Monograph |
Eric L. Quinter‡, Anthony E. McBride§
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Corresponding author: Eric L. Quinter ( protapamea@yahoo.com ) © 2025 Eric L. Quinter, Anthony E. McBride.
This is an open access book distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Quinter EL, McBride AE (2025) The Moths of North America. Advanced Books. https://doi.org/10.3897/ab.e177039
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ABSTRACT
This is the second fascicle treating a portion of the New World apameine fauna. The North American species of the genera Amphipoea Billberg and Hydraecia Guenée are reviewed, and the genera Cherokeea Quinter & Sullivan, Franclemontia Ferguson, Papaipema Smith, Parapamea Bird, and Selicanis Smith (Noctuidae: Noctuinae: Apameini) are fully revised. Three new genera are described: Arundinariaphila Quinter (type species: Arundinariaphila hayesi Quinter), Fibigeria Quinter (type species: Gortyna erepta Grote), and Sarita Quinter (type species: Acrapex relicta Ferguson). Seventy-seven species are treated, eleven of which are described as new: Arundinariaphila hayesi Quinter; Cherokeea gabrielleae Quinter; Cherokeea odonnellae Quinter; Cherokeea slotteni Quinter; Papaipema albimontana Quinter; Papaipema amianthii Quinter; Papaipema blackorum Quinter; Papaipema hesseli Quinter; Papaipema matheri Quinter; Papaipema sablensis Quinter; Papaipema sullivani Quinter. Three new combinations are created: “Resapamea” trigona (Smith) is transferred to Cherokeea, “Amphipoea” erepta (Grote) to Fibigeria, and “Acrapex” relicta (Ferguson) to Sarita. Two neotypes and three lectotypes are designated. Three species are raised from synonymy: Papaipema rubiginosa Bird, Papaipema birdi (Dyar), and Papaipema depictata Benjamin. Gortyna aerata Lyman is deemed to be a hybrid of P. cataphracta × necopina and is placed into synonymy as an unavailable name. Numerous new life histories are reported. Adults and genitalic structures of all species are illustrated in color, and their larvae, where known, are as well. Keys to all genera are included, and species keys to adults, where practical, are included. Keys to larvae are included wherever sufficiently known. Selected images of habitats and host plant symptoms of larval damage are presented in color.
DEDICATION
Dedicated to honor the legacy of Henry Bird (October 12, 1869 – July 8, 1959) without whose enormous contributions to the knowledge of Papaipema species and their life histories a century or more ago, much would still remain unknown.
AND
To the cherished memory of his son Junius Bird (September 21, 1907 – April 2, 1982) who strove to expand upon his father’s work and accompanied Eric Quinter retracing his father’s footsteps in the field and sharing insights and family archives.
PREFACE
By Eric L. Quinter
The serendipitous discovery in 1971 by Eric L. Quinter (ELQ) of a spectacular, previously unknown species of Papaipema in Pennsylvania initiated what has become a study of near epic proportions, now finally completed in its sixth decade. In the course of this work, an astonishing body of new information has been uncovered revealing the existence of previously unknown species of Papaipema and numerous other unknown species of undescribed apameine genera, as well as myriad intricate details of their biology.
The conceptualization of this fascicle did not come about for several more years, when Richard Dominick sat in my office in New York City and suggested I pursue this goal. At the time, a number of fascicles in the Moths of North America series had already been published. He encouraged me to join what seemed a daunting undertaking, but I agreed. By this time, several additional new species of Papaipema had come to my attention through correspondence with lepidopterists around the nation. There appeared to be sufficient new information to render production of a fascicle devoted to this group of moths a worthwhile endeavor.
Long before any writing began, the first few decades of work on this project were devoted to visiting institutional and private collections and recording specimen data. An enormous amount of time was devoted to field work with the goal of elucidating details of the life histories of the new species, as well as learning more about the known species. These years were punctuated with lengthy periods of inactivity as other life priorities intervened. In retrospect it sometimes appears unfortunate that what has turned into a lifetime endeavor could not have been completed decades earlier. My greatest regret is that so many people who were influential and supportive of this work remain only cherished memories of mine, as they have not lived to see this result. Offsetting this is the knowledge that the present work is so much more complete, accurate, and richly detailed than it would have been, had it been completed sooner. The seemingly unending cascade of new information has both hindered the production of this fascicle and enriched it.
As the project continued to expand, it became necessary to include other genera believed related to Papaipema, along with the need to present new information regarding other apameine borers newly discovered in the process. It was also deemed necessary to enlist the efforts of additional individuals to bring the project to fruition. One coauthor joined the effort to complete this volume with major contributions throughout. Anthony E. McBride (AEM) undertook the daunting task of creating the distribution maps for all the species, compiled from far more than fifteen thousand specimens he and I examined and documented in nearly one hundred collections across the continent and beyond. He also photographed all the adult specimens, produced in situ images of all the larvae we have found in the field, and collaborated with me on their arrangement. Over the years, I have prepared many hundreds of genitalic dissections of these moths, but the task of making and photographing fresh genitalic preparations to facilitate comparative study during production of the fascicle was completed by AEM. In addition, he arranged the images of adult specimens, their larvae, habitats, host plants, and symptoms of larval damage for the color plates. He also contributed to many sections of the text and proofread the entire manuscript, adding many valuable suggestions. James R. Wiker (JRW), with decades of field experience in his native Midwest, contributed substantial observations of larval behavior in the field, documented new insights relative to host plant utilization, and provided much-needed fresh material for DNA analysis as well as larval vouchers for preservation. As this fascicle was in the final stages of production, it was recognized that the superb dissections created by Christi Jaeger at the CNC would significantly enhance the quality of the genitalia images for the plates, so she undertook the considerable task of preparing them for all taxa covered in the fascicle. Finally, Chris Schmidt contributed his considerable expertise relative to interpreting results of barcoding and analyses of nuclear genes, undertaken to clarify some problematic issues and provide a rationale for our arrangement of the taxa. Numerous valuable discussions with him concerning biogeographic scenarios have enriched the final product.
One of the greater challenges we faced was rendering this work optimally useful to those who may rely upon it to aid their recognition and identification of these moths. The genus Papaipema in particular is especially difficult to those attempting to identify the various species with any degree of certainty. It was particularly challenging to create keys to species that would actually yield reliable identifications. We created a workable key to Papaipema larvae which may be encountered by those interested in this genus but were not able to achieve the same for the adult moths. The best hope for achieving accurate identification of adult specimens will come through very careful comparison of the subtle details of wing pattern variation and coloration of specimens illustrated on the plates with the diagnoses provided in the text, distribution maps, and genitalic examination when necessary. We have therefore departed from the tradition in most previous fascicles of arranging the adults figured on the color plates in the same sequence as the putative phylogenetic order presented in the text, for two reasons. The first simply reflects the impossibility of replicating a branching phylogeny in a linear fashion, a perennial problem with all such attempts. The second and far more practical reason for this departure is to facilitate the identification of species, which remains a primary goal of this work. Accordingly, the adult images on the color plates were arranged so that those species most closely resembling one another appear on the same plates, wherever possible. We also slightly increased the image size from actual to 1.25 × natural size. It is hoped that these modifications will facilitate the comparison of a specimen at hand with the various choices on a single plate, without having to search through all the plates to find the best visual matches.
During the course of this study, it has become widely recognized that species of Papaipema, due to their high degree of host plant specificity, are particularly well-suited subjects as indicators of habitat diversity, environmental quality, and assessment of ecological change. Among a disturbing percentage of these species are indications of dramatic decreases in both number of occurrences and population density in recent decades. These species have become a primary focus of several ecological studies, and they are appearing with increasing frequency on state and regional lists of species considered to be of special concern, threatened, endangered, or extirpated. It is regrettable that the body of work presented in this fascicle was not available long before to guide conservation endeavors. We hope that the present work will be found so much more accurate, comprehensive, definitive, and hence more valuable to future workers.
ADDENDUM
Eric Quinter passed away on 4 July 2024 after a battle with cancer. Eric had a lifelong interest in the Apameini and made huge contributions to our understanding of these moths. He especially enjoyed discussions about these moths, both learning from fellow enthusiasts and passing new information along with his usual cheery disposition. He helped with difficult identifications, entertained with funny stories of times spent in the field with fellow lepidopterists, and provided coveted clues needed to search for larvae. As his friend and colleague Alberto Zilli remarked, “Eric was one of the great noctuid guys.” Eric’s spirit never waivered after his diagnosis, and he bravely pushed on to finish this work. While he lived to finish the first draft of this fascicle and saw the draft in its completed form with maps and plates, he unfortunately did not get to see this final published result. He did, however, take comfort that he was able to complete what constituted his life’s work. He is greatly missed.
AUTHORS’ ACKNOWLEDGEMENTS
ELQ:
It comes as no surprise that a study which has thus far occupied the entirety of my adult lifetime has involved the participation and contributions, both large and small, of a very large number of persons. To acknowledge only those who have made major contributions and dismiss all others as being too numerous to mention would be demeaning to so many who have done much to make this fascicle possible. I shall endeavor to list, in alphabetical order, all whom I can recall, past and present, living and deceased, without regard to the size or relative importance of their contributions, and beg forgiveness from any I may have omitted. This work would have been impossible without their combined contributions. The list is formidable, and a large number of these people are long deceased, or presumed to be by now, so I am not indicating their status, as I don’t actually know in numerous instances.
I wish to personally and gratefully acknowledge all the following persons who directly assisted me in the earlier decades, and more recently through the involvement of my coauthor, for their participation in this endeavor, their support, facilitating or assisting with field work, valuable discussions, technical assistance, the gift of personally collected specimens, or for the loan of institutional specimens under their care: James Adams, Dave Baggett, George Balogh, John Bauer, Glen Belyea, Ken, Elaine, and Judd Bernard, Jim Bess, Junius and Peggy Bird, Bill and Nancy Black, Bill Boscoe, Bea Brewster, Vernon and Charlotte Brou, Maurice Broun, Ed Brower, Richard Brown, Linda Butler, John Cadbury, Ann Carter, Harry Clench, Charlie Covell, Lars Crabo, Wilfred Craig, Don Davis, Liz Day, George Derkovitz, Phil DeVries, Dick Dominick, Julian Donahue, Lance Durden, Günter Ekis, Graham Fairchild, Les Ferge, Doug Ferguson, Michael Fibiger, Hermann Flaschka, Scott Francis, John Franclemont, Tom Franks, Larry Gall, Karl Gardner, Loran Gibson, Jocelyn Gill, James Gillingham, George Godfrey, Paul Goldstein, Jean Graber, Paul Grey, Elmer Griepentrog, Steve Hall, Louis Hanfield, Nancy Hanson, Dave Hardwick, Chuck Harp, Alan Hayes, Richard Heitzman, Richard Henderson, Tom Henry, Ted Herig, Sid Hessel, Ron Hodges, Dick Holland, Martin Honey, John Hornbrook, Bill Howe, Jeff Ingraham, Joel Johnson, Kurt Johnson, Steve Johnson, Judith Jones, Ken Kadow, Rick Kergosien, Warren Kiel, Colin Kilmer, Ke Chung Kim, Charley Kimball, Ian Kitching, Leroy Koehn, Vicki Kondo, Hugo Kons, Larry Kopp, Richard Kruzansky, Bill Lamond, Joe Lankalis, Ellis Laudermilk, Irwin Leeuw, Claude Lemaire, Rae Letsinger, Ron Leuschner, Robert Lewis, Zheiwei Liu, Eric LoPresti, Jim Madenjian, Bryant Mather, Tim McCabe, Peter McEvoy, Noel McFarland, Hugh McGuinness, Mark Mello, Charles Merker, Eric Metzler, Don Miller, Charles Mitter, John Morton, David Morton, Joe Muller, Ken Neil, Mike Nelson, John Newman, Alfred Newton, Jr., Mo Nielsen, John Nordin, Phillip Nordin, Mark O’Brien, Paul Opler, John Oswald, Ron Panzer, George Patterson, Ricky Patterson, Mike Pogue, Bob Poole, Jerry Powell, Ray Pupedis, Daniel Quinter, Janice Quinter, Sarah Quinter, Brett Ratcliffe, John Rawlins, Charles Remington, Fred Rindge, Steve Roble, Tom Rocheleau, Ted Sargent, Dale Schweitzer, Yale Sedman, Mat Seidensticker, Andrey and Alla Sharkov, Paul Shealer, Jon Shepard, A.C. Sheppard, Bill Sieker, Ed Simons, George Sleesman, Jeff Slotten, Joe Smaglinski, Jeff Smith, Norman Smith, Ray Stanford, Ken Stead, Fred Stehr, H.G. Stevenson, Gayle Strickland, Bo and Ashley Sullivan, Gaby Tannor, Richard Thurston, Steve Thurston, Ed Todd, Asher Treat, Jim Troubridge, Paul Tuskes, Leonard Uttal, Jim Vargo, Pierre Viette, Ed Voss, Dave Wagner, Herb Wagner, Dave Wahl, Virgil Warczynski, Allan Watson, Reggie Webster, Jason Weintraub, Bill Westrate, Jim Wiker, Ben Williams, Darryl Willis, Judy Wink, Dave Winter, Brooke Worth, Barry Wright.
Special thanks go to Lloyd Culp and the staff of the Great Dismal Swamp National Wildlife Refuge who graciously facilitated my extensive work there. Royce Hayes provided accommodations, extensive logistical support, and much helpful advice during my nineteen separate visits to St. Catherines Island, Georgia where much of the early research for this project was conducted during the years 1989 through 1994. Financial support for my work on St. Catherines Island was provided by a grant from The Edward J. Nobel Foundation, Bio Insect Genera Fund. Several of a multitude of field trips conducted by me over the years were made together with Paul Goldstein, supported in part by funds from Sigma Xi and an AMNH Theodore Roosevelt Memorial Fund travel grant. Special acknowledgement goes to The American Museum of Natural History and its staff for use of its superb facilities and its supportive stance for my work over the many years I was employed there.
Special recognition and gratitude is extended to the following persons: Erik Neff for his superb drawings of the larvae illustrated on monochrome plates 4–7 and for habitat photographs taken on St. Catherines Island, Georgia; Erica Chiao for her drawings of the genitalia illustrated in
AEM:
Over the thirteen years that I worked on this fascicle with Eric, many of the individuals listed above and a number of others, kindly offered assistance on this project, important data and information, specimens for study, and companionship in the field, for which I am immensely grateful. I wish to recognize the following individuals: James Adams, Ken, Elaine, Judd, and Lucas Bernard, Kyhl Austin, Dwayne Badgero, Bill and Nancy Black, Rick Boscoe, Stéphanie Boucher, Steve Bransky, Aaron Brees, Gerald Burnett, Michael Butler, Janine Caira, Frank Carle, Lars Crabo, Paul Dennehy, Jill Dodds, Jason Dombroskie, Jim Durbin, Becky Elkin, Les Ferge, Dave Flitcroft, Tom French, Betsy Froyd, Larry Gall, Joe Garris, Loran Gibson, Paul Goldstein, Jerry Goth, David Grimaldi, Chris Grinter, Steve Hall, Chuck Harp, Chris Heckscher, Richard Henderson, Ted Herig, Jeff Hooper, John Hyatt, Dave Iftner, Bonnie Isaac, Kyle Johnson, Liz Johnson, Steve Johnson, Matt Kenne, Brian Klassanos, Stephen Kloiber, Leroy Koehn, Vicki Kondo, Ellis Laudermilk, Betsy Leppo, Raymond Little, Eric LoPresti, Alan Loudell, Lydia McBride, Steven McBride, Tim McCabe, Megan McCarty, Hugh McGuinness, Paul McKenzie, Jerry McWilliams, Mark Mello, Eric Metzler, Tom Middagh, John Morton, David Morton, Mike Nelson, Tam Nguyen, Mo Nielsen, Jane O’Donnell, Jim Petranka, Mike Pogue, Derick Poindexter, Suzanne Rab Green, John Rawlins, Hartmut Rietz, Steve Roble, Peter Rosenbaum, Mike Sabourin, Craig Segebarth, Kevin Segebarth, Danny Shurey, Karen Sime, Jeff Slotten, Sam Smith, Sarah Smith, Bo and Ashley Sullivan, Jason Tesauro, Jim Troubridge, Jim Vargo, Vanessa Verdecia, Dave Wagner, Matt Wallace, Sharon and Wade Wander, Jason Weintraub, Dave Wikle, Darryl Willis, Ann-Marie Woods, Peter Woods, Jim Zamos, and Harry Zirlin.
Special recognition and thanks are extended to Jocelyn Gill, who provided me invaluable assistance with the techniques needed to produce the color plates and maps; Andrew Bennett, Jose Fernandez-Triana, and James O’Hara for their detailed and thorough work providing determinations for our reared parasitoid specimens; Jane O’Donnell for her thorough review of the manuscript, assitance with taxonomic issues, and unwavering support of the project; Christi Jaeger for her preparation of the beautiful genitalic plates; Chris Schmidt for facilitating our work on this project, providing his expertise on barcode data and insights throughout, and guiding and assisting me throughout the review process; the reviewers of this fascicle for their thorough and detailed work, adding many valuable insights, corrections, and suggestions; my good friend Jim Wiker, with whom I’ve spent many fun and important days afield; Eric Quinter, my co-author, mentor, and good friend, from whom I learned so much about the study of Lepidoptera; and also to my good friend Dale Schweitzer, who is responsible for igniting my interest in Papaipema and related borers.
MATERIALS AND METHODS
The senior author examined more than 15,000 specimens deposited in 96 institutional and private collections, most of which are listed under Notes in the appendix. AEM documented significant numbers of additional specimens. In the preparation of the maps for this fascicle, only those records of specimens with identities that were personally verified by at least one of the authors were included. Literature records as well as records available online but unverified by us were excluded due to the rampant misidentification pervasive among these taxa. Accordingly, the maps are not to be construed as range maps, but rather maps of distribution of specimens examined. Nonetheless, coverage of material preserved in collections has been comprehensive, and the maps should approximate actual geographic distribution of the species. Reports of occurrences significantly beyond these distributions might best be regarded with suspicion.
AEM prepared all photographs of pinned adults and live larvae using a Canon 750D SLR camera with a Tamron 90mm macro lens. Images were taken in a light box with a circline fluorescent bulb rated at 6500K (Halco Lighting Technologies ProLume 22W). Photographs of adults were taken using focus stacking procedures with final composites produced using Adobe Photoshop CS5 software. Photographs of larvae were also produced using focus stacking after larvae were positioned in the light box within their larval burrows. For the images of adults, Adobe Photoshop CS5 software was used to digitally remove insect pins, repair physical defects on specimens such as pin holes, missing scales, small tears, and chips, and to adjust the position of wings and abdomens as needed. All illustrated adult primary types were left completely unaltered and are represented on the plates in their actual condition. The monochrome plate images consisting of adult thoracic tufting and larval damage were produced using paper negatives made from drawings by Henry Bird that were given to ELQ by Junius Bird. These negatives were scanned and converted to positive images using Adobe Photoshop CS5. AEM compiled the map distributional data and then generated the maps using SimpleMappr (http://www.simplemappr.net/).
Genitalia were photographed by Christi Jaeger using a Leica DFC450 camera mounted on a Leica M205 C stereo microscope. Stacked images were assembled using the Leica Application Suite Version 4.12 and edited using Adobe Photoshop CS6. Images were edited to remove the background and, in non-type material, torn or extraneous membranous tissue and taxonomically uninformative setae around the ventral edge of the valves in males.
A reference library of DNA barcodes for most North American noctuids was previously published by
All plates and maps were assembled using Adobe Photoshop. Plates of adults were produced using a background of 15% gray. The adult specimens figured each have a digital shadow added, and all specimens are illustrated at 1.25 × natural size.
Introduction and overview of genera
superfamily NOCTUOIDEA (CONTINUED)
family NOCTUIDAE (continued)
subfamily Noctuinae (part)
tribe Apameini Guenée, 1841
Type genus Apamea Ochsenheimer, 1816
note—Synonymy of family-group names is given in
subtribe Apameina Guenée, 1841 (part)
A historical review of the noctuine tribe Apameini was provided in the preceding apameine Fascicle 26.9 (
The monophyly of Apameini as currently defined is firmly established (
This fascicle addresses two additional groups of genera. The polyphyletic Arundinaria-feeding genera are an ecologically associated group of four genera, the larvae of which are borers specializing on the single plant genus Arundinaria Michaux (Poaceae: Bambusoideae: Arundinarieae), a genus of monopodial, temperate-forest bamboos colloquially called “cane” in the United States. These genera are regarded as being of eastern Asian origin, more closely related to each other than to any North American genera yet still rather distantly related to each other, comprising the descendants of several ancestral Asian lineages. This group is covered in this fascicle because nearly all included species were previously unknown, and a large body of information concerning them is presented here for the first time. Their inclusion in this fascicle does not imply they are closely related to the dicot feeders treated here, so they should not be construed as an immediate sister group to the Papaipema group genera, as their placement in the check list might imply. It is likely that they are more basal clades within Apameini. The second group treated in this fascicle is the Papaipema group, comprising six genera with endophagous larvae that are primarily dicot specialists. The Papaipema group belongs to a larger assemblage of genera, the Holarctic Gortyna group, that includes a number of strictly Palearctic genera, and two genera of Holarctic distribution, Amphipoea and Hydraecia. Taken together, the genera comprising the Gortyna and Papaipema groups form a monophyletic unit with a hypothetical common ancestor which likely arose somewhere in the Holarctic during the Tertiary Period.
The Holarctic Apameina are strictly univoltine, one of the defining biological characteristics of the subtribe. The majority of the included genera, and all but one treated in this fascicle, overwinter in the egg stage. In some other genera of Apameini, including Apamea, the eggs hatch later in summer or autumn and the larvae hibernate, usually as first instars. These larvae resume feeding whenever winter or early spring temperatures favor feeding activity. This is characteristic of certain grass- and sedge-feeding genera where the dormant meristematic tissues of their host plants remain viable during winter. As might be expected with these genera, larval development and pupation occur earlier than in genera that overwinter as eggs, and the adults typically fly earlier in the year.
Overwintering apameine eggs hatch very early in spring, coinciding with the first new growth of potential host plants. Among the numerous Papaipema species that are widely distributed in North America, the result is that their larvae commence feeding as much as two months earlier in southern populations than those in the northern portions of their ranges. One might naturally conclude that adults in southern populations of these species would fly comparably earlier in the season than their northern counterparts, but the exact opposite is true. Flights of southern species begin even later. The potential for a second annual generation would seemingly increase with decreasing latitude, as is typically the case with many other noctuid groups. Nonetheless, the life cycle of all these species requires an entire year to complete, regardless of where they occur. Univoltinism is obviously highly conserved in Apameina. The apparent gap between complete maturation of the final instar and adult eclosion is occupied by a period of larval aestivation. The pupal period averages 28 days for nearly all Papaipema species regardless of where they occur. In northern populations, there is no aestivation, and pupation occurs within a few days of cessation of larval feeding. In the southern United States, larvae may mature as soon as early summer in some cases and enter a period of aestivation, the length of which is dependent upon latitude. Such larvae often abandon their galleries, if they are species which occupy aboveground portions of their host plants, and spend the hot summer months aestivating in the cooler moist soil.
Univoltinism among Apameini is not entirely a global phenomenon. The curious situation in the southern hemisphere demands explanation. Most, if not all, Afrotropical apameines, represented by the subtribe Sesamiina Fibiger and Goldstein are either multivoltine or appear continuously throughout the year in great numbers (
There is a tendency among Papaipema species which occur in the southern United States to exhibit adult flight periods extending farther into late autumn than northern populations, but such flight seasons also begin later here and remain compressed into little more than one month on average in most instances. Several Papaipema species as well as a number of species of the cane-feeding genera are distributed on the southern Atlantic Coastal Plain, across northern Florida, and along the Gulf Coast. A few apameine species such as Papaipema polymniae, P.stenocelis, and Parapamea buffaloensis extend more than halfway down the Florida peninsula, but one species, Papaipema speciosissima, notably ranges all the way to the Florida Keys, representing the southernmost distribution of any known Papaipema species. In extreme southern Florida, speciosissima has been recorded flying in nearly every month of the year. Records for P.polymniae and P.stenocelis are too few to assess how much of the year they may be on the wing. The uniform climatic conditions prevailing throughout the year at this essentially tropical latitude accounts for the apparent breakdown of voltinism, possibly precluding extensive apameine radiation into the Neotropics. Brou & Brou 2020 reported a greatly expanded flight period for Parapamea buffaloensis in southern Louisiana, with dates ranging from June to early November. In southern Mississippi, this species has been recorded flying as early as mid-July. In southern Georgia records range from early July to mid-November, and in Florida from late April to late October. Elsewhere throughout the majority of its extensive range, this species has a limited flight season from late August through early October, with most records falling in September. This pattern may represent further evidence of the same phenomenon. It is important to distinguish multivoltinism typical of many lepidopteran species, or even continuously-breeding species lacking distinct broods, from staggered emergences of strictly univoltine species such as in Apameina, individuals of which nonetheless require an entire year to complete their life cycle. Population densities of monophagous or oligophagous species such as most Papaipema are often quite low, as they are clearly limited by the local abundance of their particular host plant, which is frequently low as well. Species such as P.speciosissima typically maintain low population densities, and emergence of the relatively few individuals in such populations, staggered throughout the year, would seemingly present a potential reproductive disadvantage. Why the parallel situation in the Old-World tropics has led to a very different evolutionary result remains a mystery.
All the taxa treated in this fascicle belong to the subtribe Apameina, commonly regarded as true borers. Significant disparity exists with regard to the actual feeding modes employed by the immature stages of these moths beyond the distinction of whether they are stem borers feeding primarily above ground or borers of roots, rhizomes, and other subterranean structures. In both the Arundinaria-feeding genera and the Papaipema group genera, stem boring involves two independent extremes of feeding modes with some degree of intermediate conditions among different species. At one extreme, a large volume of the succulent, green, inner stem parenchyma is consumed throughout the larval development period, resulting in the production of a substantial amount of expelled frass. The larva in this case may bore through half a meter or more of plant stem or root. In such instances, the required nutrition is clearly derived from sugars and other vital substances present in the masticated food passing through the larval gut. At the opposite extreme, very little plant tissue is consumed by the larva throughout its entire development. Instead, the walls of plant stems, especially hardened, woody stems or galls formed thereon, are not consumed but are girdled internally by the developing larva. The larva apparently obtains its entire nutritional requirements by simply sucking the sap flowing through the walls of the hardened stem, which then oozes from the girdling wounds. Very little dried frass is produced throughout larval development, and the entire larval burrow may be scarcely longer than the larva itself. Either mode is characteristic of the species employing it and never varies within that species. Nutrition obtained by either method is clearly sufficient to produce adults of equally large size.
What may have driven the evolution of internal feeding and when it may have occurred with regard to Apameini is a matter of considerable interest. Endophagy is a primitive feeding mode that evolved in numerous lepidopteran lineages and is relatively rare in the Macrolepidoptera. Endophagy is considered to represent a derived condition among the Noctuidae, probably having arisen from exophagous lineages such as climbing cutworms and evolving independently within the noctuine tribes Apameini and Arzamini. Stem boring in turn appears to have led to endophagous feeding within roots, rhizomes, and tubers. Seed capsule feeding in other noctuid lineages is probably independently derived. Internal feeding in the larval stages appears to be an important adaptation among the many other mechanisms employed by Lepidoptera to avoid predation and parasitism. If so, it is much more successful in the case of predation than in the case of parasitism, a subject dealt with later in this fascicle. By remaining concealed within stems, rhizomes, or roots for the duration of larval development, borers effectively avoid exposure to both avian predators and the usual predatory insects such as vespine and polistine wasps, asopine pentatomids, etc. Within many of the apameine genera specializing upon grasses, it may also enable young larvae feeding internally upon tender embryonic tissues to avoid the foliage of these plants which have notoriously high silica content. Even some large herbivorous mammals such as deer completely avoid grasses, which they are not adapted to process, perhaps for the same reason. If one compares same instar larval head capsule size, indicative of musculature required to masticate food, of lepidopteran groups specializing on grass foliage (e.g., many hesperiids, satyrine butterflies, or Leucania Ochsenheimer species) with those of endophagous apameines feeding on the very same plants, the apameine head capsules are consistently smaller. A shift from external foliage feeding to endophagous behavior is accompanied by a parallel shift from the cryptic, longitudinally striped patterns prevalent among larvae of many external feeders, especially those feeding on grasses and herbaceous forbs, to a reduction of longitudinal striping prevalent among stem borers (which often must relocate externally to additional plants). Larvae of root-boring specialists remain concealed for the duration of their development within the plants and tend to lose pigmentation and longitudinal stripes in favor of a grub-like appearance. These features provide potential phylogenetic information as well as diagnostic characters for larval identification.
It has even been suggested that the endophagous behavior exhibited by these borers may be a mechanism to avoid the toxic allelochemicals present in the foliage of many of the host plants specifically preferred by a number of apameine species, especially in the genus Papaipema.
Dicotyledonous plants appear in the fossil record during the mid-Cretaceous and grasses a bit later in the late Cretaceous and early Tertiary, along with Asteraceae. The huge diversification in the fossil record of asters and their kin – major host plants for many of the dicot-feeding borers – along with that of Poaceae appears to have rapidly followed the mass extinctions that occurred at the Cretaceous / Paleogene boundary. Molecular, phylogenetic, and other data indicate that disjunct distributions of many plant groups between eastern Asia and eastern North America occurred during the Tertiary when temperate forests achieved their maximum development (
Some of the ideas presented here are admittedly speculative but are suggested as hypotheses to encourage further investigation of the phylogenetic relationships both within the tribe Apameini as well as among the other noctuid lineages. These relationships are not yet well understood, although serious progress is being made in the realm of molecular systematics. Sequencing of mitochondrial DNA has provided resolution of some closely related species where traditional morphology has not enabled establishment of species boundaries. Recent work employing nuclear DNA appears encouraging. Understanding the relationships between species groups and genera remains challenging. It is hoped that further work with nuclear genes already in progress may provide further insights into these relationships and ultimately contribute to a stable classification.
The Arundinaria-feeding Genera
In the course of a comprehensive survey of the terrestrial arthropod fauna of St. Catherines Island, Georgia conducted from June 1989 through September 1992 by staff of the American Museum of Natural History, ELQ documented the presence of several previously unrecognized species of moths associated with cane (Arundinaria) with generic affinities which were uncertain, other than their obvious placement within the noctuid tribe Apameini. He later undertook a much more detailed investigation of the bionomics of these entities on St. Catherines Island and elsewhere, including the species described below.
The large perennial grasses of the genus Arundinaria, together with closely related genera indigenous to East Asia, form a group of temperate woody bamboos which spread by running rhizomes (i.e., monopodially) and are only distantly related to the great diversity of giant, clump-forming bamboos indigenous to the New- and Old-World Tropics. The highly disjunct occurrence of the three native North American species comprising this genus distributed across the southeastern United States (
The Arundinaria-feeding genera, currently known solely from southeastern North America and having no obvious close relatives elsewhere in the Nearctic or the western Palearctic, represent ancient Arcto-Tertiary lineages arising from common ancestors which may be shared with some as yet unrecognized but likely extant eastern Asian genera also sharing an endophagous specialization on Arundinarieae (
Extensive phylogenetic analyses conducted by Triplett et al. show that Arundinaria is closely related to the bamboos Sasa, Pseudosasa, Sasamorpha, and Pleioblastus of Japan and that either Arundinaria or its immediate ancestor migrated across the Bering land bridge into North America. This introduction most likely occurred during the Miocene, an extended period when climatic conditions were conducive to migration of temperate plants across Beringia 23–16 million years ago, as detailed by
Among the Apameini, the specialized adaptation to and utilization of cane as a sole food resource is not limited to the Arundinaria-feeding genera. Two species of Papaipema and two species of Protapamea Quinter share this resource in the southeastern United States. One of the Papaipema requires tender, newly emerged culms to complete its larval development; the other exclusively utilizes woody, second year and older culms. The two species have an allopatric distribution, occupying different cane species. Both species of Protapamea share a uniquely different exophagous feeding mode as climbing cutworms feeding exclusively upon tender, unexpanded, axillary shoots of cane. The host plant utilized by Loscopia roblei Quinter and Lafontaine is unknown but believed to be switch cane, based on habitat association. Laboratory-reared larvae of roblei have features consistent with related apameine cutworms. Franclemontia interrogans (Walker), Sarita relicta (Ferguson), and the non-apameine noctuids Leucania calidior Forbes and Argillophora furcilla Grote are additional competing cane specialists. Franclemontia spends most of its one-year life cycle in the larval stage feeding internally upon second-year and older culms of low nutritional value; consequentially, larval development is extremely slow. Sarita relicta larvae bore in tender, young petioles of the terminal foliage. Leucania calidior is an external foliage feeder (ELQ, unpublished data). The immature stages of Argillophora furcilla have not yet been documented, but females have been observed ovipositing under new leaf sheaths at the tops of new culms of hill cane by AEM, and a single, unidentified, mature larva collected on giant cane by JRW produced an adult A.furcilla. It is suspected that the arctiine Leucanopsis longa (Grote) also may be a specialist on switch cane, as the moth is regularly found in various habitats where this grass is abundant. It is cryptically patterned and colored to match the canebrake microhabitat. Collectively all these, together with one pyralid (Crocidophora pustuliferalis Lederer), two of the three North American species of the satyrine butterfly genus Enodia Hübner, the closest relatives of which occur in southeastern Asia (
The Arundinaria-feeding group of genera in North America comprises: Arundinariaphila Quinter, a new genus described below, Cherokeea Quinter and Sullivan, Franclemontia Ferguson, and Sarita Quinter, also described below. All eight species included here share a large majority of bionomic attributes, the same host plant genus, and a North American distribution confined to the southeastern United States. These do not constitute a monophyletic unit but rather an ecologically associated group, the ancestral lineages of which had presumably already diverged prior to their dispersal into North America during the Miocene. It might seem reasonable to have included Arundinariaphila and Cherokeea in a single genus. As the relationships between the species of these two genera became better understood, such action became increasingly untenable. Combining the two genera into a single genus would render it difficult to characterize, due to the extreme diversity and complexity of the genitalic characters defining each. Further, doing so would possibly obscure historical biogeographical relationships within this group, and potentially create a paraphyletic assemblage with respect to hypothesized Asian congeners, discussed elsewhere. It is possible that a close phylogenetic relationship with eastern Palearctic species currently placed in Arenostola Hampson may be discovered. The inclusion of Franclemontia and Sarita is purely tentative, and more a matter of expedience than any evidence of close phylogenetic relationships among these genera. Until such a time as a comprehensive global assessment of the phylogenetic relationships of all the genera within the Apameini has been completed, placement here seems a reasonable solution.
With the recent recognition of three indigenous species of cane in North America, intrageneric phylogeographic events can also be estimated.
Due to extreme polymorphism among adults of Arundinariaphila and Cherokeea, recognition of both the species and genera based upon superficial appearance is problematic, but the genitalic characters are diagnostic and the genera are well defined on that basis. The genitalic characters of both sexes are impressive in their complexity and reliable consistency. Use of the keys provided here will prove initially necessary for identification purposes and will facilitate recognition when consulting the color plates, which will have limitations if used alone.
The complexity of genitalic structures in the Arundinaria-feeding genera, especially those of the male, present special problems. There has been much confusion regarding homology and terminology applied to some of these structures, due to varying interpretations by different authors.
The clasper is derived from a basal sclerite located on the inner face of the central part of the valve from which the ampulla, if present, arises. The clasper may be represented by nothing more than the appressed basal sclerite itself or may terminate in a freestanding process which may be oriented dorsally, ventrally, or distally. In all cases, the clasper originates ventrally relative to the digitus. The digitus, if present, is derived from the sclerotized costa of the valve, dorsally relative to the clasper. Its freestanding parts are usually oriented ventrodistally, but occasionally otherwise. The clavus is a heavily sclerotized process arising between the juxta and the base of the valve, seemingly attached to the sacculus.
In the species descriptions of the Arundinaria-feeding genera as well as the Papaipema group genera, use of the term terminalia refers to the genitalia in the traditional sense and also includes other diagnostic characters of the terminal segments of the abdomen.
KEY TO THE ARUNDINARIA-FEEDING GENERA (ADULTS)
1. Small (forewing length 8–10 mm), slender moth with long, narrow forewing 2.5 × as long as wide Sarita
— Larger moth (forewing length typically greater than 9.6 mm); forewing normal, broad, about 2.0 × as long as wide 2
2. Male genitalia lacking clavus Cherokeea
— Male genitalia with long, slender, sclerotized, dorsally projecting clavus arising from base of sacculus 3
3. Male genitalia with prominent, symmetrical, freestanding, dorsally oriented clasper Arundinariaphila
— Male genitalia lacking freestanding, prominent clasper Franclemontia
GENUS Arundinariaphila Quinter, NEW GENUS
Gender: feminine. Type species: Arundinariaphila hayesi Quinter.
note—Generic name derived from the name of the host plant, Arundinaria [from Latin arundo, a reed, or cane] + Greek philos, loving.
Arundinariaphila is closely related to but abundantly distinct from Cherokeea, the members of which are more specialized in their occurrence and specific choice of Arundinaria host species. The sole species of the genus Arundinariaphila, A.hayesi, inhabits most of the range of its host plant genus Arundinaria throughout southeastern North America, with larvae utilizing these grasses almost everywhere they grow, regardless of habitat.
DIAGNOSIS OF GENUS ARUNDINARIAPHILA. The reinforcing rod-like structures between the anal papillae in the female, the s-shaped pleurite joining the tegumen and vinculum in the male, and the ventrally reflexed cucullus are all characters among numerous others which unambiguously place the genus within the Apameini. The following features define the genus Arundinariaphila: 1) specialized adaptation to endophagous utilization solely of Arundinaria; 2) forewing venation as figured in
DESCRIPTION OF GENUS ARUNDINARIAPHILA. Head: antennae of both sexes ciliate, laminate, less deeply in female; dorsum scaled; ventral lamination obscured by densely pubescent sensilla; eye smooth, rounded; haustellum unreduced; labial palpus laterally flattened, upcurved; first segment swollen basally, arching slightly upward, slightly more than half as long as second segment; second segment straight, about as long as eye width, scales broad; third segment ⅓ × length of second segment, and projecting slightly anteriorly, scales narrow; frons convex, unmodified, with central dense tuft of converging spatulate scales overlaying fine, converging hairs. Thorax: vestiture of prothoracic collar, patagia, mesothorax, and metathorax concolorous, a mixture of coarse spatulate, thin apically spatulate, and hairlike scales; mesoscutellar crest prominent, metascutellar tuft less so. Wings: forewing elongate, length 2.0 × width, apex acutely rounded, venation normal apameine as figured in
Arundinariaphila hayesi Quinter, NEW SPECIES (Common Cane Borer)
Arundinariaphila hayesi Quinter.
Type locality: St. Catherines Island, Liberty Co., Georgia, USA. [CNC]
note—Named in recognition of Royce Hayes, Superintendent of St. Catherines Island, Georgia. An accomplished naturalist himself, Royce offered many valuable suggestions, and his cheerful and enthusiastic logistical support of my work greatly facilitated my extensive studies in the canebrake habitats there.
Despite being a common species in its canebrake habitat, Arundinariaphila hayesi was virtually unknown until ELQ’s research on canebrake moth fauna began in the 1980s. The single known historic specimen was found unidentified in the American Museum of Natural History, having been acquired with the J. B. Smith collection from Rutgers University. It is a male labelled simply “Tenn.” “4.25” [an impossibly early date], and “Collection S.T. Kemp” without a year given. This is the same person who collected the four syntypes of Luperina trigona in 1899 which Smith described in 1902. It is very likely this specimen was collected around the same year and at the same place in Smith County, Tennessee.
Adults of Arundinariaphila hayesi are superficially similar to some species of Cherokeea which may co-occur with them. Aside from the phenotypic variability of forewing coloration and pattern exhibited in both genera, a modest degree of sexual dimorphism exists in A.hayesi not present in any Cherokeea species. Males of A.hayesi typically have a smooth appearance lacking a contrasting or mottled pattern, with unicolorous liver-brown forewings; females most frequently tend to be tan with more prominent pattern features. The species most likely to be confused with A.hayesi is Cherokeea gabrielleae, although their distributions barely overlap. For distinguishing features, refer to the discussion in the diagnosis of C. gabrielleae. Very large, tan A.hayesi individuals might be confused with unusually small individuals of the plain phenotype of Cherokeea trigona having similar coloration. In all instances however, the dramatic differences between the two genera in their male genitalia will eliminate any confusion, as will the presence of prominent, externally visible lodices in females of A.hayesi.
Arundinariaphila hayesi is a medium-sized species with a forewing length of 11.5 to 17 mm. Many of the adult specimens examined were obtained as wild-collected larvae and reared in captivity under unnatural conditions, often on an artificial medium. Many, but not all, of these individuals were stunted during their larval development and produced smaller than normal adults. Such specimens have been excluded from measurement. Measurements provided here are largely based upon wild-collected adult specimens. Individuals from midwestern populations of A.hayesi utilizing giant cane may attain considerably larger size than those of the coastal plain populations boring in the smaller switch cane. A female from Ballard Co., Kentucky (
Head and Thorax: antennae of both sexes ciliate, laminate, less deeply in female; dorsum scaled, ventral lamination obscured by densely pubescent sensilla; otherwise as in genus description. Forewing: length similar in both sexes = 11.5–17.0 mm; dorsal ground color in both sexes variable but usually rather uniform, light tan to gray brown, wood brown, or dull, dark reddish brown, the reddish phase mostly restricted to males; filling of antemedial and postmedial lines concolorous with ground; both lines ranging from obscure in many individuals to defined with blackish scales in some but never especially prominent; antemedial line double, evenly scalloped, straight, meeting the inner margin diagonally; postmedial line finely scalloped basally between veins, defined with darker scales in some and marked outwardly by series of short, dark dashes interrupted with light points on veins, moderately sinuous, excurved around reniform, then meeting inner margin perpendicularly; medial shade obscure in most specimens, otherwise defined with darker scales, angled below reniform and meeting inner margin perpendicularly; subterminal line usually obscure, otherwise generally paler than ground color and preceded by series of darker chevrons filling spaces between veins; terminal line a series of dark spots between veins with narrow, obscure adterminal line concolorous with or paler than ground; fringe concolorous or occasionally slightly reddish; orbicular and reniform stigmata of highly variable coloration and definition, black and strongly contrasting in some, obscure (particularly the orbicular) in others, but usually at least outer margin of reniform light yellowish-cream that contrasts with ground, rarely both reniform and orbicular surrounded by diffuse dark scales in medial area such that both stigmata contrast in their entirety; suborbicular, subreniform, and claviform stigmata absent. The phenotype with dark reniform and orbicular stigmata more prevalent in midwestern populations, occurring much more frequently among females. Hindwing: dorsal surface uniform light brown to dark fuscous with a satiny sheen; discal spot and postmedial line usually obscure, sometimes faintly visible; terminal line a series of fuscous chevron spots, sometimes prominent; fringe pale straw to pinkish buff, conspicuous and strongly contrasting in individuals with dark fuscous hindwings. Abdomen: basal coremata absent. Male genitalia: as in genus description. Female terminalia: as in genus description.
Mature larva approximately 2.5 cm long; head capsule brown, lacking any reticulate patterning, darker than uniformly honey-colored prothoracic shield, which may bear some slightly darker pigmentation at anterior angle of lateral margin; pinacula on T1–T3 as well as those on A1–A7 small, brown, inconspicuous; D2 pinacula at least twice as large as D1 on segment A8, D1 pinacula on A9 widely spaced, D2 pinacula 2 × size of D1, narrowly spaced; anal shield honey-colored, similar to the prothoracic shield; somatic pigmentation of wide pinkish to purplish-brown dorsal bands on each segment terminates at spiracular line, more heavily pigmented on abdominal segments than thoracic segments; remainder of body lacks pigmentation; longitudinal lines absent.
TYPES. Holotype: ♂. Forewing length 14.5 mm [alar expanse 31 mm]. St. Catherines Island, Liberty Co., Georgia; bred ex larva in Arundinaria tecta; pupate: 26 April 1994; emerge: 19 May 1994; E. L. Quinter. [CNC] Paratypes: 167 ♂, 138 ♀. Arkansas. Madison Co., Aurora, War Eagle Creek, 1,360´ [415 m] jct. Rts. 23 &127; 29 Jun 1993; E. L. Quinter (3 ♂, 3 ♀). Georgia. Bartow Co., 5 mi. ese Fairmount, east of Salacoa Rd., south of Salacoa Ck., 34°24´03-12˝N, 84°40´12-16˝W; light traps, 13–14 Jun 2009; James K. Adams (1 ♂). Same locality; light traps, 12–13 Jun 2020; James K. Adams (1 ♂). Liberty Co., St. Catherines Island; 10–15 May 1991; E. Quinter & A. Sharkov (11 ♂, 3 ♀). Same locality; 31 May–3 Jun 1992; E. Quinter & A. Sharkov (12 ♂). Same locality; 18–24 May 1993; Eric Quinter & Erik Neff (3 ♂). Same locality; ex larva in Arundinaria tecta, emerge: 11–31 May 1993, E. L. Quinter (5 ♂, 5 ♀). Same locality; ex larva in Arundinaria tecta; emerge: 18 May–13 Jun 1994; E. L. Quinter (23 ♂, 18 ♀). Same locality; ex larva in Arundinaria tecta; emerge: 15–26 May 1995; E. L. Quinter (9 ♂, 12 ♀). McIntosh Co., Sapelo Island, Airport Rd., 31°25´31˝N, 81°17´00˝W; cane north of hangar; light trap 14–18 May 2020; J. K. Adams & L. Durden (3 ♂). Illinois. Alexander Co., 3 mi. w Tamms, canebrake at McCrite Farm cemetery, T15S-R2W, Sec. 3; coll. w/flashlight in canebrake; 28 Jun 1997; James R. Wiker (5 ♀). Same locality; ex larva coll. 8 Jun 1999 in Arundinaria gigantea; pupate: 5 Jul, emerge: 13 Jul 1999; James R. Wiker (1 ♀). Same locality; coll. w/flashlight in canebrake; 29 Jun 1999; J. R. Wiker (1 ♂, 2 ♀). Same data; 12 Jul 1999; J. R. Wiker (3 ♂). Jackson Co., 2 mi. e Pomona, east end Co. Rd. 379, Cedar Lake boat ramp area, T10S-R2W, Sec. 26; coll. w/flashlight in canebrake; 28 Jun 1999; J. R. Wiker (1 ♀). Same locality; ex larva coll. 21 May 1997 in Arundinaria gigantea; pupate: early Jun, emerge: 26 Jun–1 Jul 1997; J. R. Wiker (4 ♂, 1 ♀). Jackson Co., Cave Valley, old GM&O row, T10S-R2W, Sec. 21 north of Pomona; ex larva coll. 21 May 1997 in Arundinaria gigantea; emerge: 20 Jun 1997; J. R. Wiker (1 ♀). Same locality; ex larva in Arundinaria gigantea; emerge: 19 Jun 2000; E. L. Quinter & J. R. Wiker (1 ♂). Pope Co., east end Avery Lake, cane site #3; UV light trap 22 Jul 2000; Wm. R. Black, Jr. (2 ♂). Union Co., Pine Hills campground area, T11S-R3W, Sec. 34; coll. w/flashlight in canebrake; 22 Jun 1997; J. R. Wiker (3 ♂). Kentucky. Ballard Co., Rt. 121 SE Wickliffe, Stovall Creek; netted w/headlamp in canebrake; 15 Jun 1999; E. L. Quinter & J. R. Wiker (26 ♂, 19 ♀). Same locality; ex larva in Arundinaria gigantea; emerge: 25 Jun–4 Jul 1999; E. L. Quinter (7 ♂, 4 ♀). Boone Co., Big Bone Lick State Park, 38°53´14.3˝ N 84°45´04.8˝W; UV trap #2, 22 Jul 2022; Loran Gibson (2 ♂, 1 ♀). Same locality, 38°53´16.7˝ N, 84°44´52.1˝ W; UV trap #6, 26 Jul 2022; L. Gibson (2 ♂). Same locality; 38°53´17.6˝ N, 84°44´54.3˝ W; UV trap #27, 10 Jul 2023; L. Gibson (3 ♂). Same locality; 38°53´12.92˝ N, 84°45´01.11˝ W; UV trap #8, 10 Jul 2023; L. Gibson (1 ♂). Same locality; 38°53´08.0˝ N, 84°45´03.8˝ W; UV trap #5, 10 Jul 2023; L. Gibson (1 ♂). Carlisle Co., Sandy Branch, E old GM&O right of way, Westvaco WMA, 5 mi. sw Bardwell; 14 Jun 1999; E. Quinter, W. Black & J. Wiker (1 ♀). Same locality; ex larva in Arundinaria gigantea; emerge: 23–29 Jun 1999; E. L. Quinter (2 ♂, 5 ♀). Fulton Co., Rt. 94 E of Cayce; ex larva in Arundinaria gigantea; emerge: 24–27 Jun 1999; E. L. Quinter (2 ♂, 1 ♀). Graves Co., Rt. 849, railroad near Boaz, east of Rt. 45; ex larva in Arundinaria gigantea; emerge: 1 Jul 1999; E. L. Quinter (2 ♀). Livingston Co., Dyer Hill Creek, Rt. 60 sw Buma, east of Birdsville; ex larva in Arundinaria gigantea; emerge: 25 May–6 Jul 1999; E. L. Quinter (4 ♂, 2 ♀). McCracken Co., Massac Creek, 0.5 mi. s Rt. 60; ex larva in Arundinaria gigantea; emerge: 24 Jun–2 Jul 1999; E. L. Quinter (4 ♀). Louisiana. St. Tammany Parish, 4.2 mi. ne Abita Springs, Sec. 24 T6S, R12E; 24 May 2004; V. A. Brou (1 ♀). Database # CNC LEP 00057808. West Feliciana Parish, Weyanoke; UV light 15 Jun 1991; V. A. Brou (1 ♀). Mississippi. Hinds Co., Bolton; 13 Jun 1960; Edna Lancaster (1 ♀). Missouri. Barry Co., Roaring River State Park; Ozark Plateau flora; at blacklight 3 Jul 1988; Rae Letsinger (1 ♀). Same locality; 1,000´ [305 m] 26 Jun 1993; E. Quinter & R. Letsinger (5 ♀). Wayne Co., Mark Twain National Forest, 400´ [122 m] jct. Rt. 49 & Black River, 2.7 mi. w Williamsville, Browns Crossing; 19–23 Jun, 1993; E. L. Quinter (2 ♂, 5 ♀). Same locality; larva collected in Arundinaria gigantea 21–22 Jun 1993; emerge: 18 Jul 1993; E. L. Quinter (1 ♀). Same locality; larvae collected in Arundinaria gigantea 20–25 May 1994; pupate: 2–11 Jun, emerge: 19–26 Jun 1994; E. L. Quinter (2 ♂, 1 ♀). Same locality; ex larva in Arundinaria gigantea; emerge: 25 May–4 Jun 1995; E. L. Quinter (4 ♂, 7 ♀). Same locality; ex larva in Arundinaria gigantea; emerge: 30 May–22 Jun 1996; E. L. Quinter (10 ♂, 9 ♀). Same locality; ex larva in Arundinaria gigantea; emerge: 8–22 Jun 2000; E. L. Quinter & J. R. Wiker (1 ♂, 3 ♀). North Carolina. Cabarrus Co., Concord; 10 Jun 2015; D. Willis (1 ♀). Buncombe Co., French Broad River, Walnut Island River Park, 1,730´ [527 m] 35.7339, -82.6579; ex larva coll. 7 May 2021 on Arundinaria gigantea; pupate: 4 Jun 2021, emerge: 23 Jun 2021; Quinter, McBride, Petranka & Elkin (1 ♂). Macon Co., Rose Creek at Furman Welch Rd., Welch property, 1,946´ [593 m] 35.2559, -83.438; 7 Jul 2021 at 15w BLT; A. E. McBride (1 ♂). Macon Co., along Little Tennessee R. at Hall Farm Rd., 1,953´ [595 m] 35.2599, -83.4184; 7 Jul 2021 at 15w BLT; A. E. McBride (2 ♂). Madison Co., French Broad River at Hot Springs Spa, 1,319´ [402 m] 35.8952, -82.8243; ex larva collected 7 May 2021 on 1 year old culm of Arundinaria gigantea; pupate: 28 May 2021, emerge: 14 Jun 2021; McBride, Quinter, Petranka & Elkin (1 ♀). Swain Co., Little Tennessee River, Lower Needmore Road, 1,794´ [547 m] 35.3247, -83.5212; 8 Jul 2021, netted in flight; A. E. McBride & J. Petranka (1 ♀). Swain Co., Tuckasegee River, Ela Public Access Area, 1,784´ [544 m] 35.4527, -83.3923; ex larva collected 2–6 May 2021 on 1 year old culm of Arundinaria gigantea; pupate: 23 May–1 Jun 2021, emerge: 9–20 Jun 2021; A. E. McBride & E. L. Quinter (2 ♂, 3 ♀). Tennessee. “Tenn. 4.25” / “Collection STKemp” / “J. B. Smith Collection Rutgers” (1 ♂). Jackson Co., Cordell Hull Lake, Big Bottom; ex larva in Arundinaria gigantea; emerge: 4–23 Jun 1995; E. L. Quinter (1 ♂, 1 ♀). Same locality; MV light 1–5 Jul 1996; Eric Quinter & Scott Francis (1 ♂, 3 ♀). Same locality; ex larva in Arundinaria gigantea; pupate: 19 May–10 Jun 1998, emerge: 4–24 Jun 1998; E. L. Quinter (2 ♂, 2 ♀). Virginia. Lee Co., Dry Creek Barrens, The Cedars Natural Area; 19 Jun 2000; C. S. Hobson (2 ♂). AMNH, CMNH, CNC, INHS, LACM, MCLB, NHML, NYSM, UCD, ULK, USNM, VADNH, YPM, AEM, ELQ, GDW, JBS, JKA, JRW, LDG, VAB.
