! recognition Variegated with dark, crudely triangular spots over dorsum; these best developed over A1–A8. In some forms dorsal spots have forward-directed spurs—their shape bears some resemblance to a bat or fleur-de-lis. Ground color varies from rather pale to dark with subtle lavender flush. Thin, pale middorsal, subdorsal, spiracular, and subspiracular stripes. Body pale below spiracular stripe with greenish tint beneath thorax. Prothoracic shield dark and shiny above subdorsal stripe, poorly differentiated below; white middorsal stripe dividing shield, flaring outward behind head. Head dark and shiny with broad, nearly black band between eyes, across lower half of triangle, joined by dark bar that drops down from each lobe; labrum white. Larva to 3 cm. ! occurrence Forests from central Canada, south in East to northern New Jersey and Great Lakes states. Single generation with mature caterpillars in early to midsummer. Common northward. ! common foodplants Poplar primarily and presumably willow; Robinson et al. (2002) list birch and oak. ! remarks Adults fly in midspring. The mother of the figured individual laid her eggs in a refrigerator at temperatures too cool for most moths to be active. Early and middle instars commonly fashion a sparse shelter; new leaves are preferred. Our late instars rested by day in the vertical flutes of the coffee filters lining their rearing containers, suggesting that larvae rest in bark fissures by day. Their coloration is consistent with that of other bark-resting quakers. Caterpillars may feign death when handled. This ruse was so credible in one larva that we set it aside to be preserved, but before we could bring water to a boil, the “cadaver” righted itself and wandered off.

484 Quakers, Woodgrains, and Kin Subfamily Noctuinae: Tribe Orthosiini

Norman’s Quaker Crocigrapha normani ! recognition Last instar reddish brown to tan brown, without obvious striping and mostly undistinguished. Ground color pale below subtle white spiracular stripe (when present). Prothoracic shield brown, shiny, becoming pale laterally. A8 somewhat humped with transverse line connecting posterior dorsal (D2) setae (visible in main image). Spiracles tan, rimmed with black. Head shiny brown with one (larger) or two smaller dark spots over each lobe (lower left inset). Larva to 4 cm. Penultimate instars conspicuously bicolored; dark above and white to cream below, with ground color darkest between white subdorsal and spiracular stripes; head bearing prominent black spot(s) over each lobe (upper inset). Middle instars waxy green with prominent head spots (lower right inset). ! occurrence Woodlands and forests from southern Canada to Georgia and Mississippi. One generation with mature caterpillars from late April through June, early July northward. Very common to abundant. ! common foodplants American hornbeam, apple, ash, aspen, basswood, bayberry, birch, blueberry, cherry, currant, elm, hazel, hophornbeam, huckleberry, maple, oak, rose, spirea, sweet fern, Virginia and white pine, walnut, witch hazel, willow, and many other woody plants. ! remarks This is one of the most ubiquitous caterpillars in late spring beating samples, and liable to turn up on almost any shrub or small tree. The distinctive dark spots on the head, present in both young and mature larvae, are diagnostic. Adults have a long spring flight (end of March through May in New Jersey)—occasionally you will see adults and nearly grown caterpillars in the same woodland on the same day. Disturbed larvae release from their perch and draw themselves into a “knot.” Adults rarely visit bait.

Quakers, Woodgrains, and Kin Subfamily Noctuinae: Tribe Orthosiini 485

Intractable Quaker Himella fidelis (= Himella intractata) ! recognition Pale to blue-green with thin, white middorsal; thick, white subdorsal; four creamy, wavy, lateral stripes. Diagnostic set of wavy lateral stripes composed of two supraspiracular, one spiracular, and one subspiracular pinstripes. Segments T1–A7 with single white spot to either side of midline; A8 with two such spots. Spiracles yellow-orange. Head pearly to sea green, shiny, and unmarked. Larva to 3.5 cm. Middle instars more lime green, but otherwise similarly patterned (lower inset). Compare with Distinct Quaker (Achatia distincta), another common spring owlet, with which it frequently co-occurs. ! occurrence Woodlands and forests from southern Canada to Florida and Texas. One generation with mature caterpillars from late April through June. Very common. ! common foodplants American hornbeam, apple, aspen, blackberry, blueberry, cherry, elm, gale, hackberry, hawthorn, hazel, hickory, honey locust, hophornbeam, oak, viburnum, witch hazel, and many other woody plants. ! remarks Like many forest species, it has boom years but in other years can be scarce. In June 2005, it was the most common owlet caterpillar across much of the Northeast, occurring on most any woody plants examined. The caterpillar dislodges easily from its perch and drops to the ground. If further molested, it will thrash about, drawing itself into a C-shape and then snap back and forth. Adults visit flowers but not bait.

486 Quakers, Woodgrains, and Kin Subfamily Noctuinae: Tribe Orthosiini

Distinct Quaker Achatia distincta ! recognition Sea or yellow to blue-green with only subdorsal stripe well developed. Middorsal, spiracular, and subspiracular stripes thin and broken. Supraspiracular stripe represented by scattered spots. Dorsum and sides with faint speckling, especially obvious under magnification. Dorsum of A1 to A7 with one white spot to either side of midline; some individuals with smaller dorsal spot obliquely situated at leading edge of segment, just above subdorsal stripe. Spiracles yellow-orange. Venter with waxy blue-green patches but otherwise mostly unmarked. Head sea green, shiny, and unmarked except for white line through eyes (stemmata). Larva to 3.5 cm. Middle instars with more glossy aspect and more lime green (lower inset). ! occurrence Woodlands and forests from southern Canada to northern Florida and Texas. One generation with mature caterpillars from middle April to early July. Very common to abundant. ! common foodplants Polyphagous on woody plants: alder, amorpha or false indigo, apple, ash, basswood, birch, blackberry, blueberry, buckeye, butternut, cherry, currant, hickory, honey locust, hophornbeam, maple, ninebark, oak, poplar, redbud, sumac, tupelo, viburnum, walnut, willow, witch hazel, and many other shrubs and trees. ! remarks Caterpillars are common to abundant in spring across a variety of woodland and forest types. Like other vernal orthosiines, it is reliant on new foliage. It is sometimes confused with the Intractable Quaker (Himella fidelis), which has a stronger, whiter subdorsal stripe, and an additional lateral stripe. Egg rafts of Achatia contain 30 or more ova. Caterpillars feed mostly at night. By day, the larva sometimes spins a weak shelter between adjacent leaves, or more commonly seeks refuge in rolls and other shelters spun by microlepidopterans. When disturbed, the caterpillar rolls into a ball. Prepupae take on a dull reddish-green coloration.

Quakers, Woodgrains, and Kin Subfamily Noctuinae: Tribe Orthosiini 487

Confused Woodgrain Morrisonia confusa ! recognition Last instar pale to reddish or orange-brown, densely speckled with minute vague spots (upper left). Faint middorsal, subdorsal, spiracular, and subspiracular stripes. Pale spiracles ringed with black. Head caramel-colored laterally and above but paler across front; shiny, with dark patch over vertex (inset). Antenna with white base and black apex. Larva to 4 cm. Early and middle instars (lower left and right) pale to waxy green with conspicuous, maroon supraspiracular spot on T1–A8; all striping more obvious than in last instar; head with or without dark patch over each lobe. ! occurrence Woodlands and forests from Canada, south to Florida and Texas. One generation with mature caterpillars from June to November. Very common. ! common foodplants Many shrubs and trees including both broadleaf (e.g., ash, basswood, birch, blueberry, cherry, hickory, huckleberry, maple, oak, rhododendron, sumac, sycamore, viburnum, walnut, and willow) and occasionally coniferous trees; also forbs such as purple loosestrife and bush clover. ! remarks Wood and Butler (1991) provide biological notes on the parasites and preferred hosts of the Confused Woodgrain in West Virginia forests. The caterpillar fashions a silken nest, often between two leaves. The handsome, red-spotted, middle instars draw considerably more attention than do the drab last instars. Although earlier instars develop normally, the last instar is greatly protracted, spanning three or four months—in Connecticut, larvae can be found into September and October. Much silk is deposited by the last instar about the inner walls of the shelter—the construction is more fortified with silk than the cocoons of most owlets.

488 Quakers, Woodgrains, and Kin Subfamily Noctuinae: Tribe Orthosiini

Bicolored Woodgrain Morrisonia evicta ! recognition Smoky pink above narrow spiracular stripe and somewhat greenish tan below. Middorsal and subdorsal stripes fine, broken into small spots; subspiracular stripe weakly developed. Dorsum with minute white speckling (visible with lens). Prothoracic and anal plates light brown, both clearly delineated from rest of body. Head tan to dark red-brown, shiny, and often with large, brown patch above eyes (upper inset); triangle paler. Larva to 3.5 cm. Early and middle instars undistinguished “green fruitworms” (lower inset). ! occurrence Woodlands and forests from southern Canada to Georgia and Texas. One generation with mature caterpillars in early summer. Common. ! common foodplants Blueberry, cherry, chokeberry, dogwood, hazel, hophornbeam (Ostrya), waterleaf (Hydrophyllum) (Tim McCabe), mountain laurel, spirea, viburnum, and presumably many other hosts. ! remarks The caterpillar figured above was collected at night, feeding from the underside of a leaf near the apex of a hophornbeam shoot. It was quick to release its grasp, drop to the ground, and roll into a ball. Caterpillars are aggressive, biting at objects that touch them; their strike is rapid and accurate. The head is craned upwards and the shiny black mandibles are left flared should repeated strikes be warranted—overall, the behavior is (amusingly) snakelike.

Quakers, Woodgrains, and Kin Subfamily Noctuinae: Tribe Orthosiini 489

Fluid Arches Morrisonia latex (= Polia latex) ! recognition Variably patterned, brown caterpillar with faint (often obscured) middorsal and subdorsal pale stripes. Fine black stripe, running just above spiracles, divides darker upper portion of body from paler venter. Transverse black line runs between posterior dorsal setae (D2) on A8. Darkened chevrons that run length of abdomen frequently become more prominent rearward (these weakly expressed in our images). T2–A7 with dorsal pinacula whitened. Head usually with dark, blurry splotch over each lobe, and much additional mottling. Larva to 4 cm. Middle instars pale green to brown with characteristic darkened patch over each lobe (lower inset). Norman’s Quaker (Crocigrapha normani) is superficially similar, but its caterpillar has two pairs of black spots on the head and finishes feeding and is underground by the end of June (page 484). ! occurrence Woodlands and forests from Canada, south to Georgia (mountains) and northern Arkansas. One generation with mature caterpillars from July to September. Very common over much of East. ! common foodplants American hornbeam, basswood, beech, birch, brackenfern, cherry, elm, hophornbeam, hazel, maple, oak, poison ivy, willow, witch hazel, and many other woody plants. We have found caterpillars on hemlock that appear to be this species. ! remarks Wood and Butler (1989) provide a detailed description of the larva and list its parasites. In many Appalachian and eastern woodlands, caterpillars of M. latex can be found on almost any woody plant, and are among the most common larvae in beating samples from August and early September. While early and middle instars are green-brown and rest on foliage, the variegated brown, late instars are bark resters, as their coloration would suggest. Middle and later instars can be recognized by the large dark patch over each lobe of the head, although these may be faint in some individuals, especially those that have recently molted. We occasionally encounter late instars feeding at sugar baits. The pupa overwinters.

490 Bronzed Cutworms Subfamily Noctuinae: Tribe Tholerini

Bronzed Cutworms

Family: Noctuidae: Subfamily Noctuinae: Tribe Tholerini Only one representative of this small tribe occurs in our region. Tholerines are related to the Orthosiini but the hypopharynx lacks a transverse groove. The fall-active adults are often seen feeding at goldenrod and other flowers.

Bronzed Turfworm Nephelodes minians ! recognition Spindle shaped, dark brown to greasy black above level of spiracles, with prominent prothoracic and anal plates cut by tan to white middorsal and subdorsal lines. Tan middorsal, subdorsal, and spiracular stripes of uniform width immediately distinguish most caterpillars. Body thickest through A4–A7. Integument smooth and shiny, often with bronzy sheen. Striping obscure in melanic forms (inset). Head orange. Larva to 4.5 cm. ! occurrence Gardens, agricultural lands, fields, and meadows from Manitoba to Newfoundland, south to Georgia and Texas. One generation with mature caterpillars from April to early July. Very common. ! common foodplants Mostly grasses, including corn and other cereals, occasionally low forbs; more exceptionally buds and new leaves of woody plants. ! remarks The Bronzed Turfworm or Bronzed Cutworm is an occasional pest of turf, grains, and other grasses. Late instars live in tunnels near the soil surface. The eggs may hatch in winter with early instars feeding through the winter months, or hatch may be delayed until spring. In some years, much damage may occur under snow cover. Larvae mature and pupate by early summer, but delay their emergence until late summer and fall. Over much of the East, the adults fly from August (northward) into October (southward). A detailed treatment of its biology was authored by Crumb (1926b). Two tachinid eggs are visible between abdominal segments six and seven.

Arches and Kin Subfamily Noctuinae: Tribe Hadenini 491

Arches and Kin

Family Noctuidae: Subfamily Noctuinae: Tribe Hadenini Only about two dozen species of this largely western tribe occur in our region. The hypopharynx is divided by a transverse groove and bears minute spines over the apical lobe; the spinneret is about twice as long as broad; additional characters are given in Hacker et al. (2002) and Fibiger and

Lafontaine (2005). Diets are diverse and include a wide range of woody plants, forbs, and grasses. Among their ranks are flower feeders, fruit and seed borers, and leaf feeders. Larvae may be climbing cutworms (ascending the host to feed at night) or remain on the foodplant by day. While most are cryptically colored, a few are handsomely, and perhaps warningly, colored.

The Nutmeg Anarta trifolii (= Discestra trifolii) ! recognition Exceptionally variable: pale green to yellow-green, blackish green, brown, or gray. Broad, white to yellow lateral stripe that passes through or dodges under lower edge of spiracles, then continues beneath spiracle on A7 and A8, and drops down anal proleg. Lateral stripe frequently suffused with yellow, pink, salmon, or orange. Whole of upper portion of body densely speckled with minute cream to yellow spots, many of which are arranged in lines along trunk. Middorsal and subdorsal stripes sometimes present. Setae inconspicuous, somewhat longer than height of spiracle on A8. Head unmarked except for white bar passing through some facets of eyes. Spiracles pale with black rims. Spinneret scarcely extending beyond seta issuing from apex of labial palpus. Larva under 3.5 cm. ! occurrence Open fields, agricultural fields, and other open habitats. Transcontinental in Canada, south to Virginia and Texas. Three or more generations with caterpillars from June onward. Common southward and westward. ! common foodplants General on low-growing forbs, including crops such as alfalfa, asparagus, beets, cabbage, celery, clover, flax, legumes, lettuce, parsley, peas, spinach, tobacco, and turnip. Many weeds also eaten: e.g., lambsquarters, pigweed, and Russian thistle. Also recorded from trees such as elm and poplar. ! remarks This is a candidate for the most variable owlet caterpillar in this guide—a full page of images would not do justice to The Nutmeg’s endless varieties. Caterpillars are most common from midsummer into the fall months. In Connecticut, patches of lambsquarters and pigweed often yield the caterpillar. While widely known in the applied entomology literature as the Clover Cutworm, the name is misnomer because the larva is an aboveground feeder that remains on its foodplant by day, and clover is but one of the insect’s many hosts. Adults frequent bait. The pupa overwinters.

492 Arches and Kin Subfamily Noctuinae: Tribe Hadenini

Stormy Arches Polia nimbosa ! recognition Brown with abundant yellowish and tan flecking, and distinctive series of oblique supraspiracular lines; one sometimes present on A1, and one per segment on A2 to A8. At rest, rear of body swollen and somewhat dorsoventrally flattened. Dorsum with thin white middorsal stripe embedded in darker, diffuse middorsal stripe (both stripes obscure on T1). Dorsum of abdominal segments with indistinct diamond; anterior angle of diamond marked by black spot (poorly differentiated in figured individual), and subdorsal area flanking diamond pale, yellowish. Oblique (supraspiracular) black lines above spiracles on abdominal segments more prominent rearward. Thin, interrupted spiracular stripe dark above and pale below. Spiracles tan with black outer rim. Head bearing coronal bar over each lobe and thin black line over eyes. Larva to 5 cm. Caterpillars of Cloudy Arches (P. imbrifera) very similar but lacking dark spot at anterior reach of dorsal abdominal diamonds (Godfrey 1972) (however, this character may prove variable within species); its range largely overlaps with that of the Stormy Arches. The violet- to yellow-gray larva of the Purple Arches (P. purpurissata), with indistinct patterning and shiny prothoracic shield, is also similar (Godfrey 1972) (page 503). ! occurrence Northern hardwood and conifer forests. Transcontinental. In the East from Manitoba to Newfoundland, south to northern New Jersey, Georgia (mountains), Great Lakes states, and North Dakota. One generation with mature caterpillars from May to July (northward). Common. ! common foodplants Apparently a generalist on shrubs and small trees. Recorded hosts include alder, birch, currant, huckleberry (records probably refer to Vaccinium), maples, willow, and presumably many other woody plants. ! remarks Females raft the pale green eggs in large clusters. The figured individual is an overwintering larva. It was fed green basswood through October, and then after leaf fall was offered a salad of fallen leaves of basswood, birch, and cherry. Our larvae continued to eat senescent leaves through December when the lot was moved outdoors. With the advent of spring, European Polia move out of the litter to feed on foliage (at night) (Porter 1997). All members of the genus have a swollen, false head at the rear of the body. While the larval stage in Polia may be nine to ten months in length, we guess that most growth occurs in but one of these, when the caterpillar is feeding on new spring growth.

Arches and Kin Subfamily Noctuinae: Tribe Hadenini 493

Hitched Arches Melanchra adjuncta ! recognition Tan, olive, or lime green with darkened, distinctive backwarddirected chevrons over A1, A2, and A8; lighter chevrons over A3–A7. A8 prominently humped with sides marked by dark triangular patch that extends back to anal proleg. Thin, white middorsal stripe. Prothoracic shield with white middorsal and subdorsal lines. Spiracles white with black rim. Head pale green to light brown with coronal bars and vague mottling over lobes. Early instars bright green with subtle patterning. Larva to 3.5 cm. ! occurrence Meadows, fields, roadsides, yards, and other open habitats. Transcontinental in Canada, south in East to Georgia and Kansas; scarce on Atlantic Coastal Plain. At least two full generations in Connecticut with mature caterpillars from June onward. Adult captures suggest three generations in southern Pennsylvania. Common. ! common foodplants Many nonwoody plants including alfalfa, asparagus, aster, blackberry, brackenfern, buttercup, clover, dandelion, fringed loosestrife, horse-balm (Collinsonia), milkweed (older leaves), parsley, pea, plantain, starry campion (Silene stellata), and stinging nettle; less commonly reported from woody taxa, e.g., alder, currant, elderberry, elm, honeysuckle, salmonberry, and willow. ! remarks The Hitched Arches is exceptionally catholic in its diet, consuming many plants that are toxic to other generalists such as bracken, elderberry, milkweed, and parsley. We wonder what plants, if any, are shunned. It is among our most asked about caterpillars, especially in the fall when second generation larvae are maturing. Like many other owlets, it regurgitates when threatened. Based on our larval encounter rates, we guess that all three of our Melanchra are considerably more common than adult captures (at lights) might indicate. The pupa overwinters.

494 Arches and Kin Subfamily Noctuinae: Tribe Hadenini

Black Arches Melanchra assimilis ! recognition Green or brown with bold yellow stripes. Broad, yellow subdorsal stripe trimmed with black. Spiracular stripe edged with black and white above but only white below. Spiracles white with black outer ring. Head green or brown and mostly unmarked except for short yellow line passing through uppermost eyes; weak coronal lines sometimes present. Larva to 3.5 cm. We are unaware of readily observable characters that will distinguish its caterpillars from those of the more boreal M. pulverulenta—but see Godfrey (1972) who describes differences in the mandibles as well as other microscopic characters for both Melanchra. ! occurrence Fields, wet meadows, fens, and other open, often mesic to wet habitats. Transcontinental in Canada, south in East to Virginia and Great Lakes states. One generation with mature caterpillars in August and September. Becoming more common northward. ! common foodplants Generalist, especially on low-growing plants; recorded hosts include alder, ash, aster, birch, brackenfern, buffaloberry, cinnamon fern, gale, goldenrod, larch, mullein, purple loosestrife, raspberry, St. Johnswort, sweet fern, willow (many records), and others. See also McCabe (1990). ! remarks Covell (2005) regarded the Black Arches as an uncommon moth, yet as a caterpillar this species can be encountered in numbers. In the fall look for caterpillars in open, moist to wet Canadian zone meadows. We have taken caterpillars of the Black Arches while sweeping goldenrod flowers in fields bordering wetlands in northwestern Connecticut. The pupa overwinters.

Arches and Kin Subfamily Noctuinae: Tribe Hadenini 495

Zebra Arches Melanchra picta (= Ceramica picta) ! recognition Unmistakable black, white, and sometimes yellow-lined caterpillar. Head, legs, and prolegs reddish orange. Black ground color cut by irregular, white, vertical lines (hence its alternative name Zebra Caterpillar) between bright white or lemon-yellow subdorsal and spiracular stripes. Spiracles pale with black outer ring. Head unmarked. Larva to 3.5 cm. ! occurrence Gardens, fields, meadows, coastal communities, and other open, usually dry, habitats. Transcontinental in Canada, occurring southward in East to at least North Carolina and northern Texas. One or two generations northward; two or three broods in New Jersey and Missouri with mature caterpillars from May onward. Curiously uncommon to rare in our experience. ! common foodplants An extreme generalist on many low-growing plants including alfalfa, blueberry, cabbage, carrot, clover, corn, dandelion, hazel, knotweed, monkeyflower, peas, pigweed, timothy, and willow. Willow most commonly listed host in Godfrey (1972). Pigweed also a favorite. ! remarks It is not clear why the caterpillars are so stunningly colored. Their bold appearance and gregarious habits are strongly suggestive that the caterpillar enjoys some level of chemical protection. If true, their chemical arsenal is surely self-generated because the larvae eat a great variety of plants, most of which are nontoxic. Collection records suggest that adults are only weakly attracted to light. DLW has only seen examples of the caterpillar twice over the past 15 years in New England and both were collections made by others from coastal sites. Older literature (e.g., Forbes 1905) suggests that the insect was once a widespread animal, familiar to many farmers. We wonder if the species might be in decline. The pupa overwinters belowground.

496 Arches and Kin Subfamily Noctuinae: Tribe Hadenini

Grand Arches Lacanobia grandis ! recognition Stout, cutwormlike, pale tan or gray-brown, often with pinkish or lavender cast, lightly patterned with broadly rounded rear. Middorsal and subdorsal stripes thin, brown, and occasionally obscured. Frequently with dark middorsal spot at leading edge of each abdominal segment that increases in size to A8. Dorsum and sides densely speckled with brown spots that form irregular lines. Spiracles pale to brown with black rim. Spiracular stripe obsolescent; body mostly unpigmented below. Setae of prothorax and head relatively long. Head shiny orange-brown with conspicuous coronal bars and considerable mottling. Prolegs with a minute darkened pinaculum on outer face. Larva to 4.5 cm. ! occurrence Woodlands and forests, bogs, fens, barrens, and many other shrubby habitats. Transcontinental in Canada, south in East to southern New Jersey, North Carolina (mountains), and Great Lakes states. One generation with mature caterpillars from August to October. Common. ! common foodplants Many woody plants including alder, blueberry, cherry, dogwood, poplar, and willow. Basswood, blackberry, and oak in captivity (Crumb 1956). Less commonly recorded from nonwoody plants such as burdock. ! remarks The caterpillar of the Grand Arches is cutwormlike in its coloration and behavior, presumably spending much of its time (at least in the last instar) on or near the ground, at least during daylight hours. The caterpillar tucks itself into a ball when disturbed. Presumably the pupa overwinters.

Arches and Kin Subfamily Noctuinae: Tribe Hadenini 497

Speckled Arches Lacanobia subjuncta ! recognition Ground color usually green, tan, yellow, orange to sepia brown, with white and black speckling and subspiracular stripe. Numerous shallow incisures ring each abdominal segment. Thin middorsal stripe mostly absent, interrupted by dark intersegmental spots. Diffuse (and vague) oblique dark patch situated between dorsal setae on T2–A8, nearly engulfing (often whitened) D2 pinaculum. D1 pinaculum often encircled by small dark brown or black spot. A8 sometimes with dark transverse line that joins oblique patch to form curved band. Whitish to yellow-, peach-, orange- or red-brown lateral stripe runs along lower edge of white to tan spiracles; upper edge of stripe often edged with black. Head shiny, chestnut brown with pale snowflake spots over each lobe (inset); minute white bar arcing through upper eyes. Larva to 4.5 cm. Close to Atlantic Arches (L. atlantica), but that largely Canadian species and other western Lacanobia lack dorsal intersegmental spots. Caterpillar also similar to the Splashed Arches (Spiramater lutra), but in our few examples, the Speckled Arches tends to have weaker coronal bars, whiter spiracles, and the subspiracular stripe is not as well defined. ! occurrence Open habitats, including croplands, as well as woodlands and forests; generally scarce or absent in sandy areas. Transcontinental in Canada, south in East to North Carolina and Missouri. Two generations with mature caterpillars in June and July, then again in fall in Northeast. Common northward. ! common foodplants General feeder on many herbaceous and woody plants including alder, apple, asparagus, aspen, blueberry, cabbage, cherry, corn, dandelion, maple, meadow-rue, peach, poplar, potato, strawberry, violet, and willow. ! remarks Although host records include poplar and willow, most caterpillars are thought to mature on low, herbaceous vegetation (Forbes 1954). The record for blueberry fruits (Phipps 1930) may be based on a misidentification (Godfrey 1972). Rings (1977) regarded the Speckled Arches to be a minor pest (presumably of fruit trees, which were his specialty). Larvae of the two Lacanobia treated here are remarkably distinct in appearance and habit. If superficial larval features are considered, the Speckled Arches might be better classified with the Splashed Arches (following page); both share color polymorphism in the last instar, an arcing white line through the upper eyes, and heavy white speckling. Crumb (1956: 157) paired them in his larval key. Adults come to light and bait. Confined females readily lay eggs if held for a night or two. The pupa overwinters.

498 Arches and Kin Subfamily Noctuinae: Tribe Hadenini

Splashed Arches Spiramater lutra ! recognition Last instar bright green or more commonly yellow-green to yellow, orange, or red-brown. Dark middorsal spot towards leading edge of each abdominal segment extends onto preceding segment. Distinctively mottled with pale spots of irregular size and shape. Middorsal line hair-thin, usually broken, often more pronounced at either end of body; subdorsal line weak or absent. Most individuals also with oblique subdorsal line that may extend onto preceding segment on A1–A8; these lines lend chevroned appearance to well-marked individuals. Rump broadly rounded. Subventer speckled with creamy dots below thin, whitish to yellowish spiracular stripe that dips under spiracle on A7, and from there drops down anal proleg. Head mottled with brown over upper half, frequently with brown (coronal) bar; arcing white line through upper eyes. Prolegs often with pinkish cast. Setae fine, short, and inconspicuous. Spiracles yellowish to tan with black rim, not white as in the Speckled Arches and Atlantic Arches, Lacanobia subjuncta and L. atlantica, respectively. Larva to 5 cm. Penultimate instar greenish with more well-developed middorsal, subdorsal, and spiracular stripes and more conspicuous mottling; spots vary considerably in size and shape (inset). ! occurrence Barrens, bogs, swamps, edges of watercourses, as well as woodlands and forests from southern Canada to New Jersey, North Carolina (mountains), and Great Lakes states. One generation with mature caterpillars from August through October. Common. ! common foodplants Many woody plants, e.g., alder, blackberry, blueberry, cherry, fir, hemlock, laurel, leatherleaf, oak, spruce, tupelo, viburnum, willow, and many other shrubs and trees (see also Godfrey 1972, Rings et al. 1992, and Robinson et al. 2002). ! remarks This is a common to abundant caterpillar across much of the Northeast, especially in bogs and other acid soil habitats. We frequently run into the caterpillar when beating (especially blueberry and other heaths) in the late summer. The large black fecal pellets are highly irregular in shape. If molested, the larva readily regurgitates green vomit. Presumably the pupa hibernates, as in related moths.

Arches and Kin Subfamily Noctuinae: Tribe Hadenini 499

Striped Garden Caterpillar Trichordestra legitima ! recognition Handsome brown- and yellow-striped caterpillar. Dorsum dark brown to black, except for narrow white “V” over T1. Yellow stripes edged with white; black spiracular stripe contrasting with white spiracles. Head orange-brown, often with funnel-shaped darker brown area over each lobe that reaches eyes. Larva to 3.5 cm. Superficially similar to Black Arches (Melanchra assimilis). ! occurrence Fields and other open places, tending to do better in drier habitats. Canada south to Florida and Texas. Evidently one principal generation with mature caterpillars in late summer and fall. Common. ! common foodplants Usually low-growing plants including aster, beans, butter-and-eggs (Linaria vulgaris), dogbane, goldenrod, grasses, milkweed, lousewort (Pedicularis), tobacco, and yarrow. ! remarks Crumb (1956) regarded the insect to be principally a grass feeder—evidently he took many of his caterpillars from the seed heads of bentgrass (Agrostis) and muhly (Muhlenbergia). Most of our wild caterpillars have come from goldenrod blossoms and grasses in the fall. Although regarded as an occasional pest, population numbers rarely climb to levels where they cause economic injury. The pupa overwinters in duff or soil. See also Crumb’s (1929, 1956) accounts for this insect.