The larva utilizes both switch cane (Arundinaria tecta) and giant cane (A.gigantea). It is also the only species in its genus group to utilize both switch cane and giant cane as its larval host plants. Most other members of the genus group appear to be restricted to single species of cane. Numerous larvae of both Arundinariaphila hayesi and Cherokeea slotteni were observed feeding beneath axillary shoot sheaths 1–5 April 1994 on St. Catherines Island, Georgia. Most were apparently in the 3rd or early 4th instars. They had already dispersed from under the leaf sheaths on the primary culms where they had overwintered as eggs. A bit earlier in the season of the previous year, 1st and 2nd instars had been observed feeding gregariously upon the embryonic axillary shoots developing beneath the primary leaf sheaths appressed to the culms at the bases of the nodes. As rapidly developing green axillary shoots extend beyond the old, dried leaf sheaths, these young larvae feed both internally and externally upon the tender new growth while remaining fully concealed within the old shoot sheaths branching from the main culm. Both species exhibit identical feeding modes, at least in the earlier instars. Larvae typically relocate down the culms, feeding in synchrony with the development of the axillary shoots. Most larvae seen at this date were situated about one meter above the ground.
Axillary shoots are present on year-old culms as well as older culms, and the shoot development progresses from the terminal node toward the base of the culm, although axillary shoots are not produced at all from the several most basal nodes. By early April, a small percentage of the older larvae of both species have already dispersed to entirely new culms beginning to emerge from the soil, but these are still quite small, in short supply, and mostly limited to open, sunlit areas on the periphery of the canebrake where the youngest rhizomes are very slender. Most C. slotteni larvae are able to complete their development in the axillary shoots, but the larger A.hayesi larvae are generally obligated to locate new culms to obtain sufficient food. The adult flight period probably extends for approximately one month at any given locality. Coastal populations fly about a month earlier than interior populations, from mid-May through mid-June. Peak flight activity for interior populations occurs from mid-June to early July, with stragglers as late as the third week of July. Arundinariaphila hayesi is the most common and most widely distributed species of all the Arundinaria-feeding genera, occurring throughout almost the entire distribution of cane in the southeastern United States. Arundinariaphila hayesi is found from Missouri east to Kentucky and North Carolina, and south to Florida and eastern Texas.
GENUS Cherokeea Quinter and Sullivan
Cherokeea Quinter and Sullivan, 2014, ZooKeys 421: 183.
Type species: Cherokeea attakullakulla Sullivan and Quinter, 2014 by original designation.
Five species comprise the genus Cherokeea. The type species C. attakullakulla was described in 2014, three new species are described below, and C. trigona (Smith) is included as a new combination. The dramatic range in physical size of Cherokeea adults, from the diminutive C. attakullakulla to the robust C. trigona, is striking. The evidence that these five species comprise a monophyletic unit is best appreciated when their genitalic morphology is compared, their exclusive utilization of the uniquely North American host plant genus Arundinaria is taken into account, and their geographic distribution is analyzed. Particularly striking is the near perfect degree to which their documented occurrence coincides with the distribution of the three species of their hosts, the molecular phylogeny of which has been thoroughly detailed recently in the botanical literature, discussed above in the introduction to the Arundinaria-feeding genera. Collectively, Cherokeea occupy the entire range of cane in the southeastern United States, yet the distribution of each individual species of Cherokeea is limited to that of its singular host species, with only one known exception.
DIAGNOSIS OF GENUS CHEROKEEA. Marked bilateral asymmetry of structures in the male genitalia is uncommon in Noctuidae, which in combination with the exceptionally complex structures characteristic of many Apameini, is a distinctive but not entirely unique feature of Cherokeea. Asymmetric male genitalia are also found in two other apameine genera, Loscopia Beck and Protapamea Quinter, the latter a distantly related Arundinaria specialist. The following combination of features defines Cherokeea: 1) specialized adaptation to endophagous utilization solely of Arundinaria; 2) asymmetry in the costal lobes of the male valvae, conspicuous and extreme in some species, independent asymmetry of the digiti in others, and a distinctive non-reflexed, auriculate cucullus in all species; 3) absence of clavi on the male valves, lack of a freestanding dorsal arm of the clasper, and auriculate shape of the cucullus readily distinguish Cherokeea from Arundinariaphila, which has a trumpet-shaped cucullus and is the only other genus that is similar in appearance and sympatric with it; 4) unlike Arundinariaphila, R3+R4 is stalked for ⅓ to ½ the distance from the areole to the margin; 5) female with a rugose appendix bursae, heavily sclerotized in most, and a medially corrugated ductus bursae present in all species. Polymorphism of adult forewing pattern is dramatic in some Cherokeea species. Otherwise, the superficial appearance of adults of this genus extends across such a wide range of variability that it is difficult to characterize the group. The adults are cryptically colored and patterned and range in forewing length from 9.6 mm to 20.8 mm.
REDESCRIPTION OF GENUS CHEROKEEA. The genus Cherokeea was described in 2014 to accommodate the single new taxon C. attakullakulla. Although it was recognized that C. attakullakulla was related to several other undescribed species which had been under study for a number of years, they were not recognized as congeneric at the time. Subsequently, re-evaluation of the genitalia suggests that C. attakullakulla, despite its distinct superficial appearance, is likely a sister species of C. slotteni, described below. With this revelation, it is apparent that all five of these species are monophyletic. Cherokeea is redescribed below, modified from
Head: antennae of both sexes ciliate, laminate, less deeply in female; dorsum scaled; ventral lamination obscured by densely pubescent sensilla; eye smooth, rounded; haustellum unreduced; labial palpus laterally flattened, upcurved; first segment swollen basally, arching slightly upward, slightly more than half as long as second segment; second segment straight, about as long as eye width, scales broad; third segment ⅓ × length of second segment, and projecting slightly anteriorly, scales narrow; frons convex, unmodified, with central dense tuft of converging spatulate scales overlaying fine, converging hairs. Thorax: vestiture of prothoracic collar, patagia, mesothorax, and metathorax concolorous, a mixture of coarse spatulate, thin apically spatulate, and hairlike scales; mesoscutellar crest prominent, metascutellar tuft less so. Wings: forewing elongate, length 2.0 × width, apex acutely rounded, venation normal apameine as figured in
All five Cherokeea species share similar food resources and the same feeding mode yet have evolved means of avoiding direct competition with one another. All overwinter as eggs having been inserted by the parent female the previous season under the tightly appressed leaf sheaths arising from the base of each node of the woody culm. The timing of egg hatching varies with the species and is correlated with the development of the embryonic axillary shoots concealed within the previous season’s leaf sheaths. The smallest culms, typically situated on the periphery of the canebrake, receive more sunlight than those in the interior of the brake and consequently are the first culms to commence axillary shoot development early in the season. Since they are also the youngest and smallest culms of the colony, they produce relatively little new axillary growth, and so can support only the smallest of the Cherokeea species. The tiny C. attakullakulla is able to derive sufficient nutrition solely from the tender young axillary shoots to complete its entire larval development. The slightly larger C. slotteni commonly abandons the initial culm after devouring the few young axillary shoots available and seeks additional culms for adequate food. Both of these smaller species are the earliest to eclose as adults in their respective habitats. As the season advances, larger, older culms in the interior of the brake begin to produce more abundant new axillary growth, coinciding with egg hatching of some of the larger species of Cherokeea. Axillary shoot development always proceeds from the most terminal node of a given culm to progressively lower nodes, so on taller, older culms with more nodes, this translates to more available food on any one culm. Annual new culm production in dense, mature canebrakes of giant cane is extremely limited. Only a tiny fraction of the total number of previous years’ culms are renewed each year, and these new culms occur as single isolated culms randomly scattered among the vast numbers of older culms. It is these few tender new emerging culms, which may be 2.5 centimeters or more in diameter, that are required for the larva of the largest species, Cherokeea trigona, to complete its development. Of all the Cherokeea, eggs of C. trigona perhaps hatch latest in the season, by which time there is more abundant axillary shoot growth to feed their larvae for an extended period. These larvae develop much more slowly than those of the other species. Long after the other species of Cherokeea have emerged as adults, mated, and died, C. trigona larvae are still feeding. Those relatively few individuals that manage to locate the sparsely scattered new emergent culms survive to produce adults which fly in August and into September.
KEY TO SPECIES OF CHEROKEEA (ADULTS)
NOTE—Female genitalia of Cherokeea species are not included in the key because they are morphologically quite similar (other than size), and it is easier to distinguish females based on superficial appearance and association with males. For descriptions of female characters, refer to the text below.
1. Small species, forewing length 13.5 mm or less; male valves with dorsal lobes of sacculi strongly bilaterally asymmetrical 2
— Larger species, forewing length greater than 13.5 mm; male valves with asymmetrical digiti or with dorsal lobes of sacculi bilaterally sub-symmetrical 3
2. Forewing length 9.8–11.7 mm; forewings gray, mottled with darker gray, black, and brown in some; southern Appalachian Mountains only attakullakulla
— Forewing length 11.5–13.5 mm; forewings dull orange-red to dark red-brown; southern Atlantic and Gulf Coastal Plains and southern Appalachian Mountains slotteni
3. Southern Atlantic and Gulf Coastal Plains; male valves with symmetrical digiti gabrielleae
— Interior distribution, not on Atlantic or Gulf Coastal Plains; male valves with strongly asymmetrical digiti 4
4. Widespread species; male valves with costal lobes of sacculi bilaterally sub-symmetrical, digiti strongly asymmetrical with multiple prongs on right digitus trigona
— Restricted to higher elevation habitats in southern Appalachian Mountains; male valves with costal lobes of sacculi strongly bilaterally asymmetrical with right and left valves bearing prominent, dissimilar, dorsally oriented saccular processes; digiti asymmetrical with two prongs on right digitus odonnellae
Key to Species of Arundinariaphila and Cherokeea (Larvae)
1. Larva with pale, longitudinal mid-dorsal and subdorsal stripes 2
— Larva lacking pale longitudinal stripes 4
2. D2 setal bases closer together than D1 setal bases on segment A8; D2 and D1 pinacula on segment A8 similar in size; D2 pinacula on segment A9 fused into transverse bar Cherokeea trigona
— D2 and D1 setal bases on segment A8 the same distance apart; D2 pinacula at least twice as large as D1 on segment A8; D2 pinacula on segment A9 not fused 3
3. Coastal plain species associated with switch cane Cherokeea gabrielleae
— Southern Appalachian Mountain species associated with hill cane Cherokeea odonnellae
4. D1 and D2 pinacula on segment A9 all fused into one large plate 5
— D1 and D2 pinacula on segment A9 not fused Arundinariaphila hayesi
5. D2 pinacula slightly larger than D1 on segment A8; D2 pinacula on segment A8 sometimes fused; D1 and D2 pinacula on segments A3 to A6 very large, at least twice as large as spiracle on these segments Cherokeea slotteni
— D2 pinacula at least twice as large as D1 on segment A8; D2 pinacula on segment A8 fused; D1 and D2 pinacula on segments A3 to A6 very tiny, less than half the size of spiracle on these segments Cherokeea attakullakulla
Cherokeea attakullakulla Sullivan & Quinter (Cherokee Moth)
Cherokeea attakullakulla Sullivan and Quinter, 2014, ZooKeys 421: 181–191.
Type locality: Fontana View Estates on Lake Fontana, Swain Co., North Carolina, USA. [USNM]
note—Being the first species named in this genus, C. attakullakulla is the type species by original designation.
It is perhaps unfortunate that C. attakullakulla is the type species of Cherokeea, for it is the most divergent member of the group, differing in a number of ways yet retaining most of the shared diagnostic features. For a detailed diagnosis and description of C. attakullakulla, refer to the original 2014 publication. Otherwise, comparison with the figures of adults and their genitalia herein, together with the key to Cherokeea species, should suffice for recognition of this smallest member.
This tiny species superficially resembles some Neoligia Troubridge and Lafontaine species which are not likely to coexist with C. attakullakulla, and which are immediately distinguished by their genitalic characters. The unrelated eustrotiine moth Protodeltote muscosula (Guenée) in worn condition could also possibly be confused with it. For ease of comparison and placing the species in context with the other members of the genus treated here, the most diagnostic genitalic characters of C. attakullakulla are summarized.
Male terminalia: valves bilaterally asymmetrical with respect to shape of sacculus, costal lobe of left sacculus normal, rounded; of right sacculus, greatly expanded dorsally into free, flattened process that extends distally ½ × length of valve; saccular lobe attached at its distal base to more heavily sclerotized basal sclerite of clasper; juxta an elongated trapezoidal shield, 2 × as tall as basal width. The sclerotized keyhole-like medial protrusion at the base of the juxta mentioned in the original description as possibly autapomorphic appears to be only a variation of the tiny, sclerotized, thimble-like apodeme present in the other Cherokeea species. Like C. slotteni, this species lacks paired basal coremata on the male abdomen characteristic of the larger members of the genus. Female genitalia: The rugose, heavily sclerotized appendix bursae and medially corrugated ductus bursae characteristic of C. attakullakulla is present in all Cherokeea species except C. trigona, which lacks the heavy sclerotization; also, the sclerotized, quadrate, lamella antevaginalis with its w-shaped outline present in this and other members of the genus is diagnostic.
Mature larva approximately 21 mm long, head capsule 1.0 mm wide; mature larva very pale, pinkish-brown; relatively small, chestnut head capsule contrasts with honey-colored prothoracic shield; prothoracic shield edged anteriorly with broad, darker band; dark brown dorsal pinacula are small and inconspicuous, especially on A3 through A6, and ½ × spiracle on these segments; D2 pinacula are fused into a transverse bar on A8; on most, D1 pinacula are also fused into bar on A8, a dark mid-dorsal line connects the two bars, creating overall appearance of an uppercase letter “I;” D1 and D2 pinacula fused into large, dark brown plate on A9; anal shield similar in color to head capsule; L1 located behind and below spiracle on A7; spiracles 2 × as high as broad and light tan with dark brown outline; somatic pigmentation consists of very pale, pinkish-brown dorsal bands on each segment, fading below spiracular line.
In the spring of 2023, AEM succeeded in finding larvae and confirmed the original assumption that the host plant is hill cane (Arundinaria appalachiana). This is the only known host plant for this species.
Based on wild-collected specimen data detailed in the original description together with recent data regarding emergence dates from reared larvae, the adults appear to have a flight period of at least one month, commencing about the third week of May with captures ranging through 24 June. The average pupal period, based on a very limited sample of three larvae that produced adults, is 23 days. Cherokeea attakullakulla is presently known only from Macon, Rutherford, and Swain Counties in North Carolina, and from a single record from Rabun County, Georgia.
Cherokeea slotteni Quinter, NEW SPECIES (Lesser Switch Cane Borer)
Cherokeea slotteni Quinter.
Type locality: St. Catherines Island, Liberty Co., Georgia, USA. [CNC]
note—Named in recognition of Dr. Jeffrey Slotten, who assisted me in finding this species in Florida, and whose perseverance and passionate pursuit of some of our rarer Lepidoptera has contributed much new information.
Although it has similar forewing pattern elements and may have a pale subterminal area, Cherokeea slotteni can be distinguished easily from its sister species C. attakullakulla by its slightly larger size and reddish forewing. A number of shared genitalic characters distinguish this pair from the other members of the genus, which are significantly larger with forewings predominantly shades of tan or brown. Cherokeea slotteni bears a strong superficial resemblance to the unrelated eriopygine species Homorthodes lindseyi (Benjamin), but the ranges of the two species barely overlap, if at all, the genitalia are completely dissimilar, and the eyes of H. lindseyi are hairy.
Head and Thorax: antennae of both sexes ciliate, laminate, less deeply in female; dorsum scaled, ventral lamination obscured by densely pubescent sensilla; otherwise as in genus description. Forewing: length similar in both sexes = 11.5–13.5 mm; dorsum of both sexes rather uniform, distinctly reddish, ranging from orange red-brown to dark liver brown, orange phase tending to be restricted to males, otherwise sexual dimorphism minimal; occasional individuals with medial area toward costa somewhat darker and contrasting, and others irrorate with gray scaling; filling of antemedial and postmedial lines concolorous with ground, both lines ranging from obscure in most to weakly defined with darker scales in some, but never prominent; antemedial line double, evenly scalloped, straight, meeting inner margin diagonally; postmedial line double, defined outwardly by series of tiny black points on veins, each immediately followed by contrasting white or light point, veins occasionally highlighted with salt-and-pepper scaling, especially toward outer margin; postmedial line moderately sinuous, excurved around reniform stigma, then meeting inner margin perpendicularly; medial shade obscure in most specimens, otherwise, defined with darker scales, angled below reniform, and meeting inner margin perpendicularly; subterminal line obscure in many, otherwise, generally paler than ground color and preceded by series of darker chevrons filling spaces between veins; terminal line defined by series of minute black chevrons between veins; adterminal line and fringe concolorous with ground; reniform at least defined along its outer margin by several frequently confluent satellite spots of white or yellowish-cream that contrast with ground; in some, reniform stigma nearly filled with white or light scales with central lunule defined within by dark scales, often with contrasting dark spot in lower portion; orbicular stigma a simple, subovate ring, obscure in most individuals; suborbicular, subreniform, and claviform stigmata absent. Hindwing: dorsum dark fuscous, occasionally lighter, with satiny sheen; discal spot and postmedial line obscure in most, faintly visible in a few; terminal line a nearly confluent series of fuscous chevron spots, often obscure; fringe pale straw to pinkish buff, conspicuous and strongly contrasting in individuals with dark fuscous hindwings. Abdomen: basal coremata absent. Male genitalia: peniculus quadrangular, densely pilose; valves with costal lobe of left sacculus normal, costal lobe of right sacculus bearing very large, flat, unadorned process terminating in distally oriented blunt tip; clasper a long, straight, sclerotized bar with a small spinule (absent on left valve) at base, terminating in distally-oriented point underneath free-standing digitus; digiti symmetrical, flattened, transverse, slightly bifurcate with freestanding ventral arm 2.5 × length of appressed dorsal arm; ampulla small, sparsely setose; editum present; cucullus narrowly necked, auriculate, ventrally reflexed, lightly pilose; corona an apical series of 8–10 strong spines, anal spine absent; pollex absent; uncus linear, narrowly spatulate, with diffuse apical patch of setae, primarily oriented laterad; juxta trapezoidal, 2.5 × higher than basal width; phallus slender, curved with heavy terminal spine; vesica bearing single mesial thornlike cornutus on diverticulum and small subapical patch of slender spicules. Female terminalia: papilla analis conical with pointed apex, not dorsoventrally flattened, lateral margin straight, evenly tapered, upper surface densely covered with fine setae set in tiny membranous circles, lower surface much less setose with shorter hairs; segment A8 2 × as wide as long, anterior apophysis 1 × length of segment A8 and posterior apophysis 1.7 × length of segment A8; sternite A7 an enlarged lightly sclerotized plate, distally truncated, slightly cleft but not forming freestanding lodices; lamella antevaginalis a simple sclerotized sinuous rim, slightly cleft; antrum cup-like, quadrate with w-shaped outline; ductus bursae heavily sclerotized, slightly twisted, 3.4 × as long as wide, 0.6 × length of corpus bursae; appendix bursae an ovoid, nearly round, moderately corrugated diverticulum 0.4 × length of corpus bursae, dorsally oriented and slightly to right, ductus seminalis at posterior apex; corpus bursae oval, membranous, 2 × as long as wide with multiple long signa running length of bursa.
Mature larva approximately 2.20 cm long; head capsule brown with reticulate patterning, slightly darker than honey-colored prothoracic shield, which is edged with darker pigmentation; pinacula on T1–T3 as well as those on A1–A7 pale honey color, small, inconspicuous; D2 pinacula on A8 are slightly larger than D1 pinacula and sometimes fused, D1 and D2 on A9 fused into one large plate; anal shield brown, heavily sclerotized, about the same color as head capsule; somatic pigmentation consists of wide pinkish to purplish-brown dorsal bands on each segment terminating at the spiracular line, more heavily pigmented on the abdominal segments than on the thoracic segments; remainder of body devoid of pigmentation; longitudinal lines absent.
TYPES. Holotype: ♂. Forewing length 12 mm [alar expanse 26 mm]. St. Catherines Island, Liberty Co., Georgia, bred ex larva in Arundinaria tecta; pupate: April 12, 1994 emerge: May 1, 1994 E. L. Quinter, collector. [CNC] Paratypes: 88 ♂, 67 ♀. Florida. Liberty Co., Apalachicola National Forest, Revels Landing off Rd. 67; bred ex larva in Arundinaria tecta; pupate: 18 Apr 2001, emerge: 4 May 2001; E. L. Quinter & J. Slotten (1 ♀). Georgia. Liberty Co., St. Catherines Island; 10–15 May 1991; UV trap; E. Quinter & A. Sharkov (2 ♂, 1 ♀). Same locality; 18–24 May 1993; UV trap; E. Quinter & E. Neff (3 ♂, 2 ♀). Same locality; bred ex larva in Arundinaria tecta; emerge: 16 May 1993; E. Quinter (1 ♂). Same locality; bred ex larva in Arundinaria tecta; pupate: 10 Apr–6 May 1994, emerge: 28 Apr–28 May 1994; E. Quinter (9 ♂, 15 ♀). Same locality; bred ex larva in Arundinaria tecta; pupate: 14 Apr–1 May 1995, emerge: 3–21 May 1995; E. Quinter (12 ♂, 9 ♀). Louisiana. St. Tammany Parish, 4.2 mi. ne Abita Springs, Sec.24 T6S, R12E (UV); 18 May 1993, 15 May 1999, 24 Apr 2000, 22 May 2004, 17 May 2006, 20 May 2010, 25 May 2011 V. A. Brou (7 ♂); 18 May 1985, 26 May 1985, 1 Jun 1988. 28 May 1993, 14 May 1994, 1 Jun 2001, 15 May 2008, 20 May 2010, 21 May 2010, 27 Apr 2011; V. A. Brou (10 ♀). North Carolina. Brunswick Co., 1 mi. ne Pretty Pond; 15-watt UV trap; Leiophyllum/Slash Pine woodland; 1 Jun 1995; J. B. Sullivan & R. Broadwell (1 ♀). Brunswick Co., Boiling Springs Lakes, Half Hell Swamp; swamp forest; 15-watt UV trap; 26 May 1995; R. Broadwell & J. B. Sullivan (1 ♂). Brunswick Co., Leland; 20 Jun 1948; O. Buchholz (1 ♀). Carteret Co.; 15 Jun 1972; J. B. Sullivan (1 ♀). Craven Co., Croatan National Forest Rt. 147; 31 May 1997; E. L. Quinter (2 ♀). Same locality; 6 Jun 1996; J. B. Sullivan, 15-watt UV trap (1 ♂, 1 ♀). Same locality; 2–27 Jun 1997; J. B. Sullivan, 15-watt UV trap (9 ♂, 2 ♀). Craven Co., North Harlow; 27 May–8 Jun 1996; J. B. Sullivan (4 ♂, 4 ♀). Craven Co., County Rd. 1717; 16 Jun 1983; J. B. Sullivan (1 ♀). Craven Co., Croatan National Forest Rd. 3046, Gum Branch Rd.; 5 Jun 1998; J. B. Sullivan (1 ♂). Craven Co., Croatan National Forest Rd. 169; Little Deep Creek Rd.; 9 Jun 1999; J. B. Sullivan (1 ♂). Craven Co., Pinecliff; 17 Jun 1983; J. B. Sullivan (2 ♂). Cumberland Co., Ft. Bragg, MacPherson Creek; Neonympha type locality; borrow pit sedge marsh; 20 Jun 2001; J. B. Sullivan & S. P. Hall (1 ♂). Hoke Co., Ft. Bragg, Jumping Run Creek at FB 12; UV trap; Carex stricta bog; 20 May 2001; J. B. Sullivan & S. P. Hall (2 ♂). Jones Co., Croatan National Forest, north of Stella, Haywood Landing; 15-watt UV trap; hardwoods; 11 Jun 1996; J. B. Sullivan (7 ♂, 3 ♀). Same locality; 2 Jun 1998; J. B. Sullivan (1 ♂). Macon Co., Rose Creek at Furman Welch Rd./Welch property, 35.2559, -83.438, 1,946´ [593 m]; 7 Jul 2021 at 15w BLT, A. E. McBride, collector (2 ♂, 1 ♀). New Hanover Co., Fort Fisher Maritime Forest; coastal fringe evergreen forest; 15-watt UV trap; 17 May 1994; J. B. Sullivan, R. Broadwell & B. Smith (1 ♂). Onslow Co., Camp Lejeune, Corn Landing; 15-watt UV trap; hardwoods; 30 May 1995; J. B. Sullivan (1 ♂). Onslow Co., Verona Loop Rd.; 11-watt UV trap; mature pine, hardwoods, cane; 30 May 1995; J. B. Sullivan (2 ♂). Swain Co., Little Tennessee River, Lower Needmore Road, 1,787´ [545 m], 35.3286, -83.522; bred ex larva collected 4–5 May 2021 on 1 yr. old culms of Arundinaria appalachiana; pupate: 10–28 May 2021, emerge: 28 May–13 Jun 2021; E. L. Quinter & A. E. McBride (7 ♂, 6 ♀). Same locality, ex larva 25 Apr 2023, A. E. McBride, collector; collected on Arundinaria appalachiana and reared on ornamental bamboo, eclosed 23–27 May 2023 (3 ♂, 1 ♀). Swain Co., Little Tennessee River, Needmore Road SGL, 35.3439, -83.5225 1,750´ [533 m]; ex larva 9 May 2022, A. E. McBride, collector; collected on Arundinaria appalachiana and reared on ornamental bamboo, eclosed 27 May–1 Jun 2022 (2 ♀). Virginia. City of Suffolk, Great Dismal Swamp National Wildlife Refuge, ex larva in Arundinaria tecta; emerge: 14–30 May 1998; E. L. Quinter (8 ♂); same locality; emerge: 27 May 2000; E. L. Quinter (1 ♀). City of Virginia Beach, NAS Oceana, Oceana Pond, ca. 0.6 km se jct. Oceana Blvd. & Bells Rd.; 12 Jun 2001, UV sheet; K. L. Derge, G. M. Pisoni, & A. C. Chazai (2 ♀). AMNH, CMNH, CNC, INHS, LACM, MCLB, NHML, NYSM, UCD, USNM, VADNH, YPM, AEM, ELQ, JBS, JRW, VAB.
Cherokeea slotteni was believed to be restricted to boring in switch cane (Arundinaria tecta), inasmuch as all known distribution records for adults were from lowland habitats on the southeastern Atlantic and Gulf coastal plains. AEM and ELQ recently discovered that this species also utilizes the smaller hill cane (A.appalachiana) at elevations above approximately 500 meters in western North Carolina. There is no evidence that C. slotteni also utilizes the widespread giant cane. This higher elevation population represents an incursion across the Piedmont of this otherwise coastal species.
Discussion of the early stages of C. slotteni is partly addressed under the genus Cherokeea, and also in the treatment of Arundinariaphila hayesi. Host plant symptoms resulting from larval feeding is essentially the same for all members of the genus, discussed in the treatment of C. odonnellae; however, C. slotteni differs in that it normally completes its larval development in the axillary shoots of older culms and is not obligated to seek out tender new culms unlike its larger congeners. The pupal period of C. slotteni averages 18–20 days. Adults emerge from pupae mid-day to early evening, prior to dusk.
Coastal populations are on the wing from late April through late June, with peak flight around mid-May. The montane populations in western North Carolina fly about a month later. Records there range from 28 May to 7 July. Cherokeea slotteni is a much more widespread species than C. attakullakulla, which is apparently restricted to elevations above 500 meters in the southern Appalachians where the two species can occur together. Cherokeea slotteni is moderately common in coastal canebrakes from southeastern Virginia to southeastern Louisiana but appears to be absent from peninsular Florida where cane is uncommon. Although mainly limited to coastal plain habitats with a westward extension into and beyond the Piedmont in North Carolina, C. slotteni completely overlaps the Atlantic Coastal Plain portion of the range of Arundinariaphila hayesi. However, the range of C. slotteni extends much farther north along the coast than that of A.hayesi. Here C. slotteni is sympatric with C. gabrielleae. It is unclear what historical or ecological factors may account for the apparent exclusion of A.hayesi but not C. slotteni from the range of C. gabrielleae, since all three species share the same host plant on the Coastal Plain.
Cherokeea gabrielleae Quinter, NEW SPECIES (Switch Cane Borer)
Cherokeea gabrielleae Quinter.
Type locality: Great Dismal Swamp National Wildlife Refuge, Suffolk, Virginia, USA. [CNC]
NOTE—Named for Gabrielle Tannor, who assisted with field work involving this species, for her inspiration and exemplary dedication to her many students.
This polymorphic, highly variable species exhibits no discernible sexual dimorphism, although light morphs are significantly more prevalent among females, accounting for approximately half of the individuals. Cherokeea gabrielleae is distinguished from its congeners by being the only species without either asymmetrical digiti or dramatic asymmetry of costal saccular processes in the male valves. Its allopatric coastal plain distribution precludes confusion with C. trigona or C. odonnellae, certain phenotypes of which may resemble it superficially. Arundinariaphila hayesi could be confused with C. gabrielleae, based upon superficial appearance, were it not for the apparent near allopatry of these two species. Their genitalic characters immediately distinguish these two. It is however usually unnecessary to examine these structures due to a fortuitous subtle feature in the forewing maculation of C. gabrielleae which will consistently distinguish it from all but the very darkest individuals of A.hayesi. The near complete row of dark intervenular spots preceding the subterminal line on the forewing, characteristic of C. odonnellae, is reduced in C. gabrielleae to just two prominent, almost black spots. The first smaller spot lies between veins M1 and M2; the second, usually larger spot, lies between M2 and M3. The remainder of the row of spots is obsolescent, with only a tiny spot sometimes present in the tornal angle, and perhaps a few darkened scales preceding a vaguely defined apical patch. None of these spots are a noticeable feature of A.hayesi, and only on individuals of A.hayesi or C. gabrielleae having uniformly dark brown forewings might their presence or absence not be discernible.
Head and Thorax: antennae of both sexes ciliate, laminate, less deeply in female; dorsum scaled, ventral lamination obscured by densely pubescent sensilla; otherwise as in genus description. Forewing: length similar in both sexes = 12.3–16.2 mm; polymorphic, appearance highly variable and difficult to characterize; dorsal surface ground light tan, reddish brown, wood brown, or dark chocolate brown, mottled or uniformly smooth, with or without contrasting stigmata; filling of antemedial and postmedial lines concolorous with ground, both lines ranging from obscure in most to weakly defined with darker scales in some, but never prominent; antemedial line when visible double, evenly scalloped, excurved, meeting inner margin diagonally; postmedial line double, obscure in most, occasionally defined outwardly by series of tiny, usually obscure, black points on veins; postmedial line, when visible, sinuous, excurved around reniform, then meeting inner margin perpendicularly; medial shade obscure in most, otherwise defined with darker scales, angled below reniform, meeting inner margin perpendicularly; subterminal line obscure, defined by aforementioned diagnostic dark intervenular spots which precede it; terminal line obscure; adterminal line and fringe concolorous with ground; reniform and orbicular stigmata obscure in individuals with very dark ground color, paler than ground color and somewhat contrasting in others, and strongly contrasting black on tan background in minor percentage of individuals, which also possess prominent black claviform streak; suborbicular absent in all morphs. Hindwing: dorsal surface dark fuscous, lighter occasionally; discal spot and postmedial line obscure in most; terminal line often obscure, a faint, nearly confluent series of fuscous chevrons; fringe usually paler, tan or slightly pinkish. Abdomen: paired, lateral coremata present on 2A, fully developed. Male genitalia: peniculus quadrangular, densely hairy; valves with costal lobes of sacculus subsymmetrical, right lobe somewhat more produced and rounded than left, sclerotized costal ridge of both valves distad of lobe bearing a single blunt tooth; clasper an inconspicuous, appressed, basal sclerite terminating in distally oriented point near base of freestanding digitus; digiti symmetrical, transverse, flattened, with short, thumb-like projection caudally at base; ampulla absent; editum present; cucullus narrowly necked, ventrally reflexed, auriculate, lightly pilose centrally, more heavily so proximad of corona; corona nearly complete series of heavy spines, diminishing in length from apex toward ventral angle; anal spine and pollex absent, but three very long, strong bristles projecting distally from ventral edge of cucullus; uncus linear, narrowly spatulate, with apical dense patch of proximally oriented setae; juxta trapezoidal, 2.0 × higher than basal width; phallus slender, curved distally with single, broad, terminal tooth; vesica lacking thornlike cornuti, but with dense, elongate, ordered, mesial patch of slender spicules on dorsal surface. Female terminalia: papilla analis conical with pointed apex, somewhat dorsoventrally flattened, lateral margin straight, evenly tapered, both surfaces densely covered with fine, pubescent setae; segment A8 1.5 × as wide as long, anterior apophysis 1 × length of segment A8 and posterior apophysis 1.8 × length of segment A8; sternite A7 moderately sclerotized, deeply concave distally, but not forming freestanding lodices; lamella antevaginalis heavily sclerotized with concave posterior rim and concave anterior margin; antrum cup-like, quadrate with w-shaped outline; ductus bursae heavily corrugated, narrow distally, widening proximally to 2 × distal width, 0.85 × length of corpus bursae; appendix bursae moderately corrugated, arising posteriorly on left, 0.25 × length corpus bursae, ductus seminalis at posterior edge; corpus bursae membranous, ovoid, 1.2 × as long as wide, lacking conspicuous signa.
Mature larva approximately 2.7 cm long; dark brown to nearly black head capsule lacks distinct patterning and strongly contrasts with light honey-colored prothoracic shield, which usually bears some darker pigmentation along lateral margins; pinacula on T1–T3 and on A1–A7 light brown, very small, inconspicuous; pinacula on A8 more prominent, with setal bases of D1 and D2 same distance apart, D2 pinacula 2 × D1 on segment A8; D2 pinacula on segment A9 not fused; anal shield brown, with heavy sclerotization; somatic pigmentation consists of wide pinkish-brown dorsal bands on each segment terminating at spiracular line, more solidly pigmented on abdominal segments than thoracic segments; remainder of body lacks pigmentation; pale, longitudinal mid-dorsal and faint subdorsal lines interrupt pigmented bands on T1–A8; larva is practically indistinguishable from that of its widely allopatric sister species, C. odonnellae.
TYPES. Holotype: ♂. Forewing length 13.5 mm [alar expanse 30 mm]. Great Dismal Swamp National Wildlife Refuge, Suffolk, Virginia, bred ex larva in Arundinaria tecta, pupate: 19 June 1998, emerge: 4 July 1998, E. L. Quinter. [CNC] Paratypes: 90 ♂, 94 ♀. New Jersey. Cumberland Co., Port Norris, 1761 Main Street, 39.2449, -75.0305, 10´ [3 m]; netted at bamboo 12 Jul 2019; A. E. McBride & D. Schweitzer (8 ♂, 9 ♀). Same locality; ex larva coll. on year-old culm of Pseudosasa japonica 22 May 2021, eclosed 22 Jun–8 Jul 2021; A. E. McBride & D. Schweitzer (1 ♂, 3 ♀). North Carolina. Carteret Co., International Paper office site; 2 Jul 1972; J. B. Sullivan (1 ♀). Carteret Co., jct. St. Hwy 101 & 181; 27 Jul 1978; J. B. Sullivan (1 ♂). Carteret Co., Millis Swamp; bait trap 28 Jul 1991; J. B. Sullivan (1 ♂). Craven Co., North Harlow; 30 Jun 1990; J. B. Sullivan (1 ♀). Craven Co., Croatan National Forest Road 3046; bred ex larva in Arundinaria tecta, pupate: 2–9 Jun 1998, emerge: 20–24 Jun 1998; E. L. Quinter (3 ♂, 1 ♀). Martin Co., Bull Run Island; 17 Jul 1996; canebrake on levee next to Devil’s Gut waterway, Stephen Hall (1 ♂). Martin Co., Devil’s Gut TNC Preserve; 18 Jul 1996; cane patch on floodplain ridge next to backswamp, Stephen Hall (1 ♂). Washington Co., Pettigrew State Park, Pocosin Natural Area; 21 Jul 1993; pond pine woodland next to shoreline Lake Phelps, Stephen Hall (1 ♀). Washington Co., Pettigrew State Park, Moccasin Overlook Trail; 21 Jul 1993; cane under closed canopy, non-riverine wet hardwood forest next to Lake Phelps, Stephen Hall (1 ♂). Virginia. City of Virginia Beach, NAS Oceana, ca. 1.5 km sse jct. London Br. Road x Potters Rd; canebrake; 20 Sept. 2001, uv; K. L. Derge, S. M. Roble (2 ♂). Great Dismal Swamp National Wildlife Refuge, Suffolk, Virginia, bred ex larva in Arundinaria tecta, emerge: 5–9 Jul 1997; Eric Quinter & Steven Roble (2 ♂). Same locality; bred ex larva in Arundinaria tecta, pupate: 20 May–3 Jul 1998, emerge: 3 Jun–20 Jul 1998; Eric Quinter & Steven Roble (36 ♂, 31♀). Same locality; bred ex larva in Arundinaria tecta, emerge: 1–25 Jul 1999; Eric Quinter & Zhiwei Liu (7 ♂, 11 ♀). Same locality; bred ex larva collected 23–24 Apr 2000 in Arundinaria tecta and reared on artificial diet, emerge: 10 Jun–3 Jul 2000; E. Quinter, D. Quinter & S. Roble (11 ♂, 11 ♀). Same locality; bred ex larva collected 28–30 May 2000 in Arundinaria tecta, emerge: 6–27 Jul 2000; E. Quinter, P. Goldstein & S. Roble (11 ♂, 25 ♀). Same locality; bred ex larva collected 22 May 2001 in Arundinaria tecta, emerge: 24 Jul–3 Aug 2001; Eric Quinter & Jane O’Donnell (3 ♂). Prince George Co., James River NWR, ca. 2.2 km ne jct. 614 x Rte. 10; 8 Aug. 2001, uv; K. L. Derge (1 ♂). AMNH, CMNH, CNC, INHS, LACM, MCLB, NHML, NYSM, UCD, USNM, VADNH, YPM, AEM, ELQ, JBS, JKA, JRW, JRS.
The sole native host plant is switch cane (Arundinaria tecta). The early stages and larval behavior do not differ from the general description given for the genus. Cherokeea gabrielleae appears to have an unusually protracted flight period, with limited records of adult captures ranging from 30 June to an extreme single record from 20 September. Most records suggest a peak flight in July. Much larger samples of reared individuals showed a large majority emerging in July with a few eclosing in early June, likely due to captive indoor rearing.
Peculiarly limited in its distribution, this uncommon species appears mainly confined to the relatively small coastal portion of the eastern reaches of the Roanoke River drainage in southeastern Virginia and northeastern North Carolina and also the eastern reaches of the Santee River drainage in South Carolina, where the moth occurs in pocosins in exclusive association with switch cane, also locally referred to as maiden cane. Cherokeea gabrielleae has the northernmost distribution of all species of the Arundinaria-feeding genera. Historically, switch cane reached its northern limit on the Atlantic Coastal Plain on the lower end of the Delmarva Peninsula and in the Chesapeake Bay area to the west but was largely extirpated there since. However, the presence of ornamental cultivars of closely related exotic bamboo species beyond the range of our native species has allowed C. gabrielleae to colonize isolated patches of these, extending its range to include southern New Jersey. Dale Schweitzer (pers. comm.) has had a persistent colony of C. gabrielleae boring in a patch of what is believed to be Japanese arrow bamboo (Pseudosasa japonica) for many years in his yard in Cumberland County, New Jersey.
Cherokeea gabrielleae appears to reach its southern limit on the coast of South Carolina – with one caveat. A single male collected in Baldwin County, Alabama of what appears very similar to C. gabrielleae but with minor differences in its genitalia may either represent a highly disjunct population of C. gabrielleae, a false “disjunction” resulting from insufficient sampling of coastal populations between South Carolina and Alabama, or an undescribed sibling species. Until additional material becomes available, this issue cannot be resolved. The comprehensive sampling of canebrakes ELQ conducted on St. Catherines Island, Georgia should have revealed the presence of C. gabrielleae, if its actual range indeed spans the coastal plain from South Carolina to Alabama and beyond.
Curiously, this species fills, in almost total allopatry, a geographic niche not utilized by the much commoner and nearly ubiquitous Arundinariaphila hayesi, despite both species’ shared utilization of the same widespread host, begging a historical biogeographic explanation. Loscopia roblei manifests a similarly restricted and nearly identical core distribution to that of C. gabrielleae, possibly for the same unknown reason.
Cherokeea odonnellae Quinter, NEW SPECIES (Hill Cane Borer)
Cherokeea odonnellae Quinter.
Type locality: Lower Needmore Road, Little Tennessee River, 35.3286, -83.522, 1,787´ [545 m], Nantahala National Forest, Swain Co., North Carolina, USA. [CNC]
note—Named for Dr. Jane O’Donnell, my best friend and partner for life, who assisted in the initial discovery of the early stages of this rare species. Her understanding, patience, and unwavering support for my work has greatly facilitated the production of this fascicle.
Cherokeea odonnellae and C. gabrielleae are similar in size and general appearance. The two species might be easily confused were it not for their widely allopatric distributions and major differences in male genitalia, the valves of which bear prominent asymmetrical structures in the former; symmetrical in the latter. Cherokeea odonnellae does not exhibit the extreme polymorphism characteristic of C. gabrielleae. It is much more likely that C. odonnellae will be confused with the superficially similar, widespread, and common Arundinariaphila hayesi, which has a distribution that overlaps C. odonnellae. Arundinariaphila hayesi also utilizes a different host than C. odonnellae and has dramatically different genitalia in both sexes, which can be discerned even without dissection. One characteristic superficial but subtle feature of the forewings of C. odonnellae, best observed under magnification in fresh, unworn specimens, is the presence of pale, almost white scales highlighting the venation. This is especially pronounced beyond the reniform stigma, where the veins cross the subterminal band of dark spots, which are themselves much more prominent in C. odonnellae than in A.hayesi. Another useful superficial feature which reliably distinguishes C. odonnellae from its congeners and from Arundinariaphila hayesi is the subterminal line of the forewing which in C. odonnellae is preceded by a nearly complete series of dark intervenular spots. A very rare pale phenotype (
Head and Thorax: antennae of both sexes ciliate, laminate, less deeply in female; dorsum scaled, ventral lamination obscured by densely pubescent sensilla; otherwise as in genus description. Forewing: length similar in both sexes = 12.7–17.2 mm; dorsal surface ground brown, similar in both sexes, slightly more reddish in some; filling of antemedial and postmedial lines concolorous with ground, both lines ranging from obscure in most to weakly defined with darker scales in some, but never prominent; antemedial line double, evenly scalloped, straight, meeting inner margin diagonally; postmedial line double, defined outwardly by series of tiny, often obscure, black points on veins, each immediately followed by contrasting white or light point; mediocubital and anal veins highlighted with whitish scales, especially beyond postmedial line; postmedial line moderately sinuous, excurved around reniform, then meeting inner margin perpendicularly; medial shade obscure in most specimens, otherwise defined with darker scales, angled below reniform, and meeting inner margin perpendicularly; subterminal line prominent in some, less so in others, generally paler than ground and preceded by nearly complete series of dark intervenular spots which provide useful focus point for recognizing this species; terminal line obscure; adterminal line and fringe concolorous with ground; reniform stigma variable, defined with white scales in some , nearly concolorous, or paler than ground in others; orbicular a simple, subovate ring usually matching color of reniform but less prominent; suborbicular, subreniform, and claviform stigmata absent. Hindwing: dorsal surface dark fuscous, occasionally lighter, with satiny sheen; discal spot and postmedial line obscure in most; terminal line obscure, a faint, nearly confluent series of fuscous chevron spots; fringe pinkish, conspicuous, and strongly contrasting in individuals with dark fuscous hindwings. Abdomen: paired, lateral coremata present on 2A fully developed. Male genitalia: peniculus quadrangular, densely hairy; valves with costal lobe of left sacculus bearing large, unadorned, curved, acutely terminated, clavus-like process, arising distad of saccular base, probably not homologous with true clavus (see discussion of varying interpretations of valval structures as applied by different authors, in the introduction to the Arundinaria-feeding genera); dorsal edge of right sacculus forming sclerotized ridge terminating in acute point, about ⅓ length of valve, bearing densely setose dorsal brush; clasper an inconspicuous, appressed, basal sclerite terminating in distally-oriented point underneath free-standing digitus; heavily sclerotized asymmetrical digiti projecting ventrally; left digitus simple, terminated in three small projections, right digitus terminated in “T” resembling boat hook; ampulla absent; cucullus narrowly necked, auriculate, lightly pilose; corona an apical series of 10–12 strong spines, anal spine absent; pollex absent; uncus narrowly spatulate, with subapical patch of proximally oriented setae; juxta trapezoidal, 2.5 × higher than basal width; phallus slender, curved, lacking a terminal spine, ventral vesica bearing dense, elongate, mesial patch of slender oriented spicules. Female terminalia: papilla analis conical with pointed apex, somewhat dorsoventrally flattened, lateral margin straight, evenly tapered, both surfaces densely covered with fine, pubescent setae; segment A8 1.8 × as wide as long, anterior apophysis 1 × length of segment A8 and posterior apophysis 1.8 × length of segment A8; sternite A7 moderately sclerotized, rounded distally, deeply cleft, but not forming freestanding lodices; lamella antevaginalis heavily sclerotized with sinuous posterior rim and deeply concave anterior margin; antrum cup-like, quadrate with w-shaped outline ; ductus bursae heavily corrugated, narrow, 4.8 × as long as wide, 0.75 × length of corpus bursae; appendix bursae moderately corrugated, arising posteriorly on left, 0.3 × length of corpus bursae, ductus seminalis at posterior apex; corpus bursae membranous, pear-shaped, ovoid, 1.2 × as long as wide, lacking signa.
Mature larva approximately 2.85 cm. long; dark brown to nearly black head capsule lacks distinct patterning and strongly contrasts with light honey-colored prothoracic shield, which lacks contrasting dark pigmentation; pinacula on T1–T3 and A1–A7 light brown, small, inconspicuous; pinacula on A8 more prominent, with setal bases of D1 and D2 same distance apart, D2 pinacula 2 × D1 on segment A8; D2 pinacula on segment A9 not fused; anal shield brown with heavy sclerotization; somatic pigmentation consists of wide, pinkish-brown dorsal bands on each segment terminating at spiracular line, more solidly pigmented on abdominal segments than thoracic segments; remainder of body lacks pigmentation; pale, longitudinal mid-dorsal and faint subdorsal lines interrupt pigmented bands on T1–A8.
TYPES. Holotype: ♀. Forewing length 17 mm [alar expanse 37 mm]. Little Tennessee River, Lower Needmore Road, 35.328552, -83.522011, 1,787 ft., Swain Co., North Carolina, ex larva May 4, 2021, A. E. McBride w/Eric Quinter; collected on 1 yr. old culm of Arundinaria appalachiana, pupated May 27, 2021, eclosed June 15, 2021, larva photographed (#1). [CNC] Paratypes: 12 ♂, 28 ♀. North Carolina. Macon Co., Rose Creek at Furman Welch Rd./Welch property, 35.2559, -83.438, 1,946´ [593 m]; 7 Jul 2021, netted in flight, A. E. McBride, collector; netted in patch of Arundinaria appalachiana between 9:45 and 10:15 PM (3 ♀). Swain Co., Little Tennessee River, Lower Needmore Road, 35.3286, -83.522, 1,787´ [545 m]; ex larva 4–5 May 2021, A. E. McBride w/Eric Quinter; collected on 1 yr. old culms of Arundinaria appalachiana, pupated 23–31 May 2021, eclosed 11–22 Jun 2021 (4 ♂, 2 ♀). Same locality; 8 Jul 2021, netted in flight, A. E. McBride w/J. Petranka; netted in patch of Arundinaria appalachiana between 9:45 and 10:15 PM (9 ♀). Same locality, ex larva 25 Apr 2023, A. E. McBride, collector; collected on Arundinaria appalachiana and reared on ornamental bamboo, eclosed 7–20 Jun 2023 (7 ♂, 7 ♀). Swain Co., Fontana Lake Estates, vicinity Trails End Dr., 35.3873, -83.5515 1,782´ [543 m]; ex larva 25 Apr 2023, A. E. McBride, collector; collected on Arundinaria appalachiana and reared on ornamental bamboo, eclosed 8–13 Jun 2023 (1 ♂, 3 ♀). Same locality and collector; ex larva 8 May 2022; collected from new leaf petiole on older culm of Arundinaria appalachiana, eclosed 22 Jun 2022 (1 ♀). Swain Co., Great Smoky Mountains National Park, right bank Deep Creek, 1,840´ [561 m], 0.8 mi. nne Deep Creek campground; ex larva collected in 1st year culm of Arundinaria appalachiana, pupated 21 Jun 2001, emerged 6 Jul 2001 E. Quinter & J. O’Donnell (1 ♀). Swain Co., Great Smoky Mountains National Park, Tom Branch Road, Wiggins-Watson cemetery area, 0.8 mi. ne Deep Creek campground, 35.4652, -83.4217, 2,100´ [640 m]; ex larva collected 26 May 2001 in 1st year culms of Arundinaria appalachiana, pupated 4 Jun 2001, emerged 18-19 Jun 2001 E. Quinter & J. O’Donnell (2 ♀). AMNH, CMNH, CNC, NHML, UCD, USNM, AEM, ELQ.
This species is restricted to montane habitats of the southern Appalachians above approximately 500 meters and to the predominant cane species which occurs at these elevations, hill cane (Arundinaria appalachiana). Observations suggest C. odonnellae utilizes this single host species exclusively, which was found growing in abundance and attaining the exceptional height of two meters at the type locality, along with even taller stands of giant cane, on which no C. odonnellae larvae were found. At this site, hill cane forms thick stands in sunlit patches along the riverbank. Elsewhere in the region, growth of hill cane in wooded upland areas is sparser, with individual culms seldom exceeding one meter in height. Cherokeea odonnellae also occurs in these smaller stature stands of the host.
Like all Cherokeea species, the eggs are the diapausing stage and are presumably deposited under the leaf sheaths of the primary culm. Larvae collected in early May were in middle or penultimate instars by mid-May, suggesting that egg hatch probably had occurred about mid-April. In early May, larvae may be found fully concealed within the dried, light brown clusters of the previous year’s axillary shoot sheaths originating at nodes on the culms. New green axillary shoots developing within these sheaths were observed to have been bored by the tiny larvae living within them for perhaps the first two or three instars, after which the larvae become too large to remain inside the embryonic shoots. At this point, larvae abandon these small burrows and feed externally upon the tender bases of these new shoots, all the while remaining fully concealed within the hardened, parchment-like shoot sheaths which envelop the new growth. Early instars can also be found boring in the petioles of new leaves at the terminal ends of the culms. An accumulation of either fresh small greenish frass or dried small yellowish frass underneath the shoot sheaths or fresh, whitish frass below bore holes in new leaf petioles is an important indicator of the presence of one or more larvae on the culm. All shoot sheath clusters at each node should be carefully examined, since larvae commonly relocate to different sheaths. Presence of larger, dry, greenish frass, especially under any large primary culm sheaths persisting from the previous season, is indicative of mature Protapamea larvae which frequently rest by day under these sheaths at the same season.
Once these limited resources are exhausted, C. odonnellae larvae are obligated to leave these mature culms and seek tender new culms emerging from the ground to bore into and complete their development. Resultant mortality due to starvation or predation would appear to be very high, as new culm growth in canebrakes can be sparse, and few new culms emerge from subterranean rhizomes at this early point in the season. As with other Cherokeea, pupation likely occurs just below the soil surface, or among detritus on the surface, in a loosely constructed cell. The pupal period averages 17.4 days (n = 9).
Based upon limited captures of adults, the flight period appears to occur from 7–25 July. Eclosed adults from wild-collected larvae reared indoors emerged earlier, from 11 June to 6 July. It is likely the actual flight period begins about the last week of June.
Cherokeea odonnellae is thus far known only from two counties in western North Carolina at relatively high elevation, but it is likely the species also occurs throughout the rather limited distribution of hill cane in similar montane habitats of northwestern South Carolina, northern Georgia, northeastern Alabama, and southeastern Tennessee. Based on its extremely limited distribution and available records, Cherokeea odonnellae appears to be among North America’s rarest Lepidoptera, when in fact large numbers of its larvae can be found when one knows how and where to find them, as is so often the case with other “rare” species.
Cherokeea trigona (Smith), NEW COMBINATION (Giant Cane Borer)
Luperina trigona Smith, 1902, Journal of the New York Entomological Society, 10: 39.
Type locality: Smith County, Tennessee [USA]. [AMNH]
note—A male syntype in the American Museum of Natural History was designated lectotype by Todd in 1982.
Stanley T. Kemp collected the first specimens of this remarkable species in 1899 in Smith County, Tennessee. He relinquished them to Professor J. B. Smith of Rutgers College, who subsequently described the moth in 1902. Long regarded as among the very rarest of North American Lepidoptera, the insect was never seen or collected again until 1991, when ELQ rediscovered it in southeastern Missouri, in the course of field studies of the insect fauna of canebrakes. Since then, colleagues and resident collectors alerted to the special status of cane habitats who have specifically targeted the associated fauna have produced additional adult records over a wide geographic area, suggesting that the moth is by no means rare within its requisite habitat. The authors have experienced little difficulty locating numbers of larvae in various stages of development in virtually every suitable canebrake examined throughout the range of this species, demonstrating the importance of knowing details of a species’ biological requirements and habits before any realistic assessment of its relative abundance in an ecosystem can be made, or plans for its conservation intelligently conceived and implemented if appropriate.
This species was named for the prominent, strongly contrasting, dark triangular pattern imposed on an otherwise dull tan forewing background. All four of the original syntypes bear this distinctive pattern. Approximately 20% of the individuals in the populations sampled west of the Appalachians exhibit this phenotype. It is highly probable that the collector of those original specimens encountered some number of additional individuals at the same time and place which were likely ignored due to their general resemblance to other common dull brown moths lacking such novel appearance.
Cherokeea trigona is distinguished from its congeners by its larger size, primarily midwestern distribution, and a flight period from late summer to early autumn. The striking asymmetry of the male digiti is diagnostic and shared only with Cherokeea odonnellae, which is restricted to higher elevations in montane western North Carolina. Otherwise, C. trigona is distinguished from other Cherokeea species by characters given in the key. Females have a greatly reduced appendix bursae and much less prominent lodices than Arundinariaphila hayesi, perhaps the only other canebrake inhabitant broadly sympatric with C. trigona that might be confused with it. Inasmuch as Smith’s brief and inadequate description applies only to the phenotype with the dark, triangular patches, it is necessary to redescribe the species in greater detail, so it may be properly characterized in the context of the other members of the genus.
Head and Thorax: antennae of both sexes ciliate, laminate, less deeply in female; dorsum scaled; ventral lamination obscured by densely pubescent sensilla; otherwise as in genus description. Forewing: length similar in both sexes = 15.5–21.0 mm; dorsal surface ground of both sexes variable, ranging from light tan to wood brown to dark chocolate brown, sexual dimorphism subtle when apparent at all, females somewhat lighter in color; a striking morph (
Mature larva approximately 3.30 cm. long; dark brown to nearly black head capsule lacks distinct patterning and contrasts with honey-colored prothoracic shield, which lacks contrasting dark pigmentation but may show some mottling, especially along anterior edge among setal bases; pinacula on T1–T3 and A1–A7 are light brown, small, inconspicuous; D1 and D2 pinacula on A8 are similar in size and slightly larger than on A1–A2, with D2 setal bases closer together than D1 setal bases; D2 pinacula on A9 fused into transverse bar; anal shield light brown, with heavy sclerotization; somatic pigmentation consists of wide pinkish-brown dorsal bands on each segment terminating at spiracular line, more solidly pigmented on abdominal segments than on thoracic segments; remainder of body lacks pigmentation; pale, longitudinal mid-dorsal and subdorsal lines interrupt pigmented bands on T1–A8; these are quite prominent in early instars but become progressively fainter in later instars.
All evidence to date suggests that C. trigona only utilizes giant cane (Arundinaria gigantea) as its larval host. The moth does not occur within the range of switch cane, and hill cane does not attain sufficient size to accommodate its large larva.
As with all the members of Cherokeea, egg hatch probably occurs very early in spring. By early May, C. trigona larvae are in the second or third instars and feeding upon embryonic axillary shoots developing inside primary or secondary shoot sheaths. By mid-May these larvae appear to have their development arrested, feeding sparsely if at all, but remaining concealed within the axillary sheaths. Whether they enter a period of true aestivation or feed sporadically and simply consume minimal resources is unclear. What is clear is that until a substantial number of new cane culms emerge from the ground later in the season, the tender young growth required by these larvae remains a very limited resource, especially since Arundinariaphila hayesi utilizes new culm growth in these same canebrakes earlier in the season. It appears that C. trigona larvae do not fully resume feeding activity until sufficient time has elapsed for adequate new growth to appear. Not until late July do the larvae attain their full development. Larvae of Cherokeea gabrielleae and C. odonnellae, utilizing switch cane and hill cane respectively, are able to continue their development without interruption since new culm production in these two plants appears to commence earlier in the season.
Larvae of all Cherokeea species exhibit the same feeding mode in their early instars, described in the discussion of the genus, regardless of which of the three cane species is utilized. In later instars, feeding behavior diverges among the species, depending on host plant species and larval body mass. In the case of C. trigona, the largest species, successful completion of larval development is dependent upon the larva eventually finding a newly emerged culm of sufficient size and diameter to provide adequate food resources for the final instar. To locate larvae nearing maturity in July, one must ignore all woody culms from previous seasons as well as tall, green, healthy new culms of the current season’s growth. Evidence of larval activity at this time is easily noted by the attenuated growth of new culms which have turned yellowish tan and appear wilted, dried, or dead. These senescing culms may be several centimeters high or as much as half a meter tall. Frass ejected from the bore hole may or may not be evident on the soil below. Once the larva has matured and exhausted this final resource, it exits the culm and pupates in soil nearby. The pupal period for C. trigona averages sixteen days. Adults are on the wing from the beginning of August into early September.
This species is widely distributed throughout the range of giant cane, but primarily limited to middle latitudes west of the Appalachians. Records from Stokes and Swain counties in North Carolina east of that region suggest that the species has traversed the mountains at least near the southern end of the range, likely following rivers that flow from North Carolina into Tennessee. This uncommon species was previously known only from the four syntypes collected more than 120 years ago. The authors have documented it recently from western North Carolina and northwestern Georgia through Tennessee, Kentucky, Illinois, Missouri, Arkansas, and Mississippi.
GENUS Franclemontia Ferguson
Franclemontia
Type species: Leucania interrogans Walker, 1856.
note—Ferguson erected this monobasic genus to accommodate Walker’s species which
This peculiar apameine genus with its sole represented species reveals little in the way of an obvious relationship to any other North American genera, and probably represents another example of a unique descendant of an ancestral lineage previously established with the diversification of temperate bamboos in East Asia prior to their Miocene arrival in North America.
DIAGNOSIS OF GENUS FRANCLEMONTIA. The strongly bipectinate antenna of male Franclemontia interrogans represents a rare condition among apameines which, in combination with a prominent clavus, prominent multiple cornuti at the base of the vesica, and a small, freestanding digitus, distinguishes this genus. The conspicuous “costal sclerite” which Ferguson mentioned as a distinctive feature of Franclemontia is actually an extreme development of the basal sclerite forming the digitus, hyper-extended above the costal margin of the valve and bent into an acutely triangular, sclerotized projection. It is the same structure seen less strongly expressed in other genera such as Loscopia and Apamea.
The protracted gradual larval development and the complete absence of a true diapause in any stage of the life cycle of Franclemontia appears unique. Among those Apameini (e.g., Apamea species) which do not diapause in the egg stage but instead overwinter as early instars, development is nonetheless arrested during the winter months and rapidly completed after resumption of feeding in spring.
Franclemontia interrogans (Walker) (Franclemont’s Switch Cane Borer)
Leucania interrogans Walker, 1856. List of the Specimens of Lepidopterous Insects in the Collection of the British Museum. Vol. 9: 114.
Type locality: not given, but probably Georgia, USA. [NHML]
note—Described from two specimens from the Milne collection.
Phragmatiphila interrogans Hampson, 1910a. Catalogue of the Lepidoptera Phalaenae in the British Museum. Vol. 9: 270.
This distinctive species cannot be confused with any other. It is readily identifiable from published illustrations. Franclemontia interrogans is variable and has elongate forewings. Its ground color ranges from pale whitish to deep tan. It has a diffuse, irregular dark band that runs the length of each forewing beginning with a dark and well-marked basal dash and continuing through the orbicular-reniform complex, beyond which the band becomes faded and turns toward the wing apex. The lower lobe of the reniform spot is peculiar in that it is drawn out into an elongate point that appears smeared inwardly along the dark wing band toward the wing base.
Mature larva approximately 2.6 cm. long, and head capsule 2.0 mm wide; larva is pale and whitish-gray; small, dark brown head lacks distinct patterning and strongly contrasts with pale, whitish-gray prothoracic shield; prothoracic shield has a slight cream cast and is edged laterally with darker pigment; dark brown pinacula are large on T2-A2 and become progressively smaller through A7; D2 pinacula 4 × D1 on A8, D1 pinacula partially fused with D2 on A9; anal shield pale, whitish gray, edged laterally with darker pigment similarly to prothoracic shield; L1 located well behind and below spiracle on A7; spiracles 2-3 × as high as broad, white with black outline; somatic pigmentation consists of very pale, liver-brown dorsal bands on each segment terminating at spiracular line, more darkly pigmented on thoracic segments than abdominal segments; pale, creamy-white longitudinal mid dorsal and subdorsal lines interrupt pigmented bands on T1-A8; pale, pinkish streaks of pigment form broken, longitudinal band immediately below subdorsal lines; remainder of body lacks pigmentation.
The sole host plant is switch cane (Arundinaria tecta). Distribution records for F. interrogans suggest that the species may be restricted to canebrakes in near proximity to the coast, perhaps not occurring as far inland as many of its host plant populations.
Oviposition has not been observed in the wild, but the eggs are presumed to be deposited near the bases of tender, green, axillary shoots so that the first instars can quickly bore into the shoot base and gain access to the hollow interior of the culm. Perhaps first instars remain feeding in the green bases of the shoots until they are larger and strong enough to chew into the interior of the culm. Eggs hatch within approximately one week of deposition, atypical of most endophagous apameines, which diapause in the egg stage. Larvae can be found boring in mature culms of switch cane throughout nearly the entire year, up to within a few weeks of pupation in late March. This extremely protracted rate of development appears unique to this species. All other borers of the Arundinaria-feeding genera spend most of the summer, fall, and winter in the egg stage. It appears likely the extremely slow larval development of F. interrogans is a direct result of the very low nutritional value of its host plant. Consumption of the interior walls of the dry, woody culms is limited to girdling very small areas within the internodes. Only small accumulations of powdery sawdust-like material, evidence of internal girdling, is found within the galleries. Most of the nutrition must be derived from consumption of sparse amounts of sugary fluid flowing through the culms.
This girdling of the culms is reminiscent of the feeding mode employed by Papaipema matheri in the much larger culms of giant cane. In this latter species however, larval development is completed in several months. As P.matheri and F. interrogans have an allopatric distribution and utilize two different hosts, the girdling of the mature woody culms and subsequent breakage and loss of the top of the culm from windstorms provide an excellent opportunity for detecting the presence of either and is quite diagnostic. Even old girdled and broken off culms persisting from previous years can provide clear evidence of occurrence. An efficient technique for obtaining larvae of either species consists of walking through the canebrake, grabbing culm tops, and bending the culms downward to almost horizontal. Any culm harboring a larva will snap and break at the point of heaviest girdling, often revealing a larva, or pupa, in the appropriate season.
Prior to pupation, the larva of F. interrogans (a well as that of P.matheri) cuts an exit hole through the culm just large enough for the moth to escape and pupates head up on a platform of chewed woody particles inside the node at its base. These holes from years past can be readily detected even at some distance. Multiple round holes made from the outside of the culm indicate predation by birds such as chickadees that ELQ has observed while both they and he were searching for larvae or pupae. These and probably other birds can decimate a colony under some circumstances.
Inasmuch as nothing was known of the life history and habits of this species prior to ELQ’s work in Georgia in the canebrakes on St. Catherines Island, it is worth documenting his observations in greater detail. Resulting from a trip there on 1–5 April 1994, ELQ recorded the following data: Two eclosed pupae were found 1 April along with two partially developed live pupae. Two adult female F. interrogans flushed from low vegetation in the canebrake were confined for oviposition. The first female deposited many ova between 1–9 April and died 10 April. Most of the eggs hatched by 12 April. A second female collected 4 April deposited 35 ova in four rows (straight or staggered) of seven or eight, plus two on one leaf and two loose. The second female died 7 April. Most eggs hatched by 12 April. A female moth from one of the two pupae collected 1 April emerged 6 April in the evening.
A second trip to the island in 1994 yielded a third female, collected at 9:30 P.M. along with a heavily worn male at 10:00 P.M. EDT on 16 April. This female deposited 63 ova between 17-23 April and died 24 April. The first of these eggs hatched 25 April. All three of these females fed avidly on a dilute solution of brown sugar in captivity. No attempt to rear any of the hatchlings was made, due to unavailability of cane back in New York City. Wild-collected F. interrogans larvae are strongly cannibalistic in captivity. They proved impossible to rear without access to living cane culms throughout their extremely protracted period of development.
Another trip to St. Catherines Island was made 8–9 April 1995 to document adult activity of F. interrogans in the wild. On 8 April, eight males and eight females were collected with flashlight and net at Windmill Pond in a thick canebrake. Collecting commenced shortly after dark at 8:45 P.M. EDT. The first moth, a female, was collected at 9:00 P.M., followed by a male at almost the same instant and place, perhaps from a mating pair that had just broken up. Similarly, a male and female were collected at 9:20 P.M., nearly simultaneously. Six additional females were collected singly between 9:30 P.M. and 10:15 P.M. One was observed at 9:40 P.M. on a meter-high new culm, tippling on moist frass expelled from an Arundinariaphila bore hole. Six males were collected between 10:30 P.M. and 11:15 P.M., and numerous others seen, at which time observations were suspended. Males were observed to fly more strongly than females. No oviposition activity was observed, but three females were caged to obtain eggs. Earlier in the day a single live pupa, about a week prior to eclosion, was also collected. On 9 April, 16 males and eight females were collected employing net and headlamp as before. One additional female was caged for eggs. A male and female were observed in copulo at 11:10 P.M., and a second pair at 11:20 P.M.
Only canebrakes that have remained unburned for many years are likely to host populations of this moth. This species is highly sensitive and vulnerable to fire throughout its preferred habitat. In March of 2023, AEM collected a mature larva of F. interrogans in North Carolina. After finding no larvae during an extensive search with Bo Sullivan in coastal forests that sustained regular prescribed burns, AEM searched in canebrakes close to housing developments that were unburned, and there he found the larva along with abundant sign of larval damage.
The entire adult flight period is probably little more than two weeks at any particular location in any single year.
Franclemontia interrogans has been regarded as very rare, and at the time of Ferguson’s description of the genus, he was aware of the existence of only ten specimens. Like several other species comprising the Arundinaria-feeding genera, these moths can be quite common when sought in their highly restricted habitats. ELQ has observed dozens of individuals of F. interrogans flying on consecutive nights in less than an acre of prime, unburned habitat. The perception of rarity is furthered by the unusually brief flight season. Unless one targets the exact place at the right time, this species will indeed prove elusive.
GENUS Sarita Quinter, NEW GENUS
Gender: feminine. Type species: Acrapex relicta Ferguson, 1991.
note—Lovingly named for my daughter Sarah, who assisted me in the search for larvae of this diminutive moth when she herself was quite small.
It is clear after examination of the genitalia of both sexes that no known North American genus can accommodate the sole included species of this genus. Initially, evidence was sought for a possible placement among South American genera putatively belonging to the Amphipyrinae, which at the time encompassed a much larger assemblage of tribes than currently recognized, including the Apameini. Adults of Sarita relicta bear a superficial resemblance to those of several Brazilian genera including Acylita Hampson 1910, Amphilita Hampson 1908, Microlita Hampson 1908, Neolita Hampson 1908, and Rhyncholita Hampson 1910. Dantona Walker 1859 was also considered. The palpal characteristics alone, illustrated in
DIAGNOSIS OF GENUS SARITA. The overall habitus of Sarita is atypical of most Apameina. The thorax and abdomen are very slender, and the wings are extremely narrow and elongate, considerably more so than even other similarly small-sized apameines e.g., Photedes Lederer 1857, or Neoligia. Superficially, adult Sarita bear little resemblance to moths of these genera, other than their small size. A much stronger superficial resemblance due to the extreme elongation of the forewings and their dried-grass coloration is shared with some Asian species currently misplaced in Acrapex. Sarita shares most of the morphological features common to all Apameina and especially to Neoligia and Oligia Hübner [1821]. The genitalic morphology of Sarita appears more like that of Oligia or Neoligia than of other Apameina. The male genitalia resemble those of certain Neoligia species. A striking apparent autapomorphy in the female genitalia of S. relicta precludes placement in any available genus.
The paired lateral brushes, levers, and pockets on A1-A3 of the male, which are either fully developed, reduced, or absent in other Apameina, are absent in Sarita. They are either fully developed or reduced in Neoligia and absent in Oligia (
In the female genitalia, the heavily sclerotized dorsal surface of the sterigma (lamella postvaginalis) is produced as a flattened, spade-like extension of sternite A8 which apparently functions in assisting the insertion of the equally sclerotized, dorsoventrally flattened, acutely pointed papillae anales into the narrow crevices between the culm and the tightly appressed shoot sheaths of the host plant. This structure is unique among apameine genera; in other apameine genera, it is the ventral portion (lamella antevaginalis) of the sterigma which is often heavily sclerotized and modified to accommodate oviposition. A non-homologous structure of probable similar function occurs in Capsula Fibiger, Zilli, Ronkay, and Goldstein (2005). In that case, however, the caudally oriented lobes of the posterior margins of sternite A8 are greatly enlarged and sclerotized. Homologous structures are much reduced or absent in other Apameina. There are few additional diagnostic characters in the female genitalia. The ductus bursae is shorter in Sarita than in most Neoligia and Oligia, and the lateral margins of the anal papillae are straight in Sarita, variable in Neoligia, and convex in Oligia. Female genitalic characters in Apameina are typically quite conservative. They facilitate recognition of the tribe but offer relatively little to distinguish genera.
There are no other North American species with which S.relicta is likely to be confused although it superficially resembles a diminutive version of “Photedes” defecta (Grote), a species indigenous to northern areas of the continent far beyond the range of S.relicta.
DESCRIPTION OF GENUS SARITA. Head: antennae of both sexes ciliate, laminate, less deeply in female; dorsum scaled, ventral lamination obscured by densely pubescent sensilla; eye smooth, rounded; haustellum unreduced and fully functional, as adults have been observed imbibing water droplets; labial palpus upcurved, slender, and cylindrical, extending 0.3 × their length beyond front; first segment very short, swollen basally; second segment straight, about as long as eye width, broadly scaled with mixed light and dark scales; third segment ⅓ × length of second, narrowly scaled with mostly dark scales, projecting slightly anteriorly; frons convex, unmodified, with dense dorsal tuft of converging, spatulate, dark scales overlaying shorter, closely appressed, dark, spatulate scales. Thorax: vestiture a mixture of spatulate scales and long, hairlike scales; prothoracic collar, patagia, mesothorax, and metathorax concolorous tan with sparse, scattered, dark scales intermixed; mesoscutellar crest with contrasting, dark, spatulate scales dominant in most individuals. Legs: normal apameine; tibia lacking spiniform setae, with usual paired spurs on mesothoracic leg and two pairs on metathoracic leg (one pair ⅔ distance to distal end, one pair at end); epiphysis on prothoracic leg 0.5 × length of tibia; prothoracic tibia 1.1 × length of first tarsomere; tarsus with three rows of spiniform setae on first two tarsomeres, four irregular rows on distal three tarsomeres. Wings: forewing greatly elongated, length 2.5 × width, acutely rounded at apex, with R3+R4 stalked for ½ its length. For wing pattern features, see
Sarita relicta (Ferguson), NEW COMBINATION (Little Relict)
Acrapex relicta Ferguson, 1991, Journal of the Lepidopterists’ Society 45(3): 209–214.
Type locality: Wedge Plantation, McClellanville, Charleston Co., South Carolina [USA]. [USNM]
note—Described from eight males and three females.
In the course of ELQ’s studies in Georgia of the canebrake fauna on St. Catherines Island, hundreds of individuals of this species were observed or collected, and the immature stages were investigated. Independent collecting by Douglas C. Ferguson and others had previously revealed the presence of this species in Virginia, North Carolina, South Carolina, and Louisiana. Only about a dozen specimens were known to exist prior to the discovery of the moth on St. Catherines Island.
Seemingly unaware of ELQ’s extensive work with this insect, Ferguson named the species in 1991 from the limited material he had examined and placed it in the large Old-World genus Acrapex, putatively containing 89 species (
Given current understanding of phylogenetic relationships relative to global biogeographic patterns, it is highly improbable that any noctuid indigenous to southern India and Sri Lanka might be congeneric with another indigenous to the southeastern United States. Walker’s female holotype of A.prisca was borrowed by ELQ, along with a dissection prepared for him by Martin Honey. The holotype of A.prisca is somewhat similar in appearance to a Leucania, and this is undoubtedly why it was described as such. A drawing of that dissection (
Eggs were obtained for rearing purposes but were not preserved, so a detailed description of them is not available. It was noted that they are of the normal apameine type: flattened, lozenge shaped with vertical ribbing ornamenting the circumference. The eggs appear disproportionally large for such a tiny moth; females probably are capable of producing only a relatively small number of ova. Rearing attempts under artificial conditions failed, so a description of the early instars is unavailable. The larva of S.relicta is unique and quite unlike other apameines. The head, prothoracic shield, and anal plate are distinctively and boldly patterned with strongly contrasting, dark-pigmented patches surrounding the pinacula and superimposed on the pale background coloration. Bold longitudinal striping is heavily pigmented and contiguous across the segments, alternating with pale dorsal, subdorsal, and lateral lines. Mature larvae collected in the field provide the following brief description.
The mature larva is very slender and approximately 1.8 cm long. The head capsule is pale honey, with the Pd1 and Pd2 setae set in a single, large, strongly contrasting, dark-pigmented patch. There are two smaller peripheral dark patches at the dorsal and lateral margins of the capsule. The prothoracic shield is pale yellow with the setae set in boldly contrasting, dark-pigmented patches–four larger patches near the anterior margin, two smaller patches at the lateral margins, and four smaller patches near the posterior margin. The pinacula on T1–T3 as well as those on A1–A7 are small and inconspicuous, with dark pigmentation surrounding their bases. The D1 and D2 pinacula on A8 are more narrowly spaced than those on the other seven abdominal segments, with D1 and D2 on A9 fused into two paired plates. The anal shield is pale yellow and has two central setae set in dark-pigmented, nearly confluent patches, with two larger dark patches at the lateral margins and two smaller ones at the posterior margin. The crochets on the abdominal prolegs are a uniordinal mesoseries of the normal apameine type.
The only recorded host plant for this species is switch cane (Arundinaria tecta). ELQ’s extensive work in canebrakes throughout the southeast and lower midwestern United States failed to record the presence of S.relicta in the interior, away from the coastal plain, where another native American bamboo species, giant cane, replaces switch cane and would be a potential host.
The species is univoltine, with adults on the wing for up to one month at any given location. Across its range, adults have been collected from the second week of May through late June.
Female moths oviposit in much the same manner as other cane-feeding apameines which insert the ovipositor between the axillary shoots and their dried sheaths, which are persistent throughout the year, concealing the eggs within (
Sarita relicta occurs only in canebrake habitats from the Great Dismal Swamp in Virginia southward, apparently exclusively along the Atlantic and Gulf Coastal Plains excluding peninsular Florida, west to southeastern Louisiana (
The Papaipema group of genera
The Papaipema Group constitutes a large assemblage of approximately 70 species distributed across North America. All are known or believed to be strictly endophagous in the larval stage. Most are specialists on their select host plants. Adults of some are cryptically colored with shades of gray or brown but many are quite colorful and contrastingly marked with characteristic maculation on the forewings. All are sedentary and rarely stray far from the food sources. Many of the species are readily detected with use of ultraviolet light sources located within appropriate habitats where host plants grow in abundance, but other species are far easier to sample by searching host plants for symptoms of larval activity.
In North America this group is comprised of five closely related genera, the larvae of which are predominantly dicot-feeding specialists: Papaipema Smith, Parapamea Bird, and Selicanis Smith, solely of Nearctic distribution, along with the Holarctic genera Amphipoea Billberg and Hydraecia Guenée. A sixth genus, Fibigeria, herein described as new, is tentatively included in the Papaipema group of genera to receive the single taxon Gortyna erepta Grote, previously misplaced in Amphipoea. It is a monocot specialist that appears to belong in this genus group based on the morphology of the sub-basal part of the vesica, a synapomorphy shared with several other members of the group. These, together with several additional Palearctic genera, are in turn members of a more inclusive group, the Gortyna group of genera. Despite the strong superficial resemblance of wing patterns of adult Papaipema to those of the Palearctic genus Gortyna, Papaipema is more closely related to two other Nearctic genera, Selicanis and Parapamea, best appreciated by comparison of genitalic characters. As in the Arundinaria-feeding genera, the Papaipema group of genera is defined primarily on the basis of genitalic characters.
BIOGEOGRAPHICAL CONSIDERATIONS
A vast literature has accrued in the past century documenting and providing the basis for our present-day understanding of the geological, climatological, biological, and ecological processes influencing the evolution of the planet’s biota. Palaeobotanical evidence gleaned from both palynological studies and plant macrofossils provide firm evidence that the late Cretaceous to early Tertiary Periods (ca. 100-50 Mya) coincided with the broad establishment of many or even most of our modern plant families and genera. The early Tertiary flora is thought to have been relatively homogeneous across mid-latitudes of the Northern Hemisphere, with warm climates and land bridges facilitating floral dispersal in all directions. Although North Atlantic land bridges between North America-Greenland-Europe were long believed to have existed essentially continuously until the early Oligocene (~ 30 Mya;
The paleo-entomological evidence for this same time period is considerably less comprehensive due to the paucity of well-preserved fossil specimens, but the available evidence still suggests that modern insect taxa were in place as well. It is inconceivable that an intact, essentially modern flora existed at that time without a concomitant herbivorous insect fauna. Much concerning the present-day distribution of phytophagous insect groups can be inferred from what is already known about the evolutionary history of their hosts. Advances in molecular biology are making it possible to date divergence times of related taxa in both plants and insects, providing an increasingly clear picture of their evolutionary history.
When one contemplates present-day global distribution of the Gortyna group genera, some interesting inferences can be made regarding their historical biogeography. Species of the Holarctic genera Amphipoea and Hydraecia are roughly equally divided between the Nearctic and Palearctic regions, with no native species common to both (the Eurasian Hydraecia micacea (Esper) is introduced in North America). Relationships among species of both these Holarctic genera indicate multiple intercontinental faunal exchanges; for example, the Nearctic H.perobliqua is more closely related to the Palearctic H.micacea than it is to other Nearctic species. On the other hand, the existence of North American endemic genera such as Selicanis, Parapamea, and Papaipema, with the related genera Luperina, Gortyna, etc. endemic to Eurasia – none Holarctic – suggests these genera are derived from progenitors that became isolated earlier than Amphipoea and Hydraecia. There is little doubt that interchanges of the apameine fauna between North America and Eurasia occurred via the Bering land bridge intermittently over millions of years.
SAMPLING AND REARING
Due to the popularity of Papaipema, especially among collectors, as well as the importance members of this genus have as indicators in ecological studies of environmental quality and assessing biodiversity, it was deemed appropriate to include the following detailed discussions to enable persons undertaking the sampling of adult populations, locating their larvae in the wild, and rearing them in captivity. This information applies equally to all members of this genus group.
The following observations on sampling and surveying may prove useful to others. ELQ’s study of Papaipema began in 1971, when he initiated an intensive survey of the moth fauna of his original home county in Pennsylvania. In addition to the continuing survey of diurnal Lepidoptera which had occupied much of his attention for the previous ten years, he deployed photocell-operated, 15-watt, ultraviolet light traps in eleven differing niches within an approximate ten-mile radius, targeting habitats as distinctly different from each other as he could identify. In September of that year, one of these habitats produced specimens of the first of the new Papaipema species to come to his attention.
The following year, having ascertained the host plant of this new discovery, ELQ located an extensive colony of the species in a neighboring county. A special situation existing at this new site presented an opportunity for more detailed observations. The site is a state park situated squarely in the midst of thousands of acres of prime habitat for both the moth and its host plant. The restricted area where camping is encouraged features a number of rustic outdoor toilet facilities, equipped with incandescent lighting which at that time remained illuminated day and night throughout the entire season. As might be expected, large numbers of nocturnal insects were attracted to the lights, and since these were never turned off, many of the insects remained resting upon the wooden walls around the lights for days before ultimately expiring. That this new Papaipema species, which is a very conspicuous insect, proved by far the most abundant moth species found around these lights in mid-September was striking. One would ordinarily expect predatory songbirds to have eaten most of the daily accumulation of moths and other insects gathered upon these wooden walls, especially since the insects are not at all camouflaged there. Lack of wings on the ground suggested that predators were not feeding upon these particular moths. Could these and other Papaipema species which feed in the larval stage upon highly toxic plants have derived protection from predation by sequestering chemical compounds from their hosts? This hypothesis could neither be corroborated nor refuted over years of observation. Perhaps by September the majority of predatory birds had already vacated the area for their annual migrations.
From this unique situation came confirmation of another general observation that, for many species of moths, males are much more likely to be attracted to light sources than females. Of 58 specimens of P.amianthii randomly collected at these lights in a single season, 74% were males and 26% females. A month earlier that same season, 65 pupae had been collected at the exact same site. Of the adults which emerged, 43% were males, and 57% were females, clearly demonstrating that females are not less abundant than males in a given population.
A related observation was made repeatedly throughout the course of ELQ’s study, relative to sex ratios of sedentary moth species like Papaipema which do not normally stray significantly beyond the perimeters of their host plant colonies. The highest ratios of female to male moths attracted to light sources appear to occur at the epicenter of the host plant colony, where as much as 50% of the individuals might be expected to be females. At or slightly beyond the perimeter of such a colony, nearly 100% of the moths encountered are males. At an even greater distance, perhaps a mile or more from an isolated host plant colony, the moth species requiring that host is not likely to be encountered at all. Occasionally adult Papaipema species will be attracted to light sources miles from the nearest potential host plants, and those individuals are invariably females. As a striking example, in 1972 ELQ collected 43 males and one female of P.amianthii in an ultraviolet light trap inadvertently placed near the periphery of what was to become the type locality. That same season only two specimens of this same species were collected elsewhere, both females, three air miles from the type locality at a lower elevation and in a different habitat lacking any of the host plant. This observation was attributed to what might be called the “pioneer effect.” Papaipema species, and in fact those of most apameine genera, are largely host plant specialists highly sedentary in habit, seldom straying far from their colonies. This is in stark contrast to some dispersive noctuid genera such as Spodoptera Guenée, which may migrate a thousand miles or more. It is likely that this founder effect represents a survival mechanism in which the female moth deposits most of her eggs at the place of birth before departing on a mission to locate new food sources for her offspring. This could prove especially important in situations where host plant colonies are limited to specific habitats spotty in distribution and separated by considerable distances. Such is often the case with wetland plants typically utilized by apameine genera. Further, many host plants of Papaipema species occupy successional habitats in transition, necessitating new colonies to become established elsewhere. Numerous instances have been observed where host plant colonies persist for a decade or two before succumbing to natural habitat succession. It is worth bearing this in mind when conducting faunal surveys involving Lepidoptera. It is often more productive to seek new localities than attempt to locate target species in places where they once were known to occur.
A technique frequently employed by persons intending to breed, sample, or survey populations of other Lepidoptera such as saturniid moths is often underutilized or completely overlooked by those attempting surveys of apameine borers such as Papaipema. ELQ has attracted large numbers of rarely seen species such as Franclemontia interrogans to virgin females utilizing this method, and once successfully secured a series of Papaipema matheri, which at that time was practically unknown. A single live pupa found after an entire day of searching eclosed the following day, yielding an adult female. On a cold, clear, brightly moonlit night in late October in Mississippi, ELQ caged that moth in a canebrake and set up a black light and sheet nearby. Eight males came to his setup between the hours of 9:30 and 11:35 P.M. He remained with the sheet for an additional hour or two, but not a single other moth species or even another insect was attracted to that light on that unseasonably cold night.
Surveys conducted for assessment of populations of nocturnal Lepidoptera rely heavily on use of various types of ultraviolet light sources deployed in the habitats of the targeted species. However, perhaps the most productive method to document presence of borers at any particular site utilizes signs of larval activity. The potential for successfully employing host plant symptoms varies dramatically depending on the target species. Some species are more readily found in the larval stage than as adults; others are not.
During his master’s degree study in 2007 and 2008, AEM found that detection of larval activity was more productive than blacklighting when sampling Parapamea buffaloensis and nine species of Papaipema at known population sites. Larvae were found at these sites during the appropriate search period on all attempts for all species except Papaipema eupatorii. Using blacklights at these same sites during the appropriate adult flight period and during times of optimal blacklighting conditions, detection probability averaged 0.77 for these species. He found that for blacklighting, all-night attempts were important since many species are attracted to light both before and after midnight, but on a given night these species may only come to lights either before or after midnight. Some species, such as Papaipema astuta, P.furcata, P.impecuniosa, P.inquaesita, and P.pterisii flew to lights entirely or mostly before midnight, while P.amianthii, P.cerussata, and Parapamea buffaloensis were detected at light mostly after midnight. Papaipema appassionata, P.arctivorens, P.baptisiae, and P.speciosissima came to lights in roughly equal numbers before and after midnight when data were pooled over several nights. Larval searches should be employed for the highest detection probability using the lowest number of field visits, and that at least three all-night attempts be made under optimal conditions when sampling Papaipema and related borers using blacklights to achieve detection probabilities of 0.95 or greater.
Finding apameine borers requires a degree of familiarity with the flora of the survey area. One needs to be very observant of any signs of ill health such as wilting, drying, discoloration or other physical damage to plants, singling out and focusing upon those few among the normally much greater number of more conspicuous healthy plants. Timing is crucial. Early in the season when plants are typically small and tender, damage caused by early instars boring within them is often much more apparent than later, when vigorous surrounding plant growth obscures the earlier evidence. This is especially true of borers that move to the roots or rhizomes to continue their development. As the season advances, above-ground portions of the plant may turn brown, become dry, or even senesce entirely, leaving little or no trace of the infested plant. In the presence of large numbers of healthy plants, such evidence can be difficult to detect. Other species of borers which confine most or all of their internal feeding activity to the stems of their host plants are relatively easier to locate. In these instances, discolored foliage and stalks bearing exit holes from which larval frass is ejected, and which accumulates on the soil, can be spotted even at some distance. Recent heavy rains however can eliminate the evidence of frass accumulation on the ground beneath the plant. Crawling on hands and knees often becomes routine procedure when inspecting plants for signs of infestation. Once a larva has been found, careful excavation of the plant is necessary to avoid damaging the larva, which can quickly move from one part of its gallery to another. It will be necessary to rear the larva to maturity in captivity, whether the purpose is to obtain an adult specimen for study or simply to positively confirm its identity.
Some useful suggestions for rearing larvae of borers in captivity are provided here. The first consideration should be the type of container chosen for this purpose. Borers have sharp mandibular dentition and surprisingly strong musculature supporting their mandibles. They are capable of chewing through plastic containers to escape, especially if high humidity, temperature, or cramped space makes them uncomfortable. Given enough time, even hard plastic is no barrier, though if larvae are kept in spacious hard plastic containers, they can be successfully reared without incident. A further consideration is that larvae held in captivity for any significant amount of time should be kept in individual containers. Many borers are strongly cannibalistic, and one may end up with only one well fed individual if more than one larva is kept in the same container. A third major consideration is the food provided. If the larva was found in a root, rhizome, or other underground part of the plant, it may suffice to leave it undisturbed in the same situation it was encountered, depending on how close to maturity it may be, and place it in an appropriate container. If larvae are found boring in stems of tall plants or are only partially grown and still require weeks of additional feeding time, it is often necessary to transfer them to an alternate food supply.
The type of rearing containers ELQ employed for many years with the greatest success were glass food preservation jars of various sizes, such as mason jars. These come with a metal screw-cap lid and a removable metal insert, which can be discarded and replaced with a fine mesh stainless steel industrial filter, precut to the same diameter. Half-pint jelly jars are the most practical size for large-scale rearing of individual larvae. AEM has used various size plastic containers with lids with high success. These plastic containers can house several larvae at one time as long as enough food and space are provided. These containers should be lined with moistened paper towels which are replaced as frequently as necessary when they become excessively soiled. Larvae will complete development and pupate inside these containers, either inside the food source provided, or in the moist paper toweling. Once the pupa has hardened after a few days, it should be removed and replaced in the same jar or similar container after replacing fresh moist paper toweling and discarding the old food remnants. By doing so, it will be easier to check daily on pupal development and anticipate the emergence date. Jars or containers with freshly emerged adults should be placed under a strong light source for at least twelve hours to ensure that wings have hardened sufficiently before the specimen is dispatched, if it is being retained in a collection. The light keeps the moth quiet for the duration, preventing it from flying around in the container and quickly becoming ruined.
Regarding the food provided, unless it is the original plant tissue, there are options which have proven effective for many species, especially when larvae are collected as later instars. Potatoes, sweet potatoes, and carrots have been successfully used for this purpose. The latter is often preferrable, as potatoes tend to rot and require frequent replacement. Whichever is chosen, a hole of appropriate diameter is drilled into the vegetable and the larva is introduced. Artificial diets have also been tested. They are excellent when lab rearing larvae from eggs, but not recommended for larvae found feeding in the wild. Larvae collected in the wild will contaminate a sterile artificial medium with bacteria and yeasts from their gut, causing it to putrefy in a day. Artificial diets are expensive, and daily replacement is simply not practical.
KEY TO GENERA OF THE PAPAIPEMA GROUP (ADULTS)
NOTE—female genitalia for these genera are not included in the key because they are sufficiently similar so as to render them more difficult to characterize. It is easier to distinguish adults of these genera based on general appearance. For descriptions of female characters, refer to the text.
1. Male genitalia with triangular cucullus, bifurcate in most species 2
— Male genitalia with cucullus of various other shapes, never forked 3
2. Male genitalia with a dense patch of sensory hairs on costal lobe at base of sacculus; prominent serrate or dentate, sinuous, freestanding digitus [nearly all species]; bifurcate cucullus [most species] lacking a corona but with entire inner face densely spinose Papaipema
— Male genitalia without a dense patch of sensory hairs on costal lobe at base of sacculus; distal arm of digitus fused with cucullus, forming a ventral pollex; trigonate, bifurcate cucullus with dorsal, densely spinose crest and complete corona Selicanis
3. Male genitalia with a large, broad, pointed, distally oriented process arising from base of sacculus (clavus); cucullus paddle shaped with a small projection on the anal angle but lacking a corona Parapamea
— Male genitalia lacking these features 4
4. Male genitalia with very large, narrowly necked, cordiform cucullus with inner face densely hairy, especially basally Fibigeria
— Male genitalia lacking these features 5
5. Male genitalia with prominent, dorsally oriented process on base of sacculus (clavus); cucullus trigonate or paddle shaped, with or without densely spinose basal patch; adults orange-brown or tan, commonly with contrasting white, yellow, or orange stigmata Amphipoea
— Male genitalia without clavus; cucullus much reduced, no wider than distal portion of valves, obovate or beaked, lacking basal spinose patch; adults various shades of brown and gray without sharply contrasting stigmata Hydraecia
KEY TO GENERA OF THE PAPAIPEMA GROUP (LARVAE)
NOTE—The larva of Selicanis is unknown.
1. Larva with pale, whitish, longitudinal stripes 2
— Larva without pale, whitish, longitudinal stripes 5
2. Larva with transverse, segmental bands of pigment in addition to whitish, longitudinal stripes; darkly pigmented anterior edge on prothoracic shield Hydraecia
— Larva without transverse banding or darkly pigmented anterior edge on prothoracic shield 3
3. Pale, whitish, longitudinal subdorsal stripe is single 4
— Pale, whitish, longitudinal subdorsal stripe is double Amphipoea
4. Anal shield longer than broad; a borer in lizard’s tail Parapamea
— Anal shield broader than long; a borer in other plants Papaipema(part)
5. Darkly pigmented anterior edge on prothoracic shield; prominent transverse bands Fibigeria
— Darkly pigmented lateral edges on prothoracic shield; no transverse banding Papaipema(part)
GENUS Amphipoea Billberg
Amphipoea Billberg, 1820, Enumeratio Insectorum in Museo G. J. Billberg, 87.
Type species: Phalaena nictitans Linnaeus, 1767, by subsequent designation by Franclemont, 1950.
note— Amphipoea is derived from the Greek amphi, around + poe, grass.
Apamea of authors, not Ochsenheimer, 1816, Schmetterlinge Europa, 4: 75.
Type species: Noctua basilinea [Denis and Schiffermüller], 1775, by subsequent designation by Samouelle, 1819.
Amphipoea is a Holarctic genus of moderate size with six species indigenous to the Nearctic and perhaps four times as many occurring in the Palearctic, with no species common to both. It is an extremely homogeneous group with species so superficially similar in appearance as to render them difficult to distinguish without examination of their genitalia, which fortunately are abundantly endowed with diagnostic characters. Consequently, most of the earlier American workers considered our familiar eastern species A.americana (Speyer) to be conspecific with the Palearctic A.nictitans (Linnaeus), now regarded as synonymous with A.oculea (Linnaeus).