500 Arches and Kin Subfamily Noctuinae: Tribe Hadenini

Campion Coronet Hadena ectypa ! recognition Tan, yellow to orange-brown with dark chevrons and patterning above. Thin, weak, whitish middorsal stripe. Dark, oblique, subdorsal lines joining across midline of trailing segment, forming characteristic dorsal chevrons. Prothoracic plate with black patterning between subdorsal lines. Head with strong coronal bars. Mature larva under 3 cm. Middle and penultimate instars more pale green, yellow, or other pastel colors, with dorsal chevrons. ! occurrence Roadsides, sandy and gravelly pans, rocky slopes, quarries, ridgetops, and open forests and woodlands. While frequenting wooded communities southward (Charles Fenster, pers. comm.), most northern records are from open, well-drained sites. Great Lakes states to Massachusetts, south to northern Georgia and Kansas. Two generations in Connecticut, with mature caterpillars from late July at least into September. Local and scarce. ! common foodplants Starry campion (Silene stellata) as well as introduced bladder campion (S. vulgaris), and likely others. ! remarks In the Appalachians, the Campion Coronet is among the more frequent pollinators of starry campion; this pollination promotes the production of seeds, upon which its larvae feed (Kephart et al. 2006, Reynolds 2008). Females lay their eggs either at the base of the ovary or on the inside of the calyx of Silene flowers (Charles Fenster, pers. comm.). The caterpillars are important flower and seed predators on campion, sometimes consuming as much as 50% of the seeds produced by an inflorescence (Kephart et al. 2006). While still small enough to fit, early to antepenultimate instars enter the bell-shaped flowers, curl their bodies around, and consume the reproductive tissues within the female flowers. Late instars consume all reproductive tissues, including sepals, old petals, and maturing seed pods. Larval coloration is reflective of the caterpillar’s feeding and resting habits; the pale green, early instars tend to rest in and feed on flowers or young fruits, while the last two instars are darker (invariably some shade of brown), and are commonly found feeding externally on older inflorescences with dead (discolored) flowers. In contrast to the many native species threatened by invasive species (Wagner and Van Driesche 2010), the Campion Coronet appears to be expanding its range northward by exploiting an exotic campion (S. vulgaris). This exotic flower is the primary host for the recently documented colonies in Massachusetts (Nelson 2011) and Connecticut. All members of the genus overwinter in a cocoon spun belowground. Additional details on the moth’s distribution and life history appear in Nelson (2011) and Schweitzer et al. (2011).

Arches and Kin Subfamily Noctuinae: Tribe Hadenini 501

Wheat Head Armyworm Dargida diffusa (= Faronta diffusa)

! recognition Highly variable; green to pink, yellow or tan brown

with variously developed stripes. Body tapering rearward. When present, middorsal stripe frequently bound by diffuse brown spotting over A1–A8. Many forms with salmon-tan subdorsal stripe that is narrowly edged with white (salmon coloration usually more evident across thoracic segments). Brown supraspiracular stripe separated from darker brown spiracular stripe by thin white line; pale spiracles, ringed with black, situated along lower edge of spiracular stripe. Subspiracular stripe whitish tan to pink, running along lower edge of spiracles; whitish posterior portion running down outer face of A10 proleg. Head sometimes with brown bar to either side of midline and inconspicuous, thin, brown lateral line running to eyes; triangle pale, unmarked. Larva to 4 cm. Roseate or Pink Streaked Armyworm (D. rubripennis), shown on page 502, is associated with prairies, warm-season grasslands, dunes, dry to moist sandy fields, including disturbed, open sites (Metzler et al. 2005). Its larvae are a bit larger at maturity and less heavily patterned, but because Dargida larvae are perplexingly variable, positive identification should be based on reared adults (or other data). ! occurrence Sand plains, grain fields, grasslands, marshes, and other grassy habitats. Transcontinental in Canada, south in East to Georgia and Texas. Two generations with mature caterpillars from June onward in Connecticut and New Jersey (but see below); three or more broods in South. Common to locally abundant. ! common foodplants Grass seed heads; important cereal crops that may be damaged include barley, oats, rice, rye, sorghum, and wheat; Crumb (1956) notes timothy as a favorite. Additional host grasses mentioned below. ! remarks The body is drawn and somewhat flattened when the caterpillar is at rest. Larvae preferentially feed on seeds. We have found caterpillars (at night) in New Jersey eating the seed heads of switchgrass (Panicum virgatum) (along with much greater numbers of the Roseate Armyworm). Others were found feeding on pine barren reedgrass or sandreed (Calamovilfa brevipilis) and purpletop tridens (Tridens flavus). In New Hampshire we observed a female ovipositing on big bluestem (Andropogon gerardii). In the Midwest, first-generation caterpillars are occasional pests that hollow out maturing kernels of wheat. A September cohort of last instars collected in New Jersey eclosed in May to early June and in middle to late August of the following year, suggesting that the spring brood is sometimes partial. The pupa overwinters in an underground cell.

502 Arches and Kin Subfamily Noctuinae: Tribe Hadenini

Bog Yellow-wing Coranarta luteola

Circumarctic; in North America across northern Canada, south to northwestern Connecticut. One generation with mature caterpillars in early to midsummer. Local and uncommon. Cranberry.

Roseate Armyworm Dargida rubripennis

Minnesota to Massachusetts, south to Florida and Arizona although rare or absent over much of range. One generation with mature caterpillars from late summer into early fall. Generally scarce but sometimes locally abundant. Switchgrass (Panicum virgatum) in New Jersey. Green-form larvae common, as in D. diffusa (page 501).

Bertha Armyworm Mamestra configurata

Manitoba west to Pacific, south through western United States. One generation in Canada with mature caterpillars from late July through October. More common westward. General feeder on low plants. Crumb (1956) lists alfalfa, apricot, cabbage, geranium, gladiola, pea, and potato; major pest of canola in West.

Arches and Kin Subfamily Noctuinae: Tribe Hadenini 503 Cloudy Arches Polia imbrifera

Transcontinental in Canada, south to North Carolina (mountains), Kentucky, and North Dakota. One generation with mature caterpillars from April to July. Common northward and at higher elevations in Appalachians. Alder, birch, cherry, willow, and other woody plants (including fallen leaves).

Purple Arches Polia purpurissata

Transcontinental in Canada, south in East to New Jersey and Missouri. One generation with mature caterpillars from June to early August. Common northward. Alder, birch, blueberry, maple, poplar, spirea, and other woody plants (including fallen leaves).

The Rosewing Sideridis rosea

Transcontinental in Canada, south in East to West Virginia (mountains) and Great Lakes states. One generation with mature caterpillars presumably occurring mid to late summer. Local and uncommon. Captive larvae accept gooseberry, Russian olive, soapberry, and willow (Godfrey 1972).

504 Wainscots and Kin Subfamily Noctuinae: Tribe Leucaniini

Wainscots and Kin

Family Noctuidae: Subfamily Noctuinae: Tribe Leucaniini This cosmopolitan tribe is represented by two genera in North America: Mythimna and Leucania. The approximately 20 eastern species feed mostly or entirely on grasses or sedges. Many are migratory insects with core populations located in the Deep South (or the Caribbean and Central America). Others occur only in the North and in mountains. Identifications of both adults and larvae tend to be difficult, and some, especially in Florida, are best confirmed by examination of the genitalia. Godfrey (1972) provides a preliminary key and descriptions for caterpillars of 16 eastern species. As is commonly true of grass feeders, the caterpillars have massive mandibles. The smooth body is longitudinally striped in various shades of white, tan, and brown. Setae, usually borne from dark pinacula, are small and inconspicuous. The apex of the spinneret bears a cluster of long microspinules; mandibular teeth are thick and blunt. Coronal bars and dark reticulations form distinctive patterns over each lobe of the

head—the pattern is diagnostic for several eastern species. (A plate of the head capsule coloration of last instars would make a valuable contribution.) Because coloration varies within species and the taxonomy of the genus is difficult, names based on this treatment or the key in Godfrey (1972) should be used cautiously. Members of the tribe, and especially the migratory taxa, readily come to both light and bait. More often than not, confined females are reluctant to release eggs unless one adds grass stems with attached leaves. In the wild, females wedge their eggs in the sheath between leaf and stem (see inset for Reed Wainscot, L. linita, page 507). In nature, wainscots may feed primarily on one or a few species of grass, but in captivity they can be reared on a range of grasses. Godfrey (1972) reared many of his collections on orchardgrass (Dactylis glomerata). The Armyworm (Mythimna unipuncta) is an occasional, but sometimes serious, pest. Leucania is one of few owlet genera in which the diapausing stage differs among species, as noted below.

Armyworm Mythimna unipuncta (= Pseudaletia unipuncta) ! recognition Yellow to dark chocolate brown, smooth, with subdued to bold striping. Thin, white middorsal, subdorsal, supraspiracular, spiracular, and subspiracular stripes separate alternating, darker, and paler stripes. Middorsal and subdorsal stripes cut through prothoracic shield; spiracular and subspiracular stripes enclose broad, cream to peach or salmon subspiracular stripe. Thin, white subdorsal stripe often edged with blurry, elongate, black lines above, especially over abdominal spiracles. Setae short, less than a quarter length of segment that bears them. Dorsal abdominal pinacula minute, frequently darkened. Head with brown coronal bar to either side of midline that splits and continues down either side of triangle. Larva to 4 cm. Presumably the larvae of Wheat Armyworm (M. sequax) are similar in appearance. It occurs in grassy habitats over much of peninsular Florida (Terhune Dickel, pers. comm.), and as an occasional stray in Texas. ! occurrence Lawns, gardens, fields, wet meadows, and other grassy habitats. Distributed widely in North and South America as well as parts of Europe. Three generations over much of East with mature caterpillars from April onward. Present year-round in South; adults in every month in New Jersey but infrequent from mid-January until March. Northern populations reestablish from migrants each spring and summer. Very common to abundant.

Wainscots and Kin Subfamily Noctuinae: Tribe Leucaniini ! common foodplants Generalist on grasses (including grains), but

consuming many forbs and woody plants, including crop and garden plants. Common pest of barley, corn, oats, rye, and wheat. ! remarks The nocturnally active caterpillars are occasionally abundant enough to defoliate entire fields. The mass exodus made by caterpillars dispersing from denuded fields accounts for the insect’s common name. The Armyworm is among the most economically significant owlets, and may be especially harmful to grains, sod, and lawns, as well as home gardens. Rings (1977) noted that outbreaks are frequently reported from river valleys. Larvae associated with outbreak densities tend to be darker (almost black). In the spring and summer, adults move northward on the leading edge of storm fronts that push up from the Gulf Coast. In New England, drowned Mythimna adults occasionally wash up by the thousands on beaches of Long Island Sound. It is not known if there is a reverse, southward migration in the fall, but the related Asian M. segregata is known to have one. The Armyworm overwinters as a larva or pupa belowground, but probably, like the Asian species, only where the January mean temperature is above freezing. Adults visit flowers and baits.

MOTH EAR MITES Wainscots and their kin are often infested with mites. Our colleague and friend Asher Treat, made bats, Lepidoptera, owlet ears, and the mites that live within them, the focus of his life’s work; and according to him, the mites almost always infest just one ear, and use an aggregation pheromone to coordinate their activities (Treat 1957). Once the mites infest an ear, the tympanum is eaten and the mites reproduce in the cavity below. The ear’s functionality is lost at the time the tympanic membrane is compromised. The reason for the aggregative behavior is straightforward: if only a single ear is damaged, the moth (host) can still hear and fly away from echolocating bats (Roeder and Treat 1957; see box on page 476). Mites that infest both ears render their host deaf, and in so doing make both the moth and themselves highly vulnerable to predation. Darwinian natural selection acts against genes and individual mites that result in the infestation of both ears, and thus favors gregarious behaviors. Treat described more than a half-dozen ear mites, mostly in the Laelapidae, the most common of which is Dicrocheles phalaenodectes; the species epithet phalaenodectes translates to “moth biter” (Treat 1975). In the image to the right, the arrow points to a single female with her clutch of eggs (to her right) in the ear of an armyworm (Mythimna unipuncta).

505

506 Wainscots and Kin Subfamily Noctuinae: Tribe Leucaniini

Adjutant Wainscot Leucania adjuta ! recognition Beige to brown with mostly ill-defined striping and dorsal spotting; darker stripes include brown subdorsal and spiracular stripes, both edged with white and bisected by thin white stripe (visible with lens). Hair-thin, white middorsal stripe with dark flecking to either side. Supraspiracular stripe and subventer sometimes often infused with peach to salmon cast. Setal bases black. Minute spotting over dorsum of thorax arranged in longitudinal lines. No middorsal intersegmental “X”s over abdominal segments. Outer face of proleg with modest pigmentation, not obscuring SV setal insertions. Spiracle on A8 large, black, conspicuous, height two to two and a half times greater than that on A7. Head shiny, pale brown with blurred coronal bar that flares to either side of triangle; four thin, irregular lines on sides of head capsule. Larva to 3.5 cm. Early and middle instars pale tan, differing little from last instars. Contrast between adjacent stripes more subdued than on many Leucania. This is a large and difficult genus— identifications should be based on adults. ! occurrence Grasslands, marshes, waste lots, lawns, and other grassy habitats of Southeast, moving up coastal plain annually to Long Island, straying northward to southern Ontario and coastal New England. Presumably active nearly year-round in Deep South. Number of generations northward unstudied; in New Jersey, mature caterpillars probably from May onward, but mostly from September into November. Common southward. ! common foodplants Grasses; DFS has observed oviposition on switchgrass (Panicum virgatum) in New Jersey. ! remarks In New Jersey, adults are scarce from April through July, but become increasingly common through the fall. Newly eclosed (locally bred) adults occur in the fall, flying north at least to Long Island. In North Carolina, the Adjutant Wainscot occurs year-round. Our captive caterpillars shunned light and sought out hiding places by day, usually in a spot where they could fully extend the body (inset). Like other wainscots, they are lethargic and seemingly defenseless, simply curling up into a ball when alarmed, although some will regurgitate a clear green vomit if provoked. While all stages can survive mild freezes, it is not known if the Adjutant Wainscot has a diapausing stage that tolerates prolonged subfreezing temperatures (a statement that applies to all members of the genus that have their core populations in the Gulf states).

Wainscots and Kin Subfamily Noctuinae: Tribe Leucaniini 507

Reed Wainscot Leucania linita ! recognition Beige to yellow-brown, orange, and/or tan brown with numerous light brown stripes; dark brown stripes absent. Contrast between adjacent stripes less marked than that of most other wainscots; intersegmental areas with salmon tint. Narrow, pale middorsal stripe not edged with dark brown. Thin, straw-colored subdorsal stripe broken, running above yellow-brown supraspiracular stripe. Upper edge of broad, pale subspiracular stripe passing through small black spiracles. Body darkest between subdorsal and spiracular stripes, but only modestly darker than overall ground color. Setal bases small, black. Head with subdued markings, including narrow, subdued coronal stripe and reticulate brown mottling to either side of triangle; setae on head capsule borne from small dark areas (Godfrey 1972). Larva to 4 cm. ! occurrence Grasslands, fields, waste grounds, and various wetlands; in Ohio, closely associated with wetlands (Rings et al. 1992). Manitoba to Nova Scotia south through Gulf states. Three or more generations with caterpillars from June onward in New Jersey; presumably active nearly year-round in Deep South. Common, especially in the vicinity of common reed stands. ! common foodplants Grasses; our records from large grasses (more than 1 m tall), including switchgrass (Panicum virgatum), arrow bamboo (Pseudosasa japonica), big cordgrass (Spartina cynosuroides), and especially common reed (Phragmites australis). ! remarks This species is becoming more abundant in New England with the spread of common reed. Resting caterpillars perch fully extended and appear somewhat flattened at either end of body. The Reed Wainscot presumably positions itself along the stem, in dead leaves, or other concealed places by day (all Leucania caterpillars are nocturnal in habit). We have found the caterpillars to be very common to abundant in beating samples from common reed stands along the Chesapeake and Delaware Bays. The dislodged caterpillars were “clingy” and grabbed onto any surface with their prolegs, and often kept the body fully extended (only when handled did they coil up like other noctuids). On the sheet, several caterpillars were initially overlooked, dismissed as broken stem and leaf fragments. The pupal cell is fashioned belowground and sparsely lined with soil. Pupae overwinter.

508 Wainscots and Kin Subfamily Noctuinae: Tribe Leucaniini Wainscots (Noctuinae: Leucaniini) Ranges and seasonal phenologies should be used with caution due to confused taxonomy (and literature) for the group.

Trinity Wainscot

Mythimna oxygala (= Aletia oxygala) [Lesser Wainscot] Canada south at least through New Jersey and Great Lakes states. Two generations over much of East. Mature caterpillars in summer; overwintering stage unclarified but early larval record from Quebec (in Godfrey 1972) suggests that caterpillar overwinters. Common. Grasses and sedges.

Dusky Wainscot Leucania commoides

Transcontinental in Canada, south in East into Georgia (mountains) and Great Lakes states. One principal generation presumably with mature caterpillars in fall, following flight of adults. Common. Grasses and sedges.

Darted Wainscot Leucania insueta

Transcontinental in Canada, south in East to northern Pennsylvania and reportedly Washington, DC. Two principal generations in Northeast, presumably with mature caterpillars in midsummer, then again in fall. Uncommon. Grasses and sedges.

Many-lined Wainscot Leucania multilinea

Transcontinental in Canada, south in East to Georgia and Texas (this range possibly confused with that of L. lapidaria). One principal generation in Northeast, presumably with mature caterpillars following midsummer flight of adults. Locally common. Grasses and sedges. Our image, borrowed from George Godfrey, may actually be that of Leucania lapidaria.

Wainscots and Kin Subfamily Noctuinae: Tribe Leucaniini 509 Needle Wainscot

Leucania phragmitidicola [Phragmites Wainscot] Manitoba to Nova Scotia, south supposedly to Florida and Texas, but largely replaced by L. linda south of coastal Connecticut and central New Jersey, where both occur. Two principal generations over much of East, presumably with mature caterpillars in midsummer, then again in fall. Very common. Grasses and sedges.

Scirpus Wainscot Leucania scirpicola

Southern species that strays north to New Jersey in the fall (Godfrey 1972). Continuously brooded in Deep South, presumably. Common. Grasses and sedges.

White-dotted Wainscot

Leucania subpunctata (= latiuscula) Mostly tropical but resident in parts of Deep South, straying north some years (in fall) to New Jersey. Continuously brooded in southern Florida and Texas. Common. Grasses and sedges.

Ursula Wainscot Leucania ursula

Upper Great Plains, southern Ontario, and Massachusetts, south to Georgia and Texas. Three generations over much of East with mature caterpillars through growing season. Common. Grasses and sedges; DFS once observed larva eating dandelion in wild.

510 Wainscots and Kin Subfamily Noctuinae: Tribe Leucaniini

Hairy-legged Wainscot Leucania pseudargyria [False Wainscot]

! recognition Mostly beige with brown and white lateral striping. Dorsum

between subdorsal stripes rather evenly brown; often with dark middorsal “X”s between adjacent abdominal segments (visible in upper inset). Thin, white middorsal stripe best expressed over thoracic segments. White subdorsal stripe capping broad, beige to straw-colored supraspiracular stripe. Ventral setae on T3 from strongly pigmented spot. Dorsal setal bases often dark and contrasting with ground color. Prolegs with dark, shiny tan-brown plate along outer face. Head with strong coronal bars and prominent brown reticulation. Larva to 4.5 cm. ! occurrence Grasslands, fields, marshes, woodlands, and other grassy habitats from Manitoba to Nova Scotia, south to Georgia (mountains). One generation with mature caterpillars in spring. Common. ! common foodplants Grasses, including brome, corn, oats, orchard, redtop, timothy, wheat, and wild rye. ! remarks Unlike most wainscots, which overwinter as pupae, the Hairylegged Wainscot and Ursula Wainscot (L. ursula) pass the winter as nearly or fully mature caterpillars. The pupal stage last about a month. We adopt a common name suggested to us by Hugh McGuinness, which derives from the moth’s richly scaled legs (visible in image to right).

Groundcats Subfamily Noctuinae: Tribe Eriopygini 511

Groundcats

Family Noctuidae: Subfamily Noctuinae: Tribe Eriopygini More than 242 eriopygines have been recorded north of Mexico, mostly from drier areas of the American West. Less than 20 of these occur widely in the eastern United States. The

Bridled Groundcat

Lacinipolia lorea [Bridled Arches] Transcontinental in Canada, south in East to Georgia and Missouri; absent from much of Atlantic Coastal Plain. Evidently one generation with overwintering caterpillars maturing in spring. We wonder if lateseason adults reported in Rings et al. (1992) represent misidentifications. Common. Alfalfa, birch, blueberry, clover, dandelion, strawberry, and many other (mostly herbaceous) plants.

Olive Groundcat

Lacinipolia olivacea [Olive Arches] Transcontinental in Canada, south in East to northwestern Connecticut, Georgia (mountains), and Missouri. One generation with mature caterpillars in spring. Common northward and at high elevations. Mostly herbaceous plants including dandelion, phlox, and plantain.

Signate Groundcat

Tricholita signata [Signate Quaker] Manitoba to Nova Scotia, south to Florida and Arkansas. One generation with mature caterpillars in spring. Locally common. Thought to be polyphagous, especially on nonwoody, low-growing plants in spring (Bird 1925); eggs sometimes laid in goldenrod heads (Godfrey 1972).

hypopharynx is not divided by a transverse trough as in many related groups. The spinneret, dorsally grooved with a fringe of minute spinules at its apex, is two to three times longer than its diameter (Fibiger and Lafontaine 2005). Most of our eastern species feed near the soil (hence groundcats)—predictably, the caterpillars are often rendered in earth tones.

512 Groundcats Subfamily Noctuinae: Tribe Eriopygini

Explicit Groundcat Lacinipolia explicata [Explicit Arches] ! recognition Lacinipolia larvae may be recognized by their roughened integument—densely peppered with fine to coarse integumental granules—and short, spinelike, rearward-projecting D2 setae. Head retractable into thorax. Larva of Explicit Groundcat brown to gray, variegated in brown and black, with low, transverse, ridge over A8 and A9. Integument granules coarse, capped by minute spines that become longer rearward. Setae brown, thick, short, arising from blackened pimplelike bases. Dorsal, subdorsal, and lateral setae branched at their apex (visible with high magnification). Frons smooth (Crumb 1956). Thorax smoky, mostly without distinct markings; faint middorsal stripe, especially through shield. Middorsal stripe vague over abdominal segments, except between segments where it is edged with darker pigments that form faint X-shaped marks. Thin subdorsal stripe, especially evident on A1–A8, separates brown dorsum from dark supraspiracular region; thin spiracular stripe vague, running over broad, pale subspiracular area. A8 sometimes with pale patch above spiracle. Spiracles black. Larva to 3 cm. Caterpillar of Implicit Groundcat (L. implicata) [Implicit Arches] similar but dorsal setae deeply bifid at apex. Crumb (1956) and Godfrey (1972) authored keys that include common eastern species of Lacinipolia. ! occurrence Fields, waste lots, and woodlands from Missouri to New Jersey, south to Florida and Texas. Evidently two principal generations in New Jersey and North Carolina with mature caterpillars in spring, then again in midsummer. Common. ! common foodplants Ours ate various lawn forbs, lettuce, and clover. ! remarks This moth was first documented in southern New Jersey in 1998 and has been common there since; a few years later it was first recorded from the northwestern corner of New Jersey. Lacinipolia caterpillars are excruciating slow and seemingly incapable of rapid movement. All feign death when disturbed, rolling over but remaining extended, sometimes maintaining the ruse for 30 minutes or more. The ploy is so convincing that we once were ready to throw out a batch of caterpillars because we thought that all were “diseased.” Often, the anal prolegs are disengaged and the rear of the body is elevated above the substrate. The larval coloration and comportment suggest that the caterpillars spend most of their time on the ground and are nocturnal in habit. The body, and especially the setae, are often covered with a film of soil and litter. The granulose integument, short setae, and lethargic nature are shared by another group of ground feeders, the herminiines. Presumably the larva overwinters.

Groundcats Subfamily Noctuinae: Tribe Eriopygini 513

Bristly Groundcat Lacinipolia renigera

[Bristly Cutworm Moth]

! recognition Variegated brown cutworm with somewhat corrugated texture; integument densely peppered with granules that bear two to seven radial ridges (visible under high magnification). Thick dorsal setae borne from enlarged darkened warts. Thin, pale middorsal and subdorsal stripes; subdorsal stripe less developed over thorax, typically divided by broken, brown to red-brown pinstripe over A1–A8. Dorsum of abdomen with dark intersegmental spots that join to form diamond-shaped spots (Godfrey 1972, Fig. 527). Supraspiracular area below subdorsal stripe deep brown on A1–A9. Spiracular stripe embedded in broad, pale area that extends down to subventer; its width narrows over T1. Setal bases pimplelike; those above level of spiracles darkened. Setae thickened; dorsal setae on A8 and A9 longer than segments that bear them. Head slightly rugose, marked with dark reticulations and often one or two dark bars over each lobe. Larva to 3 cm. The Olive Groundcat (L. olivacea) [Olive Arches] (page 511) purportedly has thicker, peglike dorsal setae (Godfrey 1972), but this is not evident in our images. It is a denizen of northern transition and boreal forests. Positive identification of Lacinipolia caterpillars is best based on reared adults. ! occurrence Fields, meadows, and other open habitats, as well as woodlands. Transcontinental in Canada, south in East to northern Florida and Texas. Two or three generations in Northeast with mature caterpillars in May and June, then again from mid to late summer; probably more generations from New Jersey southward. Very common. ! common foodplants General feeder, typically on low-growing plants: alfalfa, clover, and other legumes; corn and other grasses; and cabbage, chicory, dandelion, dock, lettuce, plantain, tobacco, and turnip. Other reported hosts include apple, grape, and poplar. ! remarks The Bristly Cutworm gets its name from its short, stout setae. Across most of the East, it is the most abundant member of the genus Lacinipolia. It is a minor pest of crops, of little consequence in most years. It appears to be essentially polyphagous on grasses and other herbaceous plants. According to Crumb (1956) and Godfrey (1972), the diet may include dead leaves. The habit of consuming dead leaves appears to be common among owlets that overwinter as larvae. As in other Lacinipolia, when alarmed, the head can be fully withdrawn into the thorax. Third or fourth instars overwinter. A flimsy cocoon is spun in debris, rotten wood, or fashioned belowground.

514 Groundcats Subfamily Noctuinae: Tribe Eriopygini

Ruddy Groundcat Protorthodes oviduca [Ruddy Quaker] ! recognition Smooth, shiny, gray-brown cutworm with prominent black dorsal spots and undulating subdorsal stripe. Abdominal segments with large black spots, one above spiracle, and another behind. Lateral black spot on T2 and T3 enlarged. Prothoracic shield black anteriorly and laterally; caudal portion tan to orange. Anal shield pale with black patch to either side. White middorsal stripe obscure, best developed rearward on given segment. A8 with orange to tan subdorsal patch. Sides of body smoky with paler spots below. Larva to 3 cm. ! occurrence Fields, meadows, marshes, pastures, lawns, and other open habitats. Transcontinental in Canada, south to Florida and Mississippi. Two principal generations over much of East with mature caterpillars in early spring, then again in late summer (but see below). Very common. ! common foodplants Dandelion, grasses, mullein, plantain, and other low-growing plants. Probably generalized, but few records for wild larvae. ! remarks The moth is mostly double brooded in Connecticut, New Jersey, and North Carolina. While the two generations follow one another closely in Connecticut, southwards they are widely separated: flights of the first brood are centered about the middle of May and those of the second about mid-September in southern New Jersey. In North Carolina, the generations are further separated with peak abundances in April and mid-October. A similar pattern occurs in some double-brooded skippers, whereby the separation of the two broods increases as one moves southward. Adults come to bait, especially second-brood individuals in the late summer. Crumb (1956) made note of a sizeable larval collection made by turning withered mullein leaves in January (in Tennessee). The Ruddy Groundcat overwinters as a nearly mature caterpillar.

Groundcats Subfamily Noctuinae: Tribe Eriopygini 515

Sheathed Groundcat Ulolonche culea [Sheathed Quaker] ! recognition Shiny brown with black, elliptical, subdorsal spots over A7 and A8. Those over A8 nearly meeting over dorsum. Dorsal setae of abdomen borne from minute blackened pinacula. Thin, pale, broken middorsal stripe. Prothoracic shield distinctly lightened along its anterior margin. Tan to pinkish-brown ground color densely peppered with minute gray, brown, or black spots. Vague spiracular stripe. Setae somewhat thickened, about third of segment length. Upper half of head black and shiny; lower half chestnut brown. Larva under 4 cm. The Galena Groundcat (U. modesta) co-occurs with the Sheathed Groundcat over much of the East (inset). It can be abundant in sand plains, barrens, grassy heathlands, low shrublands, dry oak woodlands, and coastal strand woodlands of the Great Lakes states, Northeast, and Atlantic Coastal Plain. We reared a cohort of Galena Groundcat caterpillars on an initial diet of new but fully wilted and blackened oak leaves, and a mixture of downed catkins of various oaks and hickories; middle and late instars were reared on moist, partially decayed oak litter. ! occurrence Barrens and woodlands from Illinois to Quebec and Maine, south to Florida and Texas. One generation with mature caterpillars by midsummer. Locally common. ! common foodplants Captive larvae have been reared on oak. ! remarks The coloration suggests the caterpillar is a ground feeder, but Franclemont (in Godfrey 1972) reared the Sheathed Groundcat on black oak (Quercus velutina) and red oak (Q. rubra). DLW also reared ex ova larvae to maturity on oak. We suspect that the caterpillar feeds principally on downed plant debris or is a climbing cutworm that ascends to foliage at night and then hides in leaf litter by day. Since most ground-dwelling cutworms are polyphagous, it would be premature to assume the Sheathed Groundcat is an oak specialist. Clearly, much remains to be learned about the early stages of the genus. While described by Forbes (1954) as rare, the Sheathed and Galena Groundcat (U. modesta) are common in barrens and dry oak forests and woodlands of the Northeast. Early spring emergence dates are suggestive that last instars or pupae overwinter.

516 Groundcats Subfamily Noctuinae: Tribe Eriopygini

Disparaged Groundcat “Orthodes” detracta (= Polia detracta) [Disparaged Arches]

! recognition Reddish brown, flecked with tan spots; A7 and A8 swollen

(at rest). Prothoracic shield shiny and well defined; tan to whitish middorsal stripe flanked by large dark patches that extend down to subdorsal stripe. Similarly, anal plate shiny with tan to whitish middorsal line sandwiched between two dark patches. Conversely, subdorsal stripe often more developed rearward, but essentially absent on A8. Lateral setae from dark brown shiny pinacula. Setae pale, somewhat thickened and bristlelike. Head with coronal bars. Larva to 3.5 cm. ! occurrence Woodlands and forests. Transcontinental in Canada, south in East to Georgia (mountains) and Louisiana. One generation with mature caterpillars from mid-April to June. Abundant. ! common foodplants Buds of hickory and oak; birch, blueberry, dandelion (Godfrey 1972), serviceberry; also acorns (J. R. Galford in Rings et al. 1992). (See also below.) ! remarks This is a common forest moth throughout much of the Northeast and mid-Atlantic regions. Its larval habits are poorly known. At rest, the rear of the caterpillar’s body is rather headlike, drawing the eye. According to Forbes (1954), Rockburne and Lafontaine (1976), and others the larva feeds on buds of various shrubs and trees. The disparity between the abundance of adults at light versus the number of records for larvae, suggests that the caterpillar is a nocturnal creature that leaves foliage by day to secrete itself away. The coloration of the larva, its lethargic nature, and false head are indicative that it is a ground dweller. We have had captive caterpillars in the fall and winter eat and subsist on dead, browning leaves for months—the growth of these individuals was modest. With warming spring temperatures, presumably caterpillars switch over to green tissues and feed on buds and new growth, as do a number of other univoltine hadenine and noctuine cutworms. The image above is of an overwintering middle to late instar taken in October; while not fully mature, it agrees with the description provided by Godfrey (1972). Lafontaine and Schmidt (2010) used shutter quotes around the generic name because the moth is not a true Orthodes, and almost certainly will be assigned to another genus.

Groundcats Subfamily Noctuinae: Tribe Eriopygini 517

Rustic Groundcat Orthodes majuscula (= O. crenulata) [Rustic Quaker]

! recognition Brown with vague patterning and black middorsal spot at

leading edge of abdominal segments; these conspicuous on A1 and A2. Head small, often withdrawn into prothorax, held down under body and pressed against substrate. Side of head with shiny, broad, black bar and X-shaped mark centered over triangle. A8 humped; angled steeply downward, often blackened on posterior face. In some postures, rear segments pulled into A8 (inset). Broken, white middorsal and subdorsal stripes. Spiracles orange-tan or black. Larva to 4 cm. Middle and penultimate instars rather different in aspect with few of the diagnostic features noted above. A1 appears swollen and bears subdorsal white spots (below right). White subdorsal stripe usually continuous from A1 to A8. A8 frequently with vague, pale subdorsal patch. First abdominal segment is grossly swollen in first and second instars. ! occurrence Fields, woodlands, and other mostly open habitats across Canada, south to Florida and Texas. Two or more generations over much of East with mature caterpillars nearly year-round. Common. ! common foodplants General on forbs and evidently grasses as well. Our wild larvae from aster, bee balm, blackberry, dandelion, dock, goldenrod, hazel, jewelweed, thoroughwort (Eupatorium), and sensitive fern. ! remarks This is a common caterpillar in fall collections from low vegetation. Our fall larval collections fed for weeks but never matured, suggesting that the Rustic Groundcat overwinters as a larva. Dates for mature larvae in Crumb (1956) also indicate that at least some larvae overwinter. However, a penultimate instar collected by DFS pupated in November and emerged the following May. Populations of the Rustic Groundcat in the North are thought to be reinforced by migrants from the South, which would explain late fall adults and conflicting observations about how the species overwinters.

518 Groundcats Subfamily Noctuinae: Tribe Eriopygini

Cynical Groundcat Orthodes cynica [Cynical Quaker] ! recognition Stocky, glossy, brown to red-brown, sometimes with pinkish to purplish flush. A8 swollen, somewhat humped, tan posterior margin overextending A9; A9 and A10 reduced, partially withdrawn into A8. Tan middorsal stripe from posterior half of T1 to A8, often broken anteriorly or in midsegment. A8 commonly with small, black triangular spot along lower side of subdorsal stripe. Prothoracic plate edged with tan along its leading edge. Vague blackish chevrons or diamonds frequently present over dorsum (see image in Godfrey 1972). Subdorsal stripe often more developed than in our figured individual. Head small, shiny brown with broad, black band extending across lobe to base of mandible; head also with peppering of small dark spots (inset). Spiracles solid black. Larva to 4 cm. ! occurrence Fields, woodlands, forests, and variety of other habitats from Saskatchewan to Nova Scotia, south to Florida, Mississippi, and Arkansas. One generation with mature caterpillars in late spring. Very common to abundant, although local on coastal plain. ! common foodplants General feeder on forbs, e.g., dandelion and plantain. Figured individual reared on clover. McCabe (1990) collected a wild larva on developing goldenrod blossom. ! remarks Godfrey (1972) provides a detailed description and figures of the caterpillar. The rear of the body is thickened and more headlike than the anterior end, but the shape and posture are not especially convincing relative to the other ground-dwelling noctuines with false heads. Although the Cynical Groundcat is one of the most common moths seen at light in eastern woodlands during June, especially where there is a rich understory of forbs, we have never found its caterpillar in the wild. According to Forbes (1954) the pupa overwinters.

Small Brown Groundcat

Pseudorthodes vecors [Small Brown Quaker] Manitoba to Newfoundland, south to Georgia and Texas. Two generations with mature caterpillars from June onward. Very common. Aster, dandelion, jewelweed, plantain, yarrow, and other forbs; also grasses. According to Forbes (1954), the doublebrooded populations represent an unrecognized sibling species, with true P. vecors being larger, more boreal, and univoltine.

Spanish Moths Subfamily Noctuinae: Tribe Glottulini 519

Spanish Moths

Family Noctuidae: Subfamily Noctuinae: Tribe Glottulini A small tropical group represented by one species in the Southeast. The trunk is encircled with white (or cream) and black rings. The SV group is bisetose on T2 and T3; SD1 on A9 is hairlike. Our species feeds principally on Amaryllidaceae.

Convict Caterpillar Xanthopastis regnatrix [Spanish Moth] ! recognition Bold black- and white-ringed caterpillar with red-orange head, legs, and rump. White ringing posterior portion of thoracic and first eight abdominal segments. White middorsal spots on T3 and (A3) A4–A9. Orange head with black spot on either side of triangle that includes eyes. Larva to 4 cm. ! occurrence Edges of watercourses, roadside ditches, marshes, and other wetlands from southern Indiana to Long Island, south to Florida and Texas, but only as rare stray north of Carolinas along East Coast. One generation northward with mature caterpillars in August and September; year-round in southern Florida. Local and uncommon. ! common foodplants Amaryllis, narcissus, spider lily, and other plants in amaryllis, iris, and lily families; many other plants less commonly reported. Caterpillars may be reared on lettuce. ! remarks Females lay clusters containing hundreds of eggs. Early instars are gregarious. Both aboveground (leaves) and belowground (bulbs and rhizomes) tissues are eaten. The Convict Caterpillar breeds as far north as southern Indiana during summer months. Its winter range is probably limited to subtropical Florida and Texas. The caterpillar is almost certainly toxic, although we do not believe its chemical ecology has been studied. The Convict Caterpillar appears to have two heads with the one at the rear realistic enough to draw our initial attention. Xanthopastis timais, the name usually applied to North American populations of the moth, correctly refers to a South American species distinct from our North American populations (Tim McCabe, pers. comm.).

520 Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini

Cutworms and Darts

Family Noctuidae: Subfamily Noctuinae: Tribe Noctuini This is a large and taxonomically difficult group of owlets. There are about 115 species that occur east of the hundredth meridian. Contrary to the general trend of increasing species diversity as one approaches the tropics, noctuine richness peaks in temperate areas. We illustrate 51 species, a rather low proportion of the eastern fauna, because few noctuine caterpillars, with the exception of the pest species, are encountered with any regularity—they are a secretive, ground-dwelling lot that feed mostly after nightfall. Crumb (1956) treats 69 North American noctuines, including all eastern species of economic importance. Many are illustrated and/or described in Lafontaine (1998, 2004). Several European works, e.g., Porter (1997), Beck (1999, 2000), and Ahola and Silvonen (2005), are worth consulting because they illustrate species closely related to those in our fauna. Noctuine caterpillars are the classic cutworms familiar to farmer and gardener: stout, smooth, often somewhat soft and fleshy, clammy to the touch. The setae are short and thin. At rest, the caterpillars have a sizeable rump—A8 is often swollen and/or A9 and A10 are small and frequently directed downward. Most are rendered in browns, grays, and other earth tones; those that tunnel belowground by day tend to be weakly pigmented. Proportionately, the prolegs tend to be small (presumably because so many burrow and few climb). SD1 on A9 is hairlike. The apex of the short spinneret is fringed or lacerated on the upper surface (Crumb 1956, Fibiger and Lafontaine 2005). Coloration tends to be highly variable within a species, making identification difficult. Definitive larval determinations may require examination of the mouthparts under high magnification. Most noctuines feed on grasses, forbs, and low-growing plants. Northward, the proportion that includes woody plants in their diet increases—heaths, especially blueberries, are eaten by many. Some Abagrotis and Xestia feed on large trees; late instars may climb tens of meters each evening to feed. Several are serious pests of cereal and other crops. The biologies of the more common eastern species that are of economic importance were treated in detail by Crumb (1929) and Walken (1950). Damage to seedlings, buds, and spring foliage can be severe—in large measure because larvae overwinter and are sizeable at the time new growth is burgeoning in the spring. The caterpillars rest on or near the ground, often beneath low vegetation, fallen leaves, or other surface litter. Many tunnel underground by day, then emerge from the soil after sunset to feed—predictably, these tend to be more abundant in sandier soils. Some sever vegetation near ground level and then pull the

excised tissue into a vestibule or tunnel where they consume it out of harm’s way—hence the origin of their common name. When disturbed, the caterpillars drop from their host, curl into a C-shape and feign death; because most are polyphagous such a strategy is seldom a dietary risk. A large fraction have preferred soil types: e.g., Agrotis and Euxoa are more numerous in sandy habitats, whereas others are associated with acid soils (or the heaths that grow on them). Few occur in wetlands, although some specialists do so. One way to survey for cutworms is to place out bundles of cut dock, chickweed, or other forbs, and return two to three days later to look for larvae secreted beneath. Where the soil is loose and uncompacted, use a hand rake to expose buried caterpillars (Crumb 1926a). Many Noctuines are univoltine with eggs laid in late summer or early fall and hatch taking place within a few weeks; in some Euxoa eclosion may be delayed until spring. The fallhatch larvae feed as conditions permit through the winter. We sometimes encounter caterpillars basking on warm winter and spring days (see account for the Large Yellow-winged Dart, Noctua pronuba). A number of noctuines reared by DFS exhibited a marked seasonal change in diet: overwintering larvae consumed grasses and forbs, but in the spring the caterpillars preferred the foliage of woody plants. Some Xestia and Abagrotis feed on dead leaves over the winter. The majority of our noctuines mature in spring, with development timed to the availability of new growth. Females of most genera lay readily in captivity, either releasing eggs on the floor of their container (Abagrotis), or attaching the eggs singly (Agnorisma, most Xestia) or in masses (Noctua and Peridroma) to living or dead leaves and other objects. Nearly all hatchlings will do well if provided a salad of grass and forbs. Artificial diets, for example those based on wheat germ or beans, are also accepted. Single-brooded species tend to have late-summer or fall flights; this is particularly true of those associated with grasslands, barrens, and other habitats under water stress during the summer. A number emerge in early summer, then aestivate as adults until autumn. As noted above, most overwinter as larvae, but some Euxoa may do so as eggs. Dichagyris overwinter as prepupae and Cerastis, Ochropleura, and Peridroma as pupae. Pupation occurs in a cell underground, sometimes after a three- to fourmonth summer aestivation period by the prepupal larva. Both sexes come to light and some species also to bait; their ranks include many avid flower visitors. The Variegated Cutworm (Peridroma saucia) and Black Cutworm (Agrotis ipsilon) are migratory pests that sometimes arrive in destructive numbers. Detailed techniques for rearing larvae are given in Crumb (1929). Most, but not Abagrotis, are sensitive to desiccation and should not be sleeved.

Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini 521 Black Army Cutworm Actebia fennica [Finland Dart]

Transcontinental in Canada, south into northern states. One generation with mature caterpillars presumably in spring and summer. Locally common northward. General feeder on forbs, shrubs, and small trees; occasional pest of blueberry and conifer seedlings.

Early Conifer Dart

Xestia praevia [Praevia Dart] Transcontinental in Canada, south in East to Massachusetts and high elevations of Appalachians. One generation with a few caterpillars maturing in fall; most full grown in May and June. Common northward. Fir, hemlock, pine, spruce, and occasionally larch. Similar to White Pine Dart (X. badicollis) (page 542) and others. Illustrated by Duncan (2006).

Relict Prairie Dart Dichagyris reliqua

Eastern range southern Manitoba and Great Lakes states, south to Missouri with disjunct populations in New Jersey; single adult from Florida panhandle. One generation with mature caterpillars in fall with prepupae overwintering sometimes two or more times. Local and generally scarce as adult. Developing seeds of prairie dropseed (Sporobolus heterolepis), pine barren reedgrass or sandreed (Calamovilfa brevipilis), and possibly other prairie grasses. See account in Schweitzer et al. (2011).

Catocaline Dart Cryptocala acadiensis

Transcontinental in Canada, south in East to Connecticut and Great Lakes states. One generation with mature caterpillars in fall. General feeder. Locally common in wetlands, and becoming more so northward. Ours reared on lambsquarters; McCabe (1979, 1990) observed oviposition on St. Johnswort, and reared larvae on dogbane. Black pinacula wider than height of spiracle.

522 Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini

Snowy Dart Anicla illapsa (= Euagrotis illapsa) ! recognition Highly variable cutworm, ranging from green to straw colored, tan, or red-brown with prominent longitudinal striping. Many forms with black line or spot enclosing spiracles on T1 and A1. Broad, white to tan lateral stripe running beneath spiracles, commonly infused with beige, yellow, orange, salmon, or red, edged above and below with white or tan. Subdorsal stripe sometimes trimmed above with diffuse, elongate black bars on A1–A8 that may join rearward. Brown prothoracic and anal shields cut by pale middorsal and subdorsal stripes. Black lines on head extending forward to base of antennae (inset); subdorsal and supraspiracular stripes sometimes continued along cheek as diffuse lines that reach eyes. Spiracles porcelain white. Mandible with five cutting teeth (Lafontaine 2004). Larva to 3.5 cm. Early instars green with vague dorsal striping; many forms with bold subspiracular whitish stripe, red cheek line, and/or red antenna (lower right). Middle instars either green or red-brown. Forbes’s Dart (A. forbesi) and two other Anicla co-occur in the East. Identifications should be based on adults from the Great Lakes states and central New England southward. ! occurrence Fields, meadows, yards, and other open, grassy habitats from Wisconsin, southern Ontario and Quebec, and Maine, south to Florida and Texas. Two or three broods in Connecticut; flies year-round in Florida. Mature caterpillars throughout growing season. Common. ! common foodplants Grasses and sedges. ! remarks It is puzzling that the early stages and natural history of a “backyard” moth as common as the Snowy Dart are unstudied—the larva and its biology were unknown to Lafontaine when he published his noctuine revision in 2004. Ex ova rearings, with associated notes and preserved larval vouchers, are needed for nearly all North American Anicla. Lafontaine’s (2004) preliminary larval studies indicate that dentition of the larval mandible will distinguish many of our Anicla species. Larvae that we brought indoors for rearing in October and November stopped feeding as middle instars.

Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini 523

Green Cutworm Anicla infecta ! recognition As in Snowy Dart (A. illapsa) illustrated on previous page, coloration highly variable: green, yellow-brown, gray, tan, or brown, often with peppering of black flecks. Prominent pale orange, pink, or salmon subspiracular stripe touches along lower edge of white to tan, black-ringed spiracles, edged with thin white stripes above and below, continuing along side of head to antenna; upper edge sometimes margined with black, especially across thoracic segments. Spiracles often embedded in minute, elongate black spot on A1 and A2 as in Snowy Dart, but black reduced in extent. Middorsal and subdorsal stripes, when present, thin, white, and broken. Dorsum commonly with black flecking and spots; sometimes with dark subdorsal spot above each spiracle (as on A6 and A7 in main image above). Head usually with coronal bars continuing to lower corners of triangle; lateral dark lines extend to antenna. Mandibles lacking multiple distal teeth. Larva to 3.5 cm. Ground color of first three instars green. Given the variability in coloration of Anicla caterpillars, we are uncertain if our diagnoses will reliably distinguish larvae of the Green Cutworm from those of the Snowy Dart. Some forms of the Nutmeg (Anarta trifolii) also approach appearance of Anicla caterpillars. ! occurrence Fields, waste lots, croplands, marshes, wet meadows, and other open habitats from Manitoba to Maine, south to Florida and Texas (and South America). Occurs as a migrant over northern portion of range. Number of generations depends on when first immigrants arrive, apparently varying from zero to three or four in New Jersey. Mature caterpillars mostly in late summer and fall northward and throughout growing season southward. Very common, especially southward. ! common foodplants Grasses commonly, but readily consuming range of forbs and crops, including beets, clover, false mallow (Malvastrum), plantain, purslane, and tobacco (mostly from Crumb 1956). ! remarks At rest, the caterpillar is somewhat flattened with supraspiracular, longitudinal furrow tapering rearward. Our winter-reared caterpillars consumed both living and dead (brown) grass blades. We do not know which stage typically overwinters or how far north the species is able to do so. Crumb (1956) records pupae overwintering in Tennessee, but apparently the species is unable to survive year-round much farther north. Adults can be abundant at bait in autumn—in 2005, Green Cutworm adults were third only to Bicolored Sallow (Sunira bicolorago) and the Armyworm (Mythimna unipuncta) along bait trails in southern New Jersey.

524 Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini

Variegated Cutworm Peridroma saucia ! recognition Stout, smooth cutworm, ranging from gray to brown; frequently with subtle pink to orange markings. Expect forms both more and less patterned than individuals figured here. Middorsal stripe marked by small white to orange spot toward rear of abdominal segments A1–A4 (and sometimes A5–A8). Poorly differentiated subspiracular stripe, often with shades of orange along upper edge, runs just beneath black spiracles (below right). A8 broadly rounded; often with obscure yellow to orange rump patch. Vertex marked with black “M”; triangle pale tan. Larva to 4.5 cm. Larva of some Protolampra similarly patterned. Common to abundant. ! occurrence Agricultural lands, fields, and other open habitats. Worldwide migrant. Northern Canada, south into South America. Multiple generations with mature caterpillars from April onward. Crumb (1926a) records four annual broods in Tennessee. Common. ! common foodplants General feeder. Economically important plants listed by Crumb (1929) include alfalfa, apple, cabbage, carrot, clover, corn, grape, plum, potato, raspberry, rose, strawberry, tobacco, tomato, and wheat. ! remarks Crumb’s (1929) biological notes for this species are extensive. Historically, this was often the most economically important cutworm over the northern half of our region, damaging both field and orchards crops, and on occasion infesting greenhouses and cold frames. An outbreak at the turn of the last century caused more than $2,500,000 in losses (Chittenden 1902). The insect is particularly destructive in the West, Great Plains states, and across agricultural regions of Canada. Historical accounts make mention of entire fields being defoliated of essentially all green matter, be it grass, herb, shrub, or tree. Even belowground tissues such as potatoes are eaten. Females raft eggs on the underside of leaves. During the warmer summer months, larvae can pass through their six instars in 21 to 29 days (Crumb 1929). Like other noctuine pests, infestations follow the arrival of migrants from the Gulf states. In Europe, there is indication of a fall southward migration (McNeil et al. 1995). The pupa overwinters or on occasion the larva overwinters, at least southward. How far northward populations of this migrant may survive winters is not known. Adults come to light and bait.

Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini 525

Western Bean Cutworm Striacosta albicosta

(= Loxagrotis albicosta)

! recognition Variable tan, pale brown, gray-brown, to pinkish brown, with thin, whitish middorsal, subdorsal, and supraspiracular stripes. Dorsum of abdominal segments frequently with ill-defined diamond markings, with one per segment; apex of each diamond touching trailing edge of previous segment. Middorsal stripe interrupted between segments by dark spot on A1–A8. D1 pinacula sometimes embedded in small black spot on anterior abdominal segments. Shiny brown-black prothoracic shield divided by pale middorsal and subdorsal lines. Head reddish brown, shiny, varying from unpatterned to heavily patterned with black (Crumb 1956). Integument finely roughened but not granulated (under magnification); spiracles brown with black rims (Crumb 1956). Larva to 4 cm. ! occurrence Principally agricultural fields in our area, especially those with sandy soils, throughout the Great Plains from Minnesota southward into Central America; eastward in 2010 to Quebec, Pennsylvania, Long Island. One brood with mature caterpillars mostly in August and September. Locally abundant crop pest. ! common foodplants Major pest of corn but also reported from other grasses and forbs; occasional pest of lima, green, and pinto beans (Godfrey 1987, Lafontaine 2004). ! remarks Much of our information for the Western Bean Cutworm derives from Hoerner (1948). Adults are on the wing mostly from late July into August, and occasionally into September. Eggs are laid out in rafts, with clusters of up to about 200. On corn, early instars actively tunnel in tassels, the flag leaf, and other new tissues. Young caterpillars may wander as much as 3–4 m from their natal site. Middle and late instars preferentially consume reproductive tissues (e.g., corn kernels and bean fruits). Like other cutworms, the caterpillars feed mostly at night and then hide by day under litter or, more commonly, tunnel under soil. On corn, larvae may remain hidden in the husks by day. Larvae pass through five larval instars over a four- to five-week period, before tunneling deep into the ground to form a cell in which to pass the winter. Pupation occurs as early as May of the following year without additional feeding. The Western Bean Cutworm has only recently spread into our area from the West, but is becoming an increasingly serious threat to corn in the Midwest, inflicting damage mostly in late August and September. Kernels are either damaged outright or the ear is made susceptible to secondary fungal infections. Highest infestations are found in croplands with sandy soils that are conducive to the larva’s tunneling tendencies.

526 Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini

Sandhill Cutworm Euxoa detersa [Rubbed Dart] ! recognition Very pale, lacking appreciable pigment to pale brown, with faint white striping. Usually with rudimentary, thin, whitish middorsal and subdorsal stripes. Thickened supraspiracular stripe, sometimes broken, running above black spiracles. (All striping much reduced in our example.) Prothoracic shield pale brown. Head capsule without patterning or with somewhat darkened haze or freckles over each lobe. Anal proleg with 10 to 16 crochets. Larva under 3.5 cm. ! occurrence Dunes, sand plains, well-drained river shorelines and floodplains, pine barrens, and other open sandy environs. Transcontinental in Canada, south in East to Carolinas and Texas. One generation with mature caterpillars from late spring into summer. Abundant along coast; locally common inland. ! common foodplants Generalized feeder on low-growing plants; occasional pest of corn, cranberry, oats, potato, rye, strawberry, tobacco, and wheat. Probably mostly grasses. ! remarks Euxoa is the largest genus of macrolepidopterans in North America, with some 187 recognized species (Lafontaine 1987, Lafontaine and Schmidt 2010). Most are western: only about a dozen species occur in the East and many of these are northern. Eastern species are best represented in dry woodlands, barrens, grasslands, sand plains, and other open, dry, and especially sandy, habitats where their larvae can readily tunnel belowground. Most of our species fly in late summer. Eggs or young larvae overwinter. Euxoa are generalized feeders that consume almost any green plant tissue. Larvae shun light and tunnel underground if exposed; they then emerge by night to feed. The biology of three dozen Euxoa was reviewed by Hinks and Byers (1976). The Sandhill or Sand Cutworm is a largely subterranean caterpillar of sandy soils that feeds at ground level or, more commonly, belowground on roots and adjacent stem tissues. It can be especially destructive to seedlings of corn and other cereals. In coastal dunes from Massachusetts to Cape May (New Jersey), adults frequently outnumber all other macrolepidopterans at lights and on goldenrods during September.

Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini 527

Darksided Cutworm Euxoa messoria [Reaper Dart] ! recognition Dingy gray to brown, usually with side of body above level of spiracles more heavily pigmented than dorsum. Broad, white supraspiracular and subspiracular stripes separated by grayish to whitish spiracular stripe. Thin, white middorsal line embedded in narrow, diffuse field of minute dark spots. White subspiracular stripe gives way to largely unpigmented venter below level of SV setae. Prothoracic shield more heavily pigmented than adjacent cuticle. Head patterning consists of dark flecks (or freckles) (Crumb 1929, Whelan 1935). (Many other cutworms have distinct coronal bars and other stripes.) Anal proleg with 20 to 24 crochets. Larva to 3.5 cm. Until species-level characters are better known, determinations are best based on adults. Close to Striped Cutworm (E. tessellata), which has pigment flecks as well as stripelike markings extending back from eyes along “cheek.” ! occurrence Agricultural lands, waste places, lawns, fields, and grasslands, especially with sandy soils. Transcontinental in Canada, south in East to North Carolina and Missouri. One generation with mature caterpillar in late spring and early summer. Common westward and northward. ! common foodplants General feeder on grasses and low-growing plants, including many field crops: e.g., beans, cabbage, clover, corn, lettuce, onion, pea, potato, radish, strawberry, sugarbeet, sweet potato, tobacco, tomato, and turnip. Also climbs to feed on apple, currant, grape, peach, and various other woody plants. ! remarks This widespread, abundant, and destructive insect attacks a broad spectrum of field and orchard crops. It is among the more arboreal Euxoa, and will ascend shrubs and trees to seek out new foliage. Such larvae can cause significant damage to fruit trees in the spring. As with other members of the genus, most reports of economic injury are from areas with sandy soils. The eggs are laid directly into the soil, especially in areas that have been disturbed (e.g., plowed). The winter is passed as an egg or possibly young larva. We have few images of the species to study, but in some of these the dorsum is more darkly pigmented than the sides (contrary to the insect’s accepted common name).

528 Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini

Granulate Cutworm Feltia subterranea [Subterranean Dart] ! recognition Dark, mottled, appearing somewhat greasy, with short prolegs. Ground color dirty gray-brown, diffusely mottled with tan and smoky black; dorsum gray-brown with vague, smoky middorsal stripe; subdorsal area darkened, especially immediately below imaginary line connecting D2 setae. Integument peppered with numerous minute granules; those above spiracles toothlike and mostly dark, while those below spiracles rounded and pale. Prothoracic shield with pale addorsal and subdorsal areas. Venter dark. Spiracles black. Head small, only a third width of T2, partially withdrawn into thorax; dark coronal lines join over triangle and then separate, dropping down sides of largely unpigmented triangle. Dark anal shield with pale middorsal line. Larva to 4 cm. Several other members of the genus occur commonly in the East—identifications are best based on reared adults. ! occurrence Agricultural lands, waste places, lawns, fields, and grasslands from South Dakota, Michigan, and Nova Scotia (as summer and fall migrant northward), to Florida and Texas (and South America); common southward. Multiple generations with mature caterpillars from May onward in South. Locally common. ! common foodplants General feeder on grasses and low-growing forbs, including both fresh and wilted foliage. Important field and garden crops include beans, clover, corn, lettuce, peas, potato, tobacco, and wheat. ! remarks As indicated by its names, the caterpillar lives mostly underground, emerging after nightfall to feed. When handled the larva becomes turgid, cocks its body to one side, and feigns death. As might be guessed from an examination of its extremely short prolegs, the caterpillar shows little propensity to climb. Perhaps descendants of the Granulate Cutworm will one day lose the legs entirely, as has happened in numerous other subterranean animals (i.e., lizards, salamanders, and many beetle and fly larvae). Our captive caterpillars, reared on a salad of forbs, showed a penchant for clover flowers. Survivorship of pupae over winter is low in northern Tennessee (Crumb 1929), and it is unlikely any stage normally overwinters much farther north.

Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini 529

Dingy Cutworm Feltia jaculifera complex [Bent-lined Dart] ! recognition Smooth, dirty brown with weak pattern; dorsum somewhat paler between broken, chocolate-brown, subdorsal stripes. Dorsum sometimes with dull, smoky, brick-red elements; abdominal segments marked with vague, dark wedges or diamonds outwardly bounded with pale patches that run forward from D2 seta. Subdorsal stripe sometimes pale along its inner edge. Prothoracic shield brown, shiny, strongly pigmented anteriorly, cut by sharp middorsal line and broad, ill-defined subdorsal line. Small, pimplelike pinacula shiny brown. Larva to 4 cm. The Dingy Cutworm is believed to be a complex of six or more closely similar species whose members are only reliably separated by the sex pheromone blends to which they respond (Forbes 1954, Lafontaine 2004). The complex is in much need of taxonomic study—both molecular and behavioral data may be needed to tease apart this evolutionary conundrum. Further, we are unaware of larval characters that distinguish members of the jaculifera complex from other common Feltia species that co-occur over much of the East: the Master’s Dart (F. herilis), Subgothic Dart (F. subgothica), and Tricose Dart (F. tricosa) (page 533). ! occurrence Old fields, agricultural lands, orchards, gardens, grasslands, coastal communities, waste sites, and other open habitats. Transcontinental in Canada, south in East to northern Florida and Texas, becoming appreciably less common southward. One generation with mature caterpillars in early summer. Very common to abundant northward. ! common foodplants General feeder on grasses, forbs, shrubs, and low trees; crop hosts include alfalfa, apple, beans, clover, corn, dock, flax, lettuce, oats, pea, potato, raspberry, rye, strawberry, tobacco, tomato, wheat, and others. See also Remarks. ! remarks See Lafontaine (2004: 212–214) for a current summary of the (numbing) taxonomic difficulties offered by the Dingy Cutworm complex. F. jaculifera seems to be a moth in the throes of speciation with the boundaries among the component species blurred by hybridization and introgression. Females of the Dingy Cutworm, Tricose Dart (F. tricosa), and Subgothic Dart (F. subgothica) have been observed by DFS ovipositing in the flower heads of composites, especially thoroughwort (Eupatorium), ragweed (Ambrosia artemisiifolia), and camphorweed (Heterotheca subaxillaris)—a curious (and interesting) habit that presumably enhances the survivorship of the eggs relative to other darts most of which lay their eggs in soil. Gravid Feltia females confined with inflorescences of goldenrod will insert their eggs deep into the flowers with their bladelike ovipositor (John Franclemont, pers. comm. to Don Lafontaine). The biology of the early instars remains unstudied. As in most darts, winters are passed as larvae (Crumb 1929). With the arrival of spring, the polyphagous caterpillars feed up to maturity and then tunnel belowground to fashion a “plastered cell” where they aestivate for two to five months before finally pupating.

530 Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini

Black Cutworm Agrotis ipsilon [Ipsilon Dart] ! recognition Tan to brown, to virtually coal black; dorsum often lightened between subdorsal stripes. Body pudgy and soft. Integument smooth, densely peppered with minute granules and raised, blackened setal bases. Posterior dorsal setal base (D2) on abdominal segments A1–A8 is two to three times diameter of anterior base (D1). Middorsal and subdorsal stripes most evident in pale individuals. Prolegs comparatively small and weak. Head often dark with pale triangle. Anterior prolegs with 16 to 21 crochets; anal proleg with about 28 (Whelan 1935). Larva to 4.5 cm. ! occurrence Agricultural lands, waste places, lawns, fields, and grasslands. Cosmopolitan; in New World ranging from northern Canada, south to Florida, Texas, and into South America. Three generations in Ohio and Connecticut with mature caterpillars from May onward; additional generations southward. Common to occasionally abundant. ! common foodplants General feeder on grasses and low-growing forbs, including many important field and garden crops; frequently reported damaging corn. ! remarks Although the integument of the Black or Greasy Cutworm appears oily, presumably it is not, but simply is smoother than that of other caterpillars. The caterpillar tunnels belowground during the day and then emerges to feed after sundown. Larvae commonly sever seedlings and low-growing plants at ground level and then pull the plant matter into their underground burrows where they consume their cache. The Black Cutworm is one of the most chronic and widespread pests of field crops treated in this work. Infestations of economic consequence have been reported from clover, corn, lettuce, potato, tobacco, and tomato. The insect may be especially damaging to seedlings of row crops. Fields that have flooded (Crumb 1926a) or low-lying, damp areas appear to be particularly prone to attack. In addition to plant matter, the Black Cutworm will also eat other caterpillars. It is a migratory species that occurs nearly worldwide—it is not clear what portion (if any) of northern populations represent year-round residents versus migrants from the South. It is one of the modest subset of migratory moths where a reverse (southward) fall migration has been documented (Showers et al. 1989, 1993). In the Deep South, the Black Cutworm appears to be continually brooded or overwinters mostly as sexually immature adults. While Crumb (1929) notes that pupae overwinter successfully in Tennessee, we see no evidence of yearround survival in southern New Jersey or elsewhere in the Northeast.

Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini 531

Dusky Cutworm Agrotis venerabilis [Venerable Dart] ! recognition Smooth, shiny, greasy gray to nearly black (below right),

flecked with pale spots; prominent black pinacula. Dorsum often more lightly pigmented than sides of body. Thin, obscure middorsal stripe most evident over thoracic segments; rudimentary whitish stripe between dorsal setae. Spiracles black. Integument (under high magnification) cobblestonelike. Crumb (1956) describes head as “very pale brownish much obscured by the heavy black submedian arcs which are much broader dorsally, the black reticulation unusually sparse and sometimes absent.” Whelan (1935) mentions a dark line extending along sides of triangle as well as a dark spot in middle of triangle. Anterior prolegs with only five to nine crochets; anal proleg with 18 to 20 crochets. Larva under 4.5 cm. Caterpillars of the Clay-backed Cutworm (A. gladiaria) and other Agrotis are similar; Crumb (1929) provides a larval key to common eastern members of the genus. ! occurrence Old fields, waste sites, agricultural lands, gardens, grasslands, savannas, and other open habitats. Transcontinental through much of Canada, south across entirety of United States. One generation with mature caterpillars in May and June. Very common. ! common foodplants General feeder on grasses and forbs including alfalfa, barley, chickweed, clover, corn, oats, and tobacco. ! remarks While this moth may outnumber all other owlets in some agricultural fields, it only rarely causes economic injury to field and grain crops. The caterpillar’s life history is briefly summarized in Crumb (1929). Eggs hatch in the fall or winter and the caterpillars overwinter in various instars. Larvae that we held outdoors in southern New Jersey fed but did not reach second instar until very late February or early March. While the early instars accepted grasses, they showed an overwhelming preference for nonnative vetch seedlings and clovers. The caterpillars had a strong tunneling instinct, and even bored into the toweling and tissue of our rearing containers to conceal themselves. The caterpillar figured above, reared by William Forrest and Don Lafontaine on an artificial diet, is appreciably paler than the wild-collected larvae described by Crumb (1956) or the cohort we reared on forbs.

532 Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini

Voluble Dart Agrotis volubilis

Central Alberta to Newfoundland, south to Maryland and Virginia (mountains) and Great Lakes states. One generation with mature caterpillars in spring and early summer. Common. Presumably feeding generally on forbs and other low-growing plants.

Switchgrass Dart

Dichagyris acclivis [Inclined Dart] Kansas, northwestern Connecticut and Cape Cod, south to North Carolina and Texas. One brood with mature caterpillars in September and early October. Generally rare but locally common in southern New Jersey. Larvae feed at night on developing seeds of switch grass (Panicum virgatum). See species account in Lafontaine (2004).

Conglomerated Dart

Euxoa pleuritica [Fawn Brown Dart] Transcontinental in Canada, south in East to Long Island and Great Lakes states. One generation with mature caterpillars from late spring into summer. Common and widespread in Canada; protected species of sand plains and beaches in Connecticut and New York. Presumably feeding generally on lowgrowing plants.

Striped Cutworm

Euxoa tessellata [Tessellate Dart] Transcontinental in southern Canada; in East south to Georgia and Kansas. One generation with mature caterpillars from late spring into summer. Common. General feeder on wide range of plants, including many cereal and vegetable crops; rarely climbing and injuring apple, cherry, pear, and plum.

Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini 533 Fleece-winged Dart Euxoa velleripennis

Manitoba to Prince Edward Island, south in East to North Carolina (mountains) and northern Kansas. One generation with mature caterpillars in summer. More common northward. Presumably a general feeder on low-growing plants.

Knee-joint Dart Feltia geniculata

Manitoba to Nova Scotia, south to coastal North Carolina, northern Georgia, and Texas. One generation with mature caterpillars from May into June. Locally common in dry habitats. Presumably a general feeder. Black pinacula over dorsum somewhat enlarged.

Adobe Dart Feltia manifesta

Illinois, southern Ontario, and southern Maine to northern Florida and Texas. One generation with mature caterpillars from March to May. Generally rare, but sometimes locally common in dry, oak scrub and woodlands in New England and New Jersey. Feeds on herbs in spring (Crumb 1956); will eat dead leaves to some extent.

Tricose Dart Feltia tricosa

Montana, southern Manitoba, and Maine, south in East to northern Georgia and Texas. One generation with mature caterpillars in spring. Common. General feeder on low-growing plants. Dorsum lacks reddish tints of Dingy Cutworm (F. jaculifera complex).

534 Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini

Reddish Speckled Dart Cerastis tenebrifera ! recognition Green, gray, straw, brown or red-brown, with broad, welldefined, white, gray, salmon, or reddish spiracular stripe; abdomen swollen rearward. Dorsum of abdomen with dark, oblique subdorsal patches, edged with white posteriorly, which extend to anterior of trailing segment. These strongest on A7 and A8 according to Crumb (1956), although in figured individual above, these markings most evident anteriorly. Upper edge of spiracular stripe running through dark spiracles; lower edge undulate, dipping down on A3–A6, edged with darker ground coloration; spiracular stripe narrowing and better defined across thoracic segments and A9–A10, where it drops down side of anal proleg. Faint, thin, white subdorsal stripe. Some forms with oblique bar on A1–A7, white above and dark below, that runs down to and envelopes spiracle. Head small and according to Forbes (1954), “reticulate, with a dorsal stripe in front, and two short vertical bars on sides.” Larva to 4 cm. ! occurrence Woodlands from southern Ontario to Newfoundland, south to Georgia and Texas. One generation with mature caterpillars in late spring and early summer. Locally common. ! common foodplants Our lab-fed larvae started on cherry, dandelion, and lettuce, then were sleeved on cherry. McCabe (1990) reared ex ova cohort on birch, blueberry, cherry, raspberry, and willow. British species of Cerastis are polyphagous on forbs, but also consume blueberry, willow (Porter 1997), and undoubtedly others. ! remarks The Reddish Speckled Dart flies in the early spring—it is the first resident noctuine to be taken at lights over much of our coverage area. Adults do not come to bait. The White-marked [Dart], a British Cerastis species, nectars at willow blossoms (Porter 1997). The caterpillar of Reddish Speckled Dart matures in early summer. The winter is passed as a pupa in a substantial underground cocoon.

Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini 535

Flame-shouldered Dart Ochropleura implecta ! recognition Smooth, variegated yellow-brown to brown with humped eighth abdominal segment; at rest, caudal half of A8 pinched into ridge that runs from spiracle to dorsal hump. Thin, broken, white middorsal and subdorsal stripes edged with dark brown spotting. Broad, tan to white subspiracular stripe running down outer face of anal proleg; upper edge of stripe passing through lower edge of spiracles on A1–A3 and A7, through middle of spiracles on A4–A6, then dropping below spiracle on A8. In our images, caudal end of A9 and A10 blackened. Prolegs on A3–A6 with black plate. Spiracles pearly grayish to white with black rim. Larva less than 3.5 cm. ! occurrence Fields, waste lots, other open habitats, and woodlands. Transcontinental in Canada, south in East to Georgia and Texas. At least two generations in Northeast with mature caterpillars from June onward. Common. ! common foodplants Aster, clover, dock, and likely other low-growing plants (Crumb 1956, Rockburne and Lafontaine 1976); Crumb also lists willow. ! remarks We have never found a wild larva of this common moth, which suggests that it feeds on forbs and other low-growing plants mostly at night. The Flame Shoulder (O. plecta), a British congener, is thought to overwinter as a pupa (Porter 1997).

Venus Flytrap Dart

Hemipachnobia subporphyrea Rare moth restricted to wet pinelands in three counties in North Carolina and one bog near Annapolis, Maryland. One generation with mature caterpillars mostly in late spring. Venus flytrap (Dionaea muscipula). Images are of penultimate (inset) and ultimate captive-bred caterpillars. See account in Schweitzer et al. (2011).

536 Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini

Sundew Dart Hemipachnobia monochromatea ! recognition Red, yellow, or deep brown with beige to tan middorsal stripe; some individuals with frosted aspect. Middorsal stripe sometimes reduced to intersegmental spot edged with diffuse black pigmentation rearward (inset). Dorsum darkened adjacent to middorsal stripe, becoming progressively paler to subdorsal stripe. Ground color darkened through supraspiracular area; paler below spiracular line. Prothoracic shield pale, divided by middorsal and subdorsal lines. Spiracles black, small; spiracle on A8 elongate, twice height of that on A7. Larva to 3.5 cm. Too little is known about variation in our two Hemipachnobia to know if larval phenotypes overlap. Location will separate most larval collections. ! occurrence Bogs, acidic wetlands, and mesic pinelands. Northern Ontario to Labrador, to Great Lakes states and along Atlantic coast to northeastern North Carolina. One generation with mature caterpillars from fall into early spring; in Connecticut, from September to May. Generally scarce, but frequently common in bogs and at nearby flowers. ! common foodplants Early instars on sundew (Drosera); later also feeding on heaths such as cranberry, blueberry, and sheep laurel. ! remarks The Sundew Dart was formerly thought to be a subspecies of the Venus Flytrap Dart (H. subporphyrea) (Forbes 1954, Franclemont and Todd 1983), but is now regarded as a separate species (Lafontaine 1998). Early instars of the Sundew Dart feed on young Drosera leaves (Forbes 1954), consuming them from the underside—the sticky hairs that extend from leaf upper sides are shunned. Later, the caterpillar at least sometimes feeds on nearby heaths. The figured individual was reared in the lab on cranberry and sheep laurel by William Forrest and Don Lafontaine. Detailed observational studies of the feeding habits and diet breadths of our Hemipachnobia are needed. Much could be learned by observing larvae in terraria provisioned with a variety of potential hosts. See Lafontaine (1998) for a more detailed account of this moth and its caterpillar. The last instar overwinters.

Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini 537

Brick Dart Lycophotia phyllophora (= Heptagrotis phyllophora) ! recognition Red (commonly), orange-brown, or yellow-brown with

well-developed middorsal stripe and subtle smoky mottling; integument glossy. Middorsal stripe edged with diffuse black pigment, continuing across anal shield. Ground color lighter below level of spiracles. Rear somewhat thickened and broadly rounded but not humped. Crumb (1956) notes: “Oblique pale triangles indicated posterior to II [= D2].” Head reticulated with brown. Spiracles black. Larva under 4 cm. ! occurrence Heathlands, barrens, woodlands and forests, especially on acid soils from Minnesota to Nova Scotia, south to New Jersey, northern Georgia (mountains), and Great Lakes states. One generation with mature caterpillars in late spring. Common northward. ! common foodplants General feeder on woody plants; Phipps (1930) gives alder, arrowwood, birch, blueberry, cherry, meadowsweet, and willow, with cherry as a favorite. ! remarks In many wooded habitats in southern Canada and across the northern United States (e.g., the heath-rich pinelands of New Jersey), the Brick Dart can be among the most common owlets at light in late spring and early summer. Although caterpillars are often found feeding on blueberry and cherry, they do not reach densities of economic significance. Late instars overwinter.

Double Dart

Graphiphora augur (= G. haruspica) Transcontinental in Canada, south to Maryland (mountains), Great Lakes states, and South Dakota. One generation with mature caterpillars in spring and early summer. Locally common. General feeder on forbs, shrubs, and trees, including blackberry, Indian plum (Oemleria), nettle, oceanspray, salmonberry, strawberry, and willow.

538 Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini

Large Yellow-winged Dart Noctua pronuba

[Large Yellow Underwing]

! recognition Last instar brown with buff to white subdorsal stripe bordered inwardly with shallow black dashes or hemispheres on A1–A8. Subdorsal stripe rudimentary over thorax, stronger across anterior half of each segment. Dorsum of A9 and posterior half of A8 often buff. Body pale below thin, white spiracular stripe. Hair-thin, vague middorsal stripe frequently broken or absent. Spiracles tan to brown with black outer ring. Head often with black (coronal) bar over vertex continued alongside triangle; short black line under eyes. Larva to 4.5 cm. Middle instars brown (upper right page 539) to bright yellow-green (inset). Dorsal black spots on other noctuines often more triangular than those of Large Yellow-winged Dart. Lesser Yellow-winged Dart (N. comes) [Lesser Yellow Underwing], another European species, has become established in British Columbia and in the Toronto area (Crolla 2008). Its larva has the dorsal black hemispheres limited to A6–A8, and its spiracles are white (Lafontaine 2004). ! occurrence Agricultural lands, waste places, lawns, fields, grasslands, woodlands, and other, especially grassy, habitats. Transcontinental in Canada, south in East to Florida, Texas, and California. Evidently asynchronous over our region with mature caterpillars present over much of year, especially fall through early summer. Very common to abundant. ! common foodplants General feeder on grasses and forbs. ! remarks The Large Yellow-winged Dart was accidentally introduced into eastern Canada around 1979 from Europe (Neil 1981). It has since spread across the continent. The caterpillars are active throughout the winter, turning up on sidewalks, seen sauntering into garages, or crawling alongside and over snow

Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini 539 banks. Occasionally, they are encountered in considerable numbers on the surface of newly fallen snow. This account was recently sent to us by Adam Wilson: On 14 January … after a 6 in snowfall, we witnessed hundreds of Noctua pronuba on the surface of the powdery snow near Bicentennial Pond in Mansfield, Connecticut. They were mostly within 20 feet of the edge of the frozen pond in open (grassy) areas not under trees. It was cold (~32° F) and the caterpillars were hardly moving at all, so it is curious that they would have crawled up through the snow to the surface. It had been fairly warm the previous few days (in the 40s), so perhaps they had been active and were surprised by the snow. Curiously, we have heard of several reports of dogs regurgitating small numbers (less than a dozen) of cutworm caterpillars during the winter months—invariably, these turn out to be larvae of the Large Yellow-winged Dart. Since its establishment in Connecticut in 1993, the moth has become one of the most common owlets in the state. It is routine to flush adults while mowing lawns anywhere in the Northeast from May through September. In flight the (colorful) hindwings and flight are reminiscent of some grasshoppers. The grayish eggs, laid in clusters of several hundred (below), are deposited on foliage (frequently of trees and other nonhost plants), dead sticks, grass stalks, clotheslines, pine needles, wire fences, or almost any other surface. Neonates fall to the ground where they graze on grasses and forbs. The prolegs in full grown larvae strike us as comically small relative to the abdominal mass. Pupation occurs belowground in a slight cocoon.

Smaller Pinkish Dart Diarsia jucunda

Southern Ontario to Newfoundland, south in East to northern New Jersey, North Carolina (mountains), and through Great Lakes states. One generation with mature caterpillars from late spring into summer. More common northward and at high elevations in Appalachians. Evidently a general feeder; captive larvae have been reared on grasses (Rockburne and Lafontaine 1976) and dandelion (McCabe 1990). Yellow-brown dorsum with yellow predominating rearward.

540 Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini

W-marked Cutworm Spaelotis clandestina

[Clandestine Dart]

! recognition Brown to gray with latter often predominating; blackish W-shaped middorsal spots over abdominal segments (but see below). Thin, white middorsal stripe sometimes broken, often at midsegment. White to tan subdorsal stripe best developed along abdominal segments. Inner side of subdorsal stripe edged with black spots that are narrow and crisply delineated at anterior of each abdominal segment, but which broaden and become less well defined rearward. Thick, white subspiracular stripe infused with gray or brown mottling. Patterning noticeably obscure across thoracic segments with exception of prothoracic shield, which is cut by fine middorsal and subdorsal stripes. White to yellowish spiracles embedded in black spot. Head with strong black “X,” formed by fusion of coronal bars over triangle; lower arms separate and drop down each side of triangle. Larva to 4 cm. ! occurrence Fields, coastal communities, sand plains, and edges of woodlands and forests. Transcontinental in Canada, south to North Carolina (mountains) and Texas. Evidently one generation with mature caterpillars in spring that yields adults in early summer; these then diapause until late summer and fall (see also below). Occasional pest westward, but uncommon over much of our area. ! common foodplants General feeder on both herbaceous and woody plants, including apple, beans, blueberry, cabbage, maple, pine, and strawberry. ! remarks This occasional pest is one of our most easily and reliably determined cutworms, but be forewarned that not all individuals have the W-marks as prominent as those in the image above. The moth is ecologically generalized, present from sea level to mountaintops, at home in grasslands as well as woodlands and forests, yet curiously, and true to its name, seldom seen in number in any of these habitats. While published literature indicates the Clandestine Dart has two annual generations (e.g., Lafontaine 1998, Handfield et al. 1999), we think it more likely that there is only a single generation with a summer, adult diapause. The egg stage may be as long as two months (Crumb 1929). The W-marked Cutworm overwinters as a partly to nearly full-grown larva.

Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini 541 Green Arches Dart Anaplectoides prasina

Transcontinental in Canada, south in East to at least New Jersey, Georgia (mountains), and Great Lakes states. One generation with mature caterpillars from late spring into summer. More common northward. General feeder on forbs, shrubs, and other lowgrowing plants. Spots surrounding dorsal setae absent or obscure (Lafontaine 1998).

Dappled Dart

Anaplectoides pressus Transcontinental in Canada, south in East to northern New England and Great Lakes states; replaced by A. brunneomedia in southern Appalachians. One generation with mature caterpillars from late spring into summer. More common northward. General feeder on herbs, shrubs, and trees, including larch. Spots surrounding dorsal setae well defined (Lafontaine 1998).

Great Brown Dart Eurois astricta

Transcontinental in Canada, south in East to central New England and Great Lakes states. One generation with mature caterpillars from late spring into summer. Common northward. Many woody plants including alder, birch, blueberry, cherry, maple, meadowsweet, poplar, strawberry, and viburnum. Orange lines through prothoracic shield; head more than 3.75 mm in width.

Great Gray Dart Eurois occulta

Transcontinental in Canada, south in East to Virginia (mountains) and Great Lakes states. One generation with mature caterpillars in summer. Common northward. Many forbs and woody plants, including birch, blueberry, cedar, cherry, gale, larch, maple, meadowsweet, snowberry, and willow. Orange lines through shield; head less than 3.75 mm in width. Outbreaks may have played a role in collapse of Viking settlements in Greenland (Handfield et al. 1999).

542 Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini

White Pine Dart Xestia badicollis [Northern Pine Dart, Northern Variable Dart]

! recognition Ground coloration varies from green to gray-green and/or

brown. Usually with bright white subdorsal stripe, about twice width of middorsal stripe. Upper margin of subspiracular stripe often edged with black; in well-marked individuals lines may join to form black spiracular stripe. Dorsal pinacula white, inconspicuous. Larva to 4 cm. Perhaps indistinguishable from caterpillars of Southern Pine Dart (X. elimata) and Early Conifer Dart (X. praevia). Lafontaine (1998) notes that the Early Conifer Dart has a broader middorsal stripe (nearly as wide as the subdorsal stripe), and the black spiracular line (when present) is narrower than that of the White Pine Dart and less likely to be swollen across the intersegmental membrane. ! occurrence Woodlands and forests. Southern Ontario to Labrador, south to Georgia and Great Lakes states. One generation with mature caterpillars from fall into following spring (most), and early summer northward. Very common. ! common foodplants Eastern white pine, but also reported from fir, hemlock, larch, other pines, and spruce, possibly in error. See below. ! remarks This species is widely known to foresters as the White Pine Cutworm. Hosts and habitat, phenology, and range provide helpful clues to the identities of our conifer-feeding darts. The Southern Pine Dart (next page) is associated with hard pines from Maine and Missouri southward. The Early Conifer Dart (X. praevia) is recorded from jack and other hard pines, as well as fir, hemlock, larch, and spruce (Lafontaine 1998). It begins flying in June and continues into August, while adults of the White Pine Dart appear in August and fly through September, and those of Southern Pine Dart mostly in September or later. According to Don Lafontaine, collection records for the White Pine Dart are almost exclusively from locations where white pine is present, and host records from fir, hemlock, larch, spruce, and even other pines, may refer to the Early Conifer Dart. It is not known if the brown forms are inducible—it would be worthwhile to raise cohorts on different conifer hosts and under different larval densities. All of our collections of wild larvae, as well as those of Chris Maier (pers. comm.) from across the Northeast, have been of the green form.

Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini 543

Southern Pine Dart Xestia elimata [Southern Variable Dart] ! recognition Ground color gray to brown or green (rarely). Gray and brown forms heavily speckled with pale spots. Faint, middorsal stripe usually present. Dorsal setae from minute white pinacula. Gray and brown forms frequently with dark middorsal spot toward anterior of A1–A8. Subdorsal stripe broken into white spots that may be embedded in broader black stripe. Spiracular stripe, when present, black, broken over thoracic segments, and reduced on T1. Often with pinkish or reddish cast below level of spiracles. Green form with prominent white to creamy subspiracular stripe and whitish subdorsal stripe, both of which may be thinly edged with black, especially on their upper side (lower left). Larva to 4 cm. Earlier instars are described below. See also White Pine Dart (X. badicollis). ! occurrence Barrens and pine woodlands from Missouri to southern Maine, south to Florida and eastern Texas; common along Atlantic Coastal Plain. One generation with mature caterpillars in late fall to spring. Common. ! common foodplants Loblolly, longleaf, pitch, red, shortleaf, Virginia, and probably other hard pines. Usually associated with pitch pine north of New Jersey. Last instars occasionally observed by DFS eating ericaceous (e.g., Lyonia) buds in fall. ! remarks The larva is also known as the Variable Climbing Caterpillar. The caterpillar feeds on mature foliage, even in spring when new needles are flushing. The first two instars are green and rest with the body appressed along a needle. Third instars are usually gray-green to brown with a white subspiracular stripe, although a few individuals remain green through the fifth (lower right) and sixth (lower left) instars. In contrast to the first two instars, the third is more likely to wedge itself head down into needle fascicles by day. By the fourth instar, and especially the fifth and sixth instars, many caterpillars leave the foliage to shelter in bark crevices or on the ground. Caterpillars ascending from the ground are commonly encountered at bait in pine barrens. Modest frass accumulation in our outdoor rearing containers indicates that the caterpillars feed during midwinter warm spells. Some larvae reach the last instar in November in Massachusetts, and most do in New Jersey, but few complete feeding before April in either state. Larvae construct a silk-lined cell belowground in the spring, in which they will remain as prepupae until late summer when pupation takes place.

544 Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini

Reddish Heath Dart Xestia dilucida [Dull Reddish Dart] ! recognition Tan to smoky brown with diffuse black and white markings. Broken, whitish middorsal stripe; subdorsal stripe sometimes reduced to little more than white midsegmental spot on A2–A7. Dorsum frequently marked with oblique, black subdorsal spots that form chevrons over midline; these alternate with tan to brown patches. Spots on A8 truncated, ending in crisp, black line across dorsum. Black spiracles sit atop broad, variously expressed, subspiracular stripe. Head dusky orange brown, shiny, with weak coronal bars and reddish-brown reticulation over each lobe. Larva to 4 cm. Larva inseparable from that of Young’s Dart (X. youngii) (page 551). The two moths are sympatric from southern New Jersey to coastal North Carolina. Young’s Dart inhabits bogs range-wide; southward along the Atlantic Coastal Plain the moth is also a denizen of mesic to soggy pinelands, while northward it also occurs in dry barrens, heathlands, etc., essentially replacing X. dilucida. ! occurrence Pine or oak woodlands and forests, balds, barrens, heathlands, and dry pine woodlands from Ontario to Nova Scotia, south to Florida and eastern Texas (including range of X. youngii). One generation with mature caterpillars mostly in late spring. Common. ! common foodplants Blueberry, huckleberry, and likely other heaths; captive early instars ignored oak. ! remarks The eggs hatch in late fall. Early instars are thought to feed initially on senescing and fallen leaves. Over the winter months, our larvae sporadically consumed dead blueberry leaves when temperatures climbed above 5° C (~ 40° F). In the spring, the caterpillars ascend their hosts at night to feed on buds, leaves, and flowers. Crumb’s (1956: 99) records for larch and spruce probably apply to another species. Maier et al. (2004) had no records of either the Reddish Heath or Young’s Darts from any conifer. Our images are of caterpillars swept at night by Mark Mello from patches of lowbush blueberry and black huckleberry on Nantucket Island. The larva in the inset was collected with other Reddish Heath Dart caterpillars, but did not yield an adult. Prepupal caterpillars have more subdued markings and take on a pinkish flush. The two moths are sympatric in southern New Jersey. In North Carolina X. dilucida occurs in the Piedmont, whereas X. youngii is known only from coastal plain locales. Young’s Dart inhabits bogs range-wide; southward it is also a denizen of mesic to boggy pinelands, while northward it also occurs in dry barrens, heathlands, etc., essentially replacing X. dilucida.

Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini 545

Greater Spotted Cutworm Xestia dolosa [Greater Black-lettered Dart]

! recognition Green (lower right), gray, or (most commonly) brown with variable patterning and thin, indistinct middorsal and subdorsal stripes. White to tan, black-ringed spiracles situated along upper edge of pale spiracular stripe that divides darker, upper portion of body from paler venter; subspiracular stripe dipping well below spiracle on A8 and continuing down anal proleg. Gray and brown forms, usually with dark, oblique, subdorsal spots that become more prominent rearward to A8; these nearly black over A7–A8. Dark coronal bars over vertex run to either side of triangle; considerable mottling present over lobes. Proleg on A3 with about 25 crochets. Larva to 4 cm. Inseparable as a caterpillar from Spotted Cutworm (X. c-nigrum), a species of agricultural lands, waste places, fields, gardens, and grasslands, which occurs continent-wide in Canada, and in the East, south to North Carolina. Xestia as currently defined is a large and ecologically varied genus with more than 50 North American species (Lafontaine 1998)— identifications are best based on adults. ! occurrence Fields, barrens, woodlands, forests, and a variety of closed and open habitats from Saskatchewan to Nova Scotia in Canada, south to Florida and Texas. Two or three principal generations over much of East with mature caterpillars from May onward. Common. ! common foodplants General feeder on grasses, forbs, and low-growing woody plants, but also reported from apple, currant, and pear. ! remarks Up until 1980, the two aforementioned spotted cutworms were confused as a single species. Lafontaine (1998) suggests that the two moths are ecologically partitioned, characterizing X. dolosa as a species of mesic woodlands and X. c-nigrum (= X. adela) as a denizen of disturbed and grassy landscapes. Caterpillars of the latter species are commonly reported damaging field and garden crops, although rarely do their numbers warrant intervention. Feeding damage to buds can be severe. The best way to see spotted cutworms is to go out at night with a flashlight and search vegetation within a meter of the ground. There is scarcely a garden, woodland edge, or weedy lot in eastern North America that does not host larvae at some point during the spring or summer months. The winter is passed as a nearly full grown larva.