When comparing the male genitalia of the six Nearctic species of Amphipoea, one is immediately struck by the fact that they fall into two obvious groups which are characterized in each section below: the Amphipoea americana group [A.americana + A.interoceanica]; and the Amphipoea pacifica group [A.pacifica + A.keiferi + A.senilis]. Perhaps significantly, this dichotomy coincides exactly with the eastern and western distributions of the North American species. An apparently similar but perhaps less well-defined situation, with deviation, seems to occur with regard to the Palearctic Amphipoea. Both the Palearctic A. fucosa (Freyer) and A. lucens (Freyer) belong to the A.americana group. Unfortunately, the authors were unable to compare the male genitalia of all the Amphipoea species, lacking many of those from the eastern Palearctic, and so it cannot be confirmed whether this North American distribution pattern may be replicated to a degree in Eurasia.
Three species recently associated with Amphipoea in the North American literature are excluded from the genus. “Amphipoea” velata (Walker) was transferred to Loscopia Beck by Quinter & Lafontaine in
DIAGNOSIS OF GENUS AMPHIPOEA. Species of Amphipoea present a uniform appearance which, by having similar forewing maculation features and coloration, to a degree resemble the habitus of a small Papaipema species. They could only be confused with that genus, or possibly with the sole species of Fibigeria which features a very large, narrowly necked, cordiform cucullus, a freestanding clasper, and a ventrally projecting spine-like digitus. The most obvious diagnostic features distinguishing Amphipoea from Papaipema are in the male genitalia: 1) a prominent, heavily sclerotized clavus arising from the base of the sacculus in all Amphipoea species, which is never present in Papaipema; 2) cucullus triangular or paddle-shaped with a prominent corona, which is absent in all Papaipema; other distinguishing features include 3) absence of the transversely flattened, wedge-shaped, prothoracic crest in adults of Amphipoea; and 4) larvae without the contrasting, dark, cuticular pigmentation encircling the first four abdominal segments characteristic of Papaipema.
A more detailed description of the genitalic features of Amphipoea follows, with autapomorphies italicized. Male genitalia with peniculus subquadrate or rounded, densely hairy; valves bilaterally symmetrical; clavus prominent, of various shapes; clasper reduced to appressed, v-shaped, basal sclerite; ampulla present in some species; prominent, oblique digitus projecting distally or subventrally; cucullus broadly or narrowly necked, trigonate or paddle shaped, with or without basal, densely spinose patch; corona present; juxta + anellus variable, base wider than high; abdomen with paired basal coremata in some species, absent in others. In the female, papillae anales dorsoventrally flattened with two sclerotized rods between anal papillae, lyriform, with lateral margins prominently constricted toward anterior end, narrowly convex toward posterior end, acute apex, surface of papilla covered with setae set in tiny membranous circles, both dorsally and ventrally; segment A8 2 × as wide as long, anterior apophysis 0.7 × length of segment A8 and posterior apophysis 1.7 × length of segment A8; sternite A7 is normal, not expanded into functional lodices; lamella antevaginalis uniquely hemicordate, heavily sclerotized distally, more membranous proximally, with sinuous, deeply cleft posterior margin; ostium bursae membranous ventrally, heavily sclerotized dorsally; ductus bursae moderately long, narrow, lightly sclerotized in longitudinal ridges; bilobed appendix bursae membranous, viewed ventrally, oriented dorsally with diverticulum on right, ductus seminalis at posterior apex; corpus bursae obovate, membranous, with four signa.
Species of Amphipoea, like those of Hydraecia and Papaipema, overwinter in the egg stage. The early stages of the North American Amphipoea are known for only three of the six species, thus no key to larvae can be presented. Further, the known larvae are so much alike as to preclude easy recognition of any distinguishing features. Refer to the key to larvae of Papaipema group genera to distinguish Amphipoea larvae from those of other genera included in the group.
KEY TO NORTH AMERICAN SPECIES OF AMPHIPOEA (ADULTS)
1. Cucullus of male genitalia triangular, the inner face of which bears a dense medial patch of dorsally oriented hairs, basad of which is a large field of stout, proximally oriented spines 2
— Cucullus of male genitalia paddle-shaped, the entire inner face covered with dorsally oriented hairs; field of spines absent 3
2. Digitus of male genitalia long, angled then straight, subparallel to distal portion of valve and extending to anal angle of cucullus; prominent clavus on base of sacculus bifurcate, about ½ length of digitus americana
— Digitus of male genitalia short, curved ventrally but parallel to distal portion of valve and only extending halfway to cucullus; clavus acuminate, extremely long, about 2 × length of digitus interoceanica
3. Clavus on base of sacculus very long, nearly straight but slightly curved apically, a much smaller secondary tooth distally near base of clavus on costa of sacculus; digitus long, slender, sinuous or slightly curved, parallel or subparallel with cucullus and oriented ventrally 4
— Clavus on base of sacculus shorter, slightly curved or sharply angled, no secondary tooth beyond base of clavus; digitus stout, shorter, doubly angled, oriented distally with tip partially overlapping cucullus 5
4. Digitus somewhat sinuous, oriented subparallel with cucullus, but with tip touching cucullus; adult rich reddish-brown with crisp forewing pattern elements; San Bernardino Mountains of southern California cottlei
— Digitus straight or slightly curved ventrally, oriented parallel with and not touching cucullus; adult dull orange-red with less sharply defined forewing pattern elements, predominantly coastal and west-central California and Oregon pacifica
5. Clavus on base of sacculus slightly curved; a prominent tooth-like editum present near middle of valve dorsad of clasper; Colorado south to New Mexico and Arizona senilis
— Clavus on base of sacculus somewhat more acutely angled; editum very slight, broadly rounded; northern Great Basin keiferi
Amphipoea americana GROUP
In North America, this group includes A.americana and A.interoceanica, and also the Palearctic A. lucens and A. fucosa. The cucullus of the male genitalia is triangular, the inner face of which bears a dense medial patch of dorsally oriented hairs, basad to which is a large field of stout, proximally oriented spines, and the male abdomen bears fully functional, paired, basal coremata.
Amphipoea interoceanica (Smith) (Strawberry Cutworm Moth)
Hydroecia interoceanica Smith, 1899, Transactions of the American Entomological Society, 26: 17.
Type locality: Winnipeg, Manitoba [Canada]. [USNM]
note—Smith’s description was based upon three specimens from the same source.
Deceptively similar to A.americana and broadly sympatric with it, yet with practice A.interoceanica can often be distinguished from A.americana by the slightly narrower wingspan and less acutely pointed wingtips of A.interoceanica. The reniform stigma of A.interoceanica tends to be slightly less full than in A.americana. Identification is easy and more reliable by simply brushing enough scales from the tip of the male abdomen to reveal the dramatic differences in genitalia. In addition to obvious differences in cucullus shapes of the two species, the much longer and more prominent digitus of A.americana is an immediately diagnostic feature distinguishing this pair. Females of A.interoceanica are best diagnosed by association with males, as genitalic differences in female Amphipoea are much subtler.
The reddish-tan forewings of A.interoceanica are slightly elongate and relatively narrow for the genus. The forewing coloration is paler beyond the postmedial line. The orbicular is generally similar in color to the ground but may be slightly brighter. The reniform may be either white or yellow-orange. The hindwings are fuscous with a contrasting paler fringe. The clavus of the male genitalia is very long, approximately 2 × the length of the digitus, and the digitus is short, extending only half the distance to the cucullus.
The immature stages have been exceptionally well documented and illustrated together with host plant symptoms in a detailed paper by G. L.
This species seems to occupy somewhat drier habitats than A.americana and appears to be a borer specializing in crowns of herbaceous Rosaceae. Food choices include both wild and cultivated strawberries; A.interoceanica has at times proven to be a serious economic pest of commercial strawberries. ELQ and AEM found the larvae boring in crowns of prairie smoke (Geum triflorum) on Manitoulin Island, Ontario. In Connecticut where ELQ commonly collected the adults at light on a power transmission cut, abundant creeping cinquefoil (Potentilla reptans) is the presumed host in the absence of other appropriate alternatives.
A.interoceanica flies from the middle of June through late August and is probably more common than generally recognized, due to its very close resemblance to A.americana. It has a similar distribution to A.americana but does not occur as far west. It occurs farther south in the Midwest and is perhaps less adapted to boreal habitats. Unlike A.americana, it has not been recorded from the mountainous regions of the southwestern United States.
Amphipoea americana (Speyer) (American Ear Moth)
Hydroecia nictitans var. americana Speyer, 1875, Entomologische Zeitung. Entomologischen Vereine zu Stettin, 36: 152.
Type locality: North America. [Type lost]
note—Speyer described americana as a variety of the Palearctic taxon nictitans, but subsequently these two species were shown to be distinct (
Hydroecia atlantica Smith, 1899, Transactions of the American Entomological Society, 26: 18.
Type locality [lectotype]: “Ithaca, N.Y.” [New York, USA]. [USNM]
note—Described from an unknown number of specimens from multiple localities. A male was designated lectotype by
Amphipoea americana is so similar in general appearance to both A.interoceanica and A.pacifica that these three species can be quite difficult to recognize visually. Amphipoea interoceanica averages a bit smaller than A.americana, and A.pacifica is mostly allopatric with regard to both, its range overlapping that of A.americana only slightly. The hindwings of A.americana and A.interoceanica are fuscous, while those of A.pacifica are paler and have a distinct rosy cast. Amphipoea pacifica does not have a form with a white-filled reniform and does not have a contrasting pale fringe on the hindwing as is found in A.americana and A.interoceanica. All three are easily distinguished by the male genitalia, and the females, with less obvious genitalic distinctions, by association. In A.americana, the digitus is much longer than in A.interoceanica and extends just past the tip of the cucullus. The clavus of A.americana is short compared with A.interoceanica and A.pacifica and is bifurcate. In A.pacifica, the clavus and digitus are both relatively long.
The larva of A.americana is practically indistinguishable from those of the other known North American Amphipoea species. The larva is immediately distinguishable from larvae of Papaipema because the contrasting dark-pigmented, longitudinal bands appear to be imposed upon a body that is otherwise entirely of a dull whitish coloration. In Papaipema, the opposite effect is apparent, with longitudinal white bands seemingly superimposed upon a largely dark-pigmented body. For other differences from the included genera, see the key to larvae of the Papaipema group genera.
Specifically, the larva of A.americana exhibits two prominent pinkish to purplish-brown pigmented, subdorsal, longitudinal bands as well as two similar lateral bands running through the spiracles, and a thinner, less prominent, pigmented, parallel line in between. The prothoracic shield is honey-colored, its setae surrounded by dark pigmentation, and the anterior margin accented with two larger, black-pigmented spots at the anterior lateral angles, characteristic of Amphipoea. As with the great majority of borer larvae, pigmentation of the late instars, especially the last, typically fades to near obsolescence.
Amphipoea americana has been reported boring in roots of sedges as well as corn and other large grasses (
Amphipoea americana is commonly associated with wetland habitats. The species flies from the middle of June through early September. This widespread and rather common species ranges from Newfoundland west across the upper Midwest and Canadian prairie provinces to Vancouver Island, Washington, and Oregon, and is the only species treated in this volume with a range that stretches to extreme southeast Alaska. In the eastern United States and Canada, it ranges south along the Appalachian Mountains at increasingly higher elevations to northwestern Georgia. It also occurs throughout the southern Rocky Mountains to the mountains of central Arizona. The species is perhaps more at home in boreal habitats than its congeners.
Amphipoea pacifica GROUP
This group includes A.pacifica, A.cottlei, A.keiferi, and A.senilis. The cucullus is paddle-shaped, the entire inner face of which is covered with dorsally oriented hairs and lacking a field of spines. The male abdomen lacks the basal coremata present in the A.americana group.
Amphipoea pacifica (Smith) (Pacific Ear Moth)
Hydroecia pacifica Smith, 1899, Transactions of the American Entomological Society, 26: 19.
Type locality: California [USA]. [USNM]
note—It is unclear how many specimens Smith had, but at least five syntypes have been located, all from California. A male was designated lectotype by
The adults are similar in general appearance to those of both A.americana and A.interoceanica but see the discussion of that matter under A.americana. No host plant is known, but iris is certainly a plausible candidate. The immature stages are likewise unknown. The adult flight period extends from June through September, with a peak in August.
The species appears to be confined to lower elevations east of the Coastal Ranges but west of the Cascade Range in Oregon, and has a primarily coastal distribution in California, with scattered inland records elsewhere in the state.
Amphipoea cottlei (McDunnough) (Cottle’s Ear Moth)
Apamea cottlei McDunnough, 1948, Journal of the New York Entomological Society, 56: 51–52.
Type locality: Arrowhead Lake, San Bernardino Co., California [USA]. [AMNH]
note—Described from two specimens: a male holotype and a female allotype.
Relatively little is known of this species. It could be confused with A.pacifica, but the two appear to be allopatric. See key for distinguishing features. The host plant and immature stages are unknown, but iris species are likely host plant candidates. Specimen data indicate a flight period centered in August. The species is known only from the San Bernardino Mountains of southern California with further distribution uncertain due to the paucity of records.
Amphipoea keiferi (Benjamin) (Northwestern Iris Borer)
Apamea keiferi Benjamin, 1935, Pan-Pacific Entomologist, 11(2): 55–56.
Type locality: Cedarville, Modoc Co., California [USA]. [USNM]
note—Described from three males.
Apamea kieferi; Crumb, 1956, misspelling.
This species is superficially very similar in appearance to pale, nearly immaculate specimens of the more abundant Apamea niveivenosa (Grote) which coexists with it but is readily distinguishable from it by the genitalic characters which distinguish these two genera. From other Amphipoea species, A.keiferi is difficult to distinguish from A.senilis, which has a more southern distribution and perhaps averages very slightly larger. See the discussion under A.senilis for distinguishing features of maculation. The genitalic structures of both sexes show only minor differences between these two taxa. In male A.senilis, the entire anterior edge of the distal section of the clavus is completely covered in a long patch of even, short hairs. In A.keiferi, the entire clavus (anterior and posterior sides) is smooth without a trace of these hairs.
The larva “bores in the bulbous leaf bases and later in the extremely acrid rhizomes of Iris missouriensis [Rocky Mountain iris]” (
Amphipoea senilis (Smith) (Southwestern Ear Moth)
Hydoecia [sic] senilis Smith, 1892. Entomological News, 3(10): 251.
Type locality: “Colo.” [Colorado, USA]. [AMNH]
note—Described from a male and a female, both from Colorado, and redescribed by Smith,1894, Transactions of the American Entomological Society, 26: 25. The male syntype was designated lectotype by
Apamea flavostigma Barnes and Benjamin, 1924. Contributions to the Natural History of the Lepidoptera of North America. 5(3): 157.
Type locality: High Rolls, New Mexico [USA]. [USNM]
note—Described from three males. Synonymized by
On average, A.senilis may appear slightly larger than but otherwise superficially almost indistinguishable from A.keiferi. In A.keiferi, the orbicular spot is obsolescent and the reniform nearly so. In senilis, both these spots can be faint, but are outlined with slightly darker scales, best viewed under magnification. The genitalic structures of both sexes are likewise very similar, except for minor differences given in the key. See also discussion under A.keiferi.
Adults of this species are variable, ranging from pale beige to reddish-tan, sometimes with reddish-brown overscaling in the medial area and demarcating the antemedial and postmedial lines as well as the medial shade. The orbicular stigma and especially the reniform can be a contrasting yellow or orange, most commonly in southern portions of the range, and both are outlined thinly with darker scales. The mature larva is approximately 2.5 cm long. The pinkish longitudinal bands are paler in this species than in A.americana with less well-defined edges. Two dark points project distally from the dark band at the leading edge of the prothoracic shield.
In 2023, AEM found larvae of A.senilis boring in the rhizomes of Rocky Mountain iris (Iris missouriensis) near High Rolls, New Mexico. Infested plants turn a pale, straw yellow and stand out easily against healthy plants. Larvae accept other species of iris in captivity. Adults have been recorded flying from late August through September. This species is distributed throughout Colorado, New Mexico, and Arizona. The species has not yet been reported from Utah, but very likely occurs there as well.
GENUS Fibigeria Quinter, NEW GENUS
Gender: masculine. Type species: Gortyna erepta Grote, 1881 by monotypy.
note — Named to honor the late Michael Fibiger for his outstanding contributions to our knowledge of the Old World Apameini, among many other accomplishments. Our paths crossed only briefly, but he will always be remembered.
DIAGNOSIS OF GENUS FIBIGERIA. Correctly associating this genus with an appropriate group of apameine genera has proven more difficult than might be expected. It manifests an odd mixture of features which suggest the potential for placement with either the Apamea group of genera, the Arundinaria-feeding genera, or the Papaipema group of genera. The male genitalia with its very large, narrowly necked, cordiform cucullus with densely hairy inner face, especially basally, and a complete, prominent corona is strikingly similar to the condition in Apamea. The totally dissimilar larval appearance however, reflecting the plesiomorphic apameine condition (i.e., pigmented cuticular banding encircling all segments versus longitudinal lines or stripes), and the strictly endophagous feeding mode precludes an association with the Apamea group. The prominent, freestanding clasper arising from an appressed base on the male valve is reminiscent of Arundinariaphila, but other features of the genitalia are considerably less so. The larva is perhaps a better fit here although its host plant association is not linked to the bamboo-feeding specialization which seems to loosely unite the Arundinaria-feeding genera. The tentative placement of Fibigeria with the Papaipema group is more or less by default and, as a highly specialized grass feeder it clearly remains an outlier, but a synapomorphy in the subbasal area of the vesica is shared with several other members of the group. In the female genitalia, the distal margin of the lamella is slightly cleft, a condition more prominently developed in other members of the Papaipema group. Until the entire tribe has been thoroughly studied globally, placement of this genus will remain uncertain. The combination of all the above features in a single species presents a unique condition diagnostic of this monobasic genus.
DESCRIPTION OF GENUS FIBIGERIA. Head: antennae of both sexes ciliate, inconspicuously laminate [sensu
Fibigeria erepta (Grote), NEW COMBINATION (Gamagrass Borer)
Gortyna erepta
Type locality: Douglas County, Kansas [USA]. [NHML]
note — Described from a unique specimen collected by Professor Snow of the University of Kansas. The name is derived from the Latin ereptus, snatched or taken, perhaps referring to Grote’s actions upon first seeing the specimen.
Apamea erepta var. ryensis Bird 1913, Entomological News, 24: 223.
Type locality: Rye, Westchester Co., New York, U.S.A. [AMNH]
note — Named as a variety by Bird and treated as a subspecies. It remained listed as such in
The sole species of this genus has been associated with Apamea, sensu
The adult should not be confused with any other species. It is a straw yellow moth with a white, lunate reniform. Fibigeria erepta lacks the reddish overscaling of Amphipoea and the antemedial and postmedial lines contrast more strongly and track differently than in Amphipoea. The maculation of F. erepta is relatively consistent across its range. The male genitalia of F. erepta share a number of features with both Arundinariaphila and Apamea, such as a large, narrow-necked cucullus and a well-developed corona, although these character states may simply reflect apameine tribal synapomorphies.
The larva is patterned in a distinctive manner quite dissimilar to larvae of Amphipoea, but more like larvae of Cherokeea or Hydraecia. Larvae of Amphipoea bear pinkish pigmented bands interrupted by pale longitudinal stripes; those of F. erepta have transverse segmental banding and strongly contrasting purplish cuticular pigmentation. The head capsule is honey-colored; the pale-yellow prothoracic shield is patterned with heavy black pigmentation on its anterior margin, much less so on the lateral and posterior margins.
The larval host plant is eastern gamagrass (Tripsacum dactyloides). Larvae have been observed boring in young corn, a distant relative of Tripsacum, in situations where crops border wetland habitats with gamagrass. Along the Atlantic Coast, this large wetland grass grows in dense monocultures, generally limited to woodland margins bordered by salt marsh habitats, in a narrow zone between the woodland and expanses of cordgrass (Spartina), which is increasingly being replaced with common reed (Phragmites). Elsewhere in eastern North America the host plant is associated with other types of wetlands, even perennially wet roadside ditches. Such habitats are unlikely to support a population, however. The species requires extensive gamagrass colonies that are free from disturbance (especially burning) for decades.
Eggs hatch in spring and young larvae enter the innermost part of the developing culm, near the base. Their presence is easily detected as late as mid-June by noting dried green or brown central foliage within the otherwise vigorous external growth of the culm. Larval development is confined to very short galleries within the base of the culms, which arise from stout rootstocks. A single plant typically has multiple branches, any or all of which may support larvae. As many as ten larvae have been encountered in one plant, each separated from the other. The mature larva exits its gallery and pupates in wet detritus nearby. The pupal duration is the shortest of any apameine known, averaging only 13 days. In the coastal portions of its range near salt marsh habitats, this very short pupal period may provide an adaptive advantage to avoid flooding during extreme tidal events. The adult flight period is correspondingly brief, probably not lasting more than two weeks at any location, so the species is easily missed even in its appropriate habitat. Adults fly in late June and early July. For further details concerning this species, see
Fibigeria erepta is rare in collections, probably due to a combination of factors. Like most apameines, adults are highly sedentary and not likely to be encountered outside established host plant colonies which, in the case of F. erepta, are extremely localized and presently threatened by encroachment of common reed. Fibigeria erepta has been recorded along the Atlantic Coastal Plain, and across the southern and midwestern United States. It could be expected wherever its host grass occurs in sufficiently undisturbed abundance. It should be noted that gamagrass is also the host for two other borers which occur throughout its range. The larva of another apameine moth, Meropleon titan Todd and that of the crambid moth Southern Cornstalk Borer, Diatraea crambidoides (Grote) may coexist with F. erepta at some sites, but their larvae are unmistakably different.
GENUS Hydraecia Guenée
Hydraecia Guenée, 1841, Annales de la Société Entomologique de France 10: 449.
Type species: Phalaena micacea Esper, 1789, by subsequent designation by Guenée in Boisduval & Guenée, 1852: 128.
Hydroecia Agassiz, [1847], 1846, Nomenclator Zoologici Index Universalis.
note — Hydroecia Agassiz, [1847], 1846 is an incorrect subsequent spelling of Hydraecia Guenée, 1841.
Hydrooecia Herrich-Schäffer, 1849. Systematische Bearbeitung der Schmetterlinge von Europa, Zugleich als Text, Revision und Supplement zu J. Hübner’s Sammlung Europäischer Schmetterlinge, 1843–1856.
note — Hydrooecia Herrich-Schäffer, 1849 is an incorrect subsequent spelling of Hydraecia Guenée, 1841.
Hadroecia Schaus, 1894, Transactions of the American Entomological Society, 21: 223–244.
note — Hadroecia Schaus, 1894 is an incorrect subsequent spelling of Hydraecia Guenée, 1841.
Hydraecia is a relatively homogeneous group of approximately six Nearctic and nine Palearctic species, one of which has also long been established in North America. Two native and the one naturalized species occur in eastern North America. Perhaps comparable to the situation in Amphipoea, the two eastern North American species, H.immanis and H.stramentosa, together with the western H.perobliqua, form a subgroup which appears to be closely related to the Palearctic H. ultima. This subgroup is distinct from the remaining western North American Hydraecia species which bear a strong superficial resemblance to the Palearctic Cervyna cervago, but which is not reflected in their genitalic characters. Like the species of Amphipoea, but unlike all Papaipema species which entirely lack basal abdominal coremata, some Hydraecia species have basal coremata, while others lack them entirely.
Species of Hydraecia are univoltine, like those of Amphipoea and Papaipema, and overwinter in the egg stage. The early stages of the North American Hydraecia are known for only three of the seven species, thus no key to larvae can be presented. Refer to the key to larvae of Papaipema group genera to distinguish Hydraecia larvae from those of other genera included in the group. Almost no information regarding the early stages of any of the western species is available, but all are assumed to be subterranean borers of perennial forbs. As far as is known Hydraecia larvae favor mesic habitats, feeding internally in the stems, and ultimately the roots, of herbaceous dicots. In the case of H.immanis, an economically important pest, the larvae are able to complete their development in one particular thick-stemmed monocot, corn. Entire crops have been destroyed on occasion. At least two species, H.immanis and H.micacea, are known obligate feeders as neonates in stems of various perennial grasses (
The authors have limited experience with the western North American taxa and have neither new information nor novel opinions to offer regarding them. For that reason, they accept and are following the classification of
DIAGNOSIS OF GENUS HYDRAECIA. Species of Hydraecia present a variety of appearances, yet none are likely to be confused with species of any other genera. Male genitalic characters which distinguish Hydraecia from closely related genera include: 1) basal saccular process (clavus) absent as in Papaipema, unlike the prominent clavus in Amphipoea. In Papaipema but not Hydraecia, the clavus is replaced with a dense patch of sensory hairs on costal lobe at base of sacculus; 2) digitus smooth like Amphipoea, unlike dentate digitus of most Papaipema; 3) cucullus trigonate, with prominent corona like Amphipoea, but unlike bifurcate cucullus characteristic of Papaipema which lacks a corona; 4) peniculus subquadrate, densely hairy like Amphipoea, unlike Papaipema which is rounded and sparsely hairy; 5) ductus of vesica projecting dorsally, unlike Amphipoea and Papaipema where it projects ventrally. Female genitalic features distinguishing Hydraecia from closely related genera include: 1) the lack of a deeply cleft, cordate ostial plate characteristic of Amphipoea and Papaipema; 2) a membranous, bilobed appendix bursae, viewed ventrally, oriented dorsally with diverticulum on right, similar to Papaipema, unlike Amphipoea with diverticulum on left.
A more detailed description of the terminalia of Hydraecia follows. Male: peniculus subquadrate or rounded, sparsely hairy; valves bilaterally symmetrical; clavus absent; sacculus with a densely hairy, mid-ventral patch on inner face; clasper an appressed, basal, straplike sclerite; ampulla present; digitus parallel with costa, or posteroventrally oriented; cucullus reduced, narrow, obovate in some species, trigonate in others; corona with variable numbers of spines present; juxta vase- or flask-shaped; paired, basal, abdominal coremata present in some species, absent in others. A single posterior corema on A8 is present on at least one Nearctic species, H.perobliqua. Female: papillae anales dorsoventrally flattened with two sclerotized rods between them, papilla analis lyriform, with lateral margin prominently constricted toward anterior end, narrowly convex toward posterior end, acute apex, surface of papilla covered with setae set in tiny membranous circles, both dorsally and ventrally; segment A8 2.2 × as wide as long, anterior apophysis 1.8 × length of segment A8 and posterior apophysis 2.2 × length of segment A8; sternite A7 normal, not expanded into functional lodices; lamella antevaginalis subquadrate, sclerotized, with slightly cleft posterior margin; ostium bursae membranous; ductus bursae moderately long, membranous, with sclerotized longitudinal ridges in some; appendix bursae membranous, with partial rugose sclerotization, viewed ventrally, oriented on left but expanded dorsally to the right in some, ductus seminalis at posterior apex; corpus bursae round or ovate, membranous, with four signa.
KEY TO NORTH AMERICAN SPECIES OF HYDRAECIA (ADULTS)
1. Predominantly eastern North American species 2
— Predominantly western North American species 4
2. Cucullus of male genitalia triangular, well separated from ventrally projecting, short, acutely pointed digitus micacea
— Cucullus of male genitalia an elongated, conical, obovate lobe projecting ventrally and partially overlapping the dorso-distally oriented digitus 3
3. Very large, warm brown, mainly northeastern and midwestern species, sometimes occurring as far west as the Rocky Mountains immanis
— Large, straw-colored eastern and midwestern species with disjunct populations occurring in the southwestern mountains of Colorado, Utah, New Mexico, and Arizona stramentosa
4. Cucullus of male genitalia an elongated, conical, obovate lobe projecting ventrally and partially overlapping the dorso-distally oriented digitus perobliqua
— Cucullus of male genitalia triangular, well separated from ventrally projecting, short, acutely pointed digitus 5
5. Species restricted to Pacific Coast regions west of the Sierra Nevada and Cascade ranges obliqua
— Species widely distributed in western states east of the Sierra Nevada and Cascade ranges (includes the unique intermedia) medialis
Hydraecia medialis Smith
Hydroecia medialis Smith, 1892, Entomological News, 3(10): 251.
Type locality: Colorado, USA. [AMNH]
note — Named from an unknown number of specimens of both sexes, a male and a female of which are labelled as types.
Hydroecia pallescens Smith, 1899, Transactions of the American Entomological Society, 26: 25.
Type locality: Calgary, Alberta, Canada. [USNM]
note — Described from at least seven specimens. Smith’s female “type” from Calgary was selected by
Of the western North American Hydraecia species, H.medialis is the most widespread and variable with regard to coloration of the adult forewings. In particular, southwestern populations tend to be reddish rather than gray or brown, which is typical of northern populations. Distinguishing H.medialis from the similar looking H.obliqua is discussed under that species. It is perhaps more likely that H.medialis will be confused with the smaller H.perobliqua with which it is widely sympatric. The forewing coloration of H.perobliqua is of a lighter tannish-brown shade than normally seen in H.medialis, and its whiter hindwings lack a prominent medial line. The external features of the male abdomen, together with its genitalia, readily distinguish H.perobliqua from all other western Hydraecia, as discussed under H.perobliqua.
The life history, immature stages, and host plants of H.medialis are unknown. The adults fly mainly in September but can be found from late August through early October. Specimens from the White Mountains of Arizona have been taken in late July. Hydraecia medialis is distributed from southwestern Saskatchewan to southern British Columbia, south and east of the Coast Ranges to Oregon, northeastern Nevada, and farther south to Arizona and New Mexico. It ranges throughout the Rocky Mountains and east to western Nebraska.
Hydraecia obliqua (Harvey)
Gortyna obliqua Harvey, 1876, The Canadian Entomologist, 8: 53.
Type locality: [Mendocino], California, [USA]. [AMNH]
note — The type locality was given as only “California” but the holotype specimen in the AMNH is labelled “Mendocino.”
Gortyna columbia Barnes and Benjamin, 1924, Contributions to the Natural History of the Lepidoptera of North America, 5(3): 161.
Type locality: Saanich district, British Columbia, Canada. [USNM]
note — Described from two males from British Columbia.
Gortyna ximena Barnes and Benjamin, 1924, Contributions to the Natural History of the Lepidoptera of North America, 5(3): 160.
Type locality: Truckee, California, USA. [USNM]
note — Described from two males from the Sierra Nevada Mountains in northern California.
Other than by its allopatric distribution, H.obliqua is difficult to distinguish from H.medialis. The postmedial line of H.obliqua is less steeply angled than that of H.medialis, relative to its trajectory approaching the costa of the forewing. There exist subtle differences in antennal morphology of males between H.medialis and H.obliqua. In H.medialis, antennae are bipectinate with very short fasciculate branches; in H.obliqua, the antennae are both more strongly serrate and fasciculate. Also, the ampulla on the valve of H.obliqua is longer than that of H.medialis. Paired basal coremata on A1 of males are absent in both species. In coloration of the forewings, H.obliqua is more nearly like H.perobliqua which, in addition to being smaller, has whiter hindwings than H.obliqua and lacks the prominent medial line. With regard to the course of the postmedial line, H.perobliqua more nearly resembles H.medialis.
The immature stages of H.obliqua are unknown, although a note in
Hydraecia intermedia (Barnes and Benjamin)
Gortyna intermedia Barnes and Benjamin, 1924, Contributions to the Natural History of the Lepidoptera of North America, 5(3): 158.
Type locality: Lundbreck, Municipality of Crowsnest Pass, Alberta, Canada. [USNM]
note — Chris Schmidt in
This taxon is based on its unique holotype. Hydraecia intermedia was distinguished from H.medialis by Barnes and Benjamin largely on the basis of perceived differences in the male antennal structure; specifically, the length of the fasciculation which was said to be slightly longer in H.intermedia than in H.medialis. This character might only represent individual variation. One would need to see additional specimens to determine this, and no such specimens have ever been found in the past hundred years. Other trivial differences in forewing coloration and maculation were mentioned as distinguishing H.intermedia from H.obliqua. Nothing in the male genitalia of the holotype of H.intermedia stands out as convincingly different from either H.medialis or H.obliqua, although the ampulla of H.intermedia appears more like that of H.obliqua than that of H.medialis, and the digitus of H.intermedia is less acute and shorter than those of either. The holotype male was collected in August of 1916. Nothing more regarding H.intermedia is known.
Hydraecia micacea (Esper) (Potato Stem Borer)
Phalaena micacea Esper, 1789, Die Schmetterlinge in Abbildungen nach der Natur mit Beschreibungen, 4: 466 [1803], Noct. 66, pl. 145, fig. 6.
Type locality: Tyrol, [Austria].
Noctua cypriaca Hübner, [1803], Sammlung Europäischer Schmetterlinge, 4: pl. 46, fig. 244.
Type locality: [Europe]
This Palearctic species was apparently introduced into North America well before one hundred years ago (
Like H.stramentosa, the larva of H.micacea is less contrastingly marked with regard to the purplish pigmentation of the transverse bands and whitish longitudinal lines in the late instars than is observed in H.immanis. Earlier instars are well marked. Larvae of H.micacea have been reported boring in a variety of unrelated plants, most notoriously potato (Solanum tuberosum), but numerous others including queen (Filipendula), dock (Rumex), and even horsetail (Equisetum spp.) (
As is the case with H.immanis, eggs are deposited within leaf sheaths of grasses and overwinter. Neonate larvae feed within the stems of a wide range of grasses but are obligated to vacate these stems in search of much thicker-stemmed plants which can support them throughout the remainder of their development, no doubt accounting for the long list of host plants recorded. Pupation occurs in soil and adults fly during late summer and early autumn. In North America, this species ranges from Newfoundland to southwestern Ontario, the upper Midwest, and southern Ohio, and throughout New York and New England.
Hydraecia perobliqua Hampson
Hydraecia perobliqua Hampson, 1910, Catalogue of the Lepidoptera Phalaenae in the British Museum, 9: 44.
Type locality: Winnipeg, Manitoba, Canada. [NHML]
note — Described from a male type and an unspecified number of “cotypes.”
Adults of H.perobliqua are normally yellowish-tan and smaller than those of H.medialis with which H.perobliqua is broadly sympatric and might be confused. The male terminalia of H.perobliqua features paired, basal coremata on the first abdominal segment as well as persistent (i.e., non-deciduous) lateral coremata on the eighth segment. Hydraecia medialis and H.obliqua lack the coremata on both the first and eighth abdominal segments. Although the adult moths most nearly resemble those of H.medialis and H.obliqua, the male genitalia clearly place H.perobliqua in the subgroup with H.stramentosa and H.immanis, having an elongated, conical, obovate lobe projecting ventrally and partially overlapping the dorso-distally oriented digitus in the male genitalia. Also, the peniculus of H.perobliqua is subquadrate and densely hairy.
Nothing is known of the immature stages or host plants. Adults have been recorded flying from late July through mid September. Hydraecia perobliqua ranges from northern Saskatchewan to southern British Columbia and south in the mountains to Washington, Nevada, Utah, and Colorado, and as far east as the eastern Dakotas and southern Manitoba.
Hydraecia stramentosa Guenée (Figwort Borer)
Hydroecia stramentosa Guenée, 1852, in Boisduval and Guenée, Histoire Naturelle des Insectes. Species Général des Lépidoptѐres, 5: 129.
Type locality: New York, USA. [NHML]
Hydroecia diplocyma Hampson, 1910, Catalogue of the Lepidoptera Phalaenae in the British Museum, 9: 45, pl. 138, fig. 2.
Type locality: Colorado, USA. [NHML]
note —
The name is derived from the Latin stramineus, of or pertaining to straw, in reference to the characteristic straw-like coloration of the adult forewings. This distinctive species is unlikely to be confused with any other. It, together with H.perobliqua and H.immanis, forms a group distinguished by an elongated, conical, obovate lobe constituting the cucullus which projects ventrally and partially overlaps the dorso-distally oriented digitus in the male genitalia. The paired, basal, abdominal coremata present in H.perobliqua and H.immanis are absent in H.stramentosa.
The penultimate and final instars of H.stramentosa lack both the strongly contrasting white longitudinal lines and intersegmental areas characteristic of H.immanis and are more uniformly pigmented on all abdominal segments. This pigmentation becomes obscure in the final instar. Early instars of this species have not been observed. The yellowish prothoracic shield is irregularly edged on its anterior margin with black pigmentation as in other Hydraecia. The only known host plants of H.stramentosa are various species of figwort (Scrophularia). It is highly likely that H.stramentosa completes its entire larval development as a borer in the roots of figwort and does not utilize grasses in the early instars. Unlike in most Papaipema species, there is no external evidence above ground of damage to the host plant which might serve as evidence of the presence of H.stramentosa larvae in their host. It is likely that this is true for other Hydraecia species as well and may account for the dearth of information concerning their larval stages and host plant utilization. AEM found two larvae of H.stramentosa in the mountains of Bath County, Virginia on 4 July 2015 in lanceleaf figwort (Scrophularia lanceolata). One of these was a prepupa which pupated that same evening and emerged as an adult on 2 August. He found no evidence of larvae in plants with single stalks; the two found were discovered in plants that had formed large clumps with many stems. One larva was found in a burrow within a root, and the prepupa was found in soil directly under the center of the clump within five centimeters of the surface. There was no evidence of boring activity in the stems of the plant, suggesting that larval growth is completed below ground in the roots. In a highly detailed account of discovering the host plant of H.stramentosa in 1915 near Montréal, A. F.
This species is generally uncommon but has a wide distribution from Québec to North Dakota and south to Missouri, southern Illinois, Kentucky, and farther south in the Appalachians to North Carolina and Virginia. It is also found throughout the northeastern United States. A seemingly disjunct western distribution occurs in the mountains of western Nebraska, Colorado, Utah, Arizona, and New Mexico. Nothing has been found that distinguishes these populations from eastern populations.
Hydraecia immanis Guenée (Hop Vine Borer)
Hydroecia immanis Guenée, 1852, in Boisduval and Guenée, Histoire Naturelle des Insectes. Species Général des Lépidoptѐres, 5: 128.
Type locality: New York, USA. [NHML]
The name derives from the Latin immanis, enormous or very large, and this species is indeed among the largest of the genus, comparable to the Palearctic H. petasitis which it closely resembles. It is unlikely to be confused with any other North American species of Hydraecia, except perhaps H.micacea, but H.immanis is much larger and has different genitalic features, including an elongated, conical, obovate lobe constituting the cucullus which projects ventrally and partially overlaps the dorso-distally oriented digitus. Adult males, unlike those of H.stramentosa and H.perobliqua, have paired basal coremata on the first abdominal segment.
Like the larvae of H.stramentosa and H.micacea, larvae of H.immanis share a distinctive appearance due to the prominent, violet-brown pigmentation of the transverse bands on all abdominal segments which is interrupted by whitish middorsal and subdorsal longitudinal lines. This pigmentation tends to fade in the penultimate instar and may be totally lost in the final instar and prepupal stage, making identification of these Hydraecia more difficult. In H.immanis however, both the pigmentation and the contrasting white longitudinal lines are stronger than in the other two species and tend to be retained throughout the final instar. Also, the intersegmental areas of H.immanis are contrasting white. The yellowish prothoracic shield of H.immanis is irregularly edged on its anterior margin with black pigmentation, forming a distinctive pattern which distinguishes Hydraecia larvae from those of related genera.
The larva feeds on hops (Humulus), an apparent ancestral choice, and native prairie plants such as rosinweed (Silphium) (
Pupation occurs in soil surrounding the roots of the bored host. Adults are mainly encountered late in summer and early autumn. This species ranges from Nova Scotia to southern Manitoba, south through the Dakotas, Wyoming, Utah, and Colorado, and east through Missouri and the upper Midwest to Ohio, New Jersey, New York, and New England.
GENUS Selicanis Smith
Selicanis Smith, 1900. Proceedings of the United States National Museum, No. 1203, 22: 481.
Type species: Selicanis cinereola Smith, 1900 by original designation.
DIAGNOSIS OF GENUS SELICANIS. The sole known species of this genus could be confused with some species of Papaipema, from which it is readily distinguished by wing characters and by the genitalia of both sexes. It also bears a resemblance to Plagiomimicus spumosum (Grote); see the species account, below. The male valve bears an ampulla and the digitus is fused with a triangular cucullus which features both a corona and a dorsal, densely spinose crest, unlike Papaipema. In female Selicanis the distal margin of the lamella is not cleft, unlike the cordiform condition prevalent in Papaipema. Adults lack the broad, transversely flattened crest of scales on the prothorax characteristic of Papaipema. The genitalia show a closer relationship with the single species of Parapamea (see discussion there), but the adults of both genera bear no superficial resemblance to each other.
REDESCRIPTION OF GENUS SELICANIS. Smith’s original description is inadequate for comparative purposes and lacks the crucial diagnostic genitalic features, so the genus is redescribed here in greater detail. Head: antennae of both sexes ciliate, inconspicuously laminate [sensu
Selicanis cinereola Smith (Gray Wolf Moth)
Selicanis cinereola Smith, 1900, Proceedings of the United States National Museum, No. 1203, 22: 481.
Type locality: Denver, Colorado, USA. [USNM]
note — Named from three males and three females. A lectotype was designated by
Selicanis cinereola is a seldom-collected species which appears to be associated with grasslands, barrens, and prairie remnants. The moth has a distinctive appearance that renders it unlikely to be confused with species other than Papaipema. Similar Papaipema which share a range with S.cinereola include P.beeriana, P.necopina, P.maritima, P.silphii, and P.nebris. The prominent and smoothly rounded antemedial line in S.cinereola immediately separates it from Papaipema, in which the antemedial line is usually much less prominent and is strongly bent distally near the inner wing margin. The distal arm of the digitus which is fused with the cucullus and the cluster of spines on the vesica in the male and the heavily sclerotized appendix bursae in the female of S.cinereola also separate it from Papaipema. Plagiomimicus spumosum (Grote) in the subfamily Stiriinae bears a resemblance to Selicanis cinereola, but P.spumosum flies earlier in the season, and the antemedial and postmedial lines in this species are more angularly bent and less rounded and sinuous than in S.cinereola. The genitalia of these two are completely dissimilar. The host plant and early stages are unknown, but a possible clue might be extrapolated from the cryptic coloration of the adult forewings, which is reminiscent of certain late autumn flying Papaipema species which specialize on and lay their eggs upon the frosted, scaberulous, dead foliage of particular forbs, suggesting that the host is probably not a grass. Many of these cryptically colored species are specialists on hosts belonging to Asteraceae.
That S.cinereola is a borer with close phylogenetic affinities to Parapamea and Papaipema is not in doubt. Morphology of their genitalia suggest both Selicanis and Parapamea share a common ancestor with Papaipema. The species has been recorded from the central midwestern states and westward as far as Idaho. Available specimen data indicate a flight period of mid-September through October.
GENUS Parapamea Bird
Parapamea Bird, 1927. Journal of the New York Entomological Society, 35: 181.
Type species: Ochria buffaloensis Grote, 1877, by original designation.
DIAGNOSIS OF GENUS PARAPAMEA. A more detailed description than originally provided for this genus is included here with the diagnostics. This monobasic genus is characterized by the following set of features; characters in italics represent autapomorphies. In the male genitalia, the peniculus is rounded and devoid of hairs. The valves are bilaterally symmetrical. A very large, broad, acutely terminated, distally oriented clavus arises from base of the sacculus. The clasper is reduced to an appressed basal sclerite with a straplike bar, and an ampulla is present. The digitus is inconspicuous, fused with the sclerotized ventral margin of the cucullus, terminating in a small projection on the anal angle. The cucullus is broadly necked, paddle shaped, and lacking a corona. The juxta is fused with the greatly elongated anellus, forming a slender rod-like structure. The phallus is very long, strongly downcurved, and carinate with a ventral, sclerotized, ribbon-like bar bearing a cluster of short spikes extending onto subbasal part of vesica; vesica with a prominent, elongated patch of ordered, recumbent, subapical spines. The abdomen lacks basal coremata. In the female, the papillae anales are dorsoventrally flattened, with two reinforcing sclerotized, rod-like structures between them. Their shape is lyriform, with the lateral margins prominently constricted toward the anterior end, narrowly convex toward the posterior end, and having an acute apex. The surface of the papillae is covered with setae set in tiny membranous circles, both dorsally and ventrally. Segment A8 is 1.3 × as wide as long, with the anterior apophysis 1.2 × the length of segment A8 and the posterior apophysis 2.2 × the length of segment A8. Sternite A7 is a lightly sclerotized oval plate and is not differentiated into paired lodices. The lamella antevaginalis consists of a simple, v-shaped, sclerotized, ventral rim of the ostium. The ostium is moderately sclerotized. The ductus bursae is heavily sclerotized, very long, and sharply bent to the right about ½ its total length from the ostium. It is bent sharply again 90 degrees anteriorly before proceeding to the corpus bursae. The elliptical appendix bursae, viewed ventrally, is on the left, and has heavy, rugose sclerotization at the posterior end. The ductus seminalis exits from the posterior apex of the appendix bursae. The corpus bursae is oval, membranous, bears two signa, and is about the same length as the appendix bursae. Parapamea larvae are distinguished from otherwise quite similar-appearing larvae of Papaipema by the size of the lateral pinacula IV and V on A3–A8, as well as the dimensions of the anal plate. In Papaipema, the post-spiracular pinaculum IV is significantly larger than the pre-spiracular pinaculum V; in Parapamea, the reverse is generally true or both pinacula are subequal in size. In Papaipema, the anal plate is wider than long; in Parapamea, it is longer than wide.
An argument could be made for the inclusion of Parapamea within Papaipema. A similar case might be made for Selicanis. It is tempting to lump a monobasic genus like Parapamea with its apparent closest relative since its sole species P.buffaloensis (Grote) is a Papaipema in many respects. Were it not for the uniquely bizarre genitalia of both sexes of P.buffaloensis, which are so dissimilar to those of Papaipema as to render homologizing the structures difficult, it might be appropriate to do so. A stronger argument precluding such action is that employing similar criteria might necessitate the lumping of Papaipema with the Palearctic genus Gortyna, and possibly even including Amphipoea and Hydraecia, into a single large genus. These actions are untenable. Determining generic boundaries remains a problem not yet satisfactorily resolved. Aside from creating unwieldy groups by being overly inclusive, one must recognize that important historical biogeographic patterns might be obscured by such actions.
Parapamea exhibits a curious mixture of primitive and derived characters. It appears closely related to, yet abundantly distinct from, another monobasic genus, Selicanis. The two show similar derivations in the female ductus bursae which may be synapomorphic. The partial fusion of the digitus with the cucullus in the males of both genera appears to represent a transition to the unique bifurcate condition of the cucullus characteristic of Papaipema. Parapamea and Selicanis also barcode together. Both are obvious close relatives of Papaipema. Otherwise, the genitalic characters of Parapamea reflect mainly retained plesiomorphic states, whereas its larva shares synapomorphies with Papaipema, excepting the aforementioned autapomorphy, as does the adult. The larva of Selicanis is unknown.
Parapamea buffaloensis (Grote) (Lizard’s-tail Borer)
Ochria buffaloensis Grote, 1877, The Canadian Entomologist, 9: 88.
Type locality: “Buffalo” [New York, USA]. [NHML]
note — Described from a single female.
Hydroecia latia Strecker, 1899, Lepidoptera, Rhopaloceres and Heteroceres, Indigenous and Exotic, Supplement 2: 6.
Type locality: Chicago, Illinois [USA]. [FMNH]
note — Described from a single specimen.
† Xanthoecia buffaloensis simplicissima Bird, 1916, Journal of the New York Entomological Society, 24: 90.
Type locality: Wilmington, Delaware [USA]. [AMNH]
note — Unavailable infrasubspecific name. Described from four specimens.
The white-spotted phenotype of Parapamea buffaloensis, perhaps less common than the immaculate morph, can be confused with Papaipema matheri, and both species are likely to occur together. A white suborbicular spot is never present in Parapamea buffaloensis, but almost always is in Papaipema matheri, which also features a sinuous, doubly crenulate postmedial line. In Parapamea buffaloensis, this double line is straight until immediately before curving inward to meet the costa. The genitalia of these two species are dramatically different. The abundant hadenine noctuid Nephelodes minians Guenée, which flies at the same season in more northern portions of the range of P.buffaloensis, can also be mistaken for its immaculate phenotype. The forewing pattern of N. minians is generally less contrasting in the terminal area, with the coloration less bright, and N.minians is structurally different, with narrowly pectinate antennae in the male and hairy eyes. The postmedial line of N.minians is sinuous, unlike that of P.buffaloensis.
The larva is long and slender but not readily distinguishable from those of many Papaipema species, having the middorsal white stripe more or less continuous, and the subdorsal longitudinal stripes broken across abdominal segments A1–A4.
The sole host plant is lizard’s tail (Saururus cernuus), a hygrophilous plant which typically grows in standing water. Presence of the borers in these plants is indicated early in the season by wilting or dead terminal portions evident among the surrounding vigorous growth. Whitish frass expelled from exit holes in the stems is prominently visible floating on the water. The larvae are excellent swimmers, actively moving across the water surface in a serpentine fashion from one plant to another, and eventually mine extensive portions of the root. Aside from their voracious appetites, the larvae are frequently obligated to move to new plants as a result of inundation following rainfall, which often submerges the exit holes required for air passage to the galleries occupied by the larvae. See
Adults are on the wing the entire month of September in the more northern portions of its range, with a much more protracted season in the deep South. (See discussion of this phenomenon relating to voltinism in the general introduction.) The species is only rarely encountered in the northern portions of its range but becomes much more common southward. One is struck by the irony that this insect is named for the place it was first collected, which to this day, except for the Manitoulin Island specimen, remains the northernmost record for a species that ranges south to central Florida and covers most of the eastern half of North America.