546 Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini

Smith’s Dart Xestia smithii ! recognition Tan to gray-brown, sometimes with pinkish cast, of variable patterning; thin, indistinct middorsal and subdorsal stripes. Oblique, dark, diffuse dorsal patches fuse over midline at trailing edge of each abdominal segment; these often join small, oblique “lines” that extend forward from D1 setae to form dark middorsal “X”s centered over intersegments between A1 and A8. Often with whitish subdorsal patch that includes D2 seta over abdominal segments. Some individuals with orange to salmon stripe running just below subdorsal stripe. Broad, subspiracular stripe vague or absent; lower half of body mostly unpigmented with flecking of pale spots. Spiracles pale yellow to brown, rimmed with brown-black. Prothoracic shield brown with fine middorsal stripe. Coronal bars extend forward and run down either side of triangle; fine, black line running back from second stemma. Larva to 3.5 cm. ! occurrence Fields and woodlands. Transcontinental in Canada, south in East to northern Georgia (mountains), Great Lakes states, and northern Iowa. One generation with mature caterpillars in April and May. Very common over much of East; scarce on coastal plain. ! common foodplants General feeder on forbs, shrubs, and smaller trees, including alder, apple, birch, cabbage, cherry, dock, elderberry, false hellebore, grape, passion flower, raspberry, Russian olive, salmonberry, strawberry, thimbleberry, viburnum, and violet. Late instars often on shrubs and small trees. ! remarks This common cutworm is rarely abundant enough to cause economic injury, although Crumb (1956) makes note of an outbreak on strawberry in Washington State. A female that we collected at bait laid the majority of her eggs in three masses, each with several dozen ova. The larvae, which hatched in late October, were offered a salad of various forbs and crabgrass and initially showed a preference for the tiny leaflets among the flowers on goldenrod. Caterpillars also ate aster flowers and leaves of chickweed, crabgrass, passion flower, and violets over the winter. In April the caterpillars were sleeved and reared to maturity on cherry. Many hosts cited in literature are indicative that older larvae become climbers.

Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini 547

Spotted-sided Cutworm Agnorisma badinodis

[Pale-banded Dart]

! recognition Boldly marked. Ground color grayish to pale brown; integument smooth. Pinacula minute. Thin, vague middorsal stripe most evident over thoracic segments. Black, roughly triangular to hemispherical spots over A1–A8, outwardly edged with creamy subdorsal stripe; posterior side of triangles about half length of side abutting subdorsal stripe. Oblique, black lateral spots enclose or abut spiracles along their lower reach. Spiracles black. Head with dark coronal bars that reach to clypeus, and two, thin lateral lines that include eyes. Distal end of spinneret, with numerous microscopic spinules, appearing fringed under high magnification. Larva to 3.5 cm. Middle instars slender, tapered, and striped, features common among grass feeders. The caterpillar of A. bugrai possesses dark, barlike (not triangular) subdorsal spots (Lafontaine 1998); it flies with A. badinodis over much of Canada and the northern United States, south to Massachusetts and the Great Lakes states. The range of another Agnorisma, A. bollii, has been expanding eastward in recent decades; it is now encountered along the East Coast from Georgia to northern Delaware. Its larva has not been described. ! occurrence Ecologically generalized, including fields and waste lots, floodplains, and woodlands from North Dakota, extreme southern Canada, and New Hampshire; south to Georgia (mountains), Mississippi, and central Texas. One generation with mature caterpillars in spring. Common. ! common foodplants General feeder on forbs, shrubs, and small trees. Crumb (1929, 1956) lists aster, chickweed, dock, mustard, and tobacco. We reared a cohort on cabbage, chickweed, clover, dandelion, dock, goldenrod, grass, and star of Bethlehem (Ornithogalum) over the course of winter; later in spring, we fed them cherry and oak. Rings (1977) adds alfalfa, apple, cherry, pin oak, and wheat. We have found larvae on beech and viburnum in spring. ! remarks Our autumn-born, early instars fed on grass and various forbs (see above). Throughout the winter and spring, larvae were active on warm, southern New Jersey days. Most reached the fourth instar by March. Our fifth instars refused the grasses that they ate all winter. In the spring, larvae commonly ascend shrubs and trees to feed on new growth, occasionally causing injury to economically important species, e.g., fruit trees (Rings et al. 1992). The mature larva pupates in a cell belowground in the spring but does not eclose until the late summer or fall. Adults visit flowers and fermenting berries (e.g., pokeweed).

548 Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini

Mottled Gray Cutworm Abagrotis alternata

[Greater Red Dart]

! recognition Smooth, variegated in tans and browns, with whitish

middorsal and faint subdorsal stripes. Thin middorsal stripe interrupted by black spot between adjacent segments on A1–A8. Vague, diffuse, dark patches to either side of midline include both dorsal setae on first seven abdominal segments. Thin, black, transverse line followed by diffuse pale area toward rear of A8. Supraspiracular area darkest where it borders spiracular stripe. White portion of spiracular stripe fading rearward of A7. Ground color pale below level of spiracles. Setal bases over dorsum often whitened, those below spiracular stripe darkened. Prolegs with little pigment. Coronal bars radiate outward to form networks over each lobe. Spiracles tan or brown, ringed in brown or black. Larva under 4 cm. Well-marked Cutworm (A. orbis) (= A. barnesi), as denoted by its name, is more boldly patterned (page 550). ! occurrence Woodlands and forests from Alberta to Nova Scotia, south in East to Georgia (mountains) and Texas. One generation with mature caterpillars from April to June. Very common. ! common foodplants General feeder, late instars commonly on woody plants in spring, including apple, blueberry and other heaths, cherry, hickory, oak, and walnut. Most of our records from blueberry and oak. Also reported from cabbage and strawberry. See below. ! remarks Female Abagrotis fly and lay their eggs in the fall. Our hatchlings and early instars consumed forbs, senescent leaves, and/or fallen leaves, and reached the third instar by winter. Crumb (1956) collected numerous young caterpillars of the Mottled Gray Cutworm in February among fallen oak leaves in Tennessee; these matured by April. Ours, set up in pots outdoors, fed on fallen blueberry and cherry leaves through the winter. With the onset of spring, Abagrotis move up into shrubs and trees to feed on buds, leaves, and flowers. The Mottled Gray Cutworm is among the most arboreal of our darts, commonly ascending the trunks of trees (at night) to graze on foliage. Occasionally, they are numerous enough to cause significant injury to buds of orchard trees. Larvae mature before tree leaves are fully expanded, pupate belowground, eclose in early summer, then aestivate until late summer before mating and ovipositing.

Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini 549

One-dotted Dart Abagrotis magnicupida ! recognition Smooth, variegated in red-browns, tans, browns, and less

commonly, grays. Thin, pale middorsal stripe interrupted between segments. Dorsum of abdomen with dark, oblique lines running between dorsal setae; these darker, larger, and more wedge shaped on A7 and A8 (Crumb 1956). Some forms with pale subdorsal spot at leading edge of T3–A7 (sometimes A8), which fuses with mostly obsolescent subdorsal stripe; in our collections, this spot largest on A1, then becomes progressively smaller rearward. Figured individual bears vague, rusty stripe below broken subdorsal stripe. Thin, white spiracular stripe divides darker supraspiracular area from largely unpigmented venter; upper edge sometimes lined with black. A8 with low hump and dark brown line connecting D2 setal bases. Hemispherical prothoracic plate cut by white middorsal and subdorsal stripes. Pinacula black. Head amber with broad brown coronal bars and thinner lateral line extending to eyes; area between filled with reticulations. Mature larva to 4 cm. ! occurrence Grasslands, powerline right-of-ways, fields, waste lots, and (northward) coastal strand communities from Nebraska to Massachusetts, south to Georgia, Alabama, and Texas. One generation with mature caterpillars in spring. Common. ! common foodplants General feeder on both herbaceous and, in spring, woody plants—see Remarks. Sometimes damaging to apple, grape, and peach (Crumb 1956) (reported as A. cupida). ! remarks This species was formerly misidentified in collections and literature as A. cupida (Lafontaine 1998). Females of this genus simply drop the eggs, presumably as they are crawling over the ground. The early instars of the Onedotted Dart feed on forbs or grasses. Our cohort of the One-dotted Dart hatched in mid-October and readily took to chicory, chickweed, dandelion, basal rosette leaves of aster and fleabane, and freshly fallen cherry leaves, as well as aging crab grass blades. In the spring, the nocturnal caterpillars climb into trees and shrubs to feed on buds, flowers, and leaves. Ours were reared to maturity on young cherry leaves. As interpreted by Lafontaine (1998), Crumb’s (1956) record of an Abagrotis causing injury to apple, grapes, and peaches (in the spring) are referable to this species. The biology of the moth was studied by Rings (1972). At least along the Atlantic Coastal Plain, this is often a common backyard and old field Abagrotis from Long Island southward. Adults visit sugar baits in the fall.

550 Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini

Yankee Dart

Abagrotis brunneipennis Transcontinental in Canada, south to North Carolina (mountains) and Great Lakes states. One generation with mature caterpillars in spring and early summer. Common northward and in southern New Jersey, where heaths grow in abundance. Frequently blueberry (including fallen leaves during winter), also accepting cherry and other forbs and woody plants.

Well-marked Cutworm Abagrotis orbis (= A. barnesi)

Great Plains states eastward into Great Lakes region, especially in areas of sandy soil. One generation with mature caterpillars in May and June. Common westward. General feeder; occasional pest of apple, grape, peach, and other plants.

Daggered Heath Dart

Eueretagrotis attentus [Attentive Dart] Manitoba to Nova Scotia, south to northeastern Connecticut, Georgia (in mountains), and Great Lakes states. One generation with mature caterpillars from late spring into summer. Locally common. Wild hosts in spring include birch, blueberry, elderberry, strawberry, and willow. Winter habits (of younger larvae) unknown for genus.

Brown-collared Dart Protolampra brunneicollis

Alberta to Nova Scotia, south in mountains and Piedmont to Georgia, Mississippi, and Oklahoma. Captures of adults span most of growing season; two generations or possibly one with adults aestivating. Locally common. General feeder on both herbaceous and woody low-growing plants including blueberry, clover, dandelion, sweet fern, and tobacco.

Cutworms and Darts Subfamily Noctuinae: Tribe Noctuini 551 Pink-spotted Dart

Pseudohermonassa bicarnea Saskatchewan to Nova Scotia, south to Georgia (mountains), Kentucky, and Iowa. One generation with caterpillars overwintering and maturing by early summer. Common. General feeder; Crumb (1956) lists birch, blueberry, dandelion, and sweet fern; McCabe (1990) reports grasses in genus Glyceria. Dorsal setae from black pinacula.

Norman’s Dart Xestia normanianus

Alberta to Nova Scotia, south to Georgia (mountains), Kentucky, and Great Lakes states. One generation with mature caterpillars in late spring. Very common. Phipps (1930) lists blueberry, cherry, meadowsweet, raspberry, and sweet fern. Similar to Smith’s Dart (X. smithii) (page 546).

Boomerang Dart Xestia perquiritata

Transcontinental in Canada, south in East into northern New England, and as disjunct populations at high elevations to southern Appalachians. One generation with mature caterpillars mostly in June. Common northward. Fir, spruce, and larch.

Young’s Dart Xestia youngii

Minnesota to Newfoundland, south along Atlantic Coastal Plain to Carolinas and Great Lakes states. One generation with mature caterpillars in May and June. Locally common. Blueberry, huckleberry, leatherleaf, and other heaths; our larvae ate living and fallen leaves of blueberry. Probably indistinguishable as larva from Reddish Heath Dart (X. dilucida) (page 544).

552

Glossary Abdomen  Ten segments of body that follow leg-bearing thoracic segments. Addorsal  Area adjacent to dorsal midline (Fig. 11). Adenosma  Midventral defensive gland on first thoracic segment; often projecting as a tube in preserved specimens. Adfrontal area  Narrow plate adjacent to frons between adfrontal suture and ecdysial line. Area with AF setae in Fig. 2. Adfrontal suture  Line delimiting side of triangle (frons) (Fig. 2). Present in all Lepidoptera; not present in immatures of other insect orders. Aestivate  To be inactive over summer months. Aestivating adults may continue to occasionally drink and feed, but are in a state of reproductive diapause. Anal plate  Dorsal plate or shield atop last abdominal segment (A10) (Fig. 12). Anal proleg  Proleg arising from last abdominal segment (A10) (Fig. 1). Angiosperms  Flowering plants, including all broadleaf plants, forbs, grasses. (Not gymnosperms or cone-bearing plants, which include cedars, firs, hemlocks, larches, pines, spruces, and others.) Anteclypeus  Lower portion of clypeus, adjacent to labrum. Antenna  Three-segmented sensory structure between base of mandible and eyes (Figs. 2, 6); small and inconspicuous in caterpillars. Antepenultimate instar  Instar preceding penultimate instar. Aposematic  Warningly colored. Boldly colored in yellows, oranges, and reds, often with black or white markings as well; less commonly with only black and white pattern elements. Band  Pattern element running around segments, perpendicular to body axis; broader than rings. Barcode  DNA sequence; used in this work to refer to a 648-basepair section of the mitochondrial genome (i.e., cytochrome c oxidase subunit I) shared by all animal life, and one increasingly employed by zoologists to ascertain species identities. See Barcode of Life Data Systems (http://www.boldsystems.org/views/login.php). Batesian mimicry  Evolutionary phenomenon whereby a palatable species comes to resemble a distasteful, toxic, or otherwise protected species, and thereby gains some protection from predators. Beating  Method of obtaining caterpillars by abruptly striking vegetation over sheet, umbrella, or other collection surface (pages 17–18). Biordinal  Crochets of two, often alternating, lengths. Bisetose  With two setae. In this work usually referring to number of subventral setae on first two abdominal segments. Bivoltine  Having two generations per year. Chaetotaxy  Distribution and development of larval setae, and in particular primary setae. Chaetotaxy is important in identification of larval lepidopterans and has played a significant role in moth classification. Clypeus  Elongate plate between antennae, below frons or frontal triangle. Cocoon  A construction of silk fashioned by caterpillar prior to pupation. Often incorporating leaf fragments, wood chips, larval setae, etc. Congener  Member of shared genus: e.g., all Melipotis species are each other’s congeners.

Coronal bar  Dark barlike spot (or thinner line) over each lobe of head (see inset for Green Cutworm, Anicla infecta, page 523). Coxa (coxae)  First or basal segment of true (thoracic) leg. Cremaster  Rearward extension of terminal (tenth) segment of pupa, often with set of hooklike spines that lock pupa into silk strands of cocoon wall. Crochets  Hooklike structures on abdominal prolegs (Fig. 9). Cutworm  Common name for smooth, usually ground-dwelling and ground-colored caterpillar; in narrowest sense, applied to caterpillar that severs seedlings near ground level (for subsequent feeding), and that hides during the day. Taxonomically most applicable to members of the Noctuini, but often used by applied entomologists and others for virtually any (and sometimes all) noctuids. Detritus  Dead and often partially decomposed organic matter; used in this guide primarily for nonliving plant matter. Diapause  Period of hormonally controlled inactivity. Often induced by shortening day lengths in late summer and fall and broken by warm temperatures in spring. Discal cell  Closed, centrally located, cell in both forewing and hindwing. Distal  Away from body (compare with proximal). Dorsal  Along back or upper side. Dorsum  Back or upper side. Ecdysial line  Thin areas in cuticle that rupture during a molt and allow new instar or stage to issue from old integument. In this guide used for lines that run parallel to adfrontal sutures, to either side of frons (or triangle) (Fig. 2). Eclosion (eclose)  Emergence from an egg or pupa. Ectoparasitoid  Parasitoid (larva) that feeds and develops outside of host’s body, e.g., Euplectrus in parasitoid plate (page 29). Endoparasitoid  Parasitoid that feeds and completes its larval development inside body of host. Epizootic  Disease outbreak within an animal species or group. Ex ova  Reared from egg, nearly always from eggs obtained from confined female. Facultative  Brood or diapause that may or may not occur. Commonly used to refer to a partial generation, i.e., where a fraction of the individuals in a population go through an additional brood, while others remain in a state of diapause. Feculae  Pelletlike larval excrement; droppings. Femur (femora)  Second readily discernible segment of thoracic leg. Frass  Larval scrapings, for example from larval excavation in wood or stems; used by some authors to refer to feculae or excrement. Frons  Triangular area in center of head (Fig. 2); often referred to in this work as the triangle. Gena  Side of head, rearward from eyes; “cheek.” Hair pencil  Bundle or fascicle of elongate scales or setae. Haplotype  A unique mitochondrial DNA sequence found within a population or species; reciprocally distinct haplotypes are indicative that two entities are geographically or biologically distinct lineages. Heart stripe  Middorsal stripe that results when heart (and gut) is visible through larval cuticle.

Glossary Hemolymph  Translucent yellow to green fluid that fills insect’s body. Shares functions of blood and lymphic systems, but differs in that it does not transport oxygen (in most insects). Idiobiont  Parasitoid that arrests growth and development of its host following attack and oviposition, i.e., without a delay in development of parasitoid. Compare with koinobiont. Instar  Larval stage between molts; most owlets have between five and seven instars, which can differ markedly in form, duration, and habit. Integument  Body surface, cuticle, or skin. Intersegmental  Between adjacent segments; often pigmented differently than adjacent cuticle. Koinobiont  Parasitoid that allows host to feed, grow, and continue development up to the point where the parasitoid initiates and completes its own development (thereby usually killing the host larva); in essence, delayed parasitoidism. Compare with idiobiont. Labrum  Upper lip; plate below anteclypeus that rests over jaws; often with medial cleft that engages leaf when feeding (Fig. 7). Larva (larvae)  Feeding stage of insects with complete metamorphosis—synonymous with caterpillar in this work. Lash  Elongate cluster of wispy setae: less compact than a hair pencil. Lateral  Along sides. Lobe  Rounded area of head above eyes, to either side of midline. Medial  Running along or near body midline. Melanic  Dark forms; common in adults of pantheas (Pantheinae) and daggers (Acronictinae). In larvae, melanic forms (melanics) occur in Alabama, Amyna, Orthosia, armyworms, and others; disproportionately common among outbreak species. Melanized  Darkened; brown to black, often contrasting with adjacent less melanized tissue. Roughly synonymous with sclerotized in many insects. Mesoseries  Linear or curving row of crochets (never connected in circle or nearly complete circle). Middorsal  Along dorsal midline of body (Fig. 11). See also heart stripe. Midline  Line dividing left and right sides of body; dorsal midline is above and ventral midline is below. Midventral  Along ventral midline of body (Fig. 11). Mullerian mimicry  Evolutionary phenomenon whereby distasteful, toxic, or otherwise protected species come to resemble one another. In so doing, the members of a Mullerian mimicry ring gain by more efficiently educating local predators. Natural group  Group with common origin (having a shared common ancestor). Neonates  Newly hatched larvae. Noctuidae s. l. Noctuidae sensu lato (in the broad sense); embracing the four families formerly classified in the Noctuidae, i.e., the Erebidae, Euteliidae, Noctuidae, and Nolidae. Equivalent to owlets. Noctuids  Approximately the trifid subfamilies of previous workers, i.e., Plusiinae through Noctuinae here, following Lafontaine and Schmidt (2010) and Zahari et al. (2011). Noctuoids  As presently defined by Zahari et al (2011), includes six families: the Erebidae, Euteliidae, Noctuidae, Nolidae, Notodontidae, and Oenosandridae.

Oviposit  To lay an egg or ovum. Oviposition  Act of laying an egg or ova. Ovum (ova)  Egg(s). Owlet  Common name for many noctuoids. Used to collectively refer to the four family-level taxa treated in this volume: Erebidae, Euteliidae, Noctuidae, and Nolidae. See also noctuids and noctuoids. Papillose  With numerous, minute rounded projections, i.e., papillae. Parasitoid  Predator that lives internally or externally on its host. Parasitoids are parasitelike in that they are smaller than their hosts, feed from within or on the host’s body, and often do so over a period of weeks or months; however, functionally they are predators because they nearly always kill their host (prey). Penultimate instar  Instar preceding last or ultimate larval instar. Pharate  Hidden; when one stage is fully formed but has not yet emerged from the cuticle of previous stage or instar. Pharate adults are developed (diminutive) moths that are holding within their pupae (pupal shells). (See page 476.) Phenology  Seasonal timing or progression of life cycle (as determined by climate and other factors). Phenotype  Appearance of an individual; an organism’s observable features. Pheromone  Chemical used to communicate among individuals of a single species; in moths, pheromones are (often) odorless airborne molecules (or mixtures of molecules) produced in minute quantities. See sex pheromone. Phylogenetic  Relating to phylogeny or evolutionary history; a phylogenetically informative character is one that reveals evolutionary linkages within a given lineage or group. Pinaculum (pinacula)  Base plate from which a seta is borne. Nearly always sclerotized, shiny, pigmented, or in some other way distinguished from adjacent cuticle. When raised and pimplelike used synonymously with (setal) wart. Planta  Distal, crochet-bearing portion of proleg. Plastron  Physical gill formed by trapped layer of air across which gas exchange occurs. Plastrons commonly form over integumental surfaces that are resistant to wetting (because of water’s strong cohesive forces). Polyphagous  Eating plants from several to many plant families. Prepupa (prepupae)  Postfeeding portion of the last instar. Color changes occur frequently in prepupa; while most species lose patterning and color, members of several subfamilies turn pinkish or even red as prepupae (see Acronicta lithospila, page 272). Primary setae  Those setae represented in the ground plan or ancestral lineage(s) of Lepidoptera. Primary setae are broadly homologous across lepidopteran families, and have a standardized nomenclature (Fig. 10). Number and positions of primary setae are used in the identification and classification of Lepidoptera. (Compare with secondary setae.) Prolegs  In caterpillars, the fleshy “legs” located on abdominal segments three, four, five, six, and ten, although the first two pairs are often reduced or lost in several erebid and noctuid subfamilies. Only in Lepidoptera do the larval prolegs bear crochets (hooklike spines) (Fig. 1).

553

554 Glossary Prothoracic shield  Dorsal plate over first thoracic segment (Fig. 1). Proximal  Toward center of body (compare with distal). Pseudospiracle  Darkened structure; in same position as true spiracles on second and third thoracic segments of herminiine larvae. Pupa  Stage between caterpillar and adult; often enclosed in cocoon or earthen cell. A nonambulatory stage, but one of great physiological activity in which most larval cells and tissues are replaced by those of adult. Puparium (puparia)  Specially formed pupal stage of some flies (including tachinids, page 31), i.e., where pupation occurs in the last larva skin. Shape and external features of puparium are often of considerable taxonomic value. Rootlet setae  Thickened setae that form fringe along subventer of underwing (Catocala) caterpillars. Sometimes more fleshy than hairlike. Rugose  Roughened or pitted. Sallow  Any of the mostly fall- and spring-active owlets, especially among Orthosiini, Psaphidini, and Xylenini, but also used more generally as a common name for other owlets (e.g., some Heliothinae and Oncocnemidinae). Originally applied loosely to the spring-flying owlets that nectar at sallow (willow) blossoms, especially in Great Britain. Secondary setae  Setae in excess of primary setae. (Compare with primary setae.) Seta (setae)  Hairlike outgrowth of integument. Setal base  Pinaculum; rounded, hardened plate that bears seta, often more darkly pigmented than adjacent cuticle. Setose  Covered with setae or resembling a seta or bristle. Often used in this guide to describe SD1 seta on A9; when SD1 is the same diameter as adjacent setae it is said to be setose; when it is thinner than adjacent setae it is said to be hairlike. Sex pheromone  Pheromone used to attract opposite sex or, at close range, in courtship. Most commonly referring to long-range “calling” pheromones released by virgin females. Skeletonize  To remove leaf tissue except for embedded network of veins. A spectacular example is figured on page 38. Many owlets are skeletonizers over the course of the first or early instars. Spatulate  Resembling a spatula; referring here to seta that broadens distally. Specialist  Feeding on members of a single plant genus or two closely related plant genera. Spermatophore  Reproductive packet delivered to female during copulation; in addition to sperm it may contain carbohydrates, defensive substances, hormones, proteins, salts, steroids, and other compounds, which may enhance a female’s survival or reproductive efforts. Conversely, the spermatophore may contain physical or chemical elements that interfere with the ability of the female (or other males) to obtain additional copulations. Spinneret  Elongate, silk-producing structure on lower surface of head; medial projection from labium (Figs. 4, 8). Spinule  Minute spine; sharply pointed integumental extension. Spinulose  Covered with minute spines; here referring to surface texture of larval integument.

Spiracle  Lateral, round to oval opening of respiratory system; in caterpillars found on first thoracic and first eight abdominal segments (Fig. 1). Spiracular  Adjacent to or passing through spiracles (Fig. 11). Stage  One of the four principal life stages of an insect. All Lepidoptera have four: the egg, larva, pupa, and adult. Stemma (stemmata) Lateral eye(s) of caterpillars (Fig. 3); most macrolepidopteran caterpillars have six stemma arranged in a semicircle. Stemma are numbered counterclockwise, starting from the upper end of the semicircle. Stripe  Marking that runs (longitudinally) along body axis. Subdorsal  Below level of addorsal and above supraspiracular areas (Fig. 11). Subspiracular  Below level of spiracles and above subventer (Fig. 11). Subventral  Area above legs and prolegs but below subspiracular area (Fig. 11). Supraspiracular  Above level of spiracles and below subdorsal area (Fig. 11). Taxon (taxa) General and collective term that embraces various taxonomic groupings; in this work often used to refer to species- and generic-level entities. Teneral  Recently eclosed or emerged; not yet fully hardened. Thoracic  Of or pertaining to thorax. Thorax  Body area consisting of three segments between head and abdomen, and bearing true legs with claws. Tibia (tibiae)  Segment of leg above tarsus (five-segmented foot) and below “knee.” Tonofibrillae  Internal muscle attachment points on body wall; often appearing as minute circular plates on integument. Trachea (tracheae)  Larger air-delivery tubes in insect’s respiratory system (see tracheal trunk). Tracheal trunk  Principal respiratory duct connecting spiracles, often visible through body wall. Transverse  Running from side to side, perpendicular to caterpillar’s body axis. Triangle  Triangular area between eyes (Fig. 2); also called frons or frontal triangle. Area with F1 setae in Fig. 2. Trisetose  With three setae; in this work referring to number of subventral setae on either or both of the first two abdominal segments. Truncate  Appearing cut or squared off. Trunk  Thoracic and abdominal segments collectively. Tympanum  Ear or sound-detecting membrane (below hindwing on side of thorax in owlets); sensitive to the high frequencies used by echo-locating (hunting) bats. See essay on page 476. Uniordinal  Crochets of one length. Univoltine  Species or populations with only one generation each year. Venter  Underside or belly. Vertex  Dorsal or top portion of head; upper portion of each lobe. Wart  Integumental outgrowth. Often used synonymously with pinaculum when latter is raised and pimplelike, especially for the dorsal (D) setae.