GENUS Papaipema Smith
Papaipema Smith, 1899, Transactions of the American Entomological Society, 26: 2.
Type species: Gortyna cerina Grote, 1874, by subsequent designation by Hampson, 1910a.
note — The name Papaipema is derived from the Greek papai, denoting an exclamation of enduring suffering, plus pēma, suffering, or calamity. In a private letter from Henry Bird written to August Busck at the U. S. National Museum in 1930, Henry related that J. B. Smith once told him “in serious confidence that Papaipema was the only noctuid genus he could not fathom.” It appears that Smith’s agonizing over this large group of moths, so diverse and with so many deceptively similar in appearance, was indeed a source of frustration and suffering for him and perhaps the inspiration for his choice of this name.
Ochria; Grote, 1874, not Hübner, 1821. Apamea; Grote, 1874, not Ochsenheimer, 1816. Gortyna of authors, not Ochsenheimer, 1816. Hydroecia of authors, misspelling, not Guenée, 1841. Emboloecia Hampson, 1910, Catalogue of the Lepidoptera Phalaenae in the British Museum. Vol. 9: 59.
Type species: Ochria sauzalitae Grote, 1874, by subsequent monotypy.
Paipema; Rockburne and Hardwick, 1970, misspelling.
In 1899, J. B. Smith published his revision of the genus Hydraecia wherein he first proposed the name Papaipema for what he considered a subgroup of Hydraecia with characteristics distinguishing it from his much more inclusive and diverse concept of the genus. He gave no explanation for his choice of name, and a number of years passed before other authors gradually adopted Papaipema as a genus distinct from Hydraecia and of equal status with it. Dyar (1902) was the first to treat Papaipema as a full genus, placing P.cerina first in his list.
Adult Papaipema fly from late summer throughout autumn and are cryptically colored to match the prevailing autumn foliage colors of their senescing host plants and surrounding vegetation: brick red, orange, yellow, and various shades of light to dark brown or gray. Commonly superimposed on these often-colorful backgrounds are prominent stigmata on the forewings rendering members of this large North American genus, consisting of 53 known species, immediately recognizable by a distinctive maculation pattern shared with few other genera. Collectively, this maculation constitutes the entire orbicular-reniform complex, which either may be expressed as contrasting white or yellow stigmata against a background of different coloration or as dark stigmata which may or may not contrast with the background. Frequently, both conditions occur within a single species. This particular constellation of stigmata is also found in the related Palearctic genus Gortyna, incompletely developed in the Nearctic genus Parapamea, and curiously represented in the genus Hydroeciodes. On
The reniform (meaning “kidney-shaped”) is a familiar structure in noctuid moths. In some Papaipema group genera, it is particularly prominent and consists of a crescent-shaped, vertical spot called the central lunule, often of different coloration than its surrounding satellite spots. The satellite spots, often white or yellow and six or seven in number, may all be of the same color, or the one or two outermost spots may be yellow and the others white. They are arranged around the central lunule in a pattern resembling a figure 8 when broadly spaced, but sometimes the reniform is more or less laterally compressed and may also be more slanted and less perpendicular to the inner margin. Subtle differences in reniform shape, size, color, and proximity to the postmedial line may offer important diagnostic clues to the identity of the species. Basad of the reniform in the center of the discal cell lies the orbicular spot. It is usually but not always the same color as the reniform, normally round, occasionally somewhat angular, and in a couple instances, a vertical bar. It often but not always contains a dark central spot.
In Papaipema, Gortyna, and Hydroeciodes a pair of double spots lie immediately below the orbicular in the wide space between the second cubital vein and anal vein 1A+2A. These two very closely spaced spots may be equal or subequal, and in rare instances, reduced to the degree that one is obsolescent. The pair appears divided and is superimposed exactly at its division on the fold line, which is typically obscure in many Noctuidae, but weakly expressed or even well defined in some larger species of Gortyna and Papaipema, especially large-bodied females. In addition to being equally divided above and below the fold line, these spots sometimes incorporate a small section of the double antemedial line as in Gortyna, creating the appearance of a cluster of four conjoined spots. This pair of spots is commonly referred to as the claviform but is not homologous with a true claviform. These spots are designated herein as suborbicular, meaning “below the orbicular,” and used as a noun in the same manner as the adjectives “reniform,” “orbicular,” and “claviform” are employed as nouns, rather than this word’s traditional usage as an adjective meaning “almost round.” The claviform (meaning “club-like”) is a familiar, often prominent forewing feature of numerous noctuid genera such as Euxoa Hübner, Agrotis Ochsenheimer, Feltia Walker, Apamea, and Psaphida Walker which extends horizontally across part or all of the medial space, often incorporating the basal and medial dash, if present.
During the decades since the inception of this project, the senior author examined far in excess of the 15,000 specimens of Papaipema recorded as deposited in 96 institutional and private collections listed in the appendix, and AEM and JRW have added many more to that total. In not one instance were all specimens in any of these collections correctly identified. Particularly with regard to the genus Papaipema, this is mainly due to the large number of species which closely resemble one another superficially. An additional factor contributing to rampant misidentification is the confusion prevailing in earlier literature. The reader needs to remain acutely aware that names utilized in that body of literature may refer to entirely different species than those to which the same names are applied in this work. Much of the confusion in previous literature arose from the fact that type specimens of species described by Guenée and others in the nineteenth century were deposited in European institutions and consequently never actually seen by American authors of works published subsequent to the original early descriptions. New species described well into the twentieth century were likewise named by authors who never had direct access to earlier type material. The result has been a proliferation of synonyms and named forms, contributing to the relatively large synonymy listed in this work. Detailed accounts of taxonomic problems are provided with the discussions of individual species.
Reviewing all the literature and resolving this confusion has proved a daunting task. In the process, ELQ examined all extant type material in the genus Papaipema. The resulting taxonomic changes were reflected in a checklist (
Uncertainty expressed by previous American workers resulted from their lack of opportunity to directly examine type material of early authors, some of which had already been lost. In some instances, too much faith was placed in comparisons with painted illustrations or third-party opinions provided by persons unfamiliar with these moths. Regarding lost Papaipema types, it is worth summarizing what is known about them.
The types of species collected by Doubleday in America and described by Guenée in 1852 were all believed to have been destroyed by dermestid beetles (
A more complex situation exists regarding missing Grote types, especially those jointly described by Grote and Robinson. A number of Grote’s original types are in the NHML. A few are deposited at ANSP. One is in the NYSM at Albany. Other syntypes have not been located anywhere. According to Franclemont (pers. comm.), original Grote type labels all bear red borders with the genus and species name and the handwritten word “Type.” Blue-bordered Grote labels are only his determination labels and not necessarily indicative of syntypes. Inexplicably, none of these labels are present on the ANSP Grote Papaipema types, although their authenticity is not questioned. A significant number of putative “types” in the NHML appear to be spurious, including some of those referenced in
With regard to the Papaipema type material of other authors, nearly all, if not all of which has survived to this day, a few generalities can be stated. Types of taxa described by Dyar, Benjamin, Barnes and Benjamin, and Barnes and McDunnough are deposited in the USNM. J. B. Smith’s type material is divided between the USNM and the AMNH. Lyman’s type specimens are at the LEM in Montréal. Of nearly 100 specific, infraspecific, and varietal names that have been proposed for members of the genus Papaipema, almost a third of them are attributed to Henry Bird. Over a span of about forty years of activity, Henry communicated with and exchanged or donated a multitude of specimens with all the other workers of his day who shared his interest. Accordingly, his specimens may be found today in collections throughout North America and Europe. His primary collection, including his type specimens, was given in its entirety in 1960 to the American Museum of Natural History in New York City. One might naturally assume that all his type material is deposited there as well. This would be true, were it not for his unfortunate habit of “upgrading” his type specimens on occasion, leading to genuine syntype material sometimes being given away when, after publication, he happened to acquire better specimens which he then labelled as types and kept for his collection. Bird understandably lacked a modern holotype concept, employing instead such terms as male type, female type, or co-type as was common in his time, and did not appreciate that the specimens had to be in his possession or seen by him prior to the publication of their names.
Papaipema species occur predominantly in eastern North America, occupying all faunal zones except the subarctic and subtropical extremes, with the greatest diversity occurring in mid latitudes. A much smaller number occur in western North America, and, of these, some are nearly transcontinental in distribution. The genus is poorly represented in arid regions of the southwestern United States. Throughout their range, Papaipema are preponderantly inhabitants of mesic and especially wetland habitats. Host plant specialization appears to have been a primary force driving speciation in Papaipema. This in turn may have been influenced in part by host plant chemistry, discussed in greater detail elsewhere in this fascicle. Glaciation also appears to have played an important role, if not in speciation, then certainly with regard to present-day distributions. Regarding scarabid beetles occurring in regions glaciated during the final Wisconsin ice age,
The most highly differentiated and presumably oldest species in the genus tend to have more southerly distributions. Many less well-differentiated and apparently much younger species today occupy areas that would have been largely devoid of plant and insect life during the most recent glaciation events in North America. It is possible to find at a single location species which could have become differentiated twenty or thirty thousand years ago flying together with other Papaipema species representing direct descendants of lineages perhaps several tens of millions of years old.
DIAGNOSIS OF GENUS PAPAIPEMA. In recognizing Papaipema as a group distinguished from others within his broad concept of Hydraecia, Smith gave weight to the thoracic vestiture, pointing out the broad, transversely flattened crest (
Head: antennae of both sexes ciliate, inconspicuously laminate [sensu
Larvae of Papaipema together with those of the closely related monotypic genus Parapamea are distinctive and quite unlike those of any other known North American borers. The transverse, heavily pigmented cuticular banding encircling the first four abdominal segments in combination with partial to complete longitudinal striping of the thoracic and remaining abdominal segments is diagnostic. In many species the characteristic dark purplish to pinkish brown pigmentation of the cuticle fades in late instars, in which case the following character, present throughout, serves as a further diagnostic trait. The sclerotized prothoracic shield, that is generally light, honey-brown, bears contrasting black-pigmented lateral margins which are immediately recognizable. Other larval features distinguishing Papaipema from other apameine borers are much less apparent but comparison with the figures on the plates should suffice. It should be noted that Papaipema larvae can be further categorized into subgroups which will facilitate species level recognition as provided in the key. The white longitudinal stripes in some species are interrupted on segments A1-A4, leaving the solid, pigmented bands on these segments intact. In many species, only the single dorsal longitudinal stripe traverses this band. In other species, both the dorsal and two subdorsal stripes are continuous. To a limited degree, these features can vary between early and later instars but are nonetheless quite useful.
The longitudinally striped habitus characteristic of noctuids with larvae that are associated with external feeding on grasses, or that rest by day on stems of their hosts, which is assumed to represent a plesiomorphic condition, is retained to varying degrees by larvae of Papaipema. This striped condition, whether partial or complete, is more often boldly expressed in early instars but often faded or lost in later instars, especially in those species which move to the underground parts of their hosts and complete the majority of their development there. Some species however retain conspicuous stripes through all instars, particularly obligate stem borers.
LIFE CYCLE AND BIONOMICS OF PAPAIPEMA
All Holarctic Apameini are univoltine, and like the great majority of autumn-flying apameine genera, Papaipema overwinter in the egg stage. Eggs are deposited in short rows comprising from a few to a dozen or more ova which are tucked into crevices on the dead or senescing host plant foliage. These are then carefully cemented shut with a glue-like substance secreted by the female’s colleterial glands. The eggs remain securely protected from the environment and predation until early spring when the neonate larvae emerge and commence searching for tender new growth of the host plants emerging from the soil. A statement by
Without exception, larvae of Papaipema feed internally upon the stems, rhizomes, roots, or other subterranean portions of their host plants. In the earliest instars Papaipema larvae lack their full complement of abdominal prolegs and are semi-loopers. This feature which enhances their mobility is necessitated by having overwintered as eggs within dead vegetation on the ground, requiring neonates to explore their microhabitat for emerging food sources before adopting an endophagous existence. This would appear to be a trial-and-error process. If unable to quickly locate the appropriate host, young larvae may be forced to take up temporary residence in whatever tender plant may be available. Occasionally this surrogate host may prove sufficient for larvae to complete development, but most often this is not the case. To some degree this probably accounts for multiple or even many host plant species having been recorded for certain Papaipema species. Such records can be misleading, as the large majority of Papaipema species are highly host specific and only stray into unrelated plants under unusual circumstances. After the appropriate food is detected and feeding commences, it may still be necessary for young larvae to relocate to larger stems capable of better supporting them. This is especially likely to occur with species that remain stem borers throughout most or all of their development, or those that have selected host plants that prove to be too small to allow complete development in a single plant.
Taken as a whole, Papaipema have evolved host plant specializations across an astonishing range of the plant kingdom. From fern-feeding specialists derived independently in two groups, to a few species retaining the ancestral apameine association with grasses, to a specialization in other monocots progressing to a radiation into dicotyledonous plants culminating in a large group of Asteraceae feeders, the evolution of the genus seemingly parallels that of its hosts. An intriguing observation of some species with transcontinental distributions is that, in the eastern portions of their ranges, these species tend to be monophagous or narrowly oligophagous, becoming increasingly polyphagous westward. For further discussion of this phenomenon, see
Years of observation reveal a vast discrepancy with regard to the amount of plant tissue consumed by larvae of different species of Papaipema, suggesting that some host plants or parts thereof may have much greater nutritional value than others. Species such as P.circumlucens or P.maritima spend their entire larval development within the confines of a gall, no larger than an acorn, formed by their presence in the plant. Other species like P.harrisii or P.cataphracta mine hollow stems extensively, producing copious amounts of frass, often an obvious clue to their presence. The same is true of subterranean borers. Over its entire larval development, P.amianthii consumes an amount of plant tissue smaller than a cherry; larvae of an even smaller moth, P.stenocelis, mine the rhizome of its host for up to one meter.
When larvae are mature, several pupation options are available that are typically consistent within any given species. Some stem borers e.g., P.maritima, produce a gall large enough to serve as a pupation chamber in which an exit hole is prepared by the larva in advance of pupation. Other gall makers, such as P.circumlucens and P.blackorum, abandon their galls and move to the soil for pupation. Many stem borers like P.harrisii and P.sulphurata pupate inside their galleries, after having prepared an appropriate exit hole, generally hidden by a parchment-like layer of plant epidermis which the emerging adult easily breaks through. However, the majority of Papaipema species form shallow pupation cells in soil or among detritus near the base of the host plant. Some Papaipema species, especially those which develop rapidly and mature early in the season, enter a period of larval aestivation in their galleries or in soil, delaying pupation until late summer or early autumn.
Most Papaipema form typical obtect pupae, usually of a reddish-brown, light brown, or honey color, and form their pupae in soil, detritus, or large galleries. Those species that form pupae in tightly constricted stem galleries, such as P.impecuniosa, P.cataphracta, and P.sulphurata, form pupae that are narrow and cylindrical. These same species will form narrow, cylindrical pupae even if they are not confined to the tight quarters of the larval burrow. Pupal development progresses at a uniform rate; there is no pupal diapause among Papaipema species. For those species with larvae that mature by early summer, aestivation allows larval mobility in the event of high temperatures or desiccating conditions. Once pupation occurs, that option is removed. The average pupal period for all species is approximately 28 days.
The adult moths hide by day, usually resting on matching foliage, but never on bark of trees, shrubs, rocks, etc. By night, the moths actively seek mates and oviposition sites, and most species are readily attracted to blacklight, mercury vapor, or other light sources placed among or very near the host plants. They are strong fliers, yet generally do not stray far from their breeding sites. Occasionally, adults come to baits or feed at flowers at night, but this seems relatively rare and is not a reliable method for collection or observation of these moths. The tongues of all Papaipema species are fully functional, but probably utilized primarily for imbibing moisture droplets on vegetation. The primary energy source for adults is stored fat reserves acquired during the larval stage. Specimens collected when freshly emerged and prior to having metabolized their stored fat reserves through flight activity are notoriously prone to heavy greasing in collections, much like cossids, megathymids, saturniids, and other Lepidoptera which do not feed in the adult stage, or which overwinter as hibernating adults. It is recommended that in such instances, valuable specimens routinely be degreased with an appropriate solvent if they are being retained for future reference and study. Petroleum naphtha, including white gas, is ideal and inexpensive.
For additional information on the life cycle and bionomics of Papaipema, see
ABERRATIONS AND HYBRIDS
Occasional aberrations among Papaipema are seen when sufficient numbers of specimens have been amassed. These generally consist of individuals with elements of their forewing patterns distorted, exaggerated, or asymmetrically distributed. Almost without exception, such aberrations involve elements of the reniform, orbicular, or suborbicular stigmata rather than other forewing pattern features. Exactly why this appears to be so is not clear, but perhaps may be due to the high degree of phenotypic variability characteristic of these stigmata in this genus. One example wherein an additional element is introduced to the normal pattern is seen on plate A-7, figure 9; another extreme example is figured on
Of far greater significance relative to the complex evolution of species in this large genus and in related genera is the incidence of hybridization. It seems reasonable to assume that this is a relatively rare event among insect populations under natural conditions, but in the case of Papaipema, hybridizing may occur with far greater frequency than one would imagine. There are factors which would tend to favor hybridization among these species. First, like in most other apameine genera, adults are very sedentary, and their local populations are often concentrated in quite small areas. These same habitats commonly are shared with numerous congeners to the point of becoming crowded with both numbers of individuals and diversity of species. In Pennsylvania, ELQ recorded sixteen Papaipema species coexisting in a single wet meadow of only a few acres in area. Under these conditions, opportunities for interbreeding are abundant. Second, the physical ability to interbreed merits consideration. Lepidoptera have evolved various mechanisms to thwart interspecific mating, such as employing different pheromones, stratified mating times, morphological adaptations in genitalic structures, etc. Some groups appear to favor means of achieving reproductive isolation which differ from those employed by other groups. Within the Apameini, the complex genitalic morphology that is carried to an extreme in some genera would appear to serve as a primary mechanical reproductive isolating mechanism. The so-called “lock-and-key” hypothesis (
The significance of hybridization and repeated backcrossing in a group such as Papaipema and the effects introgression and gene flow may have on the evolution of species in that group cannot be discounted. When there exist abundant opportunities for the transfer of one species’ genes to the gene pool of another, the rate of divergence between them may be greatly hindered, along with the ability to distinguish these entities. CO1 barcodes showing divergences of as little as 0.5%, and even considerably less, may obscure the true relationship between two biological entities. These may appear to be conspecific when multiple additional data clearly suggest they are not. Such is the case with at least 19 Papaipema species, which suggests that they diverged too recently (i.e., less than a million years ago) to have accumulated sufficient mutations to render them recognizably distinct with barcode data alone. Stated differently, about ten species treated in this fascicle would be reduced to synonymy in the absence of data contradicting the molecular information presently available.
Two Papaipema hybrids have received taxonomic recognition. Papaipema aerata (Lyman) was described as a species in 1901. It has remained an enigmatic and rarely collected entity but has maintained its status by default in check lists until the present. Laboratory breeding experiments conducted by Ted Herig in Michigan have confirmed, as long suspected, that it is in fact not a species but a naturally occurring hybrid between P.necopina and P.cataphracta. These experiments continued with inbreeding and backcrossing for five generations with gradually diminishing viability but with retained fertility for much of the process. Accordingly, the taxon aerata is here treated as an unavailable name. In 1911, Papaipema fluxa was named by Henry Bird as an aberration of P.cataphracta. As an infrasubspecific taxon, the name fluxa remains unavailable. It appears to represent yet another naturally occurring hybrid involving P.cataphracta and some other species.
PARASITOIDS AND PREDATORS
Predation upon larvae by the usual suspects (birds, vespine and polistine wasps, reduviid bugs, stink bugs, etc.) is apparently minimal on account of the endophagous larval feeding mode. Pupae however are highly vulnerable to rodent predation, and perhaps this is responsible for the evolution of a relatively short pupal period. A curious observation repeatedly reinforced over many years of general rearing of Lepidoptera is that little or no rodent predation upon larvae reared outdoors in sleeves or cages was noted, but high vulnerability to rodents commences as soon as pupation occurs. Skunks have been recorded as being so efficient in their predation upon both larvae and pupae of Hydraecia immanis that they have been regarded as valuable allies during the 19th century in the struggle by hop farmers to protect their crops from destruction by these borers (
While their endophagous feeding mode affords these larvae protection from many predators, larvae are still vulnerable to invertebrates within the larval burrow, including through cannibalism by individuals of the same moth species. Larvae of Papaipema often have dark and heavily sclerotized anal plates which offer them some protection from enemies that enter the burrow. Still, predation by parasitoids in the hymenopteran families Braconidae and Ichneumonidae and the dipteran family Tachinidae can take a heavy toll on these larvae. Parasitism rates over 90 percent have been documented by AEM in some species such as Papaipema stenocelis and Papaipema amianthii, but in most species, parasitism rates are much lower or even seemingly non-existent. It should be noted that most collections of larvae by the authors have been of those in middle to late instars, and so parasitoids that prey upon and complete development in ova and in early instars have largely avoided documentation herein. In most years in the woodlands of Carbon County, Pennsylvania, searches for larvae of P.amianthii can often yield ten or so larvae in about a half hour’s work after examination of about two hundred fly poison plants, and one or two of the larvae are usually parasitized by the large ichneumonid wasp Lissonota brunnea Cresson. In some years however, larval densities of P.amianthii can be very high, with over 50 percent of fly poison plants bored by caterpillars. Larvae collected during these events have shown parasitism rates by this wasp of over 90 percent. The following year it can be difficult or impossible to locate amianthii larvae in these same woodlands, and it can take a few seasons before amianthii numbers return to normal densities. The conditions that lead to these fluctuations in prey and predator levels are unknown. In contrast, collections of Papaipema rubiginosa reared from larvae from an open, calcareous fen in Sussex County, New Jersey over many of the past 17 years have not yielded a single parasitoid. This situation is even more interesting when one considers that a sister species of P.rubiginosa found in forests nearby, Papaipema astuta, is commonly attacked by the ichneumonid wasp Erigorgus ontariensis. It has been theorized that the disparity in the incidence of parasitism in these two closely related moths may be related to differences in the chemical composition of the host plants (James Adams, pers. comm.). The host plant of Papaipema rubiginosa, purplestem angelica, contains high concentrations of phototoxic furanocoumarins, while the host plant of Papaipema astuta, stoneroot, lacks toxic compounds.
Since parasitoid adults can cue in on volatiles from larval feculae to locate potential hosts, many moth species have developed the ability to launch fecal pellets away from themselves to help them avoid detection. In many Papaipema, larvae drop frass out of the burrow through a hole made in the plant stem, and the frass accumulates on the substrate below. It is unknown if the large concentration of frass below helps protect the larva from detection above. In Papaipema appassionata, frass and silk are used together to build a chimney-like structure at the entrance to the burrow, which not only affords protection for the larva against high water levels in its wet habitat, but possibly also makes it difficult for parasitoid predators to access it through the fecal pellet shield. Several other species create a mass of wet frass at the burrow entrance which may aid in protection from parasitism. Papaipema marginidens and some Hydraecia species attempt to avoid parasitism by depositing much of their frass under ground instead of maintaining a frass orifice at the surface. Parapamea buffaloensis is one species in this group that has developed the ability to launch or spray its frass from the burrow entrance, and the lizard’s tail plants adjacent to the one being bored can appear “white-washed” with feculae, possibly in order to divert the attention of potential parasitoids elsewhere.
Based on collections of mostly middle to late instars, the ichneumonids Lissonota brunnea and Erigorgus ontariensis Dasch were the two most commonly observed Ichneumonidae. Lissonata brunnea was found parasitizing Papaipema amianthii, P.lysimachiae, P.marginidens, P.rutila, P.stenocelis, and P.sullivani, and the size of the adult parasitoid varied based on the size of the Papaipema larva. Once it completes development, the larva of L. brunnea exits through the moth larva’s integument and forms a dark brown, oblong, tough, pill-shaped cocoon in which it overwinters. E. ontariensis was found to parasitize Papaipema astuta, P.arctivorens, P.cataphracta, P.cerussata, P.duovata, P.nebris, P.necopina, P.nelita, and P.rigida. In E.ontariensis, the wasp larva waits until after the host moth larva has pupated before if finishes feeding, and it ultimately pupates and overwinters within the empty moth pupa, from which it emerges in the spring. Other genera of ichneumonid parasitoids reared during the course of this work include Chasmias Ashmead, Ichneumon Linnaeus, Phobocampe Förster, and Hyposoter Förster. One of the early instar collections by AEM, that of Papaipema pterisii, yielded a wasp in the genus Campoletis Förster.
Braconid wasps in the subfamily Microgastrinae are commonly found parasitizing Papaipema and relatives. Two species were found in rearings of these groups, Lathrapanteles papaipemae (Muesebeck) and Microplitis gortynae Riley. Larvae of both species emerge en masse from the moth larva and spin cocoons in a cluster alongside it. The moth larva remains alive but immobile for days to weeks after the wasps exit its body to pupate. In L.papaipemae, the cocoons are pristine white and cottony in appearance, while in M. gortynae, the cocoons are a light grayish-tan and are corrugated and papery in appearance. Both species overwinter in the cocoons. Lathrapanteles papaipemae was reared from Papaipema matheri and M.gortynae from Papaipema cerina, P.limata, P.rutila, and P.stenocelis. While searching for pupae of Papaipema duovata in New Jersey and in Rhode Island, AEM found linear clusters of about 10 dark brown cocoons tightly packed within some of the larval burrows. The individual brown cocoons within the cluster were papery in appearance and were roughly triangular, and the parasitoid that produced them belongs to the genus Bracon Fabricius from another subfamily, Braconinae. Larval rearings in the genus Cherokeea also produced braconid wasps, these from the genus Meteorus Haliday in the subfamily Euphorinae. These wasps, which when finished feeding issue from the moth larva and suspend themselves on a silk thread where they spin a pale, warm brown cocoon, were reared from Cherokeea odonnellae and Cherokeea slotteni from the North Carolina mountains.
Dipteran parasitoids in the family Tachinidae are also important predators of Papaipema and relatives. These flies exit the larva when finished feeding and form puparia. The large, metallic, greenish-blue fly Gymnocheta ruficornis Williston was reared from Papaipema inquaesita and P.rutila in New Jersey. A small tachinid fly in the genus Lydella Robineau-Desvoidy was found in Papaipema hesseli near Albany, New York, and in this species, several fly maggots emerged from a single moth larva. Flies in the genus Lixophaga Townsend were reared from Papaipema cerussata and P.lysimachiae, and also from the cane feeder Arundinariaphila hayesi.
In the mountains of North Carolina, two species of cane-feeding moths boring in Arundinaria appalachiana were found to be parasitized by nematodes in the family Mermithidae. In each case, a single, long hairworm emerged from the moth larva. The species infected were Cherokeea attakullakulla and C. odonnellae.
Rearing parasitoids to the adult stage is relatively easy. In species with larvae that emerge from the moth caterpillar to form external cocoons and puparia, detection of these parasitoids is straightforward. For those species that complete feeding and remain within the moth pupa, detection is still possible. For example, the normally active abdominal segments of Papaipema pupae that are parasitized by the ichneumonid wasp Erigorgus ontariensis soon become rigid and immobile. These unhatched pupae can be overwintered and will produce adult wasps in spring. Many smaller hymenopteran and dipteran parasitoids will emerge within a few weeks of pupating if kept in a container with some slightly moist paper toweling. Overwintering fly puparia and wasp cocoons can be stored in a chilled location using the same techniques that are appropriate for overwintering live moth pupae.
Nonparasitic invertebrates may also take their toll, despite the fact that these larvae bore inside plants. Upon examination of thousands of larval burrows of Papaipema and relatives over years of field work, it has become apparent that these burrows can be occupied by ant colonies. Whether these colonies overtake the burrow by first consuming the resident larva, their activity causes the larva to vacate the burrow prematurely, or the ants colonize after the larva has left or succumbed for other reasons is a subject for further study.
DNA BARCODE VARIATION IN PAPAIPEMA
By B. C. Schmidt
The importance of DNA sequencing in the last decade has transformed modern taxonomy and systematics, and the merits and methods of molecular techniques have been discussed at length (e.g.
Papaipema contains numerous species with lower-than-average barcode divergence from other species, often 0 – 1% for closely related species. Although the average divergence between North American noctuid species is 2.47% (
In contrast to most noctuid genera, Papaipema have a high incidence of very similar or identical barcodes among multiple species, resulting not only in shared BINs but barcodes that are not species-diagnostic. Conversely, there are several instances of high divergence within a species, resulting in BIN splits. The reasons for this incongruence between biological / morphological species delimitation versus mtDNA variation are unstudied in Papaipema, but likely are similar to those identified in other Lepidoptera groups with such patterns, including recency of evolutionary divergence and hybridization. Shared barcodes among morphologically distinct species does not necessarily imply rampant hybridization (Schmidt and Sperling 2008), and can indicate that novel mtDNA may confer an evolutionary advantage once introduced to a species’ gene pool. Given discordance between species and DNA barcodes in Papaipema, additional molecular methods such as whole-genome approaches will be needed to confidently resolve the taxonomic and phylogenetic uncertainties that remain. Despite the limited taxonomic utility of DNA barcodes in Papaipema, some important inferences can still be drawn in certain cases.
The range of barcode variation in Papaipema is skewed to two extremes, where minimum divergence rates between species (interspecific) are either exceptionally low, or higher than average. Thirty-one Papaipema species (58%) exhibit very low minimum nearest-neighbor divergence and share BINs with at least one other species. Typically, nearest-neighbor differences of less than 1% are found in this category, and at least 12 of these 31 species can exhibit 0% divergence (haplotype sharing). Conversely, ten species show minimum nearest-neighbor differences greater than 2.5%; this category represents phylogenetically isolated, morphologically distinctive species. The remaining twelve species (~23%) have minimum divergence rates similar to those of other noctuid genera of moderate- to large size, at 1.0-2.5%. The above summary is based on barcodes for all Papaipema species, and barcode variation of a taxon is briefly summarized in each species account. The overarching pattern across Papaipema appears to mimic the evolutionary track of Apameini as a whole, characterized by many small or monotypic genera versus a few, speciose genera (Zilli et al. 2009) that presumably reflect recent species radiations.
IDENTIFICATION OF PAPAIPEMA ADULTS
A key to adult Papaipema is not presented in this fascicle. Using a dichotomous key to identify species in a genus of this size approaches or exceeds the limits of what realistically can be achieved. With the rampant polymorphism prevalent in so many Papaipema species and with numerous species so similar in appearance to others, it is impossible to construct a key that would actually work. A discussion of similar species is provided with each species treated herein. It is hoped that use of the species diagnostics provided in the text in combination with distribution maps, illustrated adults, knowledge of larval host plant possibilities, and genitalic figures, will at least make possible the reliable identification of most specimens encountered in this very difficult group. Resorting to genitalic dissections proves to be the final and best method of identifying many of the genera and species covered in this fascicle, as these are ultimately the primary characters upon which the genera are founded. Unfortunately, in the genus Papaipema, characters of the male genitalia which tend to be the most useful for species-level discrimination in many genera are of limited and restricted utility. They are extremely useful in diagnosing some taxa and of little value in distinguishing others.
Key to Species of Papaipema (Larvae)
1. Larva with pale, whitish, longitudinal mid-dorsal and subdorsal stripes found on at least the thoracic segments and posterior abdominal segments 4
— Larva lacking pale, whitish, longitudinal stripes 2
2. Larva with mid-dorsal and subdorsal whitish lunules on anterior edge of A5–A8 and dark pigmentation encircling A1–A4; larva completely white at maturity with no dark edging on prothoracic shield and tiny, pale pinacula; a borer in Midwestern ironweed species Papaipema limpida
— Larva without whitish lunules 3
3. Young larva with a segmented, “ringed” appearance; mature larva pinkish-white; A borer in meadow-rue Papaipema unimoda
— Larva with a “ringed” appearance throughout its development; integument a deep brownish-pink; a borer in blazing star Papaipema beeriana
4. Dark, purplish-brown pigment encircling A1–A4 interrupted by at least the mid-dorsal pale, whitish, longitudinal stripe 5
— Dark, purplish-brown pigment encircling A1–A4 uninterrupted by any of the pale, whitish, longitudinal stripes 29
5. Dark, purplish-brown pigment encircling A1–A4 interrupted by only the mid-dorsal pale, whitish, longitudinal stripe 6
— Dark, purplish-brown pigment encircling A1–A4 interrupted by the mid-dorsal and the subdorsal pale, whitish, longitudinal stripes 25
6. Mid-dorsal longitudinal stripe an extremely thin line traversing the dark, purplish-brown pigment encircling A1–A4; line sometimes broken or absent across these segments; a borer of the Atlantic and Gulf coastal region in Virginia chainfern Papaipema stenocelis
— Mid-dorsal longitudinal stripe robust and a similar width down the entire length of the larva 7
7. Well-marked larva with very large, dark D1 pinacula on A1–A3 that are nearly as large as the dorsal pinacula on A8, a borer in Hercules’ club Papaipema araliae
— Larva without very large D1 pinacula on A1–A3 8
8. A borer in stoneroot 9
— A borer in other plants 10
9. Larva boring in the stem and barely entering the root Papaipema astuta
— Larva boring in the lower stem and also widely in the root, forming a large cell in the root and sometimes pupating therein Papaipema duplicata(part)
10. A borer in various Apiaceae 18
— A borer in other plants 11
11. A borer in ferns 12
— A borer in other plants 13
12. A borer in brackenfern Papaipema pterisii
— A borer in ostrich fern Papaipema hesseli
13. A borer forming a gall in vines 14
— A borer in other plants 15
14. A borer in hops; an eastern species of prairie and prairie edge habitats Papaipema circumlucens
— A borer in woolly pipevine; a species of the lower Mississippi drainage Papaipema blackorum
15. A borer in hairy leafcup 16
— A borer in other plants 17
16. Larva with post spiracular accessory pinacula on A7 larger than spiracle on the same segment Papaipema polymniae
— Larval with post spiracular accessory pinacula on A7 smaller than spiracle on the same segment or lacking Papaipema duplicata(part)
17. A borer forming a swelling/gall in giant ragweed; also found in corn and many other plants in the eastern United States and in Canada Papaipema nebris
— A borer in the crown of burdock, thistle, and teasel in the northcentral and northeastern United States and in Canada Papaipema arctivorens
— A borer in owl’s-claw and sometimes in larkspur, at higher elevations in the southwestern United States Papaipema depictata
— A borer in dogbane and wild indigo in the eastern United States and in Canada Papaipema baptisiae
— A borer in dogbane in the Great Basin Papaipema limata
— A borer in mayapple in the eastern United States and in extreme southern Ontario; an austral species Papaipema rutila
— A borer in sunflowers in the eastern United States and in southern Canada; a prairie species Papaipema necopina
— A borer in prairie docks and relatives in the prairies of the midwestern United States Papaipema silphii
— A borer in loosestrife; northern and eastern species Papaipema lysimachiae
— A borer in columbine; northern and eastern species Papaipema leucostigma
18. A species of the central and eastern United States and Canada, including the Rocky Mountains region 19
— A species of the western United States and Canada from the White Mountains of Arizona and the Great Basin west 22
19. A species of the southern Great Lakes region and northwestern New Jersey 20
— A species of other central and eastern regions 21
20. A borer in purplestem angelica; occasionally in cowparsnip Papaipema rubiginosa
— A borer in water hemlock, purplestem angelica, and cowparsnip Papaipema birdi(part)
21. A borer in water hemlock, purplestem angelica, and cowparsnip, and also in some non-apiaceous plants; a species found throughout the Midwest, northeastern United States, Appalachian Mountains region, New England, and southeastern Canada Papaipema birdi(part)
— A borer in cowparsnip and related plants, also wild parsnip; a species of the Rocky Mountains, Canadian prairie provinces, northern Great Lakes region, New England, and Canadian Maritimes Papaipema harrisii
22. A borer with a preference for cowparsnip, water hemlock, angelica, and licorice-root, but also found in non-apiaceous plants such as burdock, ragwort, goldenrod, lupine, and others; a species of the Great Basin and the Pacific Northwest Papaipema insulidens
— A species found along the Pacific coast or in Arizona 23
23. A borer in Western water hemlock and less often in other plants such as larkspur; a species of the White Mountains region of Arizona Papaipema albimontana
— A species of the Pacific coast 24
24. A borer in Henderson’s angelica and also commonly in thistles; found less commonly in figworts as well as various Liliaceae, Malvaceae, and Ranunculaceae; a species of coastal areas from southern California north to northern Washington Papaipema sauzalitae
— A borer in leather-root; a species of coastal regions of southern California Papaipema angelica
25. Larva with very large D1 pinacula on A1–A3; those on A1 touching the pale, whitish, longitudinal mid-dorsal stripe 26
— Larva with smaller pinacula, not touching the mid-dorsal stripe 27
26. A borer in many different plants, favoring wingstem in the southern United States; a species of eastern regions Papaipema cataphracta
— A borer in swamp loosestrife; area of Cape Cod Massachusetts and offshore islands Papaipema sulphurata
27. A borer in various Apiaceae and other plants in the Pacific Northwest 22
— A species of eastern and northern regions 28
28. A borer in bogbean; a species of open, rich fens in the Great Lakes region and also in boreal areas Papaipema aweme
— A borer in seaside goldenrod; a species of the immediate Atlantic and Gulf coasts Papaipema duovata
— A borer in switch cane; a species of the southeastern United States Papaipema sullivani
— A borer in giant cane; a species of the lower Mississippi Valley and southern Appalachian regions Papaipema matheri
— A borer as an early instar in various grasses, including wildrye spp., switching to other plants including Liliaceae, Podophyllum peltatum, and Silene stellata in later instars Papaipema cerina
29. Larva with black lateral margins on the prothoracic shield 30
— Larva lacking black lateral margins on the prothoracic shield; a subterranean borer in wild yam Papaipema marginidens
30. Black outer margins of the prothoracic shield extending onto head; a borer in ironweed in the Midwest and eastern United States Papaipema cerussata
— Larva lacking a black line on sides of the head 31
31. A borer in ferns 32
— A borer in other plants 33
32. A borer in sensitive fern Papaipema inquaesita
— A borer in Osmunda species Papaipema speciosissima
33. A borer in flypoison; a species of the Appalachian Mountains of Pennsylvania south to North Carolina Papaipema amianthii
— A borer in pitcherplants; a species found in the Great Lakes region and in eastern Canada, south along the Atlantic coast and also along the Gulf coast Papaipema appassionata
— A borer in twigs of ash in the eastern United States and southern Canada Papaipema furcata
— A borer in rattlesnake-master, a species of Midwestern tall-grass prairies and also formerly coastal North Carolina Papaipema eryngii
— A borer in various asters, commonly in purplestem aster and flat-topped aster Papaipema impecuniosa
— A borer in white turtlehead in the North and East; verbena in the Midwest Papaipema nepheleptena
— A borer in culver’s root in prairies of the midwestern United States; formerly also found in the mid-Atlantic states Papaipema sciata
— A borer in the lower stem and roots of thinleaf sunflower, paleleaf sunflower, smooth oxeye, and golden alexanders in eastern regions Papaipema rigida
— A borer in sunflowers in the prairies of the midwestern United States and central Canada; also found along the mid-Atlantic coast; forms an egg-shaped gall at the stem base within which it pupates. Papaipema maritima
— A borer in joe-pye-weed; a northern and eastern species Papaipema eupatorii
— A borer in cutleaf coneflower and burdock; found in the North and East as well as in parts of the southwestern United States Papaipema nelita
PAPAIPEMA SPECIES GROUPS
If the living descendants of the single species Homo sapiens Linnaeus, are able to know their personal genealogy via their parents, grandparents, and great-grandparents, and could combine these genealogies with those of each and every other living person, the terminal branches of an unfathomably huge “tree” might appear. Yet to connect these terminals to one another becomes an impossibility, as genealogical information disintegrates with each successive earlier generation. So, genetic clues are relied upon. Tracing the ancestry i.e., phylogeny, of the human species, where there may be three or four generations in a century, for tens or hundreds of thousands of generations – unlike insect species such as Papaipema with a million generations in a million years – to the nearest hominid common ancestor is obviously unachievable with any current technology. To contemplate achieving anything comparable with 53 Papaipema species is even more unimaginable. That Papaipema represents a monophyletic assemblage supported by multiple character systems is not in question. It should come as no surprise, then, that no attempt to reconstruct the phylogeny of this large genus has been made herein. Even a best guess is highly problematic.
Grouping the species of this genus based on morphological synapomorphies and other factors such as ecological and biogeographical considerations may approach the limit of what can be achieved at this time. Combining these types of data with genetic information derived from molecular analyses would seem a worthy goal, if one is attempting to approximate something resembling a phylogeny, but that is somewhat like adding apples and oranges. The monophyly of the P.speciosissima group, for example, seems both intuitively obvious and well supported by multiple character systems. Nevertheless, one barcode tree indicates a sister species relationship between P.speciosissima and P.inquaesita, with P.stenocelis widely separated in a different BIN. Another places all three in different BINs, as does a third. Intuitively, it is not at all apparent whether P.inquaesita or P.stenocelis is more closely allied to P.speciosissima; each shares different character states with it. It is tempting to regard a group defined by morphological synapomorphies which happens to coincide with the manner in which the same taxa cluster in a barcode tree as definitive evidence of relatedness. It is equally tempting to disregard barcode data when there is no such congruence. When barcode data suggest genetic divergence approximating two percent or more, it is normally appropriate to accept that as evidence of species-level distinction. But what to make of an indicated divergence of, say, one-half percent – especially if morphological, ecological, or other factors taken into consideration are clearly indicative of full species status? Such is the case with numerous species recognized as distinct in this fascicle. Barcode data for Papaipema species have proven of little or no utility in distinguishing some of the closely related species, and it must be borne in mind that it represents only a part of a single gene.
The phylogenetic relationships among the many species of this large monophyletic group are not presently resolvable. The presumably vast unknown and unknowable numbers of extinctions of individual species and entire lineages which must have occurred during the evolution of groups seen today effectively precludes reconstruction of any “true” phylogeny. The innate desire to create a hypothetical phylogeny is strong but may not be worth the effort if the result is simply one of a myriad of possibilities, even if deemed the most parsimonious by any methodological contrivance. It is hoped that ongoing work with nuclear genes may eventually clarify some of these issues. Studies utilizing mitochondrial DNA, specifically examining the CO1 locus of barcode fragments, have occasionally been helpful in identifying discrete species in instances where morphological differences are not clearly apparent. In a number of cases where biogeographical or ecological information, host plant utilization, etc. have indicated distinct species in the absence of strong morphological evidence, barcode data have too often been ambivalent. Perhaps the best that can be achieved at this point is to assign apparently related species to groups, some of which are better defined than others. These species groups can then be ranked, through a combination of morphological, ecological, and other evidence, from apparent more basal groups to those believed to represent younger, more derived lineages. Seventeen of these groups are informally designated herein using the name of the earliest described included taxon.
Papaipema unimoda GROUP
To call this a “group” is perhaps a misnomer, for it consists of a single basal species so unlike all other Papaipema species that it stands alone. Superficially, the adult embodies the normal habitus of other members of the genus. However, its genital morphology strongly distinguishes unimoda from all other Papaipema, although not as extreme as the situation with Parapamea, where separate generic rank is appropriate. Papaipema unimoda lacks many of the apomorphic features defining the genus. It is the only Papaipema lacking the exceptionally large, cordate, ostial plate in the female, instead bearing a simpler, reduced version approaching the much-reduced type seen in Gortyna and Luperina. The larva also lacks some of the features commonly seen in other members of the genus. In this respect, the larva resembles that of P.beeriana, another anomalous species not thought to be closely related to the others. Like the members of the cerina group, the unimoda “group” might warrant subgeneric status.
Papaipema unimoda (Smith) (Meadow Rue Borer)
Hydroecia unimoda Smith, 1892, Entomological News, 3(10): 251.
Type locality: Colorado [USA]. [AMNH]
Hydroecia unimoda Smith, 1894, Transactions of the American Entomological Society, 21: 73.
note — Described originally in 1892 from one male and redescribed by Smith in 1894 with the addition of a second male from the same locality. The holotype is the original specimen described in 1892, although it bears an unnecessary Todd lectotype label. This error was corrected by
Hydroecia frigida Smith, 1899, Transactions of the American Entomological Society, 26: 39.
Type locality: Winnipeg, Manitoba, [Canada]. [USNM]
note — Described from a single male.
Hydroecia cerrussata; Smith, 1899, in part, misspelling, not Grote, 1864. Gortyna thalictri Lyman, 1905, The Canadian Entomologist, 37: 307.
Type locality: [vicinity of] Montréal, [Québec, Canada]. [LEM]
note — Described from seven male and ten female syntypes.
† Gortyna thalictri var. perobsoleta Lyman, 1905, The Canadian Entomologist, 37: 309.
Type locality: [vicinity of] Montréal, [Québec, Canada]. [LEM]
note — an unavailable infrasubspecific name based on the form that lacks white stigmata. note — Described from one male and four female syntypes.
Papaipema imperspicua Bird, 1908, The Canadian Entomologist, 40: 39.
Type locality: Franconia, [Grafton Co.], N. H. [New Hampshire, USA]. [AMNH]
note — Described from a female which Bird simply referred to as “the type” in his description and labelled it as such. Compared to most unimoda specimens, it is a large individual of much lighter coloration than is typical of eastern unimoda populations, more closely resembling Smith’s type from Colorado. This likely accounts for Bird’s failure to recognize the specimen for what it was. He also had seen a male from Buffalo, New York which he considered to be the same, but did not include as a syntype. This specimen was not found in his collection, and its whereabouts remains unknown. This male was obviously dissected by Bird, as he compares the structures to those of nitela [= nebris] which “follow the common design.” Clearly this male was of some other species, or Bird would not have made that comparison. He gives a measurement of 37 mm expanse, but it is unclear to which specimen he is referring. Since this measurement coincides with the alar expanse of his female type, it is believed this reference is to the same specimen. In treating imperspicua,
Papaipema frigida; Hampson, 1910, in part, not Smith, 1899; Draudt, 1926, part, not Smith, 1899. † Papaipema frigida ab. terminalis, Strand, 1915 [1916], Archiv für Naturgeschichte, 81A (11): 163.
Type locality: Rye, [Westchester Co.], N.Y., [New York, USA]. [NHML]
note — Unavailable infrasubspecific name. The male holotype was described from an illustration in
This common, widely distributed species ranges northeast to Newfoundland, beyond the ranges and farther north than almost any other Papaipema species. Both phenotypes of P.unimoda are commonly confused with the maculate and unmarked dark phenotypes of P.nelita. The postmedial line of unimoda is sinuous and excurved around the reniform; in nelita, this line is almost straight until just before it meets the costa. The maculate phenotype of nelita lacks the yellowish cream basal spots on the forewing, which are always present in maculate morphs of unimoda. In especially the northwestern portions of the range of unimoda, there is a tendency for individuals to occur with much lighter wood-brown coloration than their darker eastern counterparts. Such individuals (
The genital morphology of both sexes seemingly reflects a plesiomorphic state. It is the only Papaipema species lacking the exceptionally large, cordate, ostial plate in the female, which in part defines the genus, instead bearing the simpler, much reduced type seen in Gortyna and Luperina. In the male, the distinctive narrowing of the valve proximal to the cucullus found in other members of the genus is lacking. These combined attributes suggest that P.unimoda may represent a sole surviving member of one of the ancestral lineages linking New World Papaipema with Old World Gortyna.
DNA barcodes for 15 specimens of P.unimoda sampled from New Brunswick to Minnesota form a discrete group and unique BIN at a minimum nearest-neighbor difference of 3.8%.
The peculiar, distinctive larva of P.unimoda is unlike that of any other member of the genus. The larva more closely resembles that of a Hydraecia species, especially in the earlier instars before the transverse band of pink pigmentation on the dorsum of each abdominal segment has faded. Longitudinal lines are absent. The dark lateral margins of the prothoracic shield uniquely characteristic of Papaipema species tend to be faint or even absent in P.unimoda.
The larval hosts are species of meadow-rue (Thalictrum). Presence of borers in the meadow-rue is readily detected, first by wilting tips of the main stalks accompanied by the blackened, girdled interior of the hollow stems, as the young larvae proceed to the root where most of their development occurs. Pupation occurs in soil near the base of the plant.
Specimen capture records range from 22 August to 2 November, with the peak activity being from early September to mid-October. At any single locality, the flight period lasts about three weeks in any given year. This species occurs mainly in boreal regions west as far as Alberta and Montana but is decidedly less common south of transition zone environments. It can be found in mesic habitats with large stands of its host plant. Papaipema unimoda ranges from Newfoundland west to central Alberta and south to New Mexico and the southern Appalachian Mountains. The type locality in Colorado was situated so far west that familiar, better-known eastern populations were subsequently redescribed under other names.
Papaipema appassionata GROUP
Two closely related species comprise part of this group, P.appassionata and P.amianthii. Like the species of the P.cerina group, both retain certain primitive characters and lack some of the derived character states. Also placed here is P.furcata since it too appears to be another surviving offshoot of an ancient lineage lacking some of the apomorphic features characteristic of the majority of Papaipema species.
Papaipema appassionata (Harvey) (Pitcherplant Borer)
Gortyna appassionata Harvey, 1876, The Canadian Entomologist, 8: 155.
Type locality: “London (E. B. Reed)” [Ontario, Canada]. [NHML]
note — Described from a single female, alar expanse 35 mm. The abdomen is now missing.
Hydroecia appasionata; Smith, 1899, and authors, misspelling. Papaipema horni Strand, 1914 [1915], Archiv für Naturgeschichte, 80A (11): 160.
Type locality: “N. Carolina” [USA]. [MNHU]
note—Described from a single male, expanse 32 mm, found in the botanical garden at Marburg, Germany, which had emerged in an exhibit of insectivorous plants imported from North Carolina.
Despite a close relationship to P.amianthii, there is no species with which P.appassionata could be confused. This colorful species is among the most beautiful of the genus. The bright lemon-yellow of the medial area of the forewings in combination with the deep red of the terminal area produces an effective crypsis when the moth is at rest on its host plants, especially those species prevalent in the southern portions of its range such as yellow pitcherplant (Sarracenia flava) and yellow trumpets (S. alata). Unlike the male genitalia of most species of Papaipema, which possess a narrowing of the valve just before the cucullus, P.appassionata has a cucullus that is broadly-necked. Five barcode samples from across the geographic range of P.appassionata (Québec, New Brunswick, Louisiana) represent an isolated lineage that differs by a minimum of 2.6% from all other species.
Larvae of the genus Exyra Grote share the specialized adaptation to feeding on pitcherplant species, and the adults exhibit similar bright coloration. The irony of course is that both these insects have specifically evolved to eat plants which have themselves evolved to “eat” insects. Larvae of P.appassionata have been documented feeding upon several species of pitcherplant. In the northern portions of its range, P.appassionata utilizes purple pitcherplant (Sarracenia purpurea), the only available option. This being a rather small pitcherplant species results in adult moths that average as much as 4 mm less in forewing length than their larger southern counterparts, which have forewings that commonly measure 17 mm. The larva is typical of the genus and belongs in the group lacking the longitudinal white lines crossing the dark-pigmented bands encircling segments A1–A4. Across its extensive distribution, larvae emerging from overwintered eggs deposited around the bases of the previous season’s dried pitchers apparently first enter the plant at the base of the newly-emergent central flowering stalk, rather than elsewhere among the surrounding cluster of old leaves. The young larvae quickly abandon the flowering stalk and commence mining the rhizome, which soon results in an attenuation of the flower stalk.
This species displays a classical post-Pleistocene distribution ranging across the Great Lakes Region and St. Lawrence River system, down the Atlantic Coastal Plain, and along the Gulf Coast. It is likely to be found anywhere suitably large populations of pitcherplants occur, although it is absent from most of the northern boreal region.
Papaipema amianthii Quinter, NEW SPECIES (Flypoison Borer)
Papaipema amianthii Quinter.
Type locality: Blue Mountain, 40.5494, -76.1459, 1,604´ [489 m] Kittatinny Ridge, Appalachian Mountains, 4.2 mi. SW Auburn, Schuylkill County, Pennsylvania, USA. [CNC]
note — Named for its specialized adaptation to its sole host plant.
Probably due to its restricted distribution, this colorful and conspicuous species was unknown prior to its discovery in 1971, only a few miles from ELQ’s former home in Pennsylvania. The discovery of this new entity was solely and serendipitously responsible for igniting his interest in Papaipema, the decades-long study of which led to many additional discoveries and, ultimately, this fascicle.
Papaipema amianthii could be confused with P.marginidens, although the ranges of these two species barely overlap and the habitat requirements of the host plants are different. The rather uniform red of P.marginidens is similar to that of P.amianthii, although darker red in the latter, which is rarely as large. The reniform is much more laterally compressed in P.marginidens, relative to the fully expanded “figure 8” shape of the reniform of P.amianthii. Further, the central lunule within the reniform of P.marginidens is creamy white; that of P.amianthii is yellow and strongly contrasts with the white satellite spots. The male genitalia are totally dissimilar to those of P.marginidens which features an extremely long, sinuous digitus; that of P.amianthii is short and beak-like. The cucullus of P.marginidens exhibits a fishtail-like bifurcation; that of P.amianthii is not forked but instead bears a prominent triangular dorsal lobe. Papaipema amianthii is also sometimes confused with Papaipema birdi. In P.birdi, as in P.marginidens, the reniform is more laterally compressed than in P.amianthii. The forewings of P.birdi are less uniformly red and generally have more contrast in the ground color between the medial and postmedial areas of the wing, and there is a peppering of yellow scales in P.birdi in the area along the inner margin of the forewings that is absent in P.amianthii. The hindwings of P.birdi are gray-brown, unlike the reddish-tan hindwings of P.amianthii. The genitalia of P.birdi, with a sinuous digitus and strongly bifurcate cucullus, are also dissimilar to P.amianthii.
Head: antennae of both sexes ciliate, laminate, less deeply in female; dorsum scaled; ventral lamination obscured by densely pubescent sensilla; eye smooth, rounded; haustellum unreduced; labial palpus laterally flattened, upcurved; first segment swollen basally, arching slightly upward, slightly more than half as long as second segment; second segment straight, about as long as eye width, scales broad; third segment ⅓ × length of second segment, and projecting slightly anteriorly, scales narrow; frons unmodified, smooth, convex, with unspecialized conical vestiture consisting of spatulate scales mixed with fine, hairlike scales. Thorax: with exception of row of spatulate scales with pale yellow tips outlining dorsal edge of prothoracic collar, vestiture of prothoracic collar, patagia, mesothorax, and metathorax concolorous, a mixture of coarse spatulate, thin apically spatulate, and hairlike scales; transverse, wedge-shaped, prothoracic crest prominent, mesoscutellar tuft moderate, metascutellar tuft subequal. Forewing: length similar in both sexes = 13.5–20.0 mm, with the majority being near the maximum of this range; dorsal surface ground in both sexes uniform brick red; basal spots large, boldly conspicuous, yellow; antemedial and postmedial lines concolorous with ground, their inceptions at costa defined with white or yellowish marks; antemedial line double, nearly straight until strongly bowed outward just before meeting inner margin; postmedial line double, scalloped, excurved around reniform then nearly straight, meeting the inner margin perpendicularly; medial shade obscure in most, when faintly noticeable, touching lower loop of reniform, then angled below reniform and meeting inner margin nearly perpendicularly; subterminal line indicated by series of darker chevrons filling spaces between veins, marked near apex with two small patches of contrasting yellow scales; subterminal space filled with purplish scales; terminal space concolorous with medial area; terminal line obscure; fringe concolorous. Reniform a very large, broadly rounded, figure 8 shape, with large, bright white satellite spots surrounding narrow, cream-yellow, central lunule; orbicular and suborbicular stigmata very large, bright white, orbicular with central dark spot, upper suborbicular spot incorporating yellowish fragment of antemedial line as prevalent and more prominent in Gortyna species. Hindwing: dorsal surface uniform light fuscous, sometimes with pinkish tinge; discal spot and medial shade obscure in most; subterminal area, terminal line, and fringe undistinguished. Abdomen: basal coremata and pockets absent in male; dorsal tufts on A1–A3 variably prominent, that on A2 the largest. Male genitalia: uncus narrowly spatulate, apically acuminate with sparse, dorsal, cristate patch of recumbent setae; tegumen rounded at base of uncus, flaring laterally and forming rounded peniculi, largely devoid of hairs, before narrowing at s-shaped pleural sclerite and articulating with v-shaped vinculum; juxta + anellus elongated, height 3 × width, with divergent, paired, dorsal projections; ventral margin of valve evenly convex at base, then nearly straight until curved dorsally into notch at anteroventral edge of cucullus; base of sacculus rounded, swollen, with patch of sensory hairs extending along ridge of acutely pointed process on costal lobe; dorsal margin of valve concave basally, then slightly sinuous to broad neck of cucullus; cucullus triangular with acute apical angle, its ventral angle produced into a slight tooth, inner face of cucullus covered with patch of proximally-oriented spines; corona absent; clasper an appressed, sclerotized, crescent-shaped sclerite; ampulla absent; digitus heavily sclerotized, distoventrally curved, lacking serration on dorsal edge, acuminate, short, extending halfway to ventral margin of valve; phallus slightly curved, with terminal carina consisting of two opposing, subequal, sclerotized, ribbon-like, spined bars extending onto subbasal part of vesica; vesica with single, prominent, blunt, rose-thorn cornutus on dorso-mesial diverticulum. Female genitalia: papilla analis dorsoventrally flattened, lyriform, lateral margin prominently constricted toward anterior end, narrowly convex toward posterior end, acute apices divergent, with long setae set in tiny membranous circles on both dorsal and ventral surfaces; segment A8 2.8 × as wide as long, anterior apophysis 1.8 × length of segment A8 and posterior apophysis 2.8 × length of segment A8; lamella antevaginalis heavily sclerotized, laterally flared, cordate, with cleft on posterior rim, anterior portion of plate v-shaped; ostium bursae moderately sclerotized; ductus bursae narrow, about as long as appendix bursae, heavily sclerotized; appendix bursae membranous with light rugosity, about as long as and half the width of corpus bursae, viewed ventrally, oriented left, ductus seminalis at posterior apex; corpus bursae membranous, round, with two signa.
The unique male genitalic configuration shared by P.amianthii and P.appassionata, together with strong similarities in forewing maculation and coloration, suggests both are direct descendants of a common ancestor. If the red medial coloration of the forewing of P.amianthii were substituted with yellow, the two species would be indistinguishable superficially, other than for differences in size. Molecular data do not support this obvious close relationship however, which is inexplicable. To date only one sample of P.amianthii has been barcoded, differing by a minimum of 2.6% from all other Papaipema.
The larva is one of a group lacking the longitudinal white lines crossing the dark-pigmented bands encircling segments A1–A4. The lateral margins of the honey-colored prothoracic shield bear the usual dark, nearly black pigmentation stripes which do not extend onto the head capsule, also honey-colored in this species, as is often the case in others. The dorsal pinacula on A1–A7 are about average in size for a Papaipema larva; those on A8 are large, heavily sclerotized, with pigmentation ranging from honey to almost black. The D1 and D2 pinacula on A8 are ovoid and well separated. Those on A9 are fused into a single, sclerotized bar, honey-colored to almost black, which covers most of the dorsal surface of the segment. Both the dorsal bar on A9 and the anal shield are variably colored light to dark brown, rugose, and heavily sclerotized; the anal shield often has darker lateral margins.
TYPES. Holotype: ♂. Forewing length 20 mm [alar expanse 42 mm]. Kittatinny Ridge, Blue Mountain, 40.549394, -76.145932, 1,604´ acid bog 4.2 mi. SW Auburn, Appalachian Mts., Schuylkill Co., Pennsylvania; bred ex larva in bulb of Amianthium muscitoxicum; emerge: 7 September 1972; E. L. Quinter, collector. [CNC] Paratypes: 382 ♂, 280 ♀. North Carolina. Clay Co., Buck Creek Serpentine Barrens, 35.0838, -83.6154, 3,306´ [1,008 m]; 15 Sep 2023 15 wt. UV light. Genitalia slide No. 11,210; Sullivan, Petranka, & Elkin (1 ♂). Pennsylvania. Carbon Co., Kidder Twp., Hickory Run State Park, 41.0326, -75.6863, 1,540´ [469 m]; bred ex larva coll. 3 Aug 2013 in Amianthium muscitoxicum; emerge: 27 Aug–2 Sep 2013; Anthony E. McBride (4 ♀). Same locality and collector; bred ex pupa coll. 3 Aug 2013 in Amianthium muscitoxicum; emerge: 23–25 Aug 2013 (1 ♂, 1 ♀). Same locality and collector; bred ex larva coll. 30 Jul 2020 in Amianthium muscitoxicum; emerge: 23 Aug–12 Sep 2020 (5 ♂, 7 ♀). Same locality and collector; bred ex pupa coll. 30 Jul 2020 in Amianthium muscitoxicum; emerge: 19–20 Aug 2020 (2 ♂). Carbon Co., Hickory Run; 16 Sep 2006, J. Vargo (1 ♂). Carbon Co., Hickory Run State Park campground area, 10 Sep 2013, Steve Johnson (6 ♀). Carbon Co., Hickory Run State Park, 7.4 km SE East Side; 470 m., 75°41’06”W, 5 Oct 1994, C.W. Bier, B.J. Barton (3 ♂). Carbon Co., 1.5 mi. NE Albrightsville; 17 Sep 2005, Steven Johnson (1 ♀). Same locality and collector; 16 Sep 2006 (4 ♂, 8 ♀). Same locality and collector; 10 Sep 2007 (2 ♀). Carbon Co., 5.4 mi. W Blakeslee, acid bog 1,600´ [488 m], coll. in UV light trap, 10 Sep 1977, E. L. Quinter, collector (7 ♂). Same locality and collector; 24 Sep–4 Oct 1978 (19 ♂, 1 ♀). Same locality and collector; 31 Aug–2 Sep 1979 (1 ♂). Same locality and collector; 3–6 Sep 1979 (5 ♂). Same locality and collector; 9–12 Sep 1979 (3 ♂, 1 ♀). Same locality and collector; 16–20 Sep 1979 (8 ♂, 2 ♀). Same locality and collector; bred ex larva in bulb of Amianthium muscitoxicum; emerge: 29 Aug–1 Sep 1977 (3 ♂). Same data; emerge: 23 Aug–7 Sep 1978 (4 ♂, 7 ♀). Clinton Co., Keating Twp., 20.5 km NW Renovo, Montour Road along Cameron Co.; 21 Sep 1992, C.W. Bier, J. Wagner (11 ♂). Schuylkill Co., acid bog, Blue Mtn., 1,600´ [488 m], 4.2 mi. SW of Auburn, 14 Sep 1971; collected in UV light trap; E. L. Quinter (3 ♂). Same locality and collector; 19 Sep 1971 (3 ♂). Same locality and collector; 31 Aug–5 Oct 1972 (44 ♂, 1 ♀). Same locality and collector; bred ex larva in bulb of Amianthium muscitoxicum; emerge: 1–16 Sep 1972 (2 ♂, 7 ♀). Same locality; 40.5494, -76.1459, 1,604´ [489 m]; pupae coll. in bulb of Amianthium muscitoxicum, 15 Aug 2021, eclosed 20 Aug–7 Sep 2021; A. E. McBride (6 ♂, 11 ♀). Schuylkill Co., Summer Hill, 860´ [262 m], 1.9 mi. W of Auburn; collected in UV light trap; 14 Sep 1972, E. L. Quinter, collector (2 ♀). Carbon Co., Pocono Mts., Hickory Run State Park; 12 Sep 1971; collected at light R. E. Stanford (1 ♂, 1 ♀). Carbon Co., Hickory Run State Park, 7 Sep 1973; collected at lights, E. L. Quinter (33 ♂, 10 ♀); Same locality and collector; 9 Sep 1977 (69 ♂, 67 ♀). Same locality and collector; 9 Sep 1978 (7 ♂, 7 ♀). Same locality and collector; 16 Sep 1978 (10 ♂, 21 ♀). Same locality and collector; 23 Sep 1978 (27 ♂, 22 ♀). Same locality and collector; 22 Oct 1978 (1 ♂). Same locality and collector; 7 Sep 1979 (2 ♂, 3 ♀). Same locality and collector; 15 Sep 1979 (18 ♂, 13 ♀). Same locality and collector; 11 Sep 1982 (6 ♂). Same locality and collector; 18 Sep 1982 (7 ♂, 5 ♀). Same locality and collector; 3 Oct 1982 (6 ♂). Same locality and collector; 24 Sep 1983 (2 ♂). Same locality and collector; bred ex larva in bulb of Amianthium muscitoxicum; emerge: 25 Aug–16 Sep 1972 (14 ♂, 13 ♀). Same data; emerge: 29 Aug–17 Sep 1974 (27 ♂, 40 ♀). Same data; emerge: 2–7 Sep 1976 (1 ♂, 1 ♀). Same data; emerge: 29 Aug–6 Sep 1977 (4 ♂, 4 ♀). Same data; emerge: 30 Aug–4 Sep 1978 (1 ♂, 3 ♀). Same locality and collector; bred ex ovo on artificial diet, parent ♀ collected Sept. 1982; emerge: 3–11 Aug 1983 (5 ♂, 9 ♀). Virginia. Bath Co., Valley Springs, Warm Springs Mt. n. Ingalls Field, 37.9709, -79.8259 3,681´ [1,122 m]; 10 Sep 2023 at 175w MVS, A. E. McBride w/ Steve Roble (4 ♂). Montgomery Co., Slaughterpen Hollow off of Den Creek; hardwood forest, 31 Aug 2000 UV, J.C. Ludwig (1 ♂). AMNH, CMNH, CNC, CUIC, INHS, LACM, MCLB, NHML, NYSM, UCD, UCMS, ULK, USNM, VADNH, WVU, YPM, AEM, ELQ, JBS, JKA, JRW, JRS, JVC, SJC.
The larva of P.amianthii is restricted to a single monocotyledonous host plant species, flypoison (Amianthium muscitoxicum). Eggs hatch in early spring. The first instar bores into the base of the young plant’s emergent flower stem, quickly attenuating growth of any stalk, and proceeds to the subterranean bulb where it remains for the duration of its development. Pupation occurs within the hollowed bulb, the exposed top of which is by then capped with a distinctive, tarry, plant exudate combined with frass and silk which protects the pupa but also renders it easy to recognize infested plants, even though a surrounding rosette of green foliage may remain. When searching for the larvae or pupae, one must ignore all plants bearing flowering stalks, as these never mature in infested plants. Adults have been recorded flying from the last week in August to the first week of October, but the peak flight period occurs in mid-September.
Papaipema amianthii inhabits forested Appalachian ridge tops and plateaus, generally between 450 and 500 meters, mostly in eastern Pennsylvania west to the Allegheny Plateau wherever the host plant grows on poor, acidic, glaciated soils, often in great abundance. The moth occupies only a small portion of the range of its host plant, flypoison, which extends from eastern Pennsylvania south to the Florida panhandle and west along the Gulf Coast to Louisiana. A disjunct population of the plant occurs in the Ozark and Ouachita Mountains. The common name of this plant derives from the fact that it contains a highly toxic alkaloid concentrated primarily in the bulbs, which were blended with honey and used to kill house flies in colonial times. Papaipema amianthii is one of a number of Papaipema species which specialize upon highly toxic host plants, consuming the most toxic portions of their hosts during the larval stage. Whether or not chemical protection is conferred upon the adult moths remains unknown.
Until very recently this species had not been detected outside a very limited range, suggesting factors beyond availability of a food source were responsible for its peculiar distribution. Like other monophagous members of this genus, the distributions of their host plants often extend far beyond the ranges of the moths specializing upon them. Climate, elevation, and soil factors may well account for distribution of the host plants, but less likely for the insects utilizing them. Historical biogeographical influences such as glaciation probably played a major role in the present distribution of this moth. The moth seems to require the montane habitats where its host grows, even though flypoison appears to be as well adapted to dry sandy soils of savannas in the lower Atlantic Coastal Plain as it is to upland Appalachian habitats with poor, acidic soils.
If one considers the distribution of P.amianthii together with that of its apparent closest relative, an interesting pattern appears. The host plants of both species are broadly sympatric. The distributions of the two moths are contiguous, yet allopatric, in the small portion of range of P.appassionata where they nearly contact. Modern distributions very likely differ dramatically from those which would have prevailed during periods of maximum glaciation (
For over fifty years, this species was known only from a very limited distribution comprising the Pocono Plateau in eastern Pennsylvania and a few counties to the south and west. Recently, its occurrence in both Montgomery and Bath Counties in Virginia between 550 and 1,100 meters and in Clay County, North Carolina at 1,000 meters was documented. Although seemingly rather restricted in its occurrence, the species can be locally abundant. It might be expected in other states throughout the Appalachian Mountains with suitable habitats and poor soils at higher elevations, and there remains the potential for a disjunct occurrence in the Ozarks, yet efforts to locate this species outside Pennsylvania have only recently met with success.
Papaipema furcata (Smith) (Ash Shoot Borer)
Hydroecia furcata Smith, 1899, Transactions of the American Entomological Society, 26: 46.
Type locality [lectotype]: Urbana, [Champaign Co.], Illinois, [USA]. [USNM]
note — Described from five males from four states. Of three syntypes located by
The combined size, color, and maculation of this distinctive species could be confused with no other. The species is remarkably uniform in maculation across its broad range, though the deeper brownish-yellow of freshly emerged individuals quickly fades to a pale straw yellow as the brownish scales fall off through wear. The contrasting large, white stigmata on the yellow ground and the lack of white in the basal area save for a single spot in some specimens are diagnostic for this species. The name given by Smith undoubtedly is in reference to its most distinctive male genitalic feature, a prominent, forked digitus unique in the genus. DNA barcodes are available for nine specimens of P.furcata from across the species range, differing by at least 3.05% from any other species.
Like larvae of P.appassionata and P.amianthii, P.furcata larvae also lack longitudinal white lines crossing the dark-pigmented bands encircling segments A1–A4, and the lateral margins of the honey-colored prothoracic shield bearing the usual dark, nearly black pigmentation stripes do not extend onto the head capsule. Larvae of P.furcata have prominent black pinacula and a large, black, heavily armored anal plate, similar to some other twig and stalk boring species. For a more detailed account of the larval stages, see Bird (1915c).
This species is further unusual in its host plant choice, a tree. The larvae bore in the terminal shoots of ash (Fraxinus), both in young saplings and mature trees, and have been reported in the economic literature causing damage to ash trees in nurseries (
This species ranges from the Canadian border nearly to the Gulf of Mexico, being broadly distributed throughout eastern North America wherever ash trees are abundant, which may account for its apparent near absence from the southern Atlantic Coastal Plain. It varies from locally abundant to decidedly uncommon.
It is unknown to what extent the death of ash trees in Canada and the United States caused by the emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera), will have on populations of Papaipema furcata. On 4 July 2014, AEM found several larvae of P.furcata boring in the terminal twigs of young, four-meter-tall green ash trees growing in a wet roadside ditch near Petersburg, Michigan. All mature ash trees at this location had been previously killed by emerald ash borer damage, but P.furcata persisted here in the shoots of younger trees. Petersburg is approximately 30 miles southwest of Canton, Michigan, just west of Detroit, where the emerald ash borer was first detected in North America in 2002 (
Papaipema cerina GROUP
This group comprises three grass-feeding specialists retaining certain primitive character states and lacking some of the apomorphic features characteristic of most Papaipema. The included species are Papaipema cerina, P.sullivani, and P.matheri. As with Amphipoea and Hydraecia, each of which could be split into two subgenera, the Papaipema cerina group would be a strong candidate within Papaipema for such recognition. By some criteria, even full generic rank might be entertained. There is little value however in taking any such formal action as it would be much less apparent how to parse the large number of remaining Papaipema species into comparable ranks. Retaining the P.cerina group within Papaipema is necessitated by the fact that P.cerina is the type species. To assign this group generic rank would require the drastic measure of erecting a new genus for the balance of the species currently residing there, which is untenable.
This group is characterized by a much-reduced, non-furcate cucullus in the male genitalia, similar to that of Hydraecia and Gortyna. The digiti of the two cane-feeding members, completely lacking the characteristic serration in one species, parallel the costa for most of their length, unlike other Papaipema species. The larvae are long, cylindrical, and boldly striped with white longitudinal bands reminiscent of those of other noctuine genera which feed and rest externally on the stems of their host plants, a widespread condition among numerous noctuid lineages. Perhaps this affords some temporary cryptic protection for these borers when immatures may be forced to abandon their galleries and externally seek new culms in which to bore, as shown on
Papaipema cerina (Grote) (Wild Rye Borer)
Gortyna cerina Grote, 1874, Proceedings of the Academy of Natural Sciences of Philadelphia, 26: 200.
Type locality: [Douglas Co.], Kansas, [USA]. [NHML]
note — Described from a single female, the locality given as just “Kansas.” The source of that specimen was Professor Snow at the University of Kansas, situated in Douglas County, who also collected Grote’s type specimen of Gortyna erepta which came from Douglas County, Kansas. It is presumed that this is the type locality for both species. Grote’s name is derived from the Latin cerinus, apparently in allusion to its yellow coloration. In appearance and genital morphology, P.cerina is atypical for the genus, but is the type species nonetheless.
Only the yellow morph of P.sullivani with dark stigmata could be mistaken for this species, but these two are unlikely to be found together. The ground color of these yellow morphs of P.sullivani is typically more orange than the lemon yellow of P.cerina, and the stigmata are fuller and darker than in P.cerina. The male valve of P.cerina has a digitus that is downcurved and serrated unlike that of P.sullivani, which is smooth, straighter, and extends posteriorly. The female possesses a very large, quadrate ostial plate which differs from that of P.sullivani. Papaipema cerina is consistent in maculation throughout its range.
This species is clearly related to the only other grass-boring species of the genus, P.sullivani and P.matheri, suggesting they are all descendants of an Asian common ancestor which likely arrived in North America during the Miocene (see discussion in the Arundinaria-feeding genera). That P.cerina apparently adapted to a more boreal environment with an allopatric distribution beyond the range of the giant grasses of the genus Arundinaria might account for its peculiar larval strategy of switching hosts. Based on DNA barcodes of six specimens from Illinois, New Hampshire, North Carolina, Ontario, and Wisconsin, P.cerina has a distinct barcode BIN that differs by at least 1.9% from all other Papaipema species, despite its close morphological similarities to P.sullivani and P.matheri.
The larva of P.cerina belongs to a larger group of species in which larvae have both dorsal and subdorsal longitudinal white lines continuous across all abdominal segments, thus encompassing more than the three members of its own species group. In this respect, the larva resembles the much more commonly encountered larva of P.cataphracta which is less slender and has larger pinacula in comparable instars. Papaipema cataphracta commonly can be found using the alternate host plants of P.cerina, including lily (Lilium spp.) and mayapple (Podophyllum peltatum). In the mature larva of P.cataphracta, the D1 pinacula on segments A1–A4 are large, approximately four times larger than the D2 pinacula on these same segments, and touch or very closely approach the pale dorsal stripe. In P.cerina, the D1 pinacula on segments A1–A4 are small, scarcely larger than the D2 pinacula on these same segments, and are well separated from the pale dorsal stripe by approximately the width of the pinacula or greater. The later average maturation date of P.cataphracta larvae may offer a clue as to identity as well.
Known primary host plants of P.cerina include eastern bottlebrush (Elymus hystrix), riverbank wildrye (Elymus riparius), and probably other Elymus species (wildrye). AEM also found early instars in other grasses such as reed canarygrass (Phalaris arundinacea). Development is invariably completed in other larger monocots, e.g., lily, iris, and wild onion (Allium spp.), or occasionally in dicots such as mayapple and campion (Silene spp.). The curious apparent dependency of P.cerina on woodland grasses such as wildrye may indicate these are obligatory hosts for oviposition and development of early instars, as the moth nearly always occurs in association with such grasses. This strange larval behavior would appear maladaptive, since later instars are obligated to seek alternate hosts (typically monocots) in which they can complete maturation. Completing larval development on wildrye or other slender grasses would be impossible without switching from boring internally to feeding externally on the grass stems, and there is no evidence this ever occurs. This species is essentially oligophagous, but with the potential to be found in later instars boring in a variety of unrelated plants co-occurring with wildrye.
Young larvae only bore in the tender terminal two or three internodes of a single culm of the host grass before exiting to seek alternate hosts; they do not enter the grass bases or roots. The bored culm top becomes pale tan in color, contrasting strongly with adjacent, green culms (
The paucity of modern records of the species from the northeastern portions of its seemingly disjunct distribution suggests it has become quite rare in these areas. Recent captures in the mountains of western North Carolina are both surprising and encouraging. Although never particularly common, P.cerina is seen regularly throughout the upper Midwest, along with a single record from southern Saskatchewan.
Papaipema sullivani Quinter, NEW SPECIES (Sullivan’s Switch Cane Borer)
Papaipema sullivani Quinter.
Type locality: Great Dismal Swamp National Wildlife Refuge, City of Suffolk, Virginia, USA. [CNC]
note — Named for Dr. J. Bolling Sullivan, III who first brought the existence of this species to my attention, in recognition of his many ongoing contributions to our knowledge of the lepidopterous fauna of North Carolina.
Papaipema sullivani is a large species, but varies considerably in size, with a forewing length of 14.5–22.0 mm. In all known populations, this species exhibits the most extreme range of phenotypic variability of the genus. Without knowledge of its intermediate transitional morphs, one is initially incredulous that specimens such as those illustrated on
Head: antennae of both sexes ciliate, laminate, less deeply in female; dorsum scaled; ventral lamination obscured by densely pubescent sensilla; eye smooth, rounded; haustellum unreduced; labial palpus laterally flattened, upcurved; first segment swollen basally, arching slightly upward, slightly more than half as long as second segment; second segment straight, about as long as eye width, scales broad; third segment ⅓ × length of second segment, and projecting slightly anteriorly, scales narrow; frons unmodified, smooth, convex, with unspecialized conical vestiture consisting of spatulate scales mixed with fine, hairlike scales. Thorax: vestiture of prothoracic collar, patagia, mesothorax, and metathorax concolorous, a mixture of coarse spatulate, thin apically spatulate, and hairlike scales; transverse, wedge-shaped, prothoracic crest prominent, mesoscutellar tuft moderate, metascutellar tuft subequal. Forewing: length similar in both sexes = 14.5–22.0 mm; dorsal surface ground in both sexes highly variable, ranging from yellowish pumpkin to dark, rusty, reddish-orange in medial background; filling of antemedial and postmedial lines concolorous with ground, both lines ranging from obscure in dark individuals to well defined with rust-colored scales in yellower phenotypes; basal spots concolorous with medial area, space between basal line and antemedial line overlaid with darker, rust-colored scales; antemedial line double, evenly scalloped, straight until strongly bowed outward just before meeting inner margin; postmedial line double, scalloped, sinuous, excurved around reniform, then meeting inner margin perpendicularly; medial shade defined with darker rust-colored scales, touching lower loop of reniform, then sharply angled below reniform and meeting inner margin perpendicularly; subterminal line preceded by a series of darker charcoal chevrons filling spaces between veins, subterminal space filled with gray scales; terminal line obscure; fringe dark gray; reniform and orbicular complex stigmata strongly contrasting in both extreme phenotypes, less so in transitional morphs. In the phenotype with dark stigmata, the reniform, orbicular, and suborbicular spots are a dark, charcoal gray-brown, strongly contrasting with the pumpkin-colored medial area of the forewing. In somewhat worn specimens of this phenotype, scattered, rust colored deciduous scales overlaying the yellow ground of the medial area become lost, resulting in a much more yellow appearance than that of freshly emerged individuals. In the white-marked phenotype, the stigmata strongly contrast with a dark, rusty-orange medial area background, but in transitional morphs, the background coloration of the medial area exhibits a continuum between the two extremes. Hindwing: dorsal surface light beige basally, discal spot faint or obscure; a fuscous medial shade with lighter subterminal area; terminal line usually obscure with dull pinkish fuscous fringe. Abdomen: basal coremata and pockets absent in male; dorsal tufts on A1–A3 variably prominent. Male genitalia: uncus narrowly spatulate, apically acuminate with dorsal cristate patch of recumbent setae; tegumen rounded at base of uncus, flaring laterally and forming rounded peniculi, largely devoid of hairs, before narrowing at s-shaped pleural sclerite and articulating with narrowly v-shaped vinculum; juxta + anellus elongated, height 3 × width, with divergent, paired, dorsal projections; ventral margin of valve evenly convex at base, then nearly straight until sharply angled dorsally into notch at anteroventral edge of cucullus; base of sacculus rounded, swollen, with dense patch of sensory hairs on costal lobe; dorsal margin of valve concave basally, then nearly straight to narrow neck of cucullus; cucullus small, rounded, subtriangular, its inner face covered with a dense patch of proximally-oriented, spine-like setae; corona absent; clasper an appressed, sclerotized, crescent-shaped sclerite; ampulla absent; digitus heavily sclerotized, scabrous but not dentate, flattened and nearly straight, acuminate, subparallel with costal margin; phallus slender, slightly curved, with terminal carina consisting of two opposing, subequal, sclerotized, ribbon-like, spinulate bars extending onto subbasal part of vesica; vesica with a single, prominent, blunt, rose-thorn cornutus on a dorso-mesial diverticulum. Female genitalia: papilla analis dorsoventrally flattened, lyriform, with lateral margin prominently constricted toward anterior end, narrowly convex toward posterior end, acute apices divergent, with long setae set in tiny membranous circles on both dorsal and ventral surfaces; segment A8 2.7 × as wide as long, anterior apophysis 1.5 × length of segment A8 and posterior apophysis 2.7 × length of segment A8; lamella antevaginalis heavily sclerotized, laterally flared, cordate, with cleft on posterior rim; ostium moderately sclerotized; ductus bursae narrow, about as long as appendix bursae, heavily sclerotized; appendix bursae membranous with moderate rugosity, about as long as and half the width of corpus bursae, viewed ventrally, oriented left, ductus seminalis at posterior apex; corpus bursae membranous, round, with two signa.
Mature larva can exceed 4 cm; head capsule 3.0 mm wide; head warmer honey than paler, yellowish-brown prothoracic shield; prothoracic shield edged laterally with darker pigment, this dark line extending onto head capsule; somatic pigmentation liver-brown, darker on first four abdominal segments ; whitish dorsal and subdorsal stripes run length of body and interrupt darker somatic pigment on A1–A4; dark brown D1 pinacula on segment A1 are half size of those on A2 and well separated from whitish dorsal stripe; D1 and D2 pinacula are roughly equal in size on A8, and D1 pinacula may be partially to completely fused with D2 pinacula on A9; anal shield is pale brown and edged with darker pigment laterally; L1 located well behind and below the spiracle on A7.
TYPES. Holotype: ♀. Forewing length 21 mm [alar expanse 43.5 mm]. Great Dismal Swamp National Wildlife Refuge, Suffolk, Virginia; bred ex larva in Arundinaria tecta, emerge: 30 Sep 1999; E. L. Quinter. [CNC] Paratypes: 74 ♂, 57 ♀. Florida. Gadsden Co., Quincy; 25 Oct 1966; William B. Tappan (1 ♂). Liberty Co., Apalachicola N.F., Revell’s Landing 30.1283, -84.6729 14´ [4.3 m]; ex larva coll. 3 Jul 2021 in Arundinaria tecta; reared in potato, eclosed 3–21 Nov 2021; A. E. McBride w/ J. Slotten (3 ♂, 1 ♀). Georgia. Liberty Co., St. Catherines Island; ex larva in Arundinaria tecta, emerged 28 Oct 1991; E. Quinter and A. Sharkov (1 ♂). Same locality; ex larva in Arundinaria tecta, emerged 12–26 Oct 1992; E. Quinter and A. Sharkov (6 ♂, 3 ♀). Same locality; ex pupa collected in Arundinaria tecta, emerged 22–26 Oct 1992; E. Quinter (2 ♂). Monroe Co., Forsyth; 9 Oct 1955; F. T. Naumann (1 ♀). North Carolina. Carteret Co., Jct. St. Hwy. 101 & 181; 15 Oct 1971; J. B. Sullivan (1 ♀). Carteret Co., 10–22 Oct 1972; J. B. Sullivan (2 ♂, 1 ♀). Carteret Co., International Paper Co. office; 14 Oct–1 Nov 1972; J. B. Sullivan (6 ♂, 4 ♀). Carteret Co., Jct. St. Hwy. 101 & 181; 7 Oct–11 Nov 1972; J. B. Sullivan (4 ♂, 4 ♀). Same locality; 14–29 Oct 1973; J. B. Sullivan (8 ♂, 3 ♀). Carteret Co., Jct. St. Hwy. 101 & 181; 11 Oct–4 Nov 1978; J. B. Sullivan (6 ♂, 1 ♀). Carteret Co., 0.2 mi. S Jct. St. Hwy. 101 & 1154; bred ex larva in Arundinaria tecta, emerge: 27 Oct–6 Nov 1978; E. L. Quinter (4 ♂). Same locality; bred ex larva in Arundinaria tecta, emerge: 13–21 Oct 1980; E. L. Quinter (1 ♂, 4 ♀). South Carolina. Charleston Co., McClellanville; 12 Nov 1967; R. B. Dominick (1 ♀). Same locality; 8 Nov 1968; at light, R. B. Dominick (1 ♂). Same locality; 18 Oct 1975; R. B. Dominick (1 ♂). Virginia. City of Virginia Beach, NAS Oceana, ca. 1.5 km SSE jct. London Bridge Rd. x Potters Rd. cane woods; 19 Oct 2001, UV; K. L. Derge, A. C. Chazal (1 ♂, 1 ♀). City of Virginia Beach, North Landing River, Vaughn Tract, cane-mixed herb and forested uplands; 12 Oct 2001, UV; C. S. Hobson (1 ♀). Suffolk, Great Dismal Swamp National Wildlife Refuge; bred ex larva in Arundinaria tecta, emerge: 6 Sep–1 Oct 1997; E. L. Quinter (2 ♂, 1 ♀). Same locality; bred ex larva in Arundinaria tecta, emerge: 17–21 Sep 1999; E. L. Quinter (2 ♂). Same locality; bred ex larva in Arundinaria tecta on artificial diet, emerge: 24 Aug–10 Oct 2000; Quinter, Goldstein, and Roble (15 ♂, 16 ♀). Same locality; pupa collected 18 Sep 2000 in Arundinaria tecta, emerge: 6 Oct 2000; E. Quinter and J. O’Donnell (1 ♀). Suffolk, Great Dismal Swamp NWR, Railroad Ditch 36.6184, -76.5584 32´ [9.8 m]; ex larva coll. 8 Jun 2015 in Arundinaria tecta; reared in potato, eclosed 14 Sep–24 Oct 2015; A. E. McBride, collector (8 ♂, 13 ♀). AMNH, CMNH, CNC, INHS, LACM, MCLB, NHML, NYSM, UCD, USNM, VADNH, WPC, YPM, AEM, ELQ, JKA, JRW, VAB.
The larva bores in only the new, tender, first-year culms of switch cane (Arundinaria tecta). Older growth is never utilized. Larvae have been found as early and middle instars in late June, so eggs probably hatch in late May. Few food resources are available until new culms develop from the rhizome and emerge from the soil. The bored culms typically attain heights of 0.5-1.5 m before growth is halted due to internal damage, accompanied by a change in coloration from green to tan. In several instances, very young emerging culms only 15-25 cm tall were found recently bored, as indicated by prominent mounds of fresh white frass on the ground beneath them. These culms had single holes from which the frass had been ejected by nearly mature larvae. Such evidence suggests that at least occasionally larvae may abandon culms initially utilized, having exhausted the available food resources, and relocate to additional young culms before reaching maturity. Larval feeding can extend between several internodes, sometimes well into the rhizome, depending on the vigor and size attained by the culm prior to attenuation of its growth. The mature larva prepares an exit hole for adult emergence near the base of the dried culm, leaving a thin, parchment-like membrane covering the exit. It then constructs a one-centimeter-deep platform of chewed particles in the base of that node, upon which the pupa is formed, just below the prepared exit. Pupation occurs within the dried culms around mid-September to early October. Adult eclosion occurs approximately a month later. Recorded flight dates range from 7 September to 12 November, with peak abundance midway between these extremes.
Papaipema sullivani has not been recorded from inland habitats where switch cane is replaced with giant cane. It is mainly confined to the southern Atlantic and Gulf Coastal Plains, ranging from southeastern Virginia to southern Mississippi, and is unlikely to be found throughout the more interior portions of the range of its host plant where it meets and sometimes hybridizes with giant cane.
Papaipema matheri Quinter, NEW SPECIES (Mather’s Giant Cane Borer)
Papaipema matheri Quinter.
Type locality: McCrite Farm, 37.2418, -89.3258, 387´ [118 m], Diswood, Alexander County, Illinois, USA. [CNC]
note — Named to honor the late Dr. Bryant Mather, who collected the first specimens of this species to come to my attention, in recognition of his many contributions to knowledge of the insects of Mississippi, and in appreciation for his generous support of my work over the years.
Based on abundant morphological and ecological considerations, P.matheri is the obvious immediate sister species of P.sullivani. Inexplicably, this is not supported by molecular data. DNA barcodes are available for four specimens of P.matheri from Kentucky and Georgia, which form a discrete group and unique BIN at a minimum nearest-neighbor difference of 3.3%. Papaipema matheri is another large species but with much less extreme individual variation than that of P.sullivani; its forewing length ranges from 17–26 mm, with most individuals being closer to the maximum. In sharp contrast to P.sullivani, P.matheri exhibits almost no phenotypic variability in either coloration or maculation. Except as discussed under P.sullivani, P.matheri is unlikely to be mistaken for any other Papaipema. The forewing of P.matheri is uniformly rich maroon, darker than the darkest rust-red P.sullivani specimens. The two species are readily distinguished by their genitalic features as well. A striking resemblance however to the white-marked phenotype of Parapamea buffaloensis might cause confusion, as Parapamea buffaloensis and Papaipema matheri can occur together. The ground color of the forewing of both species can be nearly identical. The suborbicular spots are unusually small in P.matheri, tending towards obsolescence, and may be completely absent in occasional individuals; in the maculate phenotype of Parapamea buffaloensis, the suborbicular spots are always absent. Otherwise, the postmedial line of P.buffaloensis is almost straight until just before meeting the costa; in Papaipema matheri, it is sinuous and excurved around the reniform.
Head: antennae of both sexes ciliate, laminate, less deeply in female; dorsum scaled; ventral lamination obscured by densely pubescent sensilla; eye smooth, rounded; haustellum unreduced; labial palpus laterally flattened, upcurved; first segment swollen basally, arching slightly upward, slightly more than half as long as second segment; second segment straight, about as long as eye width, scales broad; third segment ⅓ × length of second segment, and projecting slightly anteriorly, scales narrow; frons unmodified, smooth, convex, with unspecialized conical vestiture consisting of spatulate scales mixed with fine, hairlike scales. Thorax: vestiture of prothoracic collar, patagia, mesothorax, and metathorax concolorous, a mixture of coarse spatulate, thin apically spatulate, and hairlike scales; transverse, wedge-shaped, prothoracic crest prominent, mesoscutellar tuft moderate, metascutellar tuft subequal. Forewing: length similar in both sexes = 17–26 mm; dorsal surface ground in both sexes relatively uniform dark maroon, the medial area in some somewhat lighter than basal and subterminal areas which are plumbeous-violet; basal spots obscure; a tiny, contrasting, white subbasal spot visible at juncture of cubital and anal veins; antemedial and postmedial lines concolorous with ground, both lines being rather obscure; antemedial line double, evenly scalloped, straight until bowed outward just before meeting inner margin; postmedial line double, scalloped, sinuous, excurved around reniform, then meeting the inner margin diagonally; medial shade defined with slightly darker scales, touching lower loop of reniform, then angled below reniform and meeting inner margin diagonally; subterminal line preceded by a series of faintly-visible, darker, charcoal chevrons filling spaces between veins, subterminal space filled with gray-violet scales; terminal line obscure; fringe dark gray-violet; reniform and orbicular complex stigmata apparently always white and strongly contrasting. Hindwing: dorsal surface uniform dull fuscous, or slightly paler basally, sometimes with a faint rosy tinge; discal spot and medial shade obscure; subterminal area, terminal line, and fringe undistinguished. Abdomen: basal coremata and pockets absent in male; dorsal tufts on A1–A3 variably prominent. Male genitalia: uncus narrowly spatulate, apically acuminate with dorsal, cristate patch of recumbent setae; tegumen rounded at base of uncus, flaring laterally and forming rounded peniculi, largely devoid of hairs, before narrowing at s-shaped pleural sclerite and articulating with narrowly v-shaped vinculum; juxta + anellus elongated, height 3 × width, with divergent, paired, dorsal projections; ventral margin of valve evenly convex at base, then nearly straight until angled dorsally into notch at anteroventral edge of cucullus; base of sacculus rounded, swollen, with patch of sensory hairs on costal lobe sparse and greatly reduced in size and density; dorsal margin of valve concave basally, then nearly straight to narrow neck of cucullus; cucullus small, rounded triangular, its inner face covered with dense patch of proximally-oriented, spine-like setae; corona absent; clasper an appressed, sclerotized, crescent-shaped sclerite; ampulla absent; digitus heavily sclerotized, smooth, strongly arched, acuminate, overall orientation subparallel with costal margin; phallus slender, slightly curved, with terminal carina consisting of two opposing, subequal, sclerotized, ribbon-like, spinulate bars extending onto subbasal part of vesica; vesica with single, prominent, blunt, rose-thorn cornutus on dorso-mesial diverticulum. Female genitalia: papilla analis dorsoventrally flattened, lyriform, with lateral margin prominently constricted toward anterior end, narrowly convex toward posterior end, acute apices divergent, with long setae set in tiny membranous circles on both dorsal and ventral surfaces; segment A8 2.5 × as wide as long, anterior apophysis 1.8 × length of segment A8 and posterior apophysis 3 × length of segment A8; lamella antevaginalis heavily sclerotized, laterally flared, fan-shaped, lacking cleft on convex posterior rim; ostium moderately sclerotized; ductus bursae narrow, about as long as appendix bursae, heavily sclerotized; appendix bursae membranous with light rugosity, about as long as and half the width of corpus bursae, viewed ventrally, oriented left, ductus seminalis at posterior apex; corpus bursae membranous, round, with two signa.
Mature larva can exceed 4 cm, head capsule is 3.0 mm wide; head is warmer honey than paler, yellowish-brown prothoracic shield; prothoracic shield edged laterally with darker pigment, this dark line does not extend onto head capsule; somatic pigmentation pale brown on A1–A4, equally pale but warmer brown on remaining abdominal segments; whitish dorsal and subdorsal stripes run length of body and interrupt somatic pigment on A1–A4; dark brown D1 pinacula on segment A1 half the size of those on A2 and separated by at least width of pinacula from whitish dorsal stripe; D1 and D2 pinacula roughly equal in size on A8, and D1 pinacula may be partially to completely fused with D2 pinacula on A9; anal shield brown, edged with darker pigment laterally; L1 located well behind and below spiracle on A7. The larva of P.matheri at maturity attains sizable proportions but is noticeably shorter and thicker than that of mature P.sullivani, which is longer and more slender, likely due to the significant differences in average culm diameter of their hosts.