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Foodplant Index Most hostplants are listed under their common name. Listings for scientific names are given for some less familiar plants. Common name usage and spellings generally follow those of the USDA PLANTS Database (http://plants.usda.gov/index.html). abutilon  76, 240–1, 346 Abutilon fruticosum  232 incanum  245 theophrasti  76 acacia  146, 166–7, 186 blackbrush 70 Acacia rigidula  70 acorns 516 Aeschynomene virginica 346 Agastache foeniculum  218 Ageratina altissima  368 Alcea rosea  240 alder  59, 67, 207, 222, 255, 260, 262–4, 268, 271, 283, 286, 292–3, 297–8, 330, 343, 401–3, 421–2, 424, 430, 435, 440–2, 453, 468, 470, 474–5, 486, 492–4, 496–8, 503, 537, 541, 546 alfalfa  70, 190, 215, 221, 385, 388, 393, 407, 491, 493, 495, 502, 511, 513, 524, 529, 531, 547 algae 89 green  87–90, 380–2 Allium canadense  363 almond 206 amaranth  234, 388, 391 amaryllis 519 Amblyolepis setigera  361 Ambrosia  244, 357, 362 artemisiifolia  243, 245, 357, 362, 415, 529 psilostachya  339 trifida  339, 352, 357, 362–3 Amianthium muscae­ toxicum 414 Amorpha 70 canescens  104–6, 359, 362 fruticosa  104, 106, 359 Ampelopsis  306, 308–9 arborea  306 Ampelothamnus philly­ reifolius  184 Amphiachyris dracun­ culoides  361 anachuita 324 andromeda  134, 443 Andropogon gerardii  409, 501 virginicus  157 angelica 415 anise 218

Annona glabra  81 Antirrhinum 321 Apeiba tibourbou  73 Aphanostephus ramo­ sissimus 361 Apocarya  114, 117, 120 Apocynum cannabinum 414 apple  136, 138–41, 172, 183, 257, 264–9, 271, 284, 288, 292, 304–5, 326, 388, 402, 423, 427, 429, 430, 433, 435, 438, 453, 456, 461–2, 464, 473, 477, 481, 484–6, 497, 513, 524, 527, 529, 532, 540, 545–50 apricot 502 Aquilegia canadensis  415 Aralia spinosa 414 Arctium  414–15 lappa  414 arrow bamboo  507 arrowhead 213 Artemisia filifolia  362 frigida  361 ludoviciana  362 artichoke 348 ash  260, 283, 299, 305, 319, 325, 336, 427–8, 430, 433, 438, 442, 457, 477, 481, 484, 486–7, 494 black 415 green 415 white 415 asparagus  216, 348, 365, 390, 392, 491, 493, 497 aspen  125, 173, 205, 218, 222, 255, 262, 286, 422, 425, 428–9, 431, 434, 441, 448, 453, 458, 468–9, 477, 484–5, 497 big-toothed  204, 434 quaking  127, 464 aster  58–60, 213, 219, 312–15, 347, 353, 367, 369, 372, 401, 411–12, 415, 448, 466, 471, 493–4, 499, 517–18, 535, 546–7, 549 eastern showy  352 flaxleaf whitetop  352 hairy white oldfield 317 skyblue 352

Aster azureus  363 concolor  363 laevis  361, 415 linariifolius  352, 363 multiflorus  361 novae-angliae  415 oolentangiense  352, 363 puniceus  361, 363, 415 spectabilis  352, 363 umbellatus  415 Atriplex  53 Aureolaria flava  342 grandiflora  342 pedicularia  342 autumn olive  57, 477 awnless bushsunflower 248 azalea  134, 273 pink 134 bahia 381 Balduina angustifolia  359, 362 balloon vine  234 banana 221 Baptisia  60, 412–13 alba  414 tinctoria  414 barberry 53 Barbados cherry  210 barley  221, 501, 505, 531 basswood  59, 65, 100, 173, 188, 240, 260, 265, 284, 288, 326, 424, 427–8, 433, 435, 442, 478, 481, 484, 486–7, 489, 496 bayberry  108–9, 484 beach pinweed  151 bean  60, 70, 215–16, 224–5, 348, 365, 388, 390, 392, 394–5, 499, 527–9, 540 green 525 kidney  215, 221 lima 525 pinto 525 see also soybean beardtongue 320 beautyberry 259 bee balm  378, 517 beech  45, 96, 188, 200, 249, 252–3, 260, 268, 277, 280–2, 292, 433, 480–1, 489, 547 beet  491, 523 beggarticks  60, 70, 344 bearded 339

bellflower 327 Bidens  364 aristosa  339 big bluestem  409, 501 bindweed  58, 243–4 birch  59, 70, 74, 172–3, 222, 229, 249, 252–3, 255, 260, 262–5, 268– 9, 271, 281, 283, 286, 292–3, 298, 330, 382, 421–2, 424, 428–30, 433–5, 441–2, 450, 453, 468, 470, 477, 480–1, 483–4, 486–7, 489, 492, 494, 503, 511, 516, 534, 537, 541, 546, 550–1 black  265, 402, 434, 438, 442, 446 gray  441, 472 paper  271, 441 river  67, 285 white  164, 468 yellow  164, 446 bitternut 110 blackbead  167, 186 catclaw 149 blackberry  58, 172, 264, 292–3, 297, 326, 402, 427, 430, 450, 453, 471, 485–6, 493, 496, 498, 517, 537 black ironwood  397 black olive  206 bladdermallow  76, 232, 240, 241 blazing star  352, 414 blue giant hyssop  218 blueberry  51, 61, 134–5, 170, 223, 225, 227–9, 260, 264, 273, 283, 292, 297, 299, 305, 326, 330, 403, 421, 423–4, 430, 432, 437, 442, 453, 455–6, 458, 462, 464–5, 471, 474, 479, 480, 482, 484–8, 495–8, 503, 511, 516, 521, 534, 536–7, 540– 1, 544, 548, 550–1 highbush  135, 473 lowbush  135, 143, 318, 449, 456, 459–60 bluegrass 237 boneset, blue  366 false 359 white-flowered 358 boxelder  183, 260, 289, 380, 430, 434

brackenfern  375, 377, 416, 489, 493–4 Brickellia eupatorioides 359, 362 broccoli 216 brome 510 broom snakeweed  242–3 Bucida buceras  206 buckeye  260, 427, 436, 441, 461, 482, 486 yellow  429, 436 buckthorn 398 buffaloberry  430, 470, 494 bulrush 408 softstem 408 burdock  327, 365, 411–12, 414–15 butter-and-eggs  322, 499 buttercup 493 butternut  112, 207, 209, 453, 486 buttonbush  62, 305, 309 button mangrove  206, 210 Byrsonima lucida  210 cabbage  216, 221, 224–5, 343, 388–9, 392, 465, 491, 495, 497, 502, 513, 524, 527, 540, 546–8 cacalia 413 Cacalia tuberosa  414–15 Calamovilfa brevipilis  501, 521 Callicarpa 259 Calylophus berlandieri  361 hartwegii  361 camphorweed  248, 313, 353, 360, 529 campion, bladder  500 starry  493, 500 canarygrass 404 reed 409 canola 502 cantaloupe 378 Cardiospermum  234 Carex pensylvanica  84 stricta  236 see also sedge Carphephorus corymbosus  361 carrot  225, 392, 410, 495, 524 cassia  167, 172, 186 cattail  300, 419 cauliflower 216 ceanothus 305 see also New Jersey tea

563

564 Foodplant Index cedar  403, 541 Atlantic white  47 northern white  443 red  260, 382, 443 white  382, 443 celery  225, 367, 391, 491 Celtis pallida  84 Centaurea maculosa  361 cereals 532 Chaenactis 347 Chelone glabra  415 Chenopodium  234 ambrosioides  217 cherry  70, 138–9, 172, 175, 183, 222, 257, 264–71, 283, 292–4, 297–8, 305, 326, 382, 403, 421–2, 427–30, 432–5, 437–8, 442, 448–50, 452–4, 456– 9, 461, 464–5, 470–4, 477–82, 484–9, 496–8, 503, 532, 534, 537, 541, 546–51 black  47, 138, 172, 175, 265, 270, 304, 382, 422, 424, 431, 434, 454, 462 pin  259, 265, 271, 286, 430–1, 441 see also apricot and plum chestnut  59, 85, 95, 173, 260, 272, 280–2, 326, 480 chickweed  531, 546–7, 549 chicory  344, 513, 549 chinquapin 245 chocolateweed 254 chokeberry  140, 172, 256, 265–7, 326, 456–8, 462, 464, 488 red 140 Chrysoma pauciflosculosa 362 Chrysopsis mariana  248 pilosa  363 subulata  352, 361–2 Cicuta maculata  414 Cirsium 414 Cladium 238 Clethra acuminata  202 alnifolia  202 climbing false buckwheat 397 climbing fetterbush  184 climbing hempvine  364 clover  39, 52–3, 70, 92, 152–3, 217, 221, 224, 359, 362, 385, 390, 393, 466, 491, 493, 495, 511–13, 518, 523–4, 527–31, 535, 547, 550

bush  70, 153, 254, 297, 487 coastal plain honeycomb­ head 359 coffeeberry 398 Collinsonia  493 canadensis  342, 414–15 columbine  231, 327, 344, 413, 415 composites 316 Comptonia peregrina  107, 297, 344, 430, 442–3, 459, 484, 494, 550–1 coneflower 415 Conocarpus erectus  206, 210 Conyza  213 canadensis  312–13, 317, 353, 360–2 Cooperia  235 Corchorus siliquosus  73 cordgrass, big  507 saltmeadow 157 smooth 152 Cordia boissieri  324 corn  60, 70, 155, 216–17, 225, 245, 348, 385, 388–95, 405, 407–9, 412–13, 417, 490, 495, 497, 505, 510, 513, 524, 526–31 Corylus americana  287 cosmos 217 cotoneaster  267, 304 cotton  76, 78, 215, 346, 348, 385, 388–95 cottonwood  126–7, 298, 325, 474 eastern 126 swamp 126 coyotillo 398 crabapple  137–41, 160, 257, 428, 431, 433–4, 462, 464, 478 crabgrass  57, 157, 236, 237, 546, 549 cranberry  299, 423, 433, 460, 502, 526, 536 large 225 cranesbill 221 Crataegus flava  142 margarettiae  139 marshallii  142 spathulata  142 viridis  142 see also hawthorn Crepis pulchra  361 Croptilon divaricatum  313 Croton californicus  361 capitatus  245 dioicus  361

texensis  361 crownbeard  58, 340, 365, 414 crucifers  385, 391, 395 cucumber 348 cucurbits  385, 391 Culver’s root  412, 416 cup plant  416 currant  292, 326, 421, 430, 441–2, 484, 486, 492–3, 527, 545 cutleaf coneflower  415 cypress 164 bald 165 Cyrilla racemiflora  159 dahlia  412, 417 daisy, confederate  341 cowpen 340 tahoka 313 Dalea canescens  362 dandelion  49–50, 221, 224, 231, 373, 402, 422, 493, 495, 497, 509, 511, 513–14, 516–18, 534, 547, 549–51 Dasistoma macrophylla 342 dead leaves see detritus dead wood see detritus Decodon  310, 412 verticillatus  416 Delonix regia  147 delphinium 347 desert-chicory 360 desert false indigo  70, 106, 359, 427 Desmodium  73 detritus (including litter)  11, 15, 39–46, 48–9, 51–7, 396, 500, 515–16, 533 dead insects 38 dead leaves  38–40, 42–60, 87, 89, 396, 402–3, 465, 513, 516, 520, 523, 533, 544 dead wood  85 dewberry 262 Digitaria  236 Dionaea muscipula  535 Diospyros kaki  308 dock  213, 373, 379, 397, 402, 413, 416, 465, 513, 517, 529, 535, 546–7 prairie 416 dogbane  91, 292, 344, 413–14, 499, 521 spreading 327 dogfennel 368

dogwood  65, 259, 271, 283, 292, 421, 479, 488, 496 Douglas-fir  48, 173, 229, 325, 443, 477 Drosera  536 dung 38 dustymaiden 347 earthball 40 ebony see blackbead  elderberry  417, 480, 493, 546, 550 elm  65, 70, 95, 173, 249, 253, 257, 260, 265, 271, 283–4, 288, 292, 296, 427–8, 433, 441–2, 452, 465, 477, 479, 481, 484–5, 489, 491, 493 American  65, 290 cedar 290 rock 65 slippery 65 winged 290 Elymus hystrix  414 Epilobium  306, 310, 466 Erigeron  312, 317, 362 annuus  362 modestus  313 philadelphicus  362 Eryngium yuccifolium  415 eryngo 415 Erythrobalanus  130 Eubotrys racemosa  135, 161, 297, 403, 432 Eupatorium  358, 363, 412, 517, 529 altissimum  358 capillifolium  368 compositifolium  358 fistulosum  358 purpureum  368, 415 evening primrose  306, 310, 354 false hellebore  546 farkleberry  134–5, 476 fern  374–7, 401, 403 chain 416 cinnamon  416, 494 hayscented 376 interrupted 416 marsh 377 New York  376 ostrich 415–16 royal  377, 416 sensitive  402, 413, 415, 517 Virginia chainfern 377 fetter-bush  403, 432 feverwort  319, 427

fir  47, 59, 227, 229, 250, 343, 382, 403, 435, 498, 521, 542, 551 balsam  47–8, 338, 343 fireweed 306 flax  389, 491, 529 fleabane  224, 312, 317, 372, 549 plains 313 prairie 313 fly poison  414 forbs  59, 172, 225, 295, 299–300, 344, 378, 388, 393–5, 423, 546 foxglove, false  342 mullein 342 Fraxinus americana  415 nigra  415 pennsylvanica  415 fringetree 319 fruit trees  385, 395 fungus  41, 44, 89–90, 381 bracket 89–90 earthball 40 mushrooms 89 gilled 384 pore-bearing 384 Scleroderma 40 smut 384 Gaillardia pulchella  361, 363 gale  59, 292, 485, 494, 541 sweet  107, 109, 423, 443, 452 Garberia fruticosa  361 heterophylla  361 gaura 310 biennial 354 Gaura  361 biennis  362 coccinea  362 filipes  362 mollis  362 geranium  327, 346, 502 gilia 347 gladiola  347, 405, 502 globemallow  63, 232 Glyceria  404, 551 golden alexanders  416 golden crownbeard  340 goldenaster, Maryland  248 scrubland 352 goldeneye 365 sunflower  245, 248 goldenrod  59, 213, 219, 245, 312, 314–15, 317, 323, 356, 367, 369, 372, 414, 448, 494, 499, 517–18, 529, 546–7

Foodplant Index seaside 414 woody 362 gooseberry  477, 480, 503 Gordonia lasianthus  163 grape  307–10, 326, 382, 513, 524, 527, 546, 549–50 grass  39, 49–52, 58, 72, 84, 87, 152–7, 224, 230–1, 236–8, 300, 348, 350, 381, 388–94, 404–9, 413, 417, 423, 490, 499, 505–10, 513–14, 517, 522–3, 525, 527–8, 529–31, 538–9, 545, 547 big bluestem  409, 501 bluegrass 237 bottlebrush 414 canarygrass 404 crabgrass  57, 157, 236, 237, 546, 549 mannagrass 404 orchard 510 pasture 155 panic 237 pine barren reedgrass 501, 521 prairie dropseed  521 purpletop tridens  350, 501 redtop 510 reed canarygrass  409 sandreed  501, 521 sawgrass 238 sorghum  155, 388–9, 392–3, 501 switchgrass 501–2, 506–7, 532 timothy  51, 392, 405, 495, 501, 510 turf  393, 407 wiregrass 152 see also oats, rye, and wheat greenbrier  191, 326, 328–9, 432, 480 gregorywood 206 Grindelia  347 lanceolata  362 nuda  361 groundcherry 346–8 gumhead 369 gumweed 347 Guteirrezia sarothrae  242– 3, 361 Gymnosperma glutinosum 369 hackberry  70, 86, 296, 454, 457, 485 spiny 84

hairypod cowpea  190 Hampea  78 Haplopappus  313 divaricatus  363 hawthorn  136–42, 160, 172–3, 256, 259, 266–7, 269, 284, 304, 326–7, 427–8, 433, 435, 454, 476, 479, 485 green 142 littlehip 142 lowland 142 parsley 142 yellowleaf 142 hazel  45, 49–50, 63, 173, 184, 232–3, 260, 263–4, 269, 281, 283, 287, 292, 297, 430, 434, 438, 443–4, 455, 484–5, 488–9, 495, 517 American  209, 259 beaked  209, 259 Helenium autumnale  415 Helianthus divaricatus  415 giganteus  415 microcephalus  341 petiolaris  361 porteri  341 strumosus  415–16 Heliopsis helianthoides  416 hemlock  45, 48, 250, 338, 382, 403, 435, 480–9, 498, 521, 542 water 414 western 227 Heptacodium miconioides 305 Heracleum lanatum  415 maximum  415 Hercules club  414 Herissantia  76 crispa  232, 241 Hermannia texana  362 Heterotheca canescens  361 graminifolia  313 subaxillaris  248, 313, 360–3, 529 villosa  361 hibiscus  75–6, 233, 240, 310 hickory  53, 85, 95, 110– 11, 114, 116, 118–22, 145, 173, 188, 207–8, 260, 264, 272, 283, 286, 326, 331, 427, 433–5, 437–8, 441–2, 449–50, 455, 457–8, 481, 485–7, 516, 548 bitternut 121–2

black 118 mockernut  110, 114, 116, 122 nutmeg 117 pignut  110–11, 114, 116, 118, 120, 122, 208 sand  110, 116, 118, 449 scrub 118 shagbark  69, 110–11, 113–17, 119–22, 208 shellbark  120, 122 see also pecan Hieracium gronovii  361 hogwort 245 holly  432, 463, 478, 480 American 463 mountain 482 hollyhock  76, 224, 240 honeysuckle  305, 318, 482, 493 Morrow’s 48 hop  68, 218, 309, 409, 414 hophornbeam  63, 173, 184, 209, 287, 432–3, 438, 443–4, 446, 480, 484–6, 488–9 hornbeam 435 American  63, 173, 209, 252–3, 260, 264, 271, 287, 422, 438, 443–4, 484–5, 489 horse-balm  342, 414–15, 493 horsemint 378 horseweed  213, 312–13, 317, 353, 360 huckleberry  61, 401, 459, 484, 487, 492, 544, 551 black 460 Hudsonia montana  151 tomentosa  151 Humulus lupulus  414 hydrangea 326 Hydrophyllum  488 Hypericum  399–400 prolificum  402 Ilex glabra  463 opaca  463 verticillata  58 indigo see desert false indigo and wild indigo indigo bush see desert false indigo inkberry  305, 463 Ipomaea  58, 243 leptophylla  363 iris  405, 408, 412, 423, 519 ironweed 413

Missouri 415 New York  414 ironwood 265 Isocoma drummondii  361 Iva  359 annua  362 frutescens  362 jewelweed 517–18 joe pye weed  358, 412, 415 sweetscented 368 Juncus  408 see also rushes Kalmia polifolia  318, 474 Karwinskia  398 knotweed  370, 495 Kosteletzkya virginica  240 Krugiodendron  397 kudzu 190 Kuhnia eupatorioides  359, 362 Labrador tea  227–8, 424 lambsquarters  491, 521 lantana  245, 318, 363 Lantana camara  245, 363 Laportea  66, 68, 71, 214, 218 larch  178–9, 250–1, 292, 297, 382, 443, 474, 477, 494, 521, 541–2, 551 larkspur 221 Lathyrus  347 laurel  297, 498 bog  318, 474 mountain 488 sheep  227, 403, 536 lavender thrift  353 leadplant  104–6, 359, 427 leafcup 416 leatherleaf  227, 443, 460, 482, 498, 551 leatherwood 159 Lechea maritima  151 legumes  134, 148, 154–5, 174, 491, 513 Lespedeza  153, 254, 297 lettuce  49–50, 215–16, 221, 224–5, 316, 385, 388, 392, 471, 491, 512–13, 519, 527–30, 534 Leucothoe  134 see also swamp doghobble Liatris  352, 414 cylindracea  352, 363 punctata  352, 363 scariosa  352, 363 spicata  352, 363

lichens  41–2, 87–9, 211 old man’s beard  39 rosette 88 lilac  305, 326 Lilium superbum  414 lily 412 rainlily 235 spider 519 turk’s-cap 414 Limonium carolinianum 353, 361 Linaria canadensis  321 dalmatica  322 vulgaris  322, 499 linden 457 litter see detritus and litter feeding lizard tail  412 loblolly bay  163 locust  70, 185–6, 190 black  158, 174, 184–5 honey  105–6, 168, 184, 485–6 water 106 locust berry  210 Lonicera dioica  318 loosestrife  412, 415 false 310 fringed  201, 493 purple  310, 487, 494 swamp  310, 412, 416 whirled 201 lousewort  231, 499 Ludwigia  310 Lyonia  61, 134 ligustrina  161, 199 mariana  161 Lysimachia ciliata  201 quadrifolia  415 terrestris  415 Lythrum  310 Machaeranthera annua 361 tanacetifolia  313, 361 tenuis  361 Madia  347 Malachra capitata  77 maleberry  161, 199 mallow  73, 217, 240 bladdermallow  76, 232, 241 false  63, 76, 233, 240–1, 245, 339, 523 malva de caballo  77 marshmallow 76 seashore saltmarsh  240 swamp rose  240 Texas Indian  232 Virginia saltmarsh  240 wax  77, 233, 241 Malpighia glabra  210

565

566 Foodplant Index malva de caballo  77 Malvastrum  63, 73, 76, 233, 240–1, 245, 339, 523 Malvaviscus  77, 233, 241 mannagrass 404 maple  45, 89, 164, 173, 183, 249, 253, 260, 283, 326, 403, 429–31, 433, 435, 442, 450, 452, 454, 458, 461, 465, 477, 479–81, 484, 486–7, 489, 492, 497, 503, 540–1 red  64, 164, 289, 428–9, 434, 437, 450, 454, 456, 462, 470, 478 silver  64, 289, 428, 470 sugar  64, 289, 437, 452 marigold 367 marsh-elder 359 marshmallow 76 Matteuccia struthiopteris 416 mayapple  412–14, 416 meadow-rue  80, 231, 413, 416, 497 meadowsweet  294, 403, 422, 537, 541, 551 white 320 see also spirea Melochia  254 mesquite  148, 186 Mexican tea  217 Mikania  364 milkweed  235, 493, 499 milkwort 93 Mimosa 254 mint  217, 378 mistletoe 324 Monarda  378 monkeyflower 495 monkshood 343–4 moonseed vine  79 Morella cerifera  109, 259 pensylvanica  108–9 mountain ash  139, 264–9, 271, 283, 286, 453, 462, 464 mountain goldenheather 151 mullein  494, 514 mushrooms 89 gilled 384 pore-bearing 384 see also fungus mustard  215, 343, 547 Myrica gale  107–9, 423, 443, 452 narcissus 519 nest (inquilines) 39

nettle  45, 66, 68, 214, 218–19, 223, 537 false 65 hedge 221 stinging  214, 327, 493 New Jersey tea  57, 70, 443 nightshade 391 black 347 ninebark  59, 486 Nuphar  419 Nymphaea odorata  419 Nyssa  162, 302–3 oak  52, 85, 95–6, 128–32, 143–4, 149, 172–3, 183, 188, 198, 200, 249, 253, 260, 264, 272, 275, 277–8, 283, 291–3, 297, 299, 326, 330, 333, 421–2, 427, 429, 430–1, 433–5, 437–8, 442, 448–50, 452–9, 461–2, 464–5, 467, 477, 480–7, 489, 496, 498, 515–16, 547–8 black  129, 132, 281, 334, 428 blackjack  131–3, 174, 200, 275, 476 bur 277 chestnut 278 dwarf 277 dwarf chinquapin  200 laurel 132 live  128, 145, 166, 170–1, 187, 199 pin 547 post  129, 277–8 red  95, 129–32, 145, 199–200, 275, 281–2, 291, 334–5, 434, 473 scarlet  129, 335 scrub  59, 128, 131–3, 174, 277, 452, 476 southern red  129 turkey  132, 149, 166, 275 water 132 white  40, 50, 95, 128, 130, 133, 145, 170, 199–200, 275, 278–9, 280–2, 291, 332, 335 willow  149, 175 oats  501, 505, 510, 526, 529, 531 oceanspray 537 Oemleria  537 Oenothera biennis  361 nuttallii  361 parviflora  361 rhombipetala  361

okra  76, 348 old man’s beard  39 olive, anacahuita  324 autumn  57, 477 black 206 Russian  503, 546 Texas 324 onion 527 Onoclea sensibilis  413, 415 orache 53 orchids 388 Ornithogalum  547 Osmunda cinnamomea 416 claytoniana  416 regalis  377, 416 oxeye 416 Palafoxia sphacelata  362 palm 189 Panicum virgatum  157, 501–2, 506–7, 532 Parasponia  234 Parkinsonia  147 parsley  216, 491, 493 parsnip, cow  415 water 414 parthenium  245, 365, 367 passion flower  546 pea  70, 216, 344, 389, 392, 491, 493, 495, 502, 527–9 sweet 347 peach  427, 465, 497, 527, 549–50 peanut  190, 389 pear  257, 532, 545 pecan  114, 117, 120–1, 207, 254 water 117 see also Carya Pedicularis  499 Pediomelum rhombifolium 362 pelotazo 245 penstemon  320, 327, 343–4 pepper  348, 392 peppervine 306 persimmon  82–3, 305, 450 Japanese 308 Phalaris  404 arundinacea  409 Phleum pratense  51 phlox  347, 511 Phlox pilosa  362 Phragmites  404 australis  409, 507 Physalis  347 Physcia  88 pickerelweed 418

pigweed  234, 391–2, 491, 495 pincushion 347 pine  47, 179, 182, 250–1, 337, 382, 435, 521, 540, 542 eastern white  178, 542 jack  47, 178, 180, 182, 226, 439, 441, 542 loblolly  177, 181, 543 lodgepole 178 longleaf  177, 181, 543 pitch  47–9, 177, 180–2, 439, 543 pond  177, 181–2 red  178, 180, 297, 439, 543 sand 176 Scotch 178 shortleaf  176, 543 Virginia  176, 180, 182, 439, 484, 543 white  337, 484 pine barren reedgrass  501, 521 pitcherplant  220, 414, 416 Pithecellobium  166–7, 186 unguis-cati  149 Pityopsis  360 falcata  363 graminifolia  362–3 pinifolia  363 plantain  221, 327, 493, 511, 513–14, 518, 523 Indian 414–15 plum  137–9, 172, 257, 265, 267–70, 292, 304, 403, 427, 431, 457, 465, 478, 524, 532 beach  138, 160, 175, 299, 455, 479 Chickasaw  139, 437 Indian 537 wild 139 Podophyllum peltatum  416 Poeppigia  147 poison ivy  193–4, 196, 489 pokeweed  39, 391, 547 Polygala  93 Polygonella  363 articulata  363 gracilis  363 Polygonum  370, 379 scandens  397 Polymnia uvedalia  416 pond apple  81 poplar  70, 74, 124–7, 173, 222, 255, 260, 264, 271, 286, 292–4, 296–8, 326, 343–4, 421–5, 428, 430, 433, 441–2, 455–6, 465,

468–70, 475, 477, 481, 483, 486, 491, 496–7, 503, 513, 541 balsam  127, 204, 468 Lombardy 126–7 Populus  474 deltoides  474 heterophylla  126 portia tree  77 potato  385, 390–2, 410, 497, 502, 524, 526–30 sweet  254, 378, 390–1, 395, 410, 527 prairie broomweed  313 prairie dropseed  521 prickly fanpetals  243 primrose-willow 310 Protococcus  87–90, 382 Prunus  138 americana  139 angustifolium  137 serotina  172 see also apricot, cherry, and plum Pseudosasa japonica  507 Psoralea  359, 362 Psoralidium lanceolatum 362 tenuiflorum  362 Pteridium aquilinum  375, 377, 416, 489, 493–4 purpletop tridens  350, 501 purslane 523 pyramidflower 254 Pyrrhopappus  360 carolinianus  362 geiseri  362 grandiflorus  362 multicaulis  362 pauciflourus  362 radish 527 ragweed  70, 217, 244–7, 357, 365, 367, 413, 415, 529 annual  243, 245–6, 357 cuman 339 false 367 giant  246, 352 great  339, 357 rainlily 235 raspberry  172, 218, 326, 494, 524, 529, 534, 546, 551 Rayjacksonia annua  361 redbud  260, 326, 486 redtop 510 reed 404 common  404, 409, 507 rhododendron  59, 134, 206, 326, 482, 487