TYPES. Holotype: ♀. Forewing length 21 mm [alar expanse 44 mm]. McCrite Farm, no. of cemetery 37.241777, -89.325793 387 ft. Diswood, Alexander County, Illinois; June 30, 2014 ex larva, A. E. McBride, collector. Ex Arundinaria gigantea reared in potato/carrot, eclosed Oct. 2, 2014. [CNC] Paratypes: 260 ♂, 64 ♀. Arkansas. Hempstead Co., Hope; 25 Oct 1933; L. Knobel (1 ♂). Madison Co., Aurora, War Eagle Creek, 1,360´ [415 m] jct. Rts. 23 &127; ex larva coll. in Arundinaria gigantea 30 Jun 1993, pupate: 29 Aug, emerge: 25 Sep 1993; E. L. Quinter (1 ♂). Georgia. Bartow Co., 8 mi. SE Fairmount, Salacoa Rd. at Salacoa Creek (southeast pasture); light trap, 18 Oct 2000; James K. Adams (1 ♂). Gordon Co., Calhoun, northwest end of Tate Bend Rd., 34° 33’ 44” N, 84° 57’ 54” W, cane habitat along Oostanaula River, west of I-75; light trap, 5–11 Oct 2002; James K. Adams (1 ♂, 4 ♀). Same locality and collector; 4 Oct 2003 (1 ♂). Same locality and collector; 8 Oct 2005 (1 ♀). Same locality and collector; 12–13 Oct 2007 (1 ♂). Houston Co., Oaky Woods WMA, ca. 4 mi. ESE Kathleen, cane habitat near Big Grocery Creek; light trap, 6 Oct 2001; James K. Adams (1 ♂). Illinois. Alexander Co., McCrite Farm 3 mi. W Tamms, T15S-R2W, Sec. 3; coll. in canebrake at cemetery w/flashlight, 4 Oct 1997; J. R. Wiker (1 ♂). Same locality and collector; MH light, creek bottom NW of cemetery, 4 Oct 1997 (3 ♂, 1 ♀). Same locality and collector; UV light, 12 Oct 1999; (2 ♂). Same locality and collector; larvae in Arundinaria gigantea, pupated 29 Aug–12 Sep 1999, emerged 27 Sep–8 Oct 1999 (2 ♂, 1 ♀). Same locality; larva in Arundinaria gigantea, emerged 22 Sep 1999; E. L. Quinter (1 ♂). Alexander Co., Diswood, McCrite Farm, no. of cemetery 37.2418, -89.3258 387´ [118 m]; 1 Jul 2009; ex larva in Arundinaria gigantea reared in potato/carrot, eclosed 30 Sep 2009; A. E. McBride, collector. (1 ♀). Same locality and collector; 30 Jun 2014; ex larva in Arundinaria gigantea reared in potato/carrot, eclosed 28 Sep–10 Oct 2014 (4 ♀). Alexander Co., Rock Springs Rd.; UV light trap in cane, 14 Oct 2003; W. R. Black, Jr. (1 ♂). Alexander Co., Rock Springs Freewill Bap. Ch.; UV light trap in cane behind church, 20 Oct 2003; W. R. Black, Jr. (4 ♂). Jackson Co., Cedar Lake 2 mi. E of Pomona, T10S-R2W, Sec. 26; larvae coll. in Arundinaria gigantea 24 Aug 1997, pupated 31 Aug, emerged 25 Sep 1997; J. R. Wiker (1 ♀). Johnson Co., Proctors Hill, N edge Cache River floodplain, 4 mi. W Belknap; UV light trap in cane, 12 Oct 2000; W. R. Black, Jr. (1 ♂). Massac Co., 0.25 mi. W Kincaid Mounds; UV light trap at cane site #2 in wooded slough, 4 Oct 2000; W. R. Black, Jr. (5 ♂). Same locality and collector; 12 Oct 2000 (2 ♂). Same locality and collector; cane site #3, 18 Oct 2000 (4 ♂). Same locality and collector; cane site #3, 19 Oct 2000 (3 ♂). Massac Co., Seven Mile Creek; UV light trap at cane site #1, 21 Oct 2006; W. R. Black, Jr. (2 ♂). Massac Co., Big Bay; UV light trap at cane site #1, 21 Oct 2006; W. R. Black, Jr. (2 ♂). Pope Co., War Bluff Sanctuary, 5 mi. N Golconda, T12S-R6E, Sec. 30; larva coll. 29 Jun 1996 in Arundinaria gigantea, pupated early Sep, emerged 5 Oct 1996; J. R. Wiker (1 ♂). Pope Co., east end Avery Lake; UV trap in cane, 12 Oct 2000; M. C. Nielsen (1 ♂, 1 ♀). Same locality; UV trap at cane site #2, 1 Oct 2000; Wm. R. Black, Jr. (2 ♂). Same locality and collector; UV trap at cane site #4, 2 Oct 2000 (8 ♂). Same locality and collector; UV trap at cane site #5, 12 Oct 2000 (6 ♂). Same locality and collector; UV trap at cane site #2, 18 Oct 2000 (3 ♂, 1 ♀). Pope Co., Alcorn Creek; UV trap at cane site #1, 19 Oct 2000; Wm. R. Black, Jr. (1 ♀). Pulaski Co., Mud Line Rd., 0.15 mi. E Hawkins Rd.; UV trap at cane, 20 Oct 2003; W. R. Black, Jr. (4 ♂, 2 ♀). Union Co., 3 mi. NE Wolf Lake, County Rd. #52 along Hutchins Creek; larvae coll. 10 Aug 1996 in Arundinaria gigantea, pupated 22 Aug–11 Sep, emerged 23 Sep–12 Oct 1996; J. R. Wiker (1 ♂, 2 ♀). Union Co., Pine Hills Campground Area, T11S-R3W, Sec. 34; pupa coll. in cane stem 24 Sep 1996, emerged 27 Sep 1996; J. R. Wiker (1 ♀). Union Co., County Rd. #19 E Rt. 127, W Walnut Grove Church, T11S-R2W, Sec. 27; larva coll. 25 Aug 1997 in Arundinaria gigantea, pupated 31 Aug, emerged 1 Oct 1997; J. R. Wiker (1 ♂). Union Co., TTSF, Clear Creek at unnamed tributary creek; UV light trap at cane, 8–9 Oct 2002; Wm. R. Black, Jr. (7 ♂, 2 ♀). Union Co., Hutchinson Creek, UV trap at cane [52° F low], 8–9 Oct 2002; Wm. R. Black, Jr. (2 ♂). Kentucky. (Unless noted otherwise, all Kentucky records are of specimens collected by William R. Black, Jr. in UV light traps) Ballard Co., Rt. 121 SE Wickliffe, Stovall Creek; larva coll. in Arundinaria gigantea 15 Jun 1999; pupated 28 Aug 1999, emerged 23 Sep 1999; J. R. Wiker (1 ♂). Ballard Co., Sugar Creek below Lovelaceville; 19 Oct 2001 (1 ♂). Ballard Co., Shelton Creek at Buckheart Creek; 21 Oct 2001 (1 ♀). Ballard Co., Mayfield Creek, 0.6 mi. W U.S. 62 bridge; 11 Oct 2005 (1 ♂, 1 ♀). Ballard Co., Mayfield Creek, E of Billington Creek; cane site #2, 9 Oct 2006 (1 ♂). Calloway Co., East Fork Terrapin Creek; cane site #1, 15 Oct 2001 (1 ♂). Calloway Co., Blood River; cane sites #6 & #8, 15–21 Oct 2003 (1 ♂, 1 ♀). Same locality: cane site #8, 19 Oct 2004 (4 ♂). Calloway Co., Beachy Creek & Knight Branch; cane site #1, 15 Oct 2003 (2 ♂, 1 ♀). Carlisle Co., Bardwell, canebrake s. of Sandy Branch 36.83924, -89.0946 367´ [112 m]; 15 Oct 2011 at 175w MV light, A. E. McBride w/ Jim Wiker (1 ♀). Carlisle Co., 5 mi. SW Bardwell, east of old GM&O ROW, canebrake along Sandy Branch; larvae coll. 14 Jun 1999 boring in stem of Arundinaria gigantea, pupate: 19 Aug–8 Sep 1999, emerge: 16 Sep–8 Oct 1999; J. R. Wiker (7 ♂, 4 ♀). Same locality: ex larva coll. 14 Jun 1999 in Arundinaria gigantea, emerge: 30 Sep–14 Oct 1999; E. L. Quinter (3 ♂, 6 ♀). Same locality: 29 Sep–16 Oct 1999 (14 ♂). Same locality: 0.25 mi. E of RR grade; at UV light, 1–2 Oct 1999; M. C. Nielsen (1 ♂). Same locality: Westvaco WMA, Sandy Branch; light trap, 8–9 Oct 1999; leg. L. Gibson (3 ♂). Carlisle Co., ca. Burkley-Fish Lk. Rd., on Mississippi River at cane site #2; 18 Oct 2001 (2 ♂). Same locality: ca. 5 mi. W of Burkley, along Mississippi River; UV trap in canebrake, 17–18 Oct 2001; J. R. Wiker (1 ♂). Carlisle Co., Rover, ca. Lower River Road, on Mississippi River at cane site #3; 18 Oct 2001 (1 ♂). Carlisle Co., Mayfield Creek; cane site #1, 19 Oct 2003 (3 ♂). Carlisle Co., unnamed creek at Burkley; cane site #5, 8 Oct 2004 (1 ♂, 2 ♀). Carlisle Co., Mayfield Creek, Hays’ woods, 0.1 mi. W U.S. 62 bridge; cane site #3, 5 Oct 2005 (1 ♂). Carlisle Co., Mayfield Creek, W U.S. 62 bridge, 36°57´03.6˝ N, 88°49´39.4˝ W; cane site #6, 10 Oct 2005 (1 ♂). Fulton Co., Hickman, Goodman’s hill top cane in woods; 13 Oct 2001; (4 ♂). Fulton Co., Kentucky Bend; cane site #2, 12 Oct 2001 (1 ♂). Graves Co., Terrapin Creek State Nature Preserve area #4, ½ mi. SW Bell City; 13–20 Oct 2000 (5 ♂). Graves Co., East Fork of Terrapin Creek, cane site #1 W of Hwy 97; 13 Oct 2000; (1 ♂). Same locality; cane site #2, 21 Oct 2000 (2 ♂). Graves Co., West Fork Clarks River, Hominy Branch; cane site #5, 5 Oct 2005 (1 ♂). Graves Co., West Fork Clarks River; pipeline cane site #7; 12 Oct 2005 (2 ♂). Hickman Co., Hurricane Branch at Hwy. 239; at cane, 21 Oct 2001 (1 ♂). Hickman Co., Bowles Creek, Holder Farm at Resin Rd. bridge S; at cane, 11–12 Oct 2002 (9 ♂, 1 ♀). Hickman Co., Bowles Creek., Morris Farm; cane sites #1,2&3, 14 Oct 2002 (12 ♂). Same data: site #5, 18 Oct 2002 (1 ♂). Livingston Co., Dyer Hill Creek, Rt. 60 SW Buma, E of Birdsville; ex larva coll. in Arundinaria gigantea; emerge: 15 Sep 1999; E. L. Quinter (1 ♂). McCracken Co., Paducah, Massac Creek, 0.5 mi. S. Rt. 60; canebrake #1; 2 Oct 1999 (1 ♂, 1 ♀). McCracken Co., Massac Creek S of Hinkleville Rd.; cane sites #1 & #2, 12–16 Oct 1999 (7 ♂, 1 ♀). Same locality; cane sites #2 & #4, 14–15 Oct 2000 (1 ♂, 1 ♀). McCracken Co., cane site #1, woods E Massac Creek; 17 Oct 1999 (1 ♀). McCracken Co., Massac Creek at Hinkleville Rd. (old U.S. 60); 3–4 Oct 1999 at UV; M. C. Nielsen (2 ♂). Same locality; 14–15 Oct 2000; M. C. Nielsen (2 ♂). McCracken Co., Massac Creek, Streetman’s; pupa coll. 7 Sep 2002, emerged 1 Oct 2002; W. R. Black, Jr. (1 ♀). McCracken Co., west fork Massac Creek, N of new U.S. 60 bridge; cane site #5, 7 Oct 2004 (2 ♂). McCracken Co., old Mayfield Creek 1 mi. W of Clinton Rd.; cane site #7, 13 Oct 1999 (1 ♂). McCracken Co., old Mayfield Creek ca. 1 mi. W of Hwy 339; UV trap in cane, 12–13 Oct 2000; M. C. Nielsen (2 ♂, 1 ♀). Same locality; cane sites #20, #21 & #23; 10–23 Oct 2000 (3 ♂, 1 ♀). Louisiana. West Feliciana Parish, Tunica WMA, N30°56.144´ W091°29.964´; 31 Oct 2010; Vernon A. Brou (4 ♂). Mississippi. Chickasaw Co., Natchez Trace Parkway mile 231.1, Dick’s Creek; 28 Oct 2014; R. Patterson (1 ♂). Claiborne Co., Rocky Springs campground, Natchez Trace Pkwy., 34 mi. SW Clinton; 29 Oct 1988; D. & M. Hildebrandt, collectors (2 ♂, 1 ♀). Lee Co., Tombigbee St. Park; 23–30 Oct 1992; R. Kergosien (1 ♂). Same locality and collector; 1–22 Oct 1993 (1 ♂). Same locality and collector; 15 Oct–3 Nov 1995 (1 ♂). Rankin Co., Pearl; 18 Oct 1971; Bryant Mather coll. (1 ♀). Warren Co., Vicksburg; 27 Oct 1999; R. Patterson (1 ♂). Warren Co., Vicksburg, USAE Water Expt. Sta.; ex larva in Arundinaria gigantea, emerge: 24 Oct–5 Nov 1977; E. L. Quinter (3 ♂, 1 ♀). Same data; emerge: 18–26 Oct 1980; E. L. Quinter (1 ♂, 1 ♀). Same locality; collected at lights 19 Oct–11 Nov 1976; Bryant Mather (3 ♂, 1 ♀). Same locality and collector; 22 Oct 1980 (1 ♂). Same locality and collector; 20 Oct–1 Nov 1982 (2 ♂). Same locality and collector; 25 Oct–1 Nov 1983 (6 ♂). Same locality and collector; 16 Oct 1984 (1 ♀). Same locality and collector; 21 Oct 1985 (1 ♂). Same locality and collector; 21 Oct 1988 (1 ♂). Same locality and collector; 3 Nov 1994 (1 ♂). Warren Co., I-20 westbound weigh station, vicinity Bovina; UV light trap, 15–17 Oct 1980; E. L. Quinter (5 ♂). Same locality; ♂♂ attracted to virgin ♀ between 9:30 and 11:35 P.M.; 21 Oct 1980; E. L. Quinter (16 ♂). Same locality; collected at lights 8 Nov 1972; Bryant Mather (2 ♀). Same locality and collector; collected at lights 16–26 Oct 1973 (2 ♂). Same locality and collector; 23 Oct 1974 (1 ♂). Same locality and collector; 28 Oct 1975 (1 ♂). Same locality and collector; 15 Oct–11 Nov 1976 (2 ♂, 2 ♀). Same locality and collector; 31 Oct–3 Nov 1977 (2 ♂). Same locality and collector; 10 Nov 1983 (1 ♀). Same locality and collector; 5 Nov 1993 (1 ♂). Missouri. Butler Co., Mark Twain Nat’l. Forest, 360´ [110 m] jct. Rt. 67 & Black River 0.6 mi. WNW Hendrickson; MV light, 14–16 Oct 2000; E. Quinter & J. Wiker (1 ♂). Butler Co., Hendrickson, canebrake n. Black River 36.9041, -90.4777 378´ [115 m]; 13 Oct 2015 at 175w MV light; A. E. McBride w/ Jim Wiker (2 ♂). Wayne Co., Mark Twain National Forest, 400´ [122 m] jct. Rt. 49 & Black River, 2.7 mi. W Williamsville, Browns Crossing; ex larva in Arundinaria gigantea, emerge 25 Sep 1994; E. L. Quinter (1 ♀). North Carolina. Madison Co., Lower Shut In Rd., 2.6 mi NW Hot Springs, 35.9209, -82.8602, 1,291´ [393 m]; ex larva 28 Jun 2023, collected by Becky Elkin and Jim Petranka; ex mature culm of Arundinaria gigantea, reared by A. E. McBride in carrot, eclosed 30 Oct 2023 (1 ♀). Tennessee. Henry Co., Blood River; UV light trap at cane site #P2, 12 Oct 2003; W. R. Black, Jr. (1 ♀). Same locality and collector; UV light trap at cane site #P2, 17 Oct 2004 (5 ♂). Same locality and collector; UV light trap at cane site #P2, 11 Oct 2006 (1 ♂). Same locality and collector; UV light trap at cane site #17, 18 Oct 2006 (2 ♂). Jackson Co., Cordell Hull Lake, Big Bottom; ex larva coll. in Arundinaria gigantea 23 Jun 1995, pupate: 27 Aug, emerge: 25 Sep 1995; E. L. Quinter (1 ♀). Lake Co., 1 mi. E Phillipy; UV light trap at cane, 14 Oct 2001; Wm. R. Black, Jr. (1 ♂). Lake Co., Reelfoot St. Park WMA, 200 m. E Cohoctaw/McKutchen Rd.; UV light trap at cane site #2, 20 Oct 2000; Wm. R. Black, Jr. (1 ♂, 1 ♀). AMNH, CMNH, CNC, INHS, LACM, MCLB, MEM, NHML, NYSM, UCD, ULK, USNM, VADNH, YPM, AEM, ELQ, JBS, JKA, JRW, MCN, RPC, VAB.
The larva bores in mature, woody, second-year and older culms of giant cane (Arundinaria gigantea). This is in stark contrast to the closely related P.sullivani, which utilizes only tender, green, newly emergent culms of switch cane. The culms of its host grass attain much greater diameter than those of the host of P.sullivani, which has a larva that is always tightly confined. A single larva, probably third instar, collected by ELQ on 27 May 1994 in Wayne Co., Missouri, molted five times between 5 June and 24 July, pupating 26 August. This suggests that P.matheri larvae could have as many as eight instars. A surprisingly small amount of woody material is consumed (if consumed at all, as opposed to merely chewed) during the entire larval development, compared to the copious amounts of tender meristem consumed by its counterpart P.sullivani. It appears likely that P.matheri larvae, like those of Franclemontia interrogans, obtain much of their energy requirements from the sugary fluids that flow through the living culms, rather than from ingesting the plant tissues themselves. This appears to be the case with some other Papaipema species as well, such as those that form woody galls but consume very little actual plant tissue, in contrast to those species which prefer to bore inside hollow stems. The presence of a P.matheri larva in a giant cane culm is revealed by internal girdling of one or more of the internodes, which weakens the culm and readily causes breakage at the point of girdling, commonly by windstorms. Pupation occurs at the base of the last-bored internode, with the pupa resting head up on the septum, oriented just below an exit hole prepared by the final instar prior to pupation. See discussion of F. interrogans for hints on finding larvae and pupae of P.matheri and F. interrogans. The following anecdote by ELQ illustrates that information new to science is not necessarily new information:
“Many years ago, I was travelling alone in a very rural part of Arkansas, hoping to discover some high-quality canebrake remnants that might enable me to document the occurrence there of larvae of several of my recently discovered species, including Papaipema matheri. On a gravel road many miles from my campsite, I acquired a flat tire. While jacking up the car to change the tire, the cheaply made jack bent and collapsed in such a manner as to become pinned underneath the car. Darkness was only a couple hours away, and I sat there frustrated and trying to remember how many miles back I had passed the last evidence of civilization. Just as I was contemplating spending the night sleeping in my disabled vehicle, I heard a distant rumbling. Less than a minute later, an ancient pickup truck with rusted side panels held together with duct tape and paint came roaring around the curve, skidding on the gravel to an abrupt stop next to me sitting there. A ruddy-looking fellow in old clothes emerged from the truck and immediately noted my predicament, followed by a kind offer to help. The only item he was carrying in the back of his truck was an enormous jack, and in short order, he removed the flat tire and replaced it with my spare. During that episode he had inquired what I, in a car with New York plates, was doing “way out here.” I explained my interest in finding insects associated with cane, locally. He promptly told me he knew some good spots nearby, inviting me to join him and his buddies at a catfish fry they were having. Around their campfire that evening, taking in a local dialect resembling English, I learned that “them there canes makes really good fishin’ poles, and they gets worms in ‘em that makes really good bass bait!” (I suspect that Native Americans knew that long before settlers ever came to Arkansas.)”
Papaipema matheri occurs throughout the lower Ohio and Mississippi River drainage, and eastward to the Appalachians, wherever giant cane grows in sufficient abundance to support populations. Papaipema matheri is not found on the Gulf Coastal Plain, where it is replaced by P.sullivani using switch cane. Although giant cane occurs east of the Appalachian Mountains, P.matheri has not established a significant presence there. AEM, along with Jim Petranka and Becky Elkin, have found several larvae in North Carolina in Madison and Buncombe Counties above 400 meters, however, on a river that flows westward into Tennessee.
Papaipema marginidens GROUP
As currently conceived, this group includes two seemingly ancient members, Papaipema marginidens and P.eryngii, which may have diverged from the same ancestral lineage. Papaipema eryngii, with its monophagous specialization on rattlesnake master, may indicate an ancestral preference for apiaceous hosts which is retained in several more recently derived groups.
Papaipema marginidens (Guenée) (Wild Yam Borer)
Gortyna marginidens Guenée, 1852, in Boisduval and Guenée, Histoire Naturelle des Insectes. Species Général des Lépidoptѐres, 5: 123.
Type locality [neotype]: “New Yorck?” [New York, USA]. [USNM]
note — Guenée’s description of this species was based upon two specimens from Illinois, a male and a female, although he expressed some doubt whether both were the same species. Both specimens are lost and presumed destroyed.
Gortyna nephrasyntheta Dyar, 1908, The Canadian Entomologist, 40: 78.
Type locality: Plummer’s Island, Maryland, [USA]. [USNM]
note — Described from a single female.
Gortyna nephasyntheta; Lyman, 1908, misspelling.
Any Papaipema species, if deprived of sufficient larval food resources, can result in dwarfed individuals in the adult stage. Papaipema marginidens varies in size in this regard, depending upon the size of its host plant rhizome, and can be quite small in some populations. Frequently though, P.marginidens individuals can also be quite large, and only a few other Papaipema species can attain an equal size. Papaipema eryngii can be similarly large, and has similar forewing maculation, but is normally dark chocolate brown. Individuals of P.eryngii from an apparently extirpated Atlantic Coast population (
This species has been vexing to all who sought to better understand it. Henry Bird, the quintessential pioneer researcher of Papaipema biology, devoted decades of field work in an attempt to discover the life history and host plant of Papaipema nephrasyntheta, as this species was then called. His efforts were unsuccessful in that regard but led to the discovery of several other previously unknown species in the process. AEM, JRW, and ELQ have devoted an additional 50 collective person-years of field work to this quest. Finally, AEM met with success in 2019. For a detailed account of that endeavor, see
The larva belongs to the group of Papaipema species in which all three dorsal and subdorsal longitudinal white stripes are interrupted by the pigmented bands encircling the first four abdominal segments. In P.marginidens, the pigmentation is lighter than most Papaipema species and of a pinkish-orange color. The larva is also relatively longer and more slender than larvae of many other Papaipema and features a characteristic prognathous head capsule, reflecting its tight confinement within the slender rhizome of its host. The lateral margins of the prothoracic shield are not darkened as in most species and thus do not extend to the brownish-orange head capsule. The dorsal pinacula are noticeably tiny in comparison to most other Papaipema species. For a more detailed larval description, refer to
Papaipema marginidens is a widespread species in eastern North America, apparently limited to austral regions. Its range strongly coincides with areas unaffected by the Wisconsin glaciation. It is found from Connecticut and extreme southern New York west through the Ohio River watershed to Illinois, where it spreads north into the unglaciated driftless area of southeastern Minnesota and southwestern Wisconsin. It ranges south to the northern parts of the gulf states of Louisiana, Mississipi, Alabama, and south to northern Georgia and South Carolina along the Atlantic coast.
This species likely received its name due to the dentate fringes of the outer margins on the forewings, a feature often preserved only on very fresh individuals. Owing to its early description and lack of unambiguous published illustrations, much uncertainty has prevailed as to the correct identity of P.marginidens. The original description is suggestive of several species, although the dimensions stated [48 mm] limit the possibilities.
Papaipema eryngii Bird (Rattlesnake-master Borer)
Papaipema eryngii Bird, 1917, The Canadian Entomologist, 49: 125.
Type locality: Chicago, Illinois, USA. [AMNH]
note — Described from multiple reared specimens ranging from 35 to 48mm, but the number not specified.
Papaipema nephrasyntheta; authors, not Dyar, 1908.
Very likely to be confused with Papaipema eryngii is P.birdi, which occurs with it. These two species are visually very similar but the highly restricted habitat requirements of P.eryngii should serve to help distinguish it from the ubiquitous P.birdi; P.eryngii will be found only in prairie habitats where large colonies of its relatively rare host plant flourish, but P.birdi will be found almost everywhere throughout its range. The forewings of P.eryngii are slightly narrower than those of P.birdi, chocolate brown in color, and the yellowish apical patch is usually more prominent. The medial area in P.eryngii tends to have fewer reddish scales and more yellow scales near the inner margin. The male genitalia offer a convenient means of distinguishing these two if in doubt. Simply by brushing the scales off the terminal part of the abdomen, P.birdi easily can be distinguished by its bifurcate cucullus. Otherwise, P.eryngii might be confused with P.marginidens, which also has distinctive genitalia. See distinguishing features of P.marginidens, treated above. DNA barcodes are available for four samples of P.eryngii from Illinois, which form an isolated barcode group differing by at least 3.37% from other Papaipema.
The larvae of these two species are also readily distinguishable; P.eryngii larvae have all three longitudinal white lines interrupted on the first four darkly pigmented abdominal segments, but P.birdi larvae have the dorsal line continuous across these segments. The dark purplish-brown larval pigmentation of P.eryngii is retained through all instars, although increasingly somewhat faded in the latter ones. The dark lateral margins of the prothoracic shield do not extend onto the head capsule, which is contrastingly orange in color.
This species is an obligate specialist borer in rattlesnake-master (
Papaipema eryngii historically has been associated with and largely confined to big bluestem prairie habitats and is presently found scattered across remnants thereof in at least seven states. This species had been considered a candidate for federal listing as an endangered species, based primarily on its documented historic occurrence within the rather limited immediate environs of Chicago, Illinois where it has long since been extirpated due to development. Recent extensive field work conducted by JRW throughout the lower Midwest has documented its wide occurrence wherever sufficient host plant resources survive in unburned prairie remnants. Perhaps not coincidentally, tall-grass prairies were once also the preferred habitat of the American bison, and it is speculated that bison may have been largely responsible for the dispersal of rattlesnake-master populations throughout these habitats, because the dried, spiny, seed-bearing portions of these plants regularly became tangled in the bison’s shaggy fur.
A remarkable discovery was made in 2001 while examining Papaipema specimens in the private collection of Dr. J. Bolling Sullivan. In a long series of P.marginidens, ELQ noticed a specimen that looked a bit different. Upon subsequent dissection of the genitalia of that male, it became immediately obvious the specimen was P.eryngii – the first known example from a highly disjunct Atlantic Coast population of this midwestern species. This specimen (
With the notable exception of the aforementioned Atlantic Coast population now believed extirpated, P.eryngii is largely restricted to unburned native prairie remnants across the central Midwest.
Papaipema cataphracta GROUP
Papaipema cataphracta, P.sulphurata, P.aweme, P.necopina, and P.duovata constitute this group. Papaipema sulphurata appears to be a recently derived sister species of P.cataphracta, possibly of Pleistocene origin, given its extremely limited distribution in a northern area that escaped glaciation. Papaipema aweme shares both genitalic and larval characters with P.cataphracta and P.sulphurata. Less obvious is the apparent close relationship with the dark-colored P.necopina, which likely hybridizes with P.cataphracta, is morphologically nearly indistinguishable, and barcodes with P.cataphracta, but has differing larval pigmentation. Papaipema duovata shares larval characteristics with P.cataphracta as well as genitalic morphology in the adult. All are stem borers.
Papaipema cataphracta (Grote) (Burdock Stem Borer)
Gortyna cataphracta Grote, 1864, Proceedings of the Entomological Society of Philadelphia, 3: 81.
Type locality: “Eastern and Middle States” [USA]. [ANSP]
note — Described from two males. An authentic Grote specimen of cataphracta in the NHML bearing only his blue-bordered label with handwritten determination is apparently not a syntype. Original Grote type labels all bear red borders with the genus and species name and the handwritten word “Type.” Grote’s types of species named prior to 1869 are deposited in the ANSP. These inexplicably lack their original labels, but their authenticity is not in question. The name is derived from the Greek kataphraktos, meaning fully enclosed, perhaps in allusion to the larval habit in this group of boring and remaining completely concealed inside the host plant.
† Gortyna aerata Lyman, 1901, The Canadian Entomologist, 33(12): 319. new synonymy; unavailable;
Type locality: Vicinity of Montréal, Québec, [Canada]. [LEM]
note — Unavailable name based on type specimens of hybrid origin. This taxon has heretofore been treated as a valid species of Papaipema. As discussed below, it is here considered to be a naturally occurring hybrid of P.necopina (Grote) and P.cataphracta (Grote). As such it is an unavailable name placed in synonymy under the oldest putative parent taxon, P.cataphracta. For more information on this phenomenon, see the section on Aberrations and Hybrids in the introductory section on the genus Papaipema and also below. note — No type locality is mentioned in the original description, but the syntypes are all labelled “Montréal.” Described from four males and five females. The “No. 1 ♀” was selected as lectotype by
† Papaipema cataphracta ab. fluxa Bird, 1911, The Canadian Entomologist, 43(2): 44.
Type locality: Buffalo, [Erie Co.], New York, [USA]. [AMNH]
note — Described from one male and one female as an aberration of P.cataphracta, reared from burdock. As discussed by
Other than with P.sulphurata, it is unlikely that P.cataphracta could be confused with any other species. Papaipema cataphracta looks somewhat similar to P.rigida, but the postmedial line in P.rigida is very straight, bending inward very close to the costa, whereas the postmedial line in P.cataphracta is sinuous. Papaipema sulphurata is separated mostly by range and by host plant association; see further discussion below under P.sulphurata. Occasionally, individuals of P.cataphracta have white stigmata, especially in parts of the Midwest.
To date, DNA barcode variation has been examined in 13 samples of P.cataphracta from across the species’ range. This species is one of few that exhibit an unexpected barcode pattern consisting of three disparate barcode groups in two BINs. It is possible that this pattern is produced by the wingstem (Verbesina alternifolia) feeding segregate described below, although at present an incongruous genetic artifact cannot be ruled out, given that the two widely divergent barcode groups are not each other’s nearest barcode neighbors. The situation is further complicated by P.sulphurata, the barcodes of which also place in both P.cataphracta BINs. Nevertheless, a comparison of the ecological and morphological traits of the P.cataphracta barcode groups are warranted, which are geographically segregated: one extending from the northeastern United States and Canada south through the Appalachians (Québec, Ontario, Virginia, and North Carolina), and the second so far strictly southern (Georgia, Kentucky, North Carolina, and South Carolina). Both BINs are present in North Carolina, the northern from Swain County and the southern one from the east-central part of the state.
The dissimilar appearance of adult P. cataphracta and P.necopina obscures the closeness of their relationship. The genitalic morphology of both sexes is identical, and the two species are suspected of interbreeding freely in situations where both coexist.
The larva of P.cataphracta is by far the most commonly encountered Papaipema of the phenotype with all three dorsal and subdorsal white longitudinal stripes continuous across the pigmented bands of the first four abdominal segments. The larva bears the usual darkened lateral margins of the prothoracic shield which extend onto the honey-colored head capsule, and the pinacula are quite prominent on this species. A character that can be used to separate larvae of P.cataphracta from all other Papaipema larvae with three continuous dorsal and subdorsal longitudinal stripes except those of P.sulphurata and P.duovata is seen through examination of the D1 pinacula, which in this species touch the margin of the white dorsal stripe on the anterior abdominal segments. This common species shares with P.nebris a nearly omnivorous diet, boring in the stems of a multitude of unrelated perennial forbs. Nearly one hundred years ago, P.cataphracta had already been recorded boring in plants distributed across seventeen families (
Like P.nebris, which accepts almost any herbaceous plant yet exhibits a strong predilection for ragweed throughout its range, P.cataphracta displays a similar preference for wingstem in more southern portions of its range where this plant is abundant. Papaipema sulphurata, a very close relative of P. cataphracta, has evolved a monophagous specialization probably as a relict population isolated in unglaciated, coastal (now offshore) refugia during the Pleistocene. It is possible that southern P.cataphracta populations selecting wingstem as their preferred choice may presently be undergoing a similar evolutionary trend, and it is hoped that additional molecular data may support this hypothesis.
A peculiar characteristic of P.cataphracta is the relatively large amount of frass expelled from the extensive larval galleries in the stems of its hosts, especially in comparison to other Papaipema, many of which tend to favor the subterranean parts of their hosts. Perhaps this is indicative of a relatively low nutritional value of plant stems in relation to food stores of roots, rhizomes, etc. The presence of copious amounts of light-colored frass accumulated on soil beneath bored plants is a clue to an infestation of this ubiquitous species. Wilting or bent stems also are typically observed. Pupation normally occurs within the host plant at the base of the larval gallery. This species is among those Papaipema that fly later in the season than many others. The majority of records indicate a main flight season beginning in late September and continuing through mid-October, but with earlier records in the northernmost portions of its range and occasional very late captures in the deep South.
Papaipema cataphracta is widely distributed throughout the eastern United States and southern Canada, where it is normally a very common species. It is apparently much less common in the southernmost portions of its range, where it appears to be limited to the coastal plain. See the discussion of this peculiar phenomenon under P.sulphurata, below.
Papaipema sulphurata Bird (Swamp Loosestrife Borer)
Papaipema cataphracta sulphurata Bird, 1926, The Canadian Entomologist, 58: 252.
Type locality: Martha’s Vineyard, [Dukes Co.], Massachusetts [USA]. [AMNH]
note — This taxon was described as a variety of P.cataphracta from fifteen specimens. It was elevated to full species status in ELQ’s 1983 check list (Hodges, 1983).
Other than with P.cataphracta, it is unlikely that P.sulphurata could be confused with any other species. However, the occurrence of a moth infrequently encountered and sporadically distributed throughout the southeastern United States on the coastal plain which strongly resembles P.sulphurata needs explanation. The sandy and acidic soils prevailing there do not appear to favor these occurrences. In the interior, individuals blend imperceptibly into P.cataphracta phenotypes which cannot be distinguished from them, and thus all such individuals are being treated as P.cataphracta, despite some serious reservations. Papaipema sulphurata tends to have a greater amount of yellowish coloration extending throughout the forewing, especially beyond the postmedial line and between the antemedial line and the wing base, creating less contrast over the surface of the wing and causing the moth to appear more yellow overall. The areas beyond the postmedial line and basad of the antemedial line are typically darker and more contrasting in most P.cataphracta, save for individuals of the more southern and south-coastal moths discussed above. DNA barcodes are available for only two specimens of P.sulphurata, both from Massachusetts. The barcode haplotype variation of P.sulphurata in relation to P.cataphracta is perplexing and cannot be easily explained without additional research. This is discussed further in the P.cataphracta account.
Unlike P.cataphracta, P.sulphurata seems to require the acidic wetland habitats of the coastal plain for its survival. With the exception of P.sablensis, P.sulphurata has the most restricted distribution of all our eastern Papaipema, apparently being largely confined to the islands of Nantucket, Martha’s Vineyard, and freshwater ponds on Cape Cod, Massachusetts. These areas represent present-day remnants of an unglaciated offshore refugium, now under water, which likely extended both much farther north and farther south along the Atlantic Coast during periods of maximum glaciation. The apparent fact that P.sulphurata shows no inclination to utilize any plant other than its specific larval host and remains recognizably distinct from mainland populations of P.cataphracta wherever the two occur in close proximity to one another seems to underscore its status as a distinct species. There is no evidence that swamp loosestrife, which is very widespread throughout eastern North America, is ever utilized by either species other than on Cape Cod and the aforementioned islands.
The larva has both its middorsal and subdorsal longitudinal lines uninterrupted by the pigmented banding of A1–A4. It shares the prominent pinacula with its sibling species P.cataphracta and is indistinguishable from it, although perhaps not reaching as great a size due to being restricted to the often very tight quarters of its slender-stemmed, woody host.
Swamp loosestrife (Decodon verticillatus) is the only recorded host for this highly localized species. These plants often form dense colonies growing in standing water in and around vernal ponds and the other wetland environments where P.sulphurata occurs, but although such colonies may be numerous on the rather limited remnants of the coastal plain, they do not appear to harbor large populations of the moths, and the actual numbers recorded at any particular site commonly vary from a few to none from year to year (
Larval activity is confined to the stem. The root is not bored. Prior to pupation, the larva prepares an exit hole for the adult to depart from the gallery. The pupa is formed in the gallery on a platform of chewed, woody fragments below the exit hole. It is notable that the exit holes are typically formed well up on the stem and are less susceptible to inundation. Mark
As currently conceived, this species has an extremely limited distribution on Cape Cod, Massachusetts, the adjacent Massachusetts mainland, and on the offshore islands.
Papaipema aweme (Lyman) (Bogbean Borer)
Gortyna aweme Lyman, 1908, The Canadian Entomologist, 40: 254.
Type locality: Aweme, Man. [Manitoba, Canada]. [LEM]
note — The species was described from a single specimen. The holotype male is illustrated in
Papaipema aveme; Hampson, 1910, Catalogue of the Lepidoptera Phalaenae in the British Museum, 9: 81. Unjustified emendation.
Papaipema aweme is among the smaller members of the genus. It is a tan to beige moth with a pinkish flesh tinge, only known to occur as a single phenotype. It could be confused with certain phenotypes of P.unimoda (
Larvae of P.aweme are a pale, warm brown with small pinacula and bear three whitish longitudinal stripes running the length of the body. At maturity, this pattern is obscured, and the larva becomes a pale, nondescript dull whitish color. This species is monophagous, the larva boring in the stems and rhizomes of the hygrophilous dicot bogbean (Menyanthes trifoliata). The larva matures by late July or early August and does not aestivate before pupation, an adaptation to the short growing season of its northern habitat. In captivity, larvae often pupate within a cell in the gallery, but it is unknown whether this occurs in nature or if larvae abandon the plant to pupate in nearby detritus. When found as a larva, this species is best reared in stems and rhizomes of the host plant, since it does not readily accept alternate food such as carrot. The adult emerges after a pupal period of approximately 18 days. This species is on the wing throughout August and into the first half of September, and large numbers of individuals can be attracted to light in prime habitat (Kyle Johnson, pers. comm.). For detailed accounts of the life history of P.aweme, see
For more than one hundred years, this species was regarded as one of North America’s rarest moths. Recent findings perfectly illustrate the importance of knowing larval host plant preferences. Since the discovery of the larva and its host by AEM in 2016, P.aweme has proven abundant in its preferred fen and peatland habitat across northern USA and southern Canada, covering a vast region of boreal North America. When considering species potentially in need of protection, it becomes clear than none could be more secure. The possibility of human activity negatively impacting most populations of P.aweme is highly unlikely. Its boreal peatland habitats are remote, inhospitable, and vast, stretching nearly across the continent. Papaipema aweme can be found in a band extending from east central Saskatchewan south through Manitoba and northern Minnesota and east through the Great Lakes region to eastern Ontario and upstate New York.
Papaipema necopina (Grote) (Sunflower Borer)
Gortyna necopina Grote, 1876, The Canadian Entomologist, 8: 25.
Type locality: Buffalo, N.Y. [Erie Co., New York, USA]. [NHML]
note — Described from one male and the female holotype. Grote’s name is derived from the Latin necopinus, unexpected.
Papaipema limpida; of authors, not Guenée, 1852. Papaipema imperspicua; of authors, not Bird, 1908. Papaipema imperturbata Bird, 1907, The Canadian Entomologist, 39: 274.
Type locality: [New Brighton, Beaver Co., Pennsylvania, USA]. [AMNH]
note — The type locality is not specified in the original description, but from the context of Bird’s writing, it is obvious he collected the larvae while visiting his friend F. A. Merrick at New Brighton, Pennsylvania.
Papaipema mulleri Buchholz, 1956, The Lepidopterists’ News, 10(5): 139.
Type locality: Highway 206, 6 miles north of Stanhope, Sussex Co., New Jersey, [USA]. [AMNH]
note — Described from seven males and three females which were reared from paleleaf woodland sunflower (Helianthus strumosus) by Joseph Muller. The specimen labelled “Holotype” by Buchholz is not part of the original type series since it is labelled as coming from Morris Co., New Jersey. The true syntypes are all from the type locality in Sussex County. It should be further noted that many Muller specimens, including these, bear his label “J. Muller, Short Hills, N.J.” This was where his home was located in the earlier years of his collecting and does not reflect the actual collecting locality. The syntypes collected 6 miles north of Stanhope are labelled simply “Waterloo.”
Papaipema necopina is likely to be confused with several other species with dark forewings, especially P.maritima, but also P.silphii, P.duplicata, and P.beeriana. Diagnostic features are discussed under each of those species. The forewing of P.necopina closely resembles the dead, dry leaves of its larval hostplants. These leaves are typically very scabrous, with light speckles on a dark-brown background, creating a salt-and-pepper effect replicated on the forewing of the moth, which presumably rests by day on the dead leaves of the host plant. This type of camouflage is employed by several other Papaipema species which feed upon composites with similar foliage characteristics, most notably P.silphii and P.beeriana. As discussed under P.cataphracta, that species and P.necopina are believed to hybridize, and subsequent hybrids were the basis of the taxon P.aerata.
DNA barcodes are available for 16 specimens of P.necopina, from Ontario and Pennsylvania. These form a discrete cluster within a larger BIN that includes the P.insulidens group. Minimum nearest-neighbor difference is 0.70% to P.insulidens– P.birdi haplotypes.
The larva bears only the single middorsal longitudinal white line crossing the dark-banded anterior abdominal segments. The dark lateral margins of the prothoracic shield extend onto the honey-colored head capsule. More details concerning the larval stages may be found under the name P.imperturbata in
Papaipema necopina is infrequently seen at lights. Adults are best secured by rearing larvae found boring in the stems of various thick-stemmed species of sunflower. In some places, both P.necopina and P.maritima larvae can be found boring in the same patch of sunflowers. Papaipema necopina larvae make a tapered cylindrical gall at the base of the stem and frequently vacate the gallery to pupate in soil nearby. Papaipema maritima larvae make a nearly round gall at the base of the stem and always pupate inside the gall. The robust sunflowers may not show obvious signs of being bored, but the cylindrical gall formed by P.necopina at the base of the plant is sufficiently distinctive to alert one to the presence of this species. This is frequently accompanied by a deposit of light-colored frass on soil beneath the exit hole of the gall. Pupation takes place in the gall or in soil nearby. Specimen capture data indicate a flight predominantly from mid-September through mid-October, with occasional earlier records.
This species is generally distributed from New England west to South Dakota and south to Missouri, Tennessee, and North Carolina.
Papaipema duovata (Bird) (Seaside Goldenrod Borer)
Hydroecia duovata Bird, 1902, The Canadian Entomologist, 34: 115.
Type locality: [Rye, Westchester Co., New York, USA.] [AMNH]
note — The type locality is not stated in the original description, and the species was described from an unknown number of syntypes. These were reared from larvae collected at Rye, New York, as recorded on the type specimen labels.
It is unlikely that Papaipema duovata could be confused with any other species due to both its distinctive coloration and its habitat along the immediate coast. Some individuals of P.rutila and P.polymniae may appear similar, but these are never as dark as P.duovata, and the habitat is completely different. There is a small amount of yellowish-tan in the forewings of P.duovata, especially near the inner margin, but the wings are heavily marked in dark brown, creating a coloration that is unique to this species. The stigmata of the orbicular-reniform complex are always white. Papaipema duovata is consistent in appearance throughout its range. Two DNA barcode samples from North Carolina are available, representing an isolated barcode group with a minimum nearest-neighbor distance of 2.5%.
This is another of the relatively few Papaipema species with a larva that features continuity of both its middorsal and subdorsal longitudinal lines across its first four abdominal segments. Like P.cataphracta, the larva has pinacula that are larger than in most other species. The dark-pigmented lateral margins of the prothoracic shield do not extend onto the honey-colored head capsule. Further details of larval morphology may be found in
This species occupies a highly specialized niche in coastal Atlantic salt marshes where its host plant flourishes in brackish drainage ditches and along roadsides surrounding these marshes. The host plant is often abundant near beaches and on dunes as well, but the larva is not found boring in seaside goldenrod growing in these hot, dry, sandy situations. On this account, P.duovata inhabits a very narrow and limited wetland habitat and will not likely be found elsewhere. Bird gave this epithet to the species because, in his experience, he usually found two larvae boring in each of the plants he examined and concluded that the parent female had deposited two eggs on these plants the previous fall. There is no indication that this is true. Usually only one larva is located in a plant, unless it is a large plant with multiple stems arising separately from the rootstock. As Papaipema larvae are cannibalistic, generally no more than one larva ever survives in the same stem, unless physical barriers preclude larval interaction in the galleries. Multiple P.nebris larvae may occur in the same ragweed stem, for example, but occupy physically separate galls. The larvae bore well down into the root but maintain an open, ventilated gallery extending high enough up the stem to minimize chances of drowning in their burrows during periods of exceptionally high tides. Aside from the exit hole on the stem and often accumulated tan frass below, there is little other evidence of infestation in these robust plants. The pupa is formed inside the larval gallery and is narrow and cylindrical in shape. In the north, the flight time begins in late September. This is among the later-emerging species of the genus, flying throughout October and well into November in the more southerly portions of its range.
Papaipema duovata is distributed from Massachusetts to Louisiana in a very narrow zone closely paralleling the coast.
Papaipema speciosissima GROUP
Three species constitute this cohesive group: Papaipema speciosissima, P.inquaesita, and P.stenocelis. All three are monophagous or oligophagous moths which specialize on one or more species of fern, undoubtedly having arisen from some ancient fern-feeding ancestral lineage. Their forewing patterns are distinctive.
Papaipema speciosissima (Grote & Robinson) (Royal Fern Borer)
Gortyna speciosissima Grote and Robinson, 1868, Transactions of the American Entomological Society, 1: 342.
Type locality: “Atlantic District. (Seekonk, R. I., Mrs. Bridgham.)” [East Providence, Providence County, Rhode Island, USA]. [ANSP]
note — Seekonk, Rhode Island never existed as a political subdivision. The type of Gortyna speciosissima was collected by Mrs. Eliza Bridgham at her family’s farm in Seekonk, Massachusetts. Mrs. Bridgham’s specimens typically bore a locality label that included only the town name, Seekonk. In 1862, the western part of Seekonk, Massachusetts (where the Bridgham Farm was located) was annexed to Rhode Island and became East Providence, thus the type locality is now East Providence, Rhode Island. Grote and Robinson’s stated locality is presumably a combination of Bridgham’s original label (Seekonk), since lost, and the state where Mrs. Bridgham resided at the time of the description in 1868 (Rhode Island). Described from a single female.
† Papaipema speciosissima form regalis Wyatt and Beer, 1927, Entomological News, 38: 215.
Type locality: Hessville, [Lake Co.], Indiana, [USA]. [CMNH]
note — Unavailable name based on an infrasubspecific form. note — Described from the male holotype as well as one male and three female paratypes.
This handsome species is a favorite among Papaipema afficionados due to its size, coloration, and the fact that it is never seen in sufficiently large numbers to become commonplace. There is really nothing that should cause confusion regarding the identity of this large, bright orange species, although in some respects it resembles a giant P.inquaesita and in others, a giant P.stenocelis. From P.inquaesita, P.speciosissima is distinguished by its larger size, yellower coloration, and the appearance of the reniform, orbicular, and suborbicular stigmata. The reniform and orbicular-suborbicular form long white or dark bars in P.speciosissima. In P.inquaesita, these stigmata are composed of individual spots that are most commonly orange, rarely white, never dark, and do not form long bars. From P.stenocelis, P.speciosissima is distinguished by the path of the postmedial and the medial lines. In P.speciosissima, these two lines mirror each other and are sharply bent outwardly; in P.stenocelis, these two lines do not run parallel to each other, the postmedial line being broadly curved and the medial line straighter. Papaipema speciosissima occurs in two forms. The common form has white stigmata. More rarely encountered is form regalis Wyatt & Beer, which has dark brown stigmata. DNA barcodes are available for nine specimens of P.speciosissima ranging from Florida to Ontario, which form a discrete group and unique BIN at a minimum nearest-neighbor difference of 2.9%.
The mature larva is among the largest of the genus and has a decidedly pinkish coloration but is otherwise largely devoid of distinct markings. Earlier instars reveal that this species belongs to the group wherein neither the mid-dorsal nor subdorsal longitudinal white lines extend across the dark pigmented bands of the first four abdominal segments. More details of the larval stages may be found in Bird (1915c).
Any of the regal fern species (Osmunda) may serve as hosts for the larvae of speciosissima, but royal fern (Osmunda regalis) is by far the species most commonly utilized. Cinnamon fern (O. cinnamomea) is less commonly utilized, and interrupted fern (O. claytoniana) generally occurs in drier habitats not frequented by the moth. The presence of larvae boring in the woody, subterranean portions of the typically large fern clusters is rarely obvious, as these robust plants do not wilt or exhibit obvious signs of being infested. Neonate larvae enter an emerging frond very early in the season, stunting its growth, and soon proceed to the rootstock. All the other fronds of the plant rapidly develop, and the initial damage is soon obscured. Later in the season the only evidence which may remain is a small amount of muddy orange frass accumulated on the soil at the base of the small, stunted, and dried frond initially bored by the young larva.
Papaipema speciosissima never strays far from the swampy wetlands where royal ferns flourish. Obtaining immaculate, fresh specimens by finding larvae and rearing them to adults may be a laudable goal, but quite challenging to say the least. Recognizing that a colony of royal ferns may be harboring borers is the first challenge, since this is seldom obvious. The rootstock of these ferns consists of fibrous, intergrown masses which apparently constitute a single plant and may measure several meters across or more in size, so digging up a clump of this enormous, woody rootstock can take hours. To sample multiple random patches of ferns in hopes of detecting larvae is simply too labor intensive and may prove to be all in vain.
With the exception of the extreme situation in southern Florida where this species has been recorded flying in nearly every month of the year (see the discussion concerning this phenomenon in the introduction where the topic of voltinism is addressed), the adult flight period in the northern parts of its range commences in early September and continues throughout all of October. Farther south, the flight period begins about a month later and continues throughout November.
Papaipema speciosissima has a remarkable distribution from Québec, south along the Atlantic Coastal Plain to the Florida Keys, and west along the Gulf to Louisiana. It also extends up the Mississippi and Ohio River Valleys and spreads throughout the Great Lakes Region. There are large areas within this broad range where the species has never been recorded and where it actually may be absent. With its occurrence in the Florida Keys, it has the southernmost distribution of all Papaipema species.
Papaipema inquaesita (Grote & Robinson) (Sensitive Fern Borer)
Gortyna inquaesita Grote and Robinson, 1868, Transactions of the American Entomological Society, 1: 344.
Type locality: “Atlantic District. (N.Y., Mr. Ridings; R. I., Mrs. Bridgham.)” [East Providence, Providence County, Rhode Island, USA, here restricted.] [Types lost.]
note — At least two specimens, one but presumably both females, were seen by Grote and Robinson, since it is stated that the Rhode Island specimen had white stigmata whereas the other had deep yellow spots. Both specimens appear to be lost. Neither can be located among material at the ANSP where at least one of them would be expected to be deposited. The provenance of the New York specimen and its whereabouts are unknown. The putative “type” in the NHML, re-labelled “U.S. America,” is spurious. The identity of the species is not in question, so a neotype designation is unnecessary. The locality of the Mrs. Bridgham specimen is hereby restricted as the type locality (ICZN Article 74.4): Seekonk, Rhode Island never existed as a political subdivision. One of the syntype specimens of Gortyna inquaesita was collected by Mrs. Eliza Bridgham at her family’s farm in Seekonk, Massachusetts. Mrs. Bridgham’s specimens typically bore a locality label that included only the town name, Seekonk. In 1862, the western part of Seekonk, Massachusetts (where the Bridgham Farm was located) was annexed to Rhode Island and became East Providence, where Mrs. Bridgham resided at the time of the description.