Foodplant Index Rhus  192 aromatica  196 copallinum  192 glabra  192 lanceolata  197 trilobata  192, 197 typhina  192 virens  197 see also sumac rice  155, 389, 392–3, 501 richweed  342, 414–15 rose  264, 268, 283, 292–3, 343–4, 421, 430, 453, 484, 524 rosemallow 76 rose of Sharon  75–6 rosinweed  413, 416 royal poinciana  147 Rubus  262 see also blackberry, raspberry, and salmonberry Rudbeckia laciniata  415 rush  408, 423 Russian olive  503, 546 rye  381, 501, 505, 510, 526, 529 sage 221 salmonberry  493, 537, 546 sandreed  501, 521 Sarracenia  414, 416 alata  220 flava  220 leucophylla  220 minor  220 purpurea  220 sassafras  82, 283 saw palmetto  189 sawgrass 238 Scirpus  408, 423 tabernaemontani  408 Scleroderma cepa  40 Scotch lovage  327 sea lavender  353 sedge  51, 230, 300, 408, 508–9, 522 long-withered 52 Pennsylvania 84 upright 236 yellowed 52 senna 167 sensitive plant  172 serviceberry  140–1, 222, 256–7, 265–6, 268–9, 286, 292, 298, 430, 453, 478, 516 seven son flower  305 Shepherdia  430, 470 sida  63, 70, 76, 233, 240–3 Silene stellata  493, 500 vulgaris  500

silkgrass  360, 362–3 narrowleaf 313 Silphium  409, 416 integrifolium  413 silverbell  472, 482 Simsia calva  248 Sium suave  414 sleepy morning  254 slender scratchdaisy  313 slippery burr  73 smartweed  295, 370, 379 snailseed vine  79 snapdragon 389 sneezeweed  367, 369, 415 snoutbean 190 snowberry 541 soapberry  222, 503 Solidago canadensis  362 gigantea  414 rigida  362 sempervirens  414 stricta  362 see also goldenrod sorghum  155, 388–9, 392–3, 501 sourwood  61, 283, 455, 482 soybean  70, 190, 215, 219, 254, 346, 348, 385, 389, 392 Spanish needle  339, 364 sparkleberry see farkleberry Spartina alterniflora  152 cynosuroides  507 patens  157 Spigelia anthelmia  235 spinach  348, 392, 491 spirea  292, 297, 305, 320, 423–4, 481, 484, 488, 503 Sporobolus heterolepis  521 spotted knapweed  353 spruce  47, 59, 173, 227, 229, 250–1, 337–8, 382, 403, 421, 435, 442, 474, 477, 498, 521, 542, 551 black 47 red 45 white  47, 226, 268 squash 348 St. Johnswort  399–400, 402, 494, 521 staggerbush  61, 134 maleberry  161, 199 piedmont 161 starfruit 81 star of Bethlehem  547 Stenosiphon linifolius  362 stoneroot  414, 415

strawberry  70, 327, 344, 347, 378, 390, 393, 408, 453, 497, 511, 524, 526–7, 529, 537, 540–1, 546, 548, 550 wild  452, 456 sugarbeet  225, 378, 391–2, 395, 527 sugarcane  155, 392 sumac  192, 196, 296, 343, 486–7 evergreen 197 poison  194, 196 prairie 197 skunkbush  192, 197 smooth 192 staghorn 192 winged  192, 197 sumpweed 359 sundew 536 sunflower  224, 248, 348, 365, 367, 385, 401, 412, 415–16 giant 415 Porter’s 341 small woodland  341 swamp candles  201 swamp doghobble  135, 161, 297, 403, 432 sweet fern  107, 297, 344, 430, 442–3, 459, 484, 494, 550–1 sweet gum  195, 197, 283 sweetpepperbush, coastal 202 mountain 202 sweet potato  254, 378, 390–1, 395, 410, 527 switchgrass  501–2, 507 sycamore  85, 260, 289, 442, 454, 487 Symphyotrichum pilosum 317 Tamaulipa azurea  366 tanseyleaf tansyaster  313 tarweed 347 tatalencho 369 tea, Labrador  227–8, 424 Mexican 217 New Jersey  57, 70, 443 teasel 411 Texas olive  324 Thespesia populnea  77 thimbleberry 546 thistle  224, 402, 411–12, 414 Russian 491 thoroughwort  358, 517, 529 tall 358 ticktrefoil  73, 343

timothy  51, 392, 405, 495, 501, 510 titi 159 toadflax 321 Canada 321 Dalmatian 322 old-field 321 yellow 322 tobacco  213, 215–17, 221, 225, 346, 348, 388, 390–1, 393, 465, 491, 499, 513, 523–4, 526–31, 547, 550 tomato  215, 216, 348, 385, 389–92, 394, 412, 524, 527, 529–30 toothleaf goldeneye  245, 248 Tridens flavus  350, 501 Trifolium  362 trillium 401 Triosteum perfoliatum  319 trumpetweed 358 tuliptree  85, 412 tupelo  162, 302–3, 431, 486, 498 Turk’s cap  77, 233, 241 turnip  390, 491, 513, 527 turtlehead  320, 415 Vaccinium  471, 492 angustifolium  135, 318, 460 arboreum  134–5, 476 myrtilloides  460 pallidum  135, 318 see also blueberry and cranberry velvetbean 190 velvet leaf  76 Venus flytrap  535 Verbena  323, 371 hastata  415 see also vervain Verbesina  58, 340, 365, 414 alternifolia  340 encelioides  340, 361 Vernonia  402 missurica  415 noveboracensis  414 texana  362 Veronicastrum virginicum 416 vervain  221, 224, 323, 371 blue 415 vetch  70, 152 viburnum  183, 225, 301, 305, 326, 452, 461, 464, 480, 485–8, 498, 541, 546–7 arrowwood  183, 301, 537

mapleleaf 464 possumhaw  301, 464 Vigna luteola  190 Viguiera dentata  245, 248 vine, balloon  234 moonseed 79 snailseed 79 violet  454, 497, 546 Virginia chainfern  377 Virginia creeper  308–9, 326 Virginia jointvetch  346 Vitis labrusca  382 walnut  69, 111–12, 114, 116–17, 120, 164, 207–8, 253, 260, 272, 283, 326, 331, 343, 427, 449–50, 454, 458, 479, 481–2, 484, 486–7, 548 black  120, 208 see also butternut Waltheria  254 water hyacinth  418 waterleaf 488 watermelon 216 water-willow 412 wax myrtle  109, 259 wedelia 367 West Indian pinkroot  235 wheat  217, 385, 393, 407, 501, 505, 510, 524, 526, 528–9, 547 white snakeroot  368 white waterlily  419 wiegela 305 wild indigo  60, 153, 412–14 wild petunia  236 wild raisin  222, 301, 403, 464 willow  70, 73–4, 94, 123– 7, 149, 172–3, 188, 204, 206, 227, 229, 255, 260–2, 264, 268, 271, 283, 286, 292–3, 296–8, 305, 327, 344, 401, 421–4, 427–8, 430–1, 433–5, 441–3, 453, 465, 468, 470, 477–8, 481–4, 486–7, 489, 492–8, 503, 534, 537, 541, 550 black 123 weeping 123 willowherb  306, 310, 466 wingstem  58, 340, 365 winterberry  58, 305 wiregrass 152 wisteria 172

567

568 Foodplant Index witch hazel  48, 150, 184, 203, 264, 274, 283, 292, 433–4, 438, 444– 5, 455, 484–6, 489 wood sorrel  383

woodnettle  66, 68, 71, 214, 218, 454 woodrat nest  39 Woodwardia virginica  377, 416

woolly croton  245 wooly beachheather  151 Xanthocephalum dracun­ culoides 313

yankeeweed 358 yarrow  499, 518 yaupon 305 yellow buttons  359 yellow pond-lily  419

zinnia 346 Zizia aurea  416

Species and Subject Index Abablemma brimleyana  88 duomaculata 88 Abagrotis alternata  548 brunneipennis  550 barnesi 548, 550 magnicupida 549 orbis 548, 550 Abrostola  213 ovalis 214 urentis 214 Abrostolini  214 Abutilon Moth  76 Achatia distincta  476, 485, 486 Achatodes zeae  417 Acontia aprica  240 Acontiinae  238–48 Acronicta afflicta  275 albarufa 31, 259, 276, 277, 280 alni 264 americana 11, 259, 260 betulae 285, 286 brumosa 275 clarescens  261, 267 connecta  261 dactylina  258, 262, 263 dolli 259, 297, 299 exempta 277 exilis 276, 278, 279 falcula 259, 271, 297 fragilis 258, 268, 269, 271, 273 funeralis  31, 261, 264 furcifera  270 grisea 271, 273 haesitata 259, 276, 277, 279, 280, 281 haesitata-increta group 276–305 hamamelis 31, 261, 274 hasta 270 hastulifera 263 heitzmani 268 impleta 78, 258, 266, 283 impressa 292, 293, 300 inclara see A. increta increta 276, 277, 280, 281 increta “beech”  277 innotata 285, 286 interrupta 265 laetifica  286, 287, 288 lanceolaria  259, 297, 299 leporina  298 lepusculina 296, 298 lithospila  259, 272 lobeliae 36, 258, 259, 291 longa 264, 292, 293 modica 276, 278, 279

morula 284 near longa  259 noctivaga 294 oblinita 295, 299, 300 ovata 276, 280, 282 pruni 267 quadrata 265, 298 radcliffei 266, 269 retardata 289 rubricoma  296 sperata 298 spinigera 288 subochrea 274 superans 269 tristis  276, 277, 280 tritona 273 vinnula 257, 290 vulpina 298 Acronictinae  258–64 Actebia fennica  521 Actinotiini  399–400 Adita chionanthi  319 Afrida minuta  211 ydatodes 211 Afridinae  211 Agaristinae  306–10 Agate, Cherry  304 The 259, 304 Agnorisma badinodis  547 bugrai 547 Agrilus planipennis  319, 336 Agriopodes fallax  37, 301 Agrotis gladiaria  531 ipsilon 10, 520, 530 venerabilis 531 volubilis 532 Alabama argillacea  73, 78, 190 Alder Moth  264 Aletia oxygala  508 Allagrapha aerea  219 Allotria elonympha  162 Alypia langtoni  306 octomaculata 308 wittfeldii 308 Ambiguous Litter Moth 58 American Bird’s-wing Moth  397 American Ear Moth  408 American Ermine Moth 436 Amolita fessa  84 roseola 84 Amphipoea americana  408 interoceanica 408 velata 406 Amphipyra, Copper  325, 326 Mouse 327 Smooth 325

Amphipyra glabella  325 pyramidoides 12, 325, 326 tragopoginis 327 Amphipyrinae  324–38 Amyna axis  190, 234 bullula 234 octo 234 Anagrapha falcifera  225 Anaplectoides brunneomedia  541 prasina 541 pressus 541 Anarta trifolii  371, 491, 523 Anathix puta  464 Androloma disparata  306 maccullochii 306 Angle Shades  401, 402, 403 American  401 Brown  403, 461 Hop see Hops Angleshade Olive  402 Small  401 Anicla forbesi  522 illapsa 522, 523 infecta 523 Anisota senatoria  266 Anomis commoda  75 editrix 73 erosa 76 exacta 77 flava 76 impasta 77 luridula 77 privata 75 Anterastria teratophora 378 Anthocharis midea  307 Anticarsia gemmatalis  21, 190 Antitypina  470 Apamea devastator  407 ophiogramma 404 Apameini  404–17 Apharetra dentata  318 Aposematic  351, 397, 463 Apterona helix  327 Archanara subflava  408 Arches  491–503 Atlantic 497, 498 Black 494, 499 Bridled 511 Cloudy 492, 503 Disparaged 516 Explicit 512 Fluid 489 Grand 496 Hitched 493 Olive 511

Purple 492, 503 Speckled 497, 498 Splashed 497, 498 Stormy 492 Zebra 31, 495 Arctia caja  292 Argyrogramma verruca 213 Argyrogrammatini 213, 215, 216 Argyrostrotis anilis  160, 161 erasa 159 flavistriaria 159, 161 quadrifilaris 161 Arizona Straw  73 Armyworm  19, 504 Androgea 13, 386, 388, 390 Beet 385, 392 Bertha 502 Black 394 Caribbean 386, 388 Dolichos 390 Fall 385, 392, 393 Garden 394 Large Cotton  390 Lateral-lined 394 Orange-striped  390, 394, 395 Rice  386, 388 Roseate 501, 502 Southern 385, 388, 391 Sweetpotato 388, 390, 394, 395 Tropical 388 Unbarred 389 Velvet 394 Western Yellow-striped 395 Wheat 504 Wheat Head  501, 502 White-spotted 389 Yellow-striped 385, 390, 393, 394, 395 Armyworms  385–95, 501, 502, 504, 505, 523 Artace cribraria  291 Arugisa latiorella  87 lutea 87 Arzamini  418, 419 Ascalapha odorata  186 Athetis tarda  396 Atlantic Graphic Moth 151 Autographa ampla  222 biloba 221 jota 223 mappa 213, 223 precationis 212, 215, 224 rubidus 231

Bagisara buxea  232 near rectifascia  232 rectifascia 232, 233 repanda 233 tristicta  232 Bagisarinae  232–4 Baileya, Doubleday’s  207 Eyed  209 Pale 208 Sleeping 207, 208 Small  208 Baileya australis  208 dormitans 207, 208 doubledayi 207 levitans 208 ophthalmica 209 Baileyas  207–9 Baiting  19–22, 68, 99, 123, 176, 420, 452 Balsa, Many-dotted  256, 257 Three-lined 257 White-blotched  256, 257 Balsa labecula  256, 257 malana 256, 257 tristrigella  257 Balsinae  256–7 Banding, burlap  16, 99, 426 Bark rester  16, 98, 143, 276, 426, 452, 489 Basilodes chrysopis  340 pepita 340 Bats  10, 476, 505 Beating  17, 18 Beautiful Eutelia  194 Belay  46, 64, 198, 242, 279, 288, 308, 366 Belle, Lesser  94 Marsh 94 Bellura anoa  418 brehmei 418 densa 418 diffusa 419 gortynoides 418, 419 melanopyga 419 obliqua 418, 419 vulnifica 418, 419 Betrothed, The  105, 106 Biological control  11, 31, 318, 363 Bird-dropping Moth, Cotton  240 Curve-lined 240 Delightful 240 Exposed 240 Four-spotted 240, 241 Half-yellow 248 Olive-shaded 246, 247, 349 Small 244, 246, 247 Turk’s Cap  241

569

570 Species and Subject Index Bird-dropping moths 238–48 Birds  9, 10, 18, 26, 33, 39, 254, 326, 383, 469, 476 Black-barred Brown  339 Black Bit Moth  158 Black-bordered Lemon Moth  237 Black-dotted Brown  11, 143, 144, 145 Black-dotted Maliattha 236 Black-eyed Cropia  324 Black Wedge-spot  370 Black Witch  186 Black Zigzag  250 Bleptina caradrinalis  53 inferior 53 sangamonia 53 Bog Deltote  236 Bog Yellow-wing  502 Boletobiinae  89–90 Bomolocha abalienalis  65 baltimoralis 64 bijugalis 65 deceptalis 65 edictalis 66 madefactalis 69 manalis 65 palparia 63 sordidula 71 Boreal Fan Moth  330 Boreal Sprawler  330 Borer, American  408 American Crescented 408 Broken-lined 408 Burdock  411, 412 Cattail 419 Columbine 413 Dead-wood 85 Double Lobed  404 Elder Shoot  417 Emerald Ash  319, 336 Glassy 407 Green-washed Culm 405 Hop Vine  409 Indigo Stem  413 Iris 405 Marbled Minor Culm 405 Meadow Rue  410, 413 Newman’s 409 Northern Burdock  410, 411 Palmetto 189 Phragmites 409 Pickerelweed 418 Pitcherplant  410, 416 Reed Canarygrass  409

Seaside Goldenrod  412 Sensitive Fern  413 Stalk 410, 413 Strawberry Root  408 Wild Indigo  413 Yellow Sedge  408 Borers  404–17 Brachionycha borealis  330 Braconidae  27, 29, 30, 107, 114, 260, 300, 326, 364, 366, 385 Braun’s Quandry  87 Bravo, The  167 Bride, The  98, 112, 120 Brindle, American  424 Goldenrod 424 Variegated 214 Brocade, Adorable  472 Broken-lined 405 Newman’s 409 Toadflax 11, 321, 322 Wandering 470 Brocades  318–23 Broken-banded Y  222 Bronzed Turfworm  490 Bronzy Macrochilo  51 Brother, The  255 Abrupt 255 Browntail Moth  31 Burlap banding  16, 99, 426 Burnished Bog Moth  236 Cabbage Moth  478 Cadbury’s Lichen Moth 303 Cadbury’s Mystique  36, 162, 302, 303 Caenurgia chloropha  152, 154 crassiuscula 154 erechtea 152, 154 Callopistria cordata  375 floridensis 374, 375 granitosa 375 mollissima 375, 376 Calophasia lunula  11, 321, 322 Calpinae  79–81 Calyptra canadensis  79, 80 Camphorweed Cucullia 313 Campion Coronet  500 cannibalism  15, 25, 189, 192, 196, 212, 342, 343, 348, 349, 351, 392, 396, 426, 429, 437, 449, 468, 477 Capsula subflava  408 Caradrinini  378, 396 Catabena Moth  318 Catabena lineolata  323

Caterpillar, Black Olive 206 Cabbage Palm  11, 20, 189 Cattail 300 Cluster 388 Convict 355, 519 Lantana Stick  318 Smartweed 295 Striped Garden  499 Variable Climbing  543 Velvetbean 21, 190 Zebra  31, 495 Catocala abbreviatella  105 agrippina 117 alabamae 137, 139, 141 alabamae titania  141 amatrix 126 amestris 104, 124 amica 98, 128, 132, 140 andromedae 134, 135 angusi 99, 115, 116, 117 antinympha 107 atocala 117 badia 108, 109 badia coelebs  108, 109 blandula 140, 141 briseis 126 cara 98, 123, 128 carissima 123 cerogama 99, 100, 138 clintonii 139 coccinata 98, 129 coelebs 108 concumbens 124 connubialis 131 consors  98, 100, 110, 118 crataegi 136, 137, 139 dejecta 118 delilah 131 dulciola 139 epione 98, 100, 110 flebilis 111, 116 gracilis 134, 135 grisatra 142 grotiana 125 grynea 139 habilis 99, 115, 117, 119, 122 herodias 131 herodias gerhardi  129, 131 ilia 98, 130 illecta 105 innubens 105, 106 insolabilis 118 jair 128, 132 judith 99, 116, 119 junctura 100, 126, 153 lacrymosa 114, 119 lincolnana 142 lineella 128, 132

louiseae 134, 135 luctuosa 120 maestosa 100, 112, 120 marmorata 126 meskei 127 micronympha 133 minuta 105, 106 mira 136, 137 miranda 142 muliercula 108, 109 near lineella  132 nebulosa 121 neogama 98, 112, 120 nuptialis 104, 106 obscura 99, 113 orba 142 palaeogama 114 parta 127 piatrix 120 praeclara 140 pretiosa 140, 141 pretiosa texarkana  140 relicta 98, 125 residua 99, 113, 121 retecta 99, 116, 121 robinsonii 98, 99, 100, 111, 115, 116, 122 sappho 122 semirelicta 127 serena 99, 115, 116 similis 133 sordida 135 subnata 122 ulalume 122 ultronia 138 umbrosa 130 unijuga 127 vidua 98, 116 whitneyi  105, 106 Catocalini  97–142 Celastrina ladon  431 Celiptera frustulum  158 Ceramica picta  495 Cerastis tenebrifera  24, 534 Cerathosia tricolor  235 Cerma cerintha  304 cora 259, 304 Chaetaglaea cerata  459 sericea  457, 458, 459 tremula 459 Chamaeclea basiochrea 339 Charadra deridens  249, 469 Chloephorinae  204–6 Chlosyne harrisii  31 nycteis 31 Chocolate, Erasa  159 Four-lined 161 Short-lined 160, 161 Yellow-lined 159, 161

Choristoneura fractivittana 436 Chrysanympha formosa 227 Chrysodeixis includens 215, 224 Chytolita morbidalis  49, 50 petrealis  50 Chytonix palliatricula  384 sensilis 384 Cirrhophanus triangulifer 339 Cissusa spadix  11, 143, 144, 145 Classification  9, 32–4 Clouded Crimson  351, 355 Cobbler, The  367 Collecting  22–3 Collomeninae  210 Colobochyla interpuncta 94 salicalis  94 Colocasia flavicornis  252, 253 propinquilinea 252, 253 Comachara cadburyi  36, 162, 302, 303 Common Arugisa  87 Common Pinkband  365 Common Ptichodis  153 Compsilura 31 Concana, Barbados Cherry 210 Locust Berry  210 Concana mundissima  210 near mundissima  210 Condica albigera  364, 366 confederata 364 cupentia  364 mobilis 364 sutor 365, 367, 368 vecors 368 videns 368, 369 Condicinae  363–73 Confederate, The  364 Conservation  22–3 Consort, The  98, 100, 110, 118 Copivaleria grotei  336 Coranarta luteola  502 Corn Earworm  11, 21, 345, 348, 393 Coronet, Campion  500 Cosmia calami  467 trapezina 467 Cotton Bollworm  349 Cotton Leafworm  190

Species and Subject Index Cotton Moth  78 Coxina cinctipalpis  166 Crambodes talidiformis 371 Crocigrapha normani  484, 489 Cropia connecta  324 ruthaea 324 templada  324 Crymodes devastator  407 Cryptocala acadiensis  521 Ctenoplusia oxygramma 213 Cucullia alfarata  311, 313 asteroides  311, 312, 314 convexipennis 315 florea 311, 312 intermedia 316 laetifica  313 omissa 311, 312 postera 311, 312 speyeri  311, 317 Cuculliinae  311–17 Cutina albopunctella  165 distincta  164, 165 Cutworm, Army  10, 12 Black 520, 530 Black Army  521 Bristly 513 Bronzed 490 Clay-backed 531 Clover 491 Darksided 527 Dingy 529, 533 Dusky 531 False 423 Glassy 407 Granulate 528 Greasy 530 Greater Spotted  545 Green  523 Mottled Gray  548 Sand 526 Sandhill 16, 526 Spotted 545 Spotted-sided 547 Strawberry  408 Striped 527, 532 Tobacco 388 Variegated 520, 524 Well-marked 548, 550 Western Bean  525 W-marked 540 White Pine  542 Cutworms  10, 11, 13, 16, 17, 21, 32, 37, 115, 378, 396, 403, 404, 422, 449, 452, 462, 465, 472, 491, 513, 514, 515, 520–51 Cydosia nobilitella  235

Dagger, Afflicted  275 American 11, 259, 260 Bantam Maple  289 Barrens  31, 259, 276, 277 Beech 277, 281 Charred 275 Cherry 270 Clear 261, 267 Connected 261 Cottonwood 296, 298 Delightful 257, 290 Doll’s 259, 297, 299 Eclipsed Oak  276, 277, 281 Epauleted Oak  280, 282 Exiled 278 Fingered 258, 262, 263 Fragile 258, 268, 269, 271, 273 Frosted 263 Funerary 264 Gray  271, 273 Greater Oak  36, 258, 259, 291 Hackberry 296 Hazel 259, 271, 297 Heitzman’s 268 Hesitant  259, 276, 277, 279, 280, 281 Hoary Alder  263 Hopeful 298 Impressive 292, 293, 300 Interrupted 265 Lanceolate 297 Lesser Cottonwood  298 Lesser Oak  276, 278, 279 Lobelia 291 Long-winged 259, 264, 292, 293 Marsh 300 Medium Oak  276, 278, 279, 280 Miller 298 Narrow-winged 259, 297, 299 Night-wandering 294 Nondescript 288 Oak group  276–305 Ochre 284 Ovate 282 Paddle 31, 261, 264 Pleasant 286, 287, 288 Quadrate 298 Radcliffe’s 266, 269 Retarded 289 River Birch  285, 286 Ruddy 296 Sad 276, 277, 280 Smeared 295, 299, 300

Speared 270 Splendid 268, 269 Square-eyed 265, 298 Streaked 259, 272 Triton 273 Unmarked 285, 286 Witch Hazel  31, 261, 274 Yellow-haired  78, 258, 266, 283 Danaus plexippus  355 Dargida diffusa  501, 502 rubripennis  501, 502 Darkwing, Consobrina’s 163 Gordonia 163 Smith’s 163 Dart, Adobe  533 Attentive 550 Bent-lined 529 Boomerang 551 Brick 537 Brown-collared 550 Catocaline 521 Clandestine 540 Conglomerated 532 Daggered Heath  550 Dappled 541 Double 537 Dull Reddish  544 Early Conifer  521, 542 Fawn Brown  532 Finland 521 Flame-shouldered 535 Fleece-winged 533 Forbes’s  522 Great Brown  541 Greater Black-lettered 545 Greater Red  548 Great Gray  541 Green Arches  541 Inclined 532 Ipsilon 530 Knee-joint 533 Large Yellow-winged 520, 538 Lesser Yellow-winged 538 Master’s 529 Norman’s 551 Northern Pine  542 Northern Variable  542 One-dotted 549 Pale-banded 547 Pink-spotted 551 Praevia 521 Reaper 527 Reddish Heath  544, 551 Reddish Speckled  24, 534 Relict Prairie  521

Rubbed 526 Smaller Pinkish  539 Smith’s 546, 551 Snowy 522, 523 Southern Pine  542, 543 Southern Variable  543 Subgothic 529 Sundew 536 Subterranean 528 Switchgrass 532 Tessellate 532 Tricose 529, 533 Venerable 531 Venus Flytrap  535, 536 Voluble 532 White-marked 534 White Pine  521, 542, 543 Yankee 550 Young’s 544, 551 Darter, The  143, 146, 147 Darts  520–51 Dasychira  78, 283 Datana  266, 398 Deer  150, 319, 320 Defoliation  11, 17, 78, 143, 144, 145, 146, 147, 148, 206, 234, 310, 329, 398, 428, 478, 505, 524 Deltote bellicula  236 Diachrysia aereoides  219 balluca 218 Diarsia jucunda  539 Diastema tigris  363 Dichagyris acclivis  532 reliqua 521 Dicrocheles phalaenodectes 505 Dilobinae  255 Diphthera festiva  254 Diphtherinae  254 Dipterygiini  397, 398 Discestra trifolii  491 Diver, Cattail  418, 419 Pickerelweed 418 White-tailed 419 White Waterlily  419 Divers  418–19 Doryodes bistrialis  152 grandipennis 152 spadaria 152 Dot-lined White  291 Drasteria, Blueberry  143 False Heather  151 Figure-seven 150 Drasteria atlantica  151 grandirena 150 graphica 151 occulta 143 Dun-bar  467 American 467

Dusky Silver Y  228 Dyar’s Lichen Moth  211 Dypterygia rozmani  397 Dypterygiini  397, 398 Ears (moth)  476, 505 Egira alternans  479, 481, 482 dolosa 483 Eight-spot  190, 234 Elaphria alapallida  380 cornutinus 380 festivoides 380 georgei 381 grata 380, 381 nucicolora 380, 381 versicolor 380, 382 Elaphriini  380–4 Elves  192–7 Emarginea percara  324 Emerald Ash Borer  319, 336 Enargia  468 decolor 11, 468 fausta 468 infumata 468 mephisto 468 Eosphoropteryx thyatyroides  231 Epiglaea apiata  460 decliva  403, 461 Erebidae  38–191 Erebinae  97–186 Eriopinae  374–7 Eriopygini  511–18 Erythroecia hebardi  342 Esther Moth  180 Euagrotis illapsa  522 Eucirroedia pampina  456 Euclidia cuspidea  153, 154 Euclidiini  152–8 Eudryas grata  309, 310 unio 309, 310 Eueretagrotis attentus  550 Eulepidotinae  187–91 Euparthenos nubilis  185 Euplectrus  27, 29, 385, 394 Euplexia benesimilis  401 lucipara 401 Euproctis chrysorrhoea  31 Eupsilia  447 cirripalea 447, 450, 451, 454 devia 425, 447, 448, 450 morrisoni 447, 452, 454 new species  451 sidus 449 transversa 449 tristigmata  447, 453 vinulenta 447, 450, 452, 454

571

572 Species and Subject Index Eurois astricta  541 occulta 541 Eusceptis flavifimbriata 241 Eustrotiinae  236–7 Eutelia adulatrix  194 pulcherrimus 194 Euteliidae  192–7 Euxoa auxiliaris  10, 12 detersa 16, 526 messoria 527 pleuritica  532 tessellata 527, 532 velleripennis  533 Exyra fax  220 ridingsii 220, 221 semicrocea  220 Eyespots (false eyes)  46, 55, 83, 163, 366, 390, 401 Fagitana littera  227 Falcate Orange Tip  307 False head  50, 309, 329, 366, 368, 396, 401, 422, 492, 516, 518 Fan-foot, Angulate  39 Slant-lined 52 Streaked 51 Faronta diffusa  501 Feculae  59, 67, 69, 76, 98, 147, 150, 220, 232, 233, 237, 239, 258, 351, 417, 456 Feltia geniculata  533 herilis 529 jaculifera 529, 533 manifesta  533 subgothica 529 subterranea 528 tricosa 529, 533 Feralia comstocki  337, 338 jocosa 337, 338 major 337, 338, 439 Fern Moth, Florida  374, 375 Marsh 377 Pink-shaded  375, 376 Silver-spotted 375 Filament Bearer  191 Fishia illocata  470, 472 Flame Shoulder  535 Flies see Tachinidae Florida Psychomorpha 307 Florida Tetanolita  52 Flower Moth, Amorpha 359 Arcigera 353 Bleeding 352 Brown  360

Camphorweed 360 Evening Primrose  354 False Boneset  344, 359 False Indigo  359 Fuller’s 359 Goldenrod 356 Impatient 352 Iva  359 Lynx 360 Matutinal 360 Northern 351 Ragweed 357 Slender  359 Thoreau’s 352, 357 Three-lined 358 Foodplants see Foodplant Index Forester, Disparate  306 Eight-spotted 308 Langton’s 306 MacCulloch’s 306 Paddled 307 Wittfeld’s 308 Fruitpiercer, Citrus  81 Moonseed 79 Fruitworm, Green  427 Speckled Green  422 Tomato 349 Fungus Moth, Black  89 Common 90 Four-spotted 89 White-lined 89 Galgula partita  383 Garella nilotica  206 Glaeas  456–67 Glottulini  519 Gluphisia septentrionis 447 Gold Moth  340 Goldenrod Stowaway  339 Gondysia consobrina  163 similis 163 smithii 163 telma 163 Gonodonta nutrix  81 sinaldus 79 unica 81 Grapevine Epimenis  307, 309 Graphiphora augur  537 haruspica  537 Grass Moth, Brass-dotted 236, 238 Common 152 Feeble 84 Large Mossy  238 Range 154 Spotted 72 Veiled 409 Gray Half-spot  399

Gray Sword Moth  422 Graylet, Black-patched  95, 96 Dotted 95, 96 White-lined 95, 96 Yellow-spotted 95, 96 Great Tiger Moth  292 Green Cloverworm  70 Greenbriers  328–9 Griffin  325, 474 Groundcat, Bridled  511 Bristly 513 Cynical 518 Disparaged 516 Explicit 512 Galena 515 Implicit 512 Olive 511, 513 Ruddy 514 Rustic  383, 396, 517 Sheathed 515 Signate 511 Small Brown  518 Groundling, Boneset  364, 366 Cobbler 365, 367, 368 Common Pinkband 365, 367 Confederate  364 Dusky 368 Lantana 363 Mobile 364 Smartweed 370 Splotched 364 Verbena  371 White-dotted 368, 369 Yellow Water-lily  370 Gypsy Moth  11, 31, 111, 128, 480 Hadena ectypa  500 Hadenini  491–503 Halfling, Brimley’s  88 Double-eyed 88 Fetching 87 Halflings  87–8 Harris’s Three-spot  258, 304, 305 Harrisimemna trisignata 258, 304, 305 Hawkmoth  11, 21, 31, 311, 315 Hebrew, The  36, 162, 259, 302, 303 Helicoverpa zea  10, 11, 21, 345, 348, 393 Heliocheilus lupatus  350 paradoxus 350 Heliothinae  342–50 Heliothis  10, 345 acesias 345 borealis 345

ononis 345 phloxiphaga 347 subflexa 346, 347 virescens 11, 345, 346, 347, 348 Helotropha reniformis  408 Hemeroplanis scopulepes 92 Hemipachnobia monochromatea  536 subporphyrea 535, 536 Henry’s Marsh Moth  300 Heptagrotis phyllophora 537 Herald, The  74 Herminiinae  38–60 Hieroglyphic Moth  254 Himella fidelis  485, 486 intractata 485 Homoglaea hircina  425, 455, 457 Homohadena badistriga 318 Homophoberia apicosa  370 cristata 370 Hooked Silver Y  327 Hops Angleshade  474 Hydraecia immanis  409 Hypagyrtis esther  180 Hypena abalienalis  65 appalachiensis 63 atomaria 63 baltimoralis 64 bijugalis 65 deceptalis 65 edictalis 66, 68, 71 eductalis 67 humuli 66, 68, 71 madefactalis 69 manalis 65 minualis 63, 233 palparia 63 proboscidalis 66 scabra 10, 64, 68, 70, 94, 233 sordidula 66, 68, 71 Hypeninae  63–71 Hypenodinae  87–8 Hypenula cacuminalis  39 Hypermetamorphosis 36, 55 Hyperstrotia aetheria  96 flaviguttata 95, 96 nana 96 pervertens  95, 96 secta 95, 96 villificans 95, 96 Hypocalinae  82–3 Hyppa, Common  466 Summer 466 Hyppa contrasta  466 xylinoides 466