[Gortyna] quaesita; Grote, 1881, misspelling. Papaipema inquaesita form wyatti Barnes and Benjamin, 1929, Bulletin of the Brooklyn Entomological Society, 24: 180.
Type locality: St. Therese Isl., St. Johns Co., Québec, [Canada]. [USNM]
note — Described from the unique female holotype, although the original description states that it is a male.
The name inquaesita is apparently derived from the Latin quaesitus, “sought”. This species somewhat resembles a diminutive P.speciosissima lacking white stigmata, but only abnormally small individuals of that much larger species could perhaps be confused with P.inquaesita. See additional details on these differences in the Papaipema speciossisma account. The stigmata of most specimens of P.inquaesita are orange, with white stigmata far less common. DNA barcodes are available for 19 specimens from Ontario, Maryland, and North Carolina, with minimum nearest neighbor divergence of 2.5%.
The larva belongs to the group where none of the longitudinal white lines cross the pigmented bands of the first four abdominal segments. The black lateral margins of the prothoracic shield extend as dark lines onto the head capsule. For a detailed account of the immature stages, see
Papaipema inquaesita is one of most abundant and commonly encountered Papaipema species, especially in the northern portions of its range, probably due to its nearly ubiquitous host fern. This species occupies most of the eastern United States as well as parts of southeastern Canada. It appears to be less common in the southern United States and has not yet been reported from Florida.
Papaipema stenocelis (Dyar) (Chain Fern Borer)
Hydroecia stenocelis Dyar, 1907, Journal of the New York Entomological Society, 15: 52.
Type locality: Baltimore, [Baltimore Co.], Maryland, USA. [USNM]
note — The name is derived from the Greek stenos, narrow, plus kelis, spot, referring to the peculiar shape of the reniform and orbicular stigmata. Described from the single female holotype.
Papaipema stenoscelis; authors, misspelling.
Papaipema stenocelis is a handsome species which is generally not common. It somewhat resembles a diminutive P.speciosissima but is unlikely to be confused with it. See additional details on the differences between these two in the Papaipema speciossisma account. Both species occur together on the Atlantic and Gulf Coastal Plains. Unlike P.speciosissima and P.inquaesita, the stigmata of P.stenocelis are apparently always white. DNA barcodes are available for four specimens of P.stenocelis from Florida and North Carolina, which form a discrete group and unique BIN at a minimum nearest-neighbor difference of 3.3%.
The larva, unlike those of the related P.speciosissima and P.inquaesita, features the pigmented bands of A1-A4 interrupted by a very narrow, mid-dorsal, white line. The black margins of the prothoracic shield do not extend onto the head capsule. Further details regarding the life history of P.stenocelis can be found in
The larval host plant is Virginia chainfern (Woodwardia virginica), but perhaps netted chainfern (W. areolata) is also sometimes utilized. The host fern colonies typically grow in wet habitats and even in standing water. Once a colony of this highly localized species has been located, it is very easy to identify an infestation, as the fronds initially bored are blackened but remain fully erect and are conspicuous among the surrounding green fronds. By this time, the larva is mining the extensive rhizome underwater, and its gallery may extend for one meter, making it challenging to successfully extract the larva, which is typically situated at the very end of its gallery. However, it often continues to utilize the dead frond’s original gallery for expelling frass, as this would be impossible under water without flooding the gallery. On that account, the larvae are all too frequently found to be parasitized, as they are constantly exposing themselves to attack. Often the orange frass may be observed floating on the water, or on the fibrous root mat when the water level is low. Fresh sections of the rhizome can be brought back, and the rearing process completed successfully in captivity. Pupation occurs in soil or detritus as near the abandoned rhizome gallery as possible, avoiding any standing water. The adults have been recorded flying from mid August to late October across much of the range, with a few anomalous very early records from Georgia and Florida in June and July.
Papaipema stenocelis is essentially confined to the Atlantic and Gulf Coastal Plains and shows little inclination to inhabit the interior, despite the widespread occurrence of its host fern there. It ranges from Maine to Louisiana, with an unexpected record from interior East Texas.
Papaipema rigida GROUP
Two very dissimilar-looking candidates for inclusion in this group, Papaipema rigida and P.eupatorii, share a number of attributes which unite them, including larval and genitalic morphology as well as genetic data. Molecular data further indicate a possible relationship of P.beeriana with P.eupatorii, but morphological features do not support this.
Papaipema rigida (Grote) (Rigid Sunflower Borer)
Gortyna rigida Grote, 1877, The Canadian Entomologist, 9: 87.
Type locality: “Penn.” [Pennsylvania, USA]. [NHML]
note — Described from a male from Pennsylvania and a female from Illinois. The “♂ type” in the NHML bearing Grote’s original handwritten, red-bordered label is the holotype.
The distinctive appearance of this species is somewhat reminiscent of the Palearctic Gortyna flavago (Schiffermüller) group of species. Papaipema rigida is unlikely to be confused with any other Papaipema with the possible exception of P.cataphracta, from which it is readily distinguished by the straight postmedial line on the forewings which gave P.rigida its name. A peculiar and prominent feature of P.rigida is the path of the antemedial line, which is pronounced and drawn into a sharp point which extends outward along the cubital vein almost to the orbicular. DNA barcodes are available for eight specimens of P.rigida from New Brunswick, Québec, and North Carolina, which form a discrete group and unique BIN at a minimum nearest-neighbor difference of 1.5%. The larva is quite similar in appearance to that of P.eupatorii, being pale and grub-like in the later instars. Although not apparent at this late stage, the dorsal and subdorsal lines do not cross the pigmented bands of A1–A4. The dark pigmentation of the lateral margins of the prothoracic shield does not extend to the head capsule. More details concerning the larval morphology are found in
Papaipema rigida is perhaps most commonly associated with smooth oxeye (Heliopsis helianthoides). It also favors thinleaf sunflower (Helianthus decapetalus).
This is a moderately common species ranging throughout the northeastern United States and southern Canada from New Brunswick to Manitoba and south to Virginia and Missouri, with more southerly extensions in the Appalachian Mountains.
Papaipema eupatorii (Lyman) (Joe-Pye-Weed Borer)
Gortyna eupatorii Lyman, 1905, The Canadian Entomologist, 37: 312.
Type locality: [Brownsburg, Québec, Canada]. [LEM]
note — Described from the female lectotype and two males (not a male and female as stated) from Montréal. No specific type locality is given, but a careful reading of the original description reveals that the type “No. 1 ♀” selected as lectotype by
Papaipema eupatorii has a distinctive forewing pattern featuring nearly parallel antemedial and postmedial lines with a fairly prominent, straight, medial shade midway between them. Like other members of the genus having dark forewings, it is cryptically patterned to match the appearance of the frosted, dried, late-season foliage of its host plant. This species is unlikely to be confused with any other Papaipema species due to the “striped” appearance created by the parallel forewing lines and associated dark and pale hues. DNA barcodes are available for eight specimens of P.eupatorii, originating from Québec to Georgia. Two other species, P.sciata and P.nepheleptena, share very similar barcodes, and all three are placed in the same BIN. The P.eupatorii nearest-neighbor divergences are 0.77% (P.sciata) and 0.78% (P.nepheleptena).
The larva of P.eupatorii is of the group wherein the dorsal and subdorsal lines do not cross the pigmented abdominal bands. The dark pigmentation of the lateral margins of the prothoracic shield do not extend onto the honey-colored head capsule. A more detailed account of the immature stages may be found in
As its name implies, P.eupatorii is a specialist borer of joe-pye-weed (Eutrochium), which was formerly included in the genus Eupatorium. Thus far, it has only been recorded boring in spotted joe-pye-weed (Eutrochium maculatum) and purple joe-pye-weed (Eutrochium purpureum). Whether it may also utilize species of the related genera is not known. The patches of joe-pye-weed which may harbor an infestation of this species are also very commonly found to contain P.cataphracta larvae boring in the stems of these plants. These latter produce copious amounts of expelled frass from their extensive galleries in the hollow stems, which is conspicuous enough to cause one to overlook the much more subtle evidence of P.eupatorii larvae working in the roots. Young larvae of P.eupatorii apparently enter the stem near its base, but, after working in the lower stem, mostly feed in the root. Little evidence of their presence other than some occasional expelled frass is readily observable, rendering this species among the more challenging to apprehend. Pupation occurs in a cell formed within the gallery at the base of the plant, and the adult emerges through an exit hole prepared by the larva. This is among the later-flying Papaipema species at any given locality, often not making its appearance before late September. It is a moderately common, widespread species, yet typically not seen in large numbers. It has been recorded throughout October into November.
Papaipema eupatorii is generally distributed throughout the eastern United States and southern Canada from Nova Scotia to Minnesota south to northern Georgia.
Papaipema nebris GROUP
This group includes Papaipema nebris, P.arctivorens, P.araliae, and P.polymniae. Larvae of all four are stem borers which notably proceed from feeding on the terminal meristem to lower portions of the stem but never enter the root.
Papaipema nebris (Guenée) (Stalk Borer)
Gortyna nebris Guenée, 1852, in Boisduval and Guenée, Histoire Naturelle des Insectes. Species Général des Lépidoptѐres, 5: 124.
Type locality: “Amérique Septentrionale, État des Illinois” [Illinois, USA]. [Types lost]
note — Described from at least two specimens. Guenée indicated both sexes were 38 mm in size. Although Guenée’s original types were apparently destroyed, the identities of both nebris and nitela are clear from his description, especially since he named nebris, nitela, and limpida in the same publication. Although apparently not authentic, the NHML “type” of nebris is correct for the species.
Gortyna nitela Guenée, 1852, in Boisduval and Guenée, Histoire Naturelle des Insectes. Species Général des Lépidoptѐres, 5: 124.
Type locality: “État des Illinois” [Illinois, USA] and “État de New-Yorck” [New York, USA]. [Types lost]
note — Described from one male from Illinois and one male from New York, based on the form lacking the white stigma. This taxon and nebris were described in the same publication, with nebris being listed first.
Gortyna nitella; Morris, 1860, misspelling. Papaipema nebris; Holland, 1903, not Guenée, 1852. Papaipema nitela; Holland, 1903, not Guenée, 1852.
The name nebris is derived from the Greek nebrias, spotted like a fawn skin. The only Papaipema species likely to be confused with P.nebris is the much less common P.limpida, as distinguished under that species. Both the maculate and immaculate phenotypes of P.nebris are equally common, yet in many populations one form may predominate, sometimes to the near total exclusion of the other. This phenomenon is usually unrelated to regional geographic distribution of the species but may reflect a local situation where one phenotype becomes increasingly rare until it disappears from a population, possibly through genetic drift. Plagiomimicus spumosum (Grote) in the subfamily Stiriinae could be confused with Papaipema nebris, but Plagiomimicus spumosum flies a bit earlier in the season, and the antemedial and postmedial lines are more angularly bent and less rounded near the costa compared to Papaipema nebris. The genitalia of these two are completely dissimilar. DNA barcodes are available for 23 specimens of P.nebris from eight states and provinces across the species’ range. Although the minimum nearest-neighbor divergence (to Papaipema arctivorens) is low at only 0.4%, barcode variation of P.nebris forms a discrete cluster within a BIN that is shared with P.arctivorens and the P.rutila group.
The larva is of the common type with only the middorsal longitudinal line uninterrupted by the pigmented banding of A1–A4. The dark pigmentation of the lateral margins of the prothoracic shield extends onto the head capsule, which is a bit browner compared to the yellowish prothoracic shield.
This familiar species has been widely documented in the economic literature, especially with regard to damage caused to seedling corn crops. Some of the damage to other cultivated plants blamed on nebris is at least occasionally likely caused by other Papaipema species which have overrun their preferred hosts. The majority of Papaipema species are visually indistinguishable in their larval stages from those of P.nebris upon cursory examination, and without rearing such larvae to adults for confirmation, many records of damage caused by nebris may be based solely on assumptions of identity. In nearly all instances, damage to garden plants can be mitigated by cutting or removing weedy hosts growing on the periphery of plantings. Perhaps as a result of the attention as an agricultural pest, the diversity of plant species recorded as P.nebris larval hosts is staggering, suggesting that the species may be truly omnivorous.
When boring in ragweed, larvae cause swollen galls to form on the stems. When the available food supply within the gall is exhausted, the larva may bore down the stem a short distance and cause a new gall to form. Sometimes several such galls may be observed on a single stem, often with each occupied by a larva. Frass ejected from the galleries may be observed on the soil beneath. Pupation occurs either within the larval gallery or sometimes in a cell formed just below the soil surface. For a highly detailed and comprehensive documentation of the biology of this species, see
This species has a wide distribution from Newfoundland to Manitoba, southwest to the foothills of the Rocky Mountains, and farther south to the Gulf of Mexico. Oddly, it has not yet been reported from Florida.
Papaipema arctivorens Hampson (Thistle Stem Borer)
Papaipema arctivorens Hampson, 1910, Catalogue of the Lepidoptera Phalaenae in the British Museum, 9: 73.
Type locality: Montréal, [Québec], Canada. [NHML]
note — Described from six males and five females.
Hydroecia rutila; authors, not Guenée, 1852. Papaipema artivorens; Muller, 1965, misspelling.
The name of this species refers to its association with the plant genus Arctium (Asteraceae), from which it had been commonly reared. It shares this same association with a number of other Papaipema species which also facultatively utilize this non-native host, likely due to the plant’s chemistry. Adults of P.arctivorens can be rather difficult to distinguish from P.harrisii, P.baptisiae, and P.rutila as evidenced by the mixed series of these species in most collections. For distinguishing features, refer to the discussion under P.rutila. The genitalic characters so useful in distinguishing many Papaipema are of relatively little value with these and some others, probably reflecting the more recent evolutionary origins of these species. DNA barcode variation of P.arctivorens based on 26 samples ranging from Manitoba to North Carolina showed discrete variation but are closely similar to P.nebris (0.37% minimum divergence) and the four species comprising the P.rutila group (1.1% minimum divergence). These three barcode groups comprise a single barcode BIN, suggesting they are closely related.
The larva of P.arctivorens belongs to the large subgroup of Papaipema characterized by the presence of a single, white, longitudinal, middorsal stripe interrupting the pigmented segments of A1–A4, and the absence of white subdorsal stripes crossing these same segments.
The strongly preferred native North American hosts of P.arctivorens are thistles, members of the genus Cirsium, also Asteraceae. It is very easy to detect P.arctivorens larvae in thistle plants by the presence of blackened leaves and frass at the tip of the stalk where the flower would have appeared, were it not for the destruction of the apical meristem. It is notable that P.arctivorens shares a behavioral trait with two other closely related species, P.araliae and P.polymniae, all three being terminal stem borers in their respective hosts – an uncommon characteristic among Papaipema. With these three, larval feeding commences with boring into the terminal bud of the stalk, and only proceeds down the stem as far as is required to complete development, never entering the base or roots of the host. Similar behavior is seen only in a distantly related species, P.furcata. The larva usually if not always vacates its gallery to pupate in soil nearby. This common species is one of the earliest Papaipema on the wing at any given locality, with a few adult captures recorded from the last week of July and extending throughout August into September, with stragglers as late as the third week of that month.
This species has a predominantly northeastern distribution, penetrating the southern United States mainly at higher elevations of the Appalachian Mountains, and extending westward across the Great Plains.
Papaipema araliae Bird and Jones (Hercules’ Club Borer)
Papaipema araliae Bird and Jones, 1921, The Canadian Entomologist, 53: 137.
Type locality: Sussex Co., Delaware, [USA]. [AMNH]
note — Described from five males and three females.
Papaipema araliae; McDunnough, 1938, attributed incorrectly to Bird.
In the midwestern part of its range, Papaipema araliae is most likely to be confused with P.rutila or P.polymniae, as all three are sympatric there. The distinguishing features are discussed in the treatment of P.polymniae. The Atlantic Coast population of P.araliae partially overlaps the distribution of P.sullivani, and there is sufficient superficial similarity to cause confusion. Aside from dramatic differences in the male genitalia of these two, P.araliae may be distinguished from the white-spotted phenotype of P.sullivani by the double postmedial line which is almost straight for most of its length in P.araliae but more sinuous and excurved around the reniform stigma in P.sullivani. All the reniform satellite spots in P.sullivani are white in this morph; in P.araliae only the more basal satellite spots are white, with the others tending to be orange. The coloration of the forewing medial area of P.sullivani is generally darker reddish orange; in P.araliae this area is more yellowish orange. Finally, where both species co-occur, P.araliae flies earlier with minimal overlap. The peak flight of P.sullivani occurs in mid to late October and continues into early November. DNA barcode samples were available from nine North Carolina specimens. The nearest-neighbor barcodes were those of P.harrisii, P.rubiginosa, P.pterisii, and P.hesseli, all differing by approximately 1.4%.
Larvae are of the predominant type in the genus having only the middorsal white longitudinal line crossing the heavily pigmented segments A1–A4. As is typical of all Papaipema, the earlier instars are more strongly and contrastingly pigmented than the final instar, which appears faded by comparison, sometimes to the extent that the pigmentation may be entirely lost. The dorsal pinacula are among the largest in the genus, likely reflecting the abrasive nature of the woody interior walls of the larval gallery. In general, the pinacula of stem borers tend to be larger than those of species in which the larval development primarily transpires within fleshy roots, tubers, or other softer plant tissues. For further details concerning the immature stages of this species, see
The sole host plant is Hercules’ club (Aralia spinosa), otherwise known as devil’s walking stick. Other species of Aralia are too small to accommodate the large larva. Larval infestations of the host, which attains the size of a small tree, are unmistakable even from a distance. Wilted brown terminal foliage is the first clue, and the presence of voluminous dark frass mixed with a gummy exudate produced by the damaged shoot confirms presence of the borer. Larval development is unusually rapid, and one is sometimes surprised to find vacant galleries as early as the end of June, by which time the larvae have matured, abandoned the shoots, and are aestivating in soil, presumably to avoid high summer temperatures and desiccation. The timing of this may vary and is dependent upon prevailing seasonal temperatures. There is a substantial discrepancy between the flight periods of the two disjunct metapopulations. On the Atlantic Coastal Plain, P.araliae flies almost a month later than the midwestern population, with peak abundance in late September and early October. In the Midwest, adult flights commence in mid-August and extend into early September.
This species occupies the middle third of the Atlantic Coastal Plain from Delaware to North Carolina, with highly disjunct populations in the central Mississippi and Ohio River valleys. This peculiar distribution of P.araliae reflects a fragmentation of the classical post-Pleistocene distribution pattern seen in other North American Lepidoptera (
Papaipema polymniae Bird (Leafcup Borer)
Papaipema polymniae Bird, 1917, The Canadian Entomologist, 49: 121.
Type locality: Roslyn, Va. [Virginia, USA]. [AMNH]
note — Described from five specimens.
In the early decades of this study, it was ELQ’s impression that P.polymniae and P.araliae might represent different color morphs of the same species, for the two are visually and morphologically indistinguishable except for consistent color differences of the forewings. The two had been routinely found flying together at the same times and places. After extensive field observations, it became apparent this was not the case. Many individuals have been reared from both host plants, and not a single specimen has ever been associated with the preferred host of the other. In places where only one of the host plants occur, only one species is ever observed. Barcode data indicate two closely related species. From P.araliae, this species is visually distinguished solely by the slightly greenish-yellow coloration of the medial area of the forewing, which replaces the much more orange coloration characteristic of P.araliae. This coloration of P.polymniae is reminiscent of that of P.cataphracta, but these two should not be confused otherwise. The stigmata of P.cataphracta are much larger, with a fully expanded reniform, and except in very rare individuals, almost never white. The reniform of P.polymniae is laterally constricted and bears white satellite spots, especially basally. A rare phenotype with dark stigmata is known (plate A8, figure 24). Perhaps the species most likely to be confused with P.polymniae is P.rutila, since they are broadly sympatric and fly at the same season. The coloration of the medial forewing area of P.rutila is about the same as in P.araliae and P.polymniae, but the stigmata of P.rutila, especially the orbicular group, are significantly larger than in either P.araliae or P.polymniae. DNA barcodes are available for two specimens of P.polymniae from Georgia, which are nested within, although distinct from, the P.harrisii- P.pterisii barcode haplotypes. The complex barcode variation of the entire group is discussed in more detail under P.harrisii.
The larva is practically indistinguishable from that of P.araliae, but has slightly smaller pinacula, and exhibits the usual white middorsal longitudinal line continuous across the deep-brown pigmented bands encircling the first four abdominal segments, the subdorsal lines being interrupted there. The mature larva finishes feeding upon tissues of the interior walls of the hollow stem and pupates in soil, following a possible brief period of aestivation there. See
This species is generally uncommon but widespread throughout the Austral Zone. Unlike the closely related P.araliae, P.polymniae occurs in both lowland and montane environments. It has been recorded from Maryland west to southern Illinois, south to northern Mississippi, and east to interior North Carolina, but is otherwise seemingly absent from the Atlantic Coastal Plain, with a disjunct population in peninsular Florida.
Papaipema rutila GROUP
Four closely related species are included: Papaipema rutila and P.baptisiae in eastern North America and two western North American species, P.limata and P.depictata, apparently long isolated from the common ancestor of the group.
Papaipema rutila (Guenée) (Mayapple Borer)
Gortyna rutila Guenée, 1852, in Boisduval and Guenée, Histoire Naturelle des Insectes. Species Général des Lépidoptѐres, 5: 123.
Type locality [neotype]: 4 miles SW of Albion, Area of Beadles Barrens NP N38.351218, -W088.126076, Edwards County, Illinois, USA. [CNC]
note — This is one of Guenée’s lost species discussed in the introduction. It was described from Illinois, apparently from a single specimen of 35 mm expanse, collected by Doubleday. It is impossible to ascertain from the original description to which of several similar species Guenée’s name might apply, and the putative type in the NHML appears spurious but is of ambiguous identity in any case. Much uncertainty has pervaded the earlier literature concerning the identity of rutila, and two additional taxa unnecessarily named on that account were synonymized by ELQ in his 1983 checklist. The identities of the similar species are not in doubt, so long-standing usage is hereby stabilized by designating a neotype for Guenée’s rutila. JRW, a resident of Illinois, has devoted considerable effort to documenting Doubleday’s 1837 activities in Illinois, locating the probable type locality, and securing the neotype specimen from that locality. The specimen, a male with forewing length of 17.8 mm [alar expanse 40.2 mm] is figured on
Papaipema merriccata Bird, 1907, The Canadian Entomologist, 39: 270.
Type locality: New Brighton, Pennsylvania, [USA]. [AMNH].
note — Described from 13 specimens including both sexes.
Papaipema placida Bird, 1921, The Canadian Entomologist, 53: 81.
Type locality: New Brighton, Pennsylvania, [USA]. [AMNH]
note — Described from a single male.
Papaipema merricata; Draudt, 1926, and authors, misspelling. Papaipema merrickata; McDunnough, 1938, unjustified emendation. Papaipema baptisiae; Forbes, 1954, in part, not Bird, 1902.
The name of this species comes from the Latin word rutilus, meaning golden red, an apt if not very diagnostic description. Adults of P.rutila have been and will continue to be confused with both P.arctivorens and P.baptisiae and might even be mistaken for P.harrisii. Until one becomes thoroughly familiar with all four, probably the most frequently misidentified Papaipema species in collections, correct identification is challenging. Papaipema rutila is an inhabitant of rich, moist woodlands where colonies of its host plant grow. Both P.arctivorens and P.baptisiae are more likely to be encountered in open habitats; the former in moist weedy meadows, the latter in old successional fields. Distribution is partially helpful in eliminating confusion of P.rutila with P.arctivorens, as P.rutila is confined to austral zone habitats and not found as far north as P.arctivorens; it also extends much farther south than P.arctivorens. Papaipema rutila averages slightly larger than P.baptisiae and P.arctivorens, but there is so much variability in size of individual specimens among all three species that size cannot be used reliably to separate them. From P.arctivorens, P.rutila may be distinguished by a slightly browner and more yellowish medial area of the forewings, less contrasting with the subterminal region which is generally darker and slightly narrower in P.arctivorens. The reniform stigma of P.rutila tends to be larger and more conspicuous than that of P.arctivorens, the outer medial lunule of which is usually concolorous with the medial area and not as contrasting white or yellowish as in P.rutila. The white orbicular spot of P.rutila is typically round; that of P.arctivorens is often triangular. The darker forewing scales of P.arctivorens are more reddish than in P.rutila, where they are browner. It is even more difficult to distinguish P.rutila and P.arctivorens from P.baptisiae. The forewings of P.baptisiae are a bit more reddish orange and the medial area is smoother and less powdered with overlying darker scales. The basal spots of P.baptisiae are orange; those of P.rutila and P.arctivorens, more yellow. The overall appearance of P.baptisiae is of a brighter, more colorful moth. The reniform stigma of P.baptisiae is typically larger and fuller than in P.rutila and far more than in P.arctivorens. The hindwings of P.baptisiae are more opalescent white than in either P.rutila or P.arctivorens. All three species can be easily separated from P.harrisii, due to its straight postmedial line. Neither male nor female genitalia offer any useful diagnostic characters. To fully appreciate the subtle differences in appearance between these species, one should rear series from larvae found in their respective host plants. The members of the P.rutila group exhibit barcodes with low inter- and intraspecific variation. In the case of P.rutila, some barcode haplotypes are identical to P.baptisiae. Six P.rutila samples have been barcoded to date, from Ontario, North Carolina, and Maryland; four of these are identical to one of the P.baptisiae haplotypes, the others are very similar (0.15-0.45% divergent).
The larva of P.rutila is typical of the group with the dark purplish brown cuticular pigmentation encircling segments A1–A4 interrupted by a single, middorsal, white, longitudinal line. For additional details concerning the immature stages of P.rutila, see the original work of Henry
This species is an obligate specialist borer in mayapple (Podophyllum peltatum). Larval feeding is readily detected early in the season by the usual wilting and yellowing of the host plant, accompanied by a hole in the stem and white frass on the ground. The window of opportunity for locating the larva, however, is short. Mayapple plants grow in expansive colonies which begin to senesce shortly after flowering, by which time the borers are already feeding underground and become increasingly difficult to detect as the season progresses. If one has previously established the presence of P.rutila and memorized the exact site of the mayapple colony, seeking larvae closer to maturity sometimes is fruitful, as light tan or orange frass may be detected on the soil surface among the detritus of dead and decaying vegetation. The pupa is formed in a loose cell among detritus at or just below the soil line. The adults fly from the end of August through mid-October. The species is widespread throughout the Austral Zone from New Jersey west to Nebraska and south to Louisiana and Georgia.
It has been a matter of considerable interest and importance to attempt to identify where Doubleday collected the type material of P.rutila that Guenée described. The authors are grateful for JRW’s efforts to reconstruct Doubleday’s activities in Illinois and attempt to locate the exact locality where the specimens originated. By studying a number of contemporary journals with Doubleday’s writings in them, JRW was able to ascertain that Doubleday visited Illinois only once, in late October of 1837. Doubleday came to Illinois through “Shawnee Town” on the Ohio River on 23 October 1837, made it quickly to Carmi, Graysville [Bon Pas, White County], and from there to Albion. About two miles west of there was the village of Wanborough, long since nonexistent, in Edwards County. Doubleday mentioned “staying here about a week” at the residence of Deborah Prichard. He further mentioned that “The Noctuae taken at Wanborough were all, or nearly all, caught at the lights.” Doubleday repeatedly talked of lighting up rooms [windows] at houses or inns for moths. It must have been fairly warm that year to enable him to collect several Papaipema species in the final week of October.
Papaipema baptisiae (Bird) (Dogbane Stem Borer)
Hydroecia baptisiae Bird, 1902, The Canadian Entomologist, 34(5): 109.
Type locality: vicinity Providence, [Providence Co.], Rhode Island, [USA]. [AMNH]
note — Described from multiple examples, the number not specified. The type locality was not exactly specified other than that the larvae were collected in the vicinity of Providence, Rhode Island but may actually have been found in Massachusetts, since Bird found the baptisiae larvae while attempting to locate Papaipema speciosissima at its type locality (Seekonk). See discussion of this matter under speciosissima.
Hydroecia circumlucens; Bird, 1900, not Smith, 1899.
This species is deceptively similar in appearance to two others with which P.baptisiae is broadly sympatric, P.rutila and P.arctivorens. These three species are almost invariably mixed in most collections featuring large series. For distinguishing characteristics, see the diagnostics given in the discussion of P.rutila. Genitalic characters do not provide reliable means of identification. Forty-five samples from across the range of P.baptisiae were closely similar to and intercalated with barcodes of other species in the P.rutila group (P.rutila, P.depictata, and P.limata), as discussed under P.rutila.
The larvae are visually inseparable from other members of their species group, yet still readily identified by their association with the preferred hosts and their conspicuous feeding habits. Papaipema baptisiae was named for the host plant in which its larva was first encountered but which represents more of a secondary food choice than the rule. Far more frequently than found boring in stems of wild indigo (Baptisia spp.), the larvae exhibit a strong preference for feeding in the unrelated host dogbane (Apocynum) across the entire range of the insect. On occasion, larvae of P.baptisiae have also been found boring in stems of a wide variety of other plants including goldenbanner (Thermopsis), meadow-rue (Thalictrum), rattlesnake-master (Eryngium), Indian plantain (Arnoglossum), burdock (Arctium), ironweed (Vernonia), and others. In most instances, these latter hosts represent less of a deliberate choice than a case of colonies having exhausted and overrun their preferred host resources and taken up boring in other suitable plants growing among them. Apocynaceae may well represent the primary ancestral host plant family of the entire P.rutila group.
The presence of P.baptisiae larvae in a patch of dogbane can be detected from a distance, due to the sharp contrast between healthy plants and bored ones. Plants occupied by a larva (never more than one larva per stem) show yellow, wilted foliage or even brown, dead foliage depending upon the date. A conspicuous hole in the stem near the base, often with the plant’s dried milky exudate dripping from it, accompanied by a pile of orange frass on the soil confirms the reason for the plant’s poor condition. Often by the time larval development nears completion, the stem may also be bent or broken, further attracting one’s attention. The pupa is sometimes formed at the bottom of the gallery but perhaps more frequently in soil around the plant’s base. Adults emerge from mid-August through mid-October, but with peak flight mainly in September.
The widespread abundance of this common species is perhaps attributable, at least in the easternmost portions of its range, to the proliferation of old fields resulting from the ever-increasing abandonment of agriculture in deference to the giant farms of the Midwest. Such habitats temporarily favor abundant growth of the primary host plant for a period during which the gradual transition of the land from cultivation back to forest – or development – provides an opportunity for large colonies of the moth to persist in the interim. ELQ has witnessed such colonies flourish for a couple of decades – only to die out when shrubs and young trees replace the forbs – and reappear nearby at habitats in earlier stages of succession. In the Midwest, baptisiae is commonly a denizen of native prairie remnants, probably reflecting the original environment in which the species evolved.
The species is predominantly northeastern, distributed from Maine to Iowa and south, but apparently not present along the Atlantic Coastal Plain south of Virginia, and entirely absent from the Gulf Coast.
Papaipema limata Bird (Western Dogbane Borer)
PL. A-7, FIGS 24–27;
Papaipema limata Bird, 1908, The Canadian Entomologist, 40(1): 30.
Type locality: Washington Experiment Station, Pullman, [Whitman Co.], Washington, [USA]. [AMNH]
note — Described from a unique male in the J. B. Smith Collection. The original description states that the specimen was collected “Sept. 25 th, 1898” but the holotype is labelled “15 Sept. 98.” In his 1983 checklist, ELQ treated limata as a subspecies of baptisiae but since became convinced that it is a closely related species, based on barcode data and other considerations. Its status as a distinct species was reinstated in
Papaipema ochroptena; Barnes and McDunnough, 1912, in part, not Dyar, 1908. Papaipema circumlucens vaha Benjamin, 1933, Pan-Pacific Entomologist, 9(3): 101.
Type locality: Provo [Utah Co.], Utah, [USA]. [USNM]
note — Described from four specimens: a male holotype and female paratype from Provo, a female from Richfield, Utah, and a male from Denver, Colorado which probably is not the same species, but that specimen was not located. Although described as a subspecies of circumlucens based on Benjamin’s misconception of the identity of that species, there is little similarity to circumlucens, and vaha is clearly identical to limata.
† Papaipema circumlucens form ochroptenoides; Bird, 1944, Journal of the New York Entomological Society, 52: 198.
note —Unavailable name; attributed to Benjamin by Bird (1944), but this appears to be a nomen nudum and an infrasubspecific name, and so is unavailable.
Although its wing pattern elements are virtually identical to those of its eastern counterpart P.baptisiae, to which it is perhaps most closely related, P.limata is allopatric in distribution. It has lighter yellow forewings together with strong pinkish tints. Bird’s name is derived from the Latin limatus, signifying polished, or refined.
Papaipema limata is most likely to be confused with P.depictata, a related western species of more southerly distribution. Populations of both species are largely allopatric, but there is some potential for overlap. Perhaps because both share common ancestral traits, occasional individuals of P.limata such as that figured on plate A-7, figure 27 may strongly resemble the forewing coloration more typical of P.depictata, having their antemedial and subterminal spaces filled with darker purplish fuscous, and with a dark fuscous bar connecting the orbicular with the base of the reniform. A consistent character that can be used to separate the two is the coloration of the reniform stigma. In P.limata, the reniform stigma is all white, similar to that of P.baptisiae. In P.depictata, the reniform stigma is slightly less prominent, and the central outer satellite spots are yellow rather than white, similar to those of P.rutila. Also likely to be confused is the somewhat browner P.insulidens with which P.limata is broadly sympatric. Papaipema insulidens features contrasting, paler, whitish-yellow basal forewing spots which are characteristic of the group to which that species belongs, compared to the relatively concolorous, less contrasting basal spots of P.limata. DNA barcodes are available for two samples from Oregon, which although distinct from others in the P.rutila group, are very close to a subset of the P.depictata barcode complement at 0.8%.
The larva of P.limata is a member of the large group of species having only the middorsal white longitudinal line continuous across the otherwise unbroken purplish-brown pigmented bands encircling the first four abdominal segments. It is superficially indistinguishable from the other three members of the rutila group.
AEM and JRW found larvae in Oregon boring in dogbane. AEM found larvae in northern Utah using this same host plant. An unconfirmed report of geranium is based on larvae found at Pullman, Washington. Papaipema limata may be a more general feeder like most other Papaipema species in western North America. Symptoms of larval boring in dogbane are similar to those reported for P.baptisiae. Collection records indicate a flight season extending from mid-August to early October across its range.
This colorful species reaches its northern limit in eastern Washington and extends south throughout the Great Basin. Its distribution southward appears to terminate at the southern boundary of the Great Basin Desert in Utah and California. The species has not been recorded from Nevada, perhaps due to insufficient sampling. It has been collected between 275 and 1,700 meters.
Papaipema depictata Benjamin, REVISED STATUS (Owl’s-claws Borer)
PL. A-7, FIGS 28–29 (adult);
Papaipema depictata Benjamin, 1933, Pan-Pacific Entomologist, 9(3): 101.
Type locality: White Mountains, [Apache Co.], Arizona, [USA]. [USNM]
note — Originally described from one male and one female. In his 1983 checklist, ELQ synonymized this taxon with Papaipema baptisiae (Bird), due to the apparent lack of significant distinguishing features in the genitalia of either sex or in adult maculation. With more study material currently available, including barcode data and details of the life history, it is necessary to reinstate its status as a distinct species.
This species has been collected only sporadically over the past one hundred years. Its absence from many collections probably reflects its habitat requirements more than its actual abundance. A moth frequently collected in the southwestern United States and regularly misidentified as a Papaipema is the eriopygine noctuid Hydroeciodes serrata (Grote), which is illustrated on plate A8, figure 12. It is almost the same size as Papaipema depictata and occurs in the same broad general area, if not the same habitat. The elaborate forewing patterns and coloration of both species are nearly identical, but these unrelated moths are structurally quite different. Males of H.serrata (and related species) have pectinate antennae, both sexes have hairy eyes, and the genitalia are completely dissimilar to those of Papaipema. Papaipema depictata could be confused with three other Papaipema species but is largely or completely allopatric with regard to all of them. It is somewhat smaller, less reddish, and with less prominent stigmata than P.baptisiae, which approaches from the East at lower elevations. It is more likely to be confused with P.limata and P.insulidens. See diagnoses provided with those species. DNA barcodes of seven P.depictata specimens from Arizona and New Mexico are available. Although no haplotypes are shared with any other species in the P.rutila group, all four species have very low barcode divergence. The minimum divergence between P.depictata and P.baptisiae is 0.15%.
The larvae are visually indistinguishable from other members of the group having only the single middorsal stripe crossing A1–A4. David Wagner (pers. comm.) reports finding the larvae boring both in owl’s-claws (Hymenoxys hoopesii) and Chiricahua Mountain larkspur (Delphinium andesicola) in the Chiricahua Mountains, as confirmed by other recent collections. This suggests the species may be oligophagous like other western Papaipema. Dogbane is abundant at the type locality in the White Mountains and remains another potential host. AEM reports that larvae can be found boring in the lower stems and roots of owl’s-claws, especially in plants of small stature, as well as higher in the stems of larger plants, and can cause wilting of the plants. The frass is light tan. Larval boring also causes wilting of stems in larkspur. Available records indicate an adult flight period extending from mid-August to early October.
This moth is a denizen of alpine meadows associated with pine/spruce/fir forests distributed across the mountainous regions of at least four southwestern states, and in the oak/juniper woodlands of the Davis Mountains of Texas, where all these populations have been isolated by perhaps several hundred thousand years. Documented occurrences are mostly from collections made above 2,000 meters and as high as 3,000 meters. Collecting at these elevations late in the season is often unproductive due to low nighttime temperatures. This species appears to be mainly limited to the aforementioned montane habitats in Colorado, Utah, Arizona, New Mexico, and the Davis Mountains of west Texas.
The remarkable resemblance between Papaipema (including P.limata) and the completely unrelated Hydroeciodes serrata could be explained simply by the genetic expression of ancestral noctuid traits conserved across independent lineages, but there may be an alternate and more compelling explanation. It could be a case of Batesian mimicry. It has long been known that adults of certain genera among the arctiine Erebidae which feed in the larval stages upon dogbane derive chemical protection from toxins in the plants, so it is quite plausible that a Papaipema associated with a group that favors dogbane might also. The highly colorful forewing patterns of some Papaipema species may actually be cases of aposematism, whereas the more subtle colors and patterns of others perhaps serve better as camouflage blending with autumn coloration. For more on this subject, see the Papaipema amianthii account, as well as the section on evolution of endophagy in the introduction to this fascicle.
Papaipema duplicata GROUP
Two closely related species, Papaipema duplicata and P.silphii, comprise this group.
Papaipema duplicata Bird (Stoneroot Borer)
Papaipema duplicatus Bird, 1908, The Canadian Entomologist, 40: 25.
Type locality: Rye, [Westchester Co., New York, USA]. [AMNH]
note —Described from seventeen specimens. After
† Papaipema duplicata ab. obsolescens Strand, 1915 [1916] Archiv für Naturgeschichte, 81A (11): 163.
Type locality: [Rye, Westchester Co.,] New York, [USA]. [NHML]
note—Described as an aberration of duplicata from a female illustrated in
Three other Papaipema species, P.silphii, P.necopina, and P.maritima, are sufficiently similar in appearance to cause confusion with P.duplicata. If not for differences in size, habitat association, host plant utilization, and largely allopatric distribution, P.duplicata is perhaps most likely to be confused with P.silphii. These two species are closely related, as suggested both by their superficial appearance and barcode data. No significant differences between P.duplicata and P.silphii are apparent in the genitalia. From P.necopina, P.duplicata is best distinguished by having a straight postmedial line, a lighter, more contrasting subterminal area, and darker hindwings. The subterminal area of P.maritima does not contrast with its medial area, unlike that of P.duplicata. All three species might occur in general proximity to each other.
Eight DNA barcode samples of P.duplicata from Pennsylvania and Virginia are available. The barcodes of P.duplicata are part of a large, invariant cluster that offers little or no taxonomic resolution. The most similar barcodes to P.duplicata belong to P.silphii, P.nelita, P.sablensis, P.impecuniosa, and even some haplotypes of P.harrisii. Most of these species can share identical barcodes, and total variation is less than 0.5%. Perhaps some or all of these species share a recent ancestry, but other genetic markers will be needed to elucidate this.
The larva is of the common type bearing only the middorsal white line continuing across the first four dark-pigmented abdominal segments. The dark-pigmented lateral margins of the prothoracic shield extend onto the head capsule, which is somewhat darker brown than the honey-colored cervical shield. Additional details concerning the larval morphology may be found in
The primary host is stoneroot (Collinsonia canadensis), although while collecting Papaipema polymniae larvae in Georgia from hairy leafcup, (Smallanthus uvedalius), AEM discovered that some of the larvae turned out to be P.duplicata. Adult specimens of P.duplicata have also been collected in Kentucky by Loran Gibson at light traps in areas where stoneroot had not been noted but where leafcup was abundant, suggesting this anomalous departure from the usual host plant may not be unique. In the mountains of Bath County, Virginia while searching for larvae of Hydraecia stramentosa in lanceleaf figwort (Scrophularia lanceolata), AEM found several larvae of P.duplicata in the lower stems of this plant. In this case, since the figwort was growing in an area where stoneroot was abundant, it is likely that the larvae had overrun the stoneroot and had bored into the figwort opportunistically.
In the early instars, the larva bores a gallery formed in the stem several centimeters above the ground and proceeds to the flattened, woody root where it remains for the duration of its development. Pupation occurs in a hollow chamber inside the root. One may be alerted to the presence of P.duplicata larvae in a colony of the host plants by wilted or drying leaves and often the presence of whitish frass on soil under the original gallery hole in the stem, which the larva apparently continues to utilize for the purpose of waste disposal.
This species generally has been regarded as uncommon or even rare, but at some locations in the southern Appalachian Mountains it can be surprisingly abundant. An inhabitant of the rich, damp woodlands its host plant requires, P.duplicata has become nearly extirpated throughout parts of its range, especially in the northeastern United States where white-tailed deer populations are uncontrolled by predators, and where these deer have decimated native understory vegetation by over browsing (
This species has a somewhat restricted distribution in comparison to many of its congeners. From southeastern New York, it extends down the Appalachian Mountains as far as Georgia. It also shows a preference for the rich lowland forests of the Ohio River Valley system.
Papaipema silphii Bird (Prairie Rosinweed Borer)
Papaipema silphii Bird, 1915, The Canadian Entomologist, 47: 110.
Type locality: Chicago, [Cook Co.], Illinois, [USA]. [AMNH]
note—Described from twenty-five specimens, including the male type, all from the suburban environs of Chicago.
Although P.silphii is the largest species among the small group of similar-looking Papaipema with which it is at least partially sympatric, it may be distinguished from P.duplicata, P.necopina, and P.maritima by its warmer brown forewings, larger size, and specifically from P.duplicata, by far less contrast between the antemedial, medial, and postmedial areas of the forewings. Subtle differences in the shapes of the cucullus of the male genitalia of P.necopina and P.maritima can also be diagnostic. It is less likely that P.silphii will be confused with the immaculate phenotype of P.beeriana with which it also occurs. This latter features a postmedial line that is much closer to the terminal area of the forewing than that of P.silphii and, like the three aforementioned species, is gray-brown rather than warm brown. DNA barcodes are available for three samples of P.silphii, closely similar to P.duplicata and discussed further under that species.
The larva of P.silphii belongs among the commonest group, those bearing only the single, middorsal, white line across the purplish-brown pigmented first four abdominal segments. The honey-colored prothoracic shield bears the usual dark-pigmented lateral margins which may or may not extend onto the reddish-brown head capsule.
The usual larval host plants include prairie rosinweed (Silphium terebinthinaceum), compassplant (S.laciniatum), and less commonly, cup plant (S. perfoliatum). Strays have been recorded from burdock (Arctium), sunflower (Helianthus), and Indian plantain (Arnoglossum). Presence of larvae may be readily detected by the dried, brown foliage of the stalk initially bored by early instars amid the clump of healthy green leaves. The young larvae soon enter the roots, where most of their development occurs. This prairie endemic is readily attracted to various light sources which is fortunate for the collector, since digging up the host plant with its immense woody root stock is no small effort if one is hoping to secure larvae to rear. However, several larvae may be found in a single mature plant distributed throughout its extensive root system. Larvae are most easily sought during late May before they bore deep into the roots.
Papaipema silphii is restricted to prairie habitats throughout the upper Midwest, ranging from northern Ohio to southern Michigan, Wisconsin, and Iowa and south to Missouri, Illinois, and Kentucky.
Papaipema cerussata GROUP
Three species are included: Papaipema cerussata, P.sciata, and P.nepheleptena. These three species share larval and genitalic similarities.
Papaipema cerussata (Grote) (Ironweed Stem Borer)
Gortyna cerussata Grote, 1864, Proceedings of the Entomological Society of Philadelphia, 2(4): 431.
Type locality: “Pennsylvania” [USA]. [ANSP]
note—Described from a single male.
Hydroecia cerrussata; Smith, 1899, misspelling. Papaipema cerussata; Dyar, 1903, misattributed to G. & R. Papaipema cerrusata; authors, misspelling.
The name is derived from the Latin cerussa, white lead, presumably in reference to the characteristic bluish-gray, metallic leaden coloration of the subterminal and terminal areas of the adult forewings. This large, handsome species is unlikely to be confused with any other with the possible exception of the smaller P.sciata, from which it most obviously differs by the presence of the yellowish basal spots of the adult forewings, which are absent in P.sciata. A rare phenotype of P.cerussata (
Neither the middorsal nor subdorsal longitudinal white lines cross the pigmented bands of the first four abdominal segments in this species. The honey-colored prothoracic shield features dark-pigmented lateral margins which extend as a black line onto the head capsule. The anal plate, which is commonly of similar color and degree of sclerotization to the prothoracic shield in many other Papaipema, is nearly black in P.cerussata and very heavily rugose, probably an adaptation to development in the rough, woody roots of ironweed. For a more thoroughly detailed account of the early stages, see
The larva is a specialist borer of various species of ironweed (Vernonia spp.), but strays have been recorded from a variety of other asteraceous plants growing in association with these ironweeds, including joe-pye-weed (Eutrochium), Indian plantain (Arnoglossum), rosinweed (Silphium), and even wild indigo (Baptisia). JRW (pers. comm.) relates that both P.cerussata and P.limpida larvae may be found together in the same large cluster of ironweeds. Papaipema limpida enters the stem at or just above ground level. Once inside, young larvae soon enter the root and eventually bore deep within it. In contrast, P.cerussata enters the stem from ten to twenty-five or more centimeters above the ground and bores about a third of a meter of the stem, only eventually entering the root and boring deeply into it. Damage of P.cerussata may be seen from a distance, since boring in the terminal meristem of the plant by the early instars subsequently causes a “witches broom” effect in the top of the plant (
Papaipema cerussata ranges from southern New England to Iowa and Kansas south to Mississippi, but apparently is absent on the southern Atlantic and Gulf Coastal Plains.
Papaipema sciata Bird (Culver’s Root Borer)
Papaipema sciata Bird, 1908, The Canadian Entomologist, 40: 28.
Type locality: Rye, [Westchester Co.], New York, [USA]. [AMNH]
note—Described from fourteen specimens including both sexes.
Hydroecia limpida; Bird, 1902, not Guenée, 1852. Papaipema limpida; Hampson, 1910, part, not Guenée, 1852.
The name is apparently derived from the Greek skia, shaded, probably in reference to the bluish-gray shading of the antemedial and terminal portions of the otherwise dark brown forewing. This handsome species is broadly sympatric with its close relative P.cerussata but is much less common. It could only be confused with the similarly colored P.cerussata, from which it is easily distinguished by its smaller size and the total absence of visible basal spots on the adult forewings. The digitus of the male genitalia of P.sciata is serrate on its dorsal edge; that of P.cerussata, smooth. DNA barcodes are available for four specimens of P.sciata from Wisconsin, Illinois, and New York. All haplotypes are very similar or identical to those of P.nepheleptena, between 0 and 0.5% divergent.
The larva is another of those having the first four darkly pigmented abdominal segments unbroken by the longitudinal white dorsal and subdorsal lines. The honey-colored prothoracic shield bears the usual dark lateral margins, which are not continued as a dark line onto the head capsule. The anal plate is also of the usual honey color, unlike the much darker and heavily sclerotized plate of P.cerussata. For a more detailed account of the immature stages, the reader is referred to
So far as is known, P.sciata is an exclusive specialist borer of Culver’s root (Veronicastrum virginicum). The young larvae enter the stalk of the host plant near its base and soon establish a gallery that leads to the roots, which are mined extensively and where most of the larval development occurs. This generally results in the bored stem and leaves withering and turning dark brown, which makes it easy to identify infested plants which, however, are usually relatively few and far between in their occurrence. When mature, the larvae exit the bored roots and pupate in soil nearby. This species is generally on the wing later than many congeners, although specimens have been captured throughout September and into late October, with a few records extending well into November.
Papaipema sciata is primarily a prairie endemic distributed in the upper Midwest from Ohio and southeastern-most Ontario to Minnesota and south to Missouri and Illinois. There is also a highly disjunct, mainly historic population which occurred in Connecticut, southeastern New York, northern New Jersey and eastern Pennsylvania. It probably reflects the remnants of a post-Pleistocene distribution wherein the isolated eastern prairie elements have become greatly diminished in the last century due to succession to forest habitats.
Papaipema nepheleptena