Hypsoropha hormos  82 monilis 82, 83 Ichneumonidae  27, 29, 30, 313 Idia, American  39, 40, 42 Common 40, 42 Forbes’s  43 Glossy Black  41 Gopher Tortoise  15, 43 Julia’s 43 Laurentine 41, 44 Major 15, 39 Orange-spotted 43 Pale-winged 40, 42 Rotund 43, 44 Smoky 44 Toothed 41, 42 Idia aemula  40, 42 americalis 39, 40, 42 concisa 40, 42 denticulalis  41, 42 diminuendis 43 forbesii 43 gopheri 15, 39, 43 julia 43 laurentii 41, 44 lubricalis 41 majoralis 15, 39 new species  40, 42 rotundalis 43, 44 scobialis 44 Introduced species  10, 11, 75, 318, 322, 327, 392, 538 Iodopepla u-album  400 Ipimorpha pleonectusa  469 subtusa 469 Isogona snowi  84, 86 tenuis 86 Joker, The  338 Jocheara 264 Lacanobia atlantica  497, 498 grandis 496 subjuncta 497, 498 Lacinipolia explicata  512 implicata 512 lorea 511 olivacea  511, 513 renigera 513 Lantana Moth  363 Lascoria ambigualis  58 Lateroligia ophiogramma 404 Latex 11, 196, 212, 215, 216, 224 Laugher, The  249, 469 Leadplant Moth  359

Species and Subject Index Leaf shelters  9, 198, 202, 203, 249, 272, 278, 307, 330, 335, 436, 447, 448, 467, 469 Leafworm, African Cotton  385 Brown Cotton  240 Cotton 73, 78, 190 Egyptian Cotton  385, 386 Oriental 388 Ledaea perditalis  62 Lempke’s Gold Spot  230 Lepipolys perscripta  321 Lesmone detrahens  167 Lesser underwings  143–51 Leucania  476 adjuta 506 commoides 508 insueta 508 lapidaria  508 linda 509 linita 504, 507 multilinea 508 phragmitidicola 509 pseudargyria  510 scirpicola  509 subpunctata 509 ursula 509, 510 Leuconycta, Green  372, 373 Marbled-green 373 Leuconycta diphteroides 372, 373 lepidula  373 Lithacodia muscosula  238 Lithomoia germana  424 solidaginis 424 Lithophane  426 abita 443 adipel 439 amanda 430, 441 antennata 427, 428, 429, 431 baileyi 432, 436, 441 bethunei  420, 433 disposita  441 fagina 426, 441 georgii 430 grotei  332, 420, 428, 429 hemina 420, 434, 435, 436, 437 hepatica  441 innominata 426, 434, 435, 436, 437 joannis 436 lanei 434 laticinerea 427, 428, 429 lemmeri  426, 443 lepida 337, 439 oriunda  426

patefacta 420, 437 petulca 437, 442 pexata 440 pruena 432 querquera 426, 438, 444 scottae 438 semiusta 426 signosa 442 tepida 432, 442 thaxteri 443 thujae 443 unimoda 431 viridipallens  432 Litoprosopus futilis  11, 20, 189 Little Nymph  133 Little Wife, The  108, 109 Live Oak Metria  166 Lomanaltes eductalis  67 Long-winged Doryodes 152 Looper, Abstruse  226 Beautiful 227 Bilobed 212, 221, 224 Boreal 228 Brown-patched 222 Cabbage 216 Celery 225 Chosen 229 Climbing 222 Clover 154 Common 212, 215, 224 Connected 230, 231 False Hemlock  382 Forage 152, 154 Golden 213, 229 Gray 212, 217 Gray Cypress  165 Green-patched 218 Green-sigma 226, 229 Inscribed 227, 229 Labrador Tea  228 Large 222 Large-spotted 230 Lined Copper  219 Little Bride  227 Maple 164 Nettle 214 Pale Shoulder Pitcherplant  220 Pink-patched 231 Pirate 227 Pitcherplant 220 Plantain 224 Putnam’s 230 Raspberry 222 Riding’s Pitcherplant 220, 221 Salt and Pepper  226, 229 Sharp-stigma 213 Soybean 215, 224

Splendid Brass  218 Spruce 226, 227 Straight-lined  231 Striped Grass  155, 156, 157 Unspotted 219 Variegated Violet  231 Vetch  152, 154 Loopers  212–13, 217–31 golden 215–16 grass 152–8 nettle 214 Loscopia velata  406 Loxagrotis albicosta  525 Lycophotia phyllophora 537 Lymantria dispar  11, 31, 111, 128, 480 Macrochilo absorptalis  52 litophora 39 orciferalis 51 Macronoctua onusta  405 Magusa divaricata  397 orbifera 397, 398 Maliattha synochitis  236, 238 Mamestra brassicae  478 configurata 502 Many-dotted Appleworm 257 Marathyssa, Dark  192, 193 Light 193 Marathyssa basalis  193 inficita 192, 193 Marimatha nigrofimbria 237 Marvel, Barrens  384 Cloaked 384 Gray  378 Green  37, 301 Pink-barred 238, 349, 379 Megalographa biloba  212, 221, 224 Meganola, Ashy  199 Coastal Plain  200 Meganola minuscula  198 new species  199 phylla 200 spodia 198, 199, 200 Melanchra adjuncta  493 assimilis 494, 499 picta 31, 495 pulverulenta 494 Melanism  78, 212, 261, 277, 281, 337, 490 Melipotini  143–51 Melipotis, Bookend  146 Indomitable 143, 148 Merry  145, 146, 149

Melipotis  146 acontioides  143, 146, 147 cellaris 146 indomita 143, 148 jucunda 145, 146, 149 Merolonche dolli  299 Meropleon ambifusca  409 Mesapamea fractilinea  405 Metalectra albilinea  89 discalis 90 quadrisignata 89 tantillus 89 Metamorphosis 36 Metaxaglaea australis  462, 464 inulta 462, 464 semitaria  462, 464 viatica 462 violacea 463 Metria amella  166 Midget, Festive  380 George’s 381 Grateful 380, 381 Northern Festive  380 Pale-winged 380 Pink-barred 379 Sugarcane 380, 381 Variegated 380, 382 Mimicry  78, 83, 86, 111, 122, 130, 172, 191, 193, 258, 264, 266, 283, 292, 296, 309, 339, 355, 426, 446 Batesian 283, 355 Mullerian 283 Miranda Moth  378 Mistletoe Marblelet  324 Mites 33, 39, 206, 505 Mitter, Charles  32, 33 Mocis, Small  155 Texas 155, 157 Withered 155, 156 Yellow 153, 155 Mocis disseverans  153, 155 latipes 155, 156, 157 marcida 155, 156 texana 155, 157 Monarch  355 Moon-lined Moth  168 Moonseed Moth  79 Morrisonia confusa  280, 476, 487 evicta 488 latex 489 mucens 476 Moths, bird-dropping see Bird-dropping moths fern  374–7 flower see Flower moths fruitpiercing 79–81 fungus 89–90 grass see Grass moths

litter 38–60 necklace 82–3 nettle see Nettle moths oak see Oak moths resplendent 235 scalloped 73–8 Spanish 519 St. Johnswort  399–400 sun 342–50 swordgrass 421–3 underwing 97–142 Motya abseuzalis  210 Multi-lined Tyrissa  167 Mythimna  10, 11, 505 oxygala 508 segregata 505 sequax  504 unipuncta 19, 504, 505, 523 Narrow-wing, Divaricate 397 Orbed 397, 398 Natural enemies  27 Necklace Moth, Large  82, 83 Small  82 Nedra ramosula  399, 400 Nemoria tuscarora  399 Neogalea sunia  318 Nephelodes minians  490 Nepytia canosaria  382 Nest (inquilines)  39 Nettle Moth, Speckled  214 White-shouldered 214 Nigetia formosalis  87 Niphonyx segregata  474 Noctua comes  538 pronuba 520, 538 Noctuidae  212–551 Noctuinae  378–551 Noctuini  520–51 Nola, Blurry-patched  198, 201 Sharp-blotched 17, 198, 199, 201 Sorghum 203 Sweet Pepperbush  202 Three-spotted 203 Nola cereella  203 cilicoides 198, 201 clethrae 198, 202 ovilla 200 pustulata 17, 198, 199, 201 sorghiella  203 triquetrana  203 Nolidae  198–211 Nolinae  198–200 Nomenclature 11, 33 Nutmeg, The  371, 491, 533

573

574 Species and Subject Index Nycteola, Ash-colored  204 Frigid 204, 205 Sienna-dotted 204 Nycteola cinereana  204 frigidana 204, 205 metaspilella 204 revayana 205 Oak Moth, Barrens  143, 144 Common 11, 143, 144, 145, 278 Oak Nycteoline  205 Ochropleura implecta  535 plecta 535 Ogdoconta cinereola  365, 367 Old Maid, The  108, 109 Oligia chlorostigma  405 illocata 470 mactata 472 strigilis  405 Olive, The  469 Ommatochila mundula  70 Oncocnemidinae  318–23 Oncocnemis saundersiana 320 Operophtera brumata  467 Ophiuche minualis  63 Ophiusini  166–85 Orange-striped Oakworm 266 Orgyia definita  296 Orthodes crenulata  517 cynica 518 majuscula 383, 396, 517 “Orthodes” detracta  516 Orthosia alurina  477, 478, 479 garmani 18, 479 hibisci 422, 477, 480 revicta 481, 482 rubescens  480 Orthosiini  476–89 Outbreaks  11, 17, 31, 73, 78, 143, 144, 145, 146, 147, 148, 189, 190, 206, 234, 260, 310, 386, 394, 398, 428, 478, 505, 524, 541, 546 Overbrowsing  319, 320 Owlet, Ash-colored  204 Bent-winged 53 Black-waved 166 Brown-hooded 315 Brown-lined 39 Buttonbush 62 Canadian 80 Curve-lined 191 Dark-banded 44 Decorated  61

Detracted 167 Distinguished Cypress 164, 165 Goldenrod Hooded  314 Long-horned 39 Lost 62 Meadow Rue  79, 80 Morbid  49, 50 Slant-lined 52 Snow’s 84, 86 Stone-winged 50 Thin-lined 86 Thin-winged 87 Owlets, hooded  311–17 wavy-lined 166–85 Oxycilla tripla  73 Pachypolia atricornis  470 Paectes, Blue-gray  196, 197 Clouded 195 Eyed  196, 197 Large 195, 197 Pygmy  196, 197 Paectes abrostolella  196 abrostoloides 195, 197 near pygmaea  196, 197 nubifera 195 oculatrix 196, 197 pygmaea 196, 197 Paint, Asteroid  311, 312, 314 Calico 315 Dashed 316 Halloween 311, 313 Overlooked 311, 312 Rusted 311, 312 Speyer’s 311, 317 Unrusted 311, 312 Pale-edged Selenisa  167 Palthis, Dark-spotted  59, 60 Faint-spotted 59, 60 Palthis angulalis  59, 60 asopialis 59, 60 Pangrapta decoralis  61 Pangraptinae  61–2 Panopoda, Hickory  188 Orange 187 Red-lined 188 Panopoda carneicosta  188 repanda 187 rufimargo 188 Panthea, Eastern  251 Tufted Pine  250, 251 Tufted Spruce  250 Panthea acronyctoides  250 furcilla 250, 251 pallescens  251 Pantheinae  249–53 Papaipema  18, 410 appassionata  410, 416

araliae 410, 414 arctivorens 410, 411 astuta 414 baptisiae 413 beeriana 410 cataphracta 411, 412 cerina 414 cerussata 411 duovata 412 duplicatus 414 furcata 410 inquaesita 413 leucostigma  413 nebris 410, 413 rutila  413 unimoda 410, 413 Parallelia bistriaris  164 Parasitoids  27, 28, 29 see also Braconidae, Ichneumonidae, Tachinidae Penitent, The  120 Peridroma saucia  10, 520, 524 Phalaenophana extremalis 44 pyramusalis 44 Phalaenostola, Blackbanded  52 Dark 52 Shadowy 52 Phalaenostola eumelusalis 52 larentioides 52 Pheromones 23, 326, 393, 475, 505, 529 Phlogophora iris  402 periculosa 403, 461 Phlogophorini  401–3 Phoberia atomaris  11, 143, 144, 145, 278 ingenua 143, 144 Phosphila, Spotted  328 Turbulent 266, 329 Phosphila miselioides  328 turbulenta 266, 329 Phylogeny  9, 32, 33 Phyprosopus callitrichoides 191 Phytometra rhodarialis  93 Phytometrinae  91–6 Pinion, Alder  440 Amanda’s 430, 441 Ashen  427, 428, 429, 431 Bailey’s 432, 436, 441 Bald Cypress  443 Bethune’s 420, 433 Branded 420, 437 Brown 420, 434, 435, 436, 437 Buckeye 436 Canadian Cedar  443

Dashed Gray  441 Dowdy  431 European Pale  441 George’s 430 Grote’s 332, 420, 426, 428, 429 Hoary 426, 441 Lemmer’s 426, 443 Nameless 426, 434, 435, 436, 437 Pale Green  432 Pine 337, 439 Russet-spotted 427, 428, 429 Shivering 426, 438, 444 Sycamore 442 Thaxter’s 443 Wanton 437, 442 White-eyed 432, 442 Pink-border Yellow  93 Pink Sedge Moth  84 Pitcher Plant Moth  220 Plagiomimicus pityochromus  339 Plain Golden Y  223 Plathypena scabra  70 Platypolia anceps  471 “Platypolia” mactata  472 Plusia  212 contexta 230, 231 magnimacula 230 putnami 230 venusta 230 Plusiinae  212–31 Plusiini  217–31 Plusiodonta compressipalpis 79 Poaphilini  159–65 Pococera robustella  59 Polia detracta  516 imbrifera 492, 503 latex 489 nimbosa 492 purpurissata 492, 503 Pollen predation  239, 242, 246 Pollination  10, 500 Polygrammate hebraeicum 36, 162, 259, 302, 303 Polyphenism  73, 191, 217, 234, 250, 376, 377, 379, 386, 478, 505, 542 Ponometia candefacta  246, 247, 349 erastrioides  244, 246, 247 semiflava 248 Predatory owlets  15, 87, 330, 349, 420, 426, 427, 429, 431, 432, 433, 437, 438, 449 see also cannibalism

pollen  239, 242, 246 seed  11, 15, 60, 236, 321, 339, 340, 342, 343, 346, 349, 350, 351, 353, 354, 356, 357, 369, 379, 393, 462, 465, 491, 499-501, 532 Primrose Moth  354 Prodeniini  385–95 Protodeltote albidula  238 muscosula 238 Protolampra brunneicollis 550 Protorthodes oviduca  514 Proxenus miranda  378 Psaphida electilis  331 grandis 330, 334 Psaphida resumens  332 rolandi 333, 335 styracis 334, 335 thaxterianus 335, 469 Psaphidini  330–8 Psectraglaea carnosa  456 Psectrotarsia hebardi  342 Pseudaletia unipuncta  504 Pseudeustrotia carneola 238, 349, 379 Pseudeustrotiini  378–9 Pseudeva purpurigera  231 Pseudohermonassa bicarnea  551 Pseudorthodes vecors  518 Psychomorpha epimenis 307, 309 epimenis euryrhoda  307 Ptichodis herbarum  153 Puddling  239, 307, 447 Punkie, Buttonwood  210 Confused Oak  198 Gulf Lichen  211 Live Oak  199 Merchant 206 Once-charred 198, 199 Two-spotted Oak  199, 200 Punkies  198–203 lichen 211 Pyreferra ceromatica  444, 445, 446 citrombra 443, 444, 445 hesperidago 426, 438, 444, 445 moffatiana 443, 444, 445 pettiti  446 Pyrrhia adela  343 aurantiago 342 cilisca 343, 344 exprimens  343, 344 umbra 343

Species and Subject Index Quadrifine (quadrifids) 32, 38, 212 Quaker, Cynical  518 Distinct 476, 485, 486 Garman’s  18, 479 Gray  477, 478, 479 Intractable 485, 486 Norman’s 484, 489 Ruby 480 Ruddy 514 Rustic 517 Sheathed  515 Signate 511 Small Brown  518 Speckled Green  477, 480 Subdued  481, 482 Quakers  476–89 Rachiplusia ou  212, 217 Raphia abrupta  255 frater 255 Redectis vitrea  57 Regal Cydosia  235 Renia, Chocolate  57 Dark-banded 55 Discolored 54 Dotted 56 Even-lined 57 Speckled  57 Undulate 38, 55 Yellow-spotted 57 Renia adspergillus  57 discoloralis 54 factiosalis  38, 55 flavipunctalis 57 nemoralis 57 salusalis 56 Rhizedra lutosa  409 Rhodoecia aurantiago  342 Risobinae  207–9 Rivula propinqualis  72 pusilla 72 sericealis  72 stepheni 72 Rivulinae  72–3 Rosewing, The  503 Sallow, Adorable  472 American Bordered  343 Anointed  444, 445, 446 Arcadian  473, 474 Barrens 449 Bicolored  454, 464, 465, 523 Blueberry 318 Bordered  343 Broad  473 Brown-lined 318 Checkered  471, 472 Chosen 331

Citrine 443, 444, 445 Comstock’s 337, 338 Connecticut 438 Cranberry 460 Crimson 456 Even-lined 469 Fawn  334, 335 Figure-eight 332 Five-lined 323 Footpath 462, 464 Franclemont’s 447, 450, 454 Fringe-tree 319 Goat 425, 455, 457 Gray  330, 334 Gray-dusted 471 Grote’s 336 Holly 463 Jocose 337, 338 Lost 425, 447, 448, 450 Major 337 Morrison’s 447, 452, 454 Mustard 426, 438, 444, 445 Orange 342 Pettit’s 446 Pine 337, 338, 439 Poplar Catkin  464 Red-winged 420, 425, 432, 455 Roadside 462 Roland’s 333, 335 Saunders’s 320 Scalloped 456 Scribbled 321, 322 Shield-marked 465 Silky 457, 458, 459 Sloping 403, 461 Southern 462, 464 Straight-toothed  447, 450, 452, 454 Thaxter’s 335, 469 Three-horned 470 Three-spotted 447, 453 Trembling 459 Variable 457, 458 Viburnum 462, 464 Wandering 470, 472 Sallows  318–23, 378, 420–73 antitypine 470 bearded 331–6 eupsilia 447 lash-eyed 330, 378 mossy 337–8 Satellite  449 Saturniidae 31 Satyr  474, 475 Scalloped Moth, Grayeyed  77

Gulf 73 Hibiscus 75 Straight-edged  77 White-pupiled 76 Windowed 77 Yellow 76 Schinia  351–63 arcigera 353 florida 354 fulleri 359 gaurae 351, 355 gracilenta 359 grandimedia 344, 359 lucens 359 lynx 360 mitis 360 nubila 360 nundina 356 petulans 352 rivulosa 357 sanguinea 352 saturata 360 septentrionalis  351 snowi 351 thoreaui 352, 357 trifascia  358 Scolecocampa liburna  85 Scolecocampinae  84–6 Scoliopteryginae  73–8 Scoliopteryx libatrix  74 Seed predation  11, 236, 339, 351, 361, 362, 363, 500 Seedcopper, Beggarticks 339 Crownbeard 340 Gilded 340 Malvastrum 339 Ragweed 339 Sunflower 341 Selenisa monotropa  167 sueroides 166, 167 Sericaglaea signata  457, 458 Shelter 15, 57, 198, 200, 202, 203, 205, 249, 252, 253, 276, 278, 280, 306, 307, 330, 331, 332, 334, 335, 351, 361, 363, 418, 425, 436, 447, 448, 467, 468, 469, 476, 477, 483, 486, 487 Sideridis rosea  503 Sigela brauneata  87 Simyra henrici  300 insularis 300 Six-spotted Gray  91 Skeletonize  38, 40, 50, 95, 187, 203, 258, 276 Sleeving  26 Slowpoke, The  396

Snout, Alder  67 Baltimore 64 Black 10, 64, 68, 70, 94, 233 Deceptive 65 Dimorphic 65 Flowing-line 65 Gray-edged 69 Hop Vine  66, 68, 71 Large 66, 68, 71 Mallow 63, 233 Sordid 66, 68, 71 The 66 Variable 53 Variegated 63 White-lined 65 Sodium 447 Sorghum Headworm  349 Sorghum Webworm  203 Soybean Podworm  349 Spaelotis clandestina  540 Spanish Moth  519 Spargaloma sexpunctata  91 Spermatophore  420, 447, 450, 454 Sphingidae  10, 21, 31, 311, 315 Spiloloma lunilinea  168 Spiramater lutra  497, 498 Spodoptera  10, 11, 385–95 albula 389 androgea 13, 386, 388, 390 dolichos 388, 390, 394, 395 eridania 385, 388, 391 exigua 385, 392 frugiperda  385, 392, 393 latifascia  390, 394, 395 littoralis 385, 386 litura 386, 388 ornithogalli 385, 390, 393, 394, 395 praefica 395 pulchella 386, 388 sunia 389 Spragueia, Black-dotted 243, 244, 245 Common 243, 244, 246, 247 Confounding 245 Southern 242, 243, 244 Spotted 239, 245 Yellow 242, 243 Spragueia apicalis  242, 243 dama 242, 243, 244 guttata 239, 245 jaguaralis 245 leo 243, 244, 246, 247 onagrus 243, 244, 245 Spring Azure  431

St. Johnswort Moth, Burgundy  400 Gray  399, 400 Stephen’s Gem  221 Stiria rugifrons  341 Stiriinae  339–41 Straw Dot  72 Straws  72–3 Striacosta albicosta  525 Sugaring  19, 452 see also baiting Sun Moth, False Foxglove 342 Ground Cherry  346, 347 Horse-balm 342 Noble  343, 344 Purple-lined 343, 344 Purple-top 350 Spotted Straw  347 Sunira bicolorago  454, 464, 465, 523 Sutyna privata  471, 472 Sweeping  17, 18 Sweetheart, The  126 Swordgrass Moth, Acadian 421 American 422, 423 Dot-and-dash 422 Sympistis badistriga  318 chionanthi 319 dentata 318 perscripta  321, 322 piffardi  320 riparia 320 saundersiana 320 Syngrapha abstrusa  226 alias 226, 227 epigaea  227, 229 interrogationis 229 microgamma 227 montana 228 octoscripta  228 rectangula 226, 229 selecta  229 u-aureum 229 viridisigma 226, 229 Tachinidae 29–31, 114, 148, 265, 294, 313, 315, 316, 327, 349 Tailpipes 166, 167 Tarache aprica  240 dacia  240 delecta  240 terminimaculata 240 tetragona 240, 241 Tentmaker, Blackcheeked 469 Smoky 468 Straw-eyed 11, 468 Tawny  468

575

576 Species and Subject Index Tetanolita floridana  52 Thermesiini  186 Thioptera nigrofimbria 237 Tholerini  490 Thysania agrippina 9, 186 Toadflax Defoliator  322 Tobacco Budworm  11, 345, 346–7, 348 Toothed Somberwing  153, 154 Tricholita signata  511 Trichoplusia ni  216 Trichordestra legitima  499 Tricolored Angel  235 Trifines (Trifids)  32, 33, 212 Tripudia rectangula  236 Tumbler, Common Sida 233 Hazel 232, 233 Rayed  232 Sandstorm 232 Tussock, Browntail  31 Definite 296 Tyrissa multilinea  166, 167 Ufeini  474–5 Ufeus plicatus  325, 474 satyricus  474, 475 Ulolonche culea  515 modesta 515 Underwing, Abbreviated 105 Agrippina  117 Alabama 137, 139, 141 Andromeda 134, 135 Angus’s 115, 116, 117 Barrens 128, 132 Bayberry 108, 109 Betrothed  105, 106 Bride  98, 112, 120 Briseis 126 Brou’s 117 Charming 140, 141 Chokeberry 140 Clinton’s 139 Clouded 121 Coastal Plain  132 Connubial 131 Consort  98, 100, 110, 118 Copper  12, 326 Darling 98, 123, 128 Dejected 118 Delilah 131 Epione 98, 100, 110 False 162 Gerhard’s 131 Girlfriend 98, 128, 132, 140 Graceful  134, 135

Grisatra 142 Grote’s 125 Habilis 115, 117, 119, 122 Hawthorn 136, 137, 139 Hoary-edged 129, 131 Ilia 98, 130 Inconsolable 118 Joined 100, 126, 153 Judith’s 116, 119 Large Yellow  538 Lincoln 142 Little 105, 106 Little Nymph  133 Little Wife  108, 109 Locust 185 Louise’s 134, 135 Magdalen 105 Marbled 126 Married 104, 106 Meske’s 127 Miranda 142 Mother 127 Mourning 111, 116 Obscure 113 Old Maid  108, 109 Oldwife 114 Once-married 127 Orba 142 Penitent 120 Pink 124 Precious 140, 141 Residua 113, 121 Robinson’s 98, 100, 111, 115, 116, 122, 153 Sad 100, 112, 120 Sappho 122 Scarlet 98, 129 Semirelict 127 Serene  115, 116 Shadowy 130 Similar 133 Sleepy 124 Sordid 135 Steely 128, 132 Sweet  139 Sweetfern  107, 109 Sweetheart 126 Tearful 114, 119 Three-staff 104, 124 Treasured 123 Ulalume 122 Ultronia 138 White 98, 125 Whitney’s 105, 106 Widow 98, 116 Wonderful 136, 137 Woody 139 Yellow-banded 99, 100, 138

Yellow-fringed 120 Yellow-gray 116, 121 Youthful 122 Underwings 97–142 lesser 143–51 Variable Tropic  92 Veiled Ear Moth  409 Veiled Grass Moth  406 Verbena Moth  371 Vetchworm 349 Voucher data  22–3, 36 Wainscot, Adjutant  506 Darted 508 Dusky 508 False 510 Hairy-legged 510 Large 409 Lesser 508 Many-lined 508 Needle 509 Phragmites 509 Reed 504, 507 Trinity 508 Scirpus 509 Ursula 509, 510 White-dotted 509 Waterlily Moth  370 Wavy-chestnut Y  213, 223 Wavy-lined Mallow Moth 233 Wedgeling, The  383 White-marked  534 White-spotted Redectis  57 Wild Petunia Seed Moth 236 Wine ropes  20 Winter Moth  467 Winter moths  20, 22, 420 Witch, Black  186 White 9, 186 Woodgrain, Bicolored  488 Confused 280, 476, 487 Gray  476 Woodling, Alternate  479, 481, 482 Cunning 483 Wood-nymph, Beautiful 309, 310 Pearly 309, 310 Xanthopastis regnatrix 355, 519 timais 519 Xerociris wilsonii  307 Xestia badicollis  521, 542, 543 c-nigrum 545

dilucida 544, 551 dolosa 545 elimata 542, 543 normanianus 551 perquiritata 551 praevia 521, 542 smithii 546, 551 youngii 544, 551 Xylena cineritia  422 curvimacula 422 nupera 422, 423 thoracica 421 vetusta 423 Xylenini  420–71 Xylomoia chagnoni  409 Xylotype arcadia  473, 474 capax 473 Xystopeplus rufago  420, 425, 432, 455 Y, Broken Banded  222 Dusky Silver  228 Hooked Silver  227 Plain Golden  223 Wavy Chestnut  223 Yellow Sunflower Moth 341 Yellowhorn  252, 253 Close-banded 252, 253 Yponomeuta multipunctella 436 Zale, Banded  178, 179 Barrens 169, 174, 175 Bethune’s 169, 180, 182 Black 174, 184 Black-eyed 169, 182 Bold-based  169, 174, 175 Boxelder  169, 183 Brown-spotted 169, 175, 179, 180 Buchholz’s 169, 177 Colorful 170, 172, 173, 174 Confused 166, 169, 176, 180 Dixie 171 Double-banded 183 Fawn Virginia Pine  169, 176 Gray-banded 181, 182 Gray Spring  169, 180, 182 Green-dusted 170 Hazel 169, 184, 444 Lunate 169, 170, 172, 173, 174 Northeastern Pine  182 Oblique 181 Okefenokee 184

One-lined 169, 174 Scaled 183 Washed-out 169, 180 Zale aeruginosa  170 bethunei 169, 180, 182 buchholzi 169, 177 calycanthata 183 confusa 166, 169, 176, 180, 181 curema 169, 182 declarans 171 duplicata 178, 179 galbanata 169, 183 helata 169, 175, 179, 180 horrida 183 intenta 169, 174, 175 largera 178 lunata 169, 170, 172, 173, 174 lunifera 174, 175 metata  169, 176, 180 metatoides 169, 180 minerea  170, 172, 173, 174 obliqua  181 perculta  184 phaeocapna 169, 184, 444 squamularis 181, 182 submediana 169, 180, 181, 182 undularis 174, 184 unilineata 174 Zales  169–84 Zanclognatha, Conifer  47 Dark 45, 46, 49 Early 45, 46 Grayish  45, 49 Lettered 48 Pine Barrens  47, 49 Toothed  47, 48 Variable  45, 48 Wavy-lined 45, 46 Zanclognatha cruralis  45, 46 dentata  47, 48 jacchusalis 45, 46 laevigata 45, 48 lituralis  48 martha 47, 49 obscuripennis  45, 46, 49 ochreipennis 45, 46 pedipilalis 45, 49 protumnusalis 47 theralis  39 theralis-minoralisgypsalis-inconspicualis 46 Zelicodes 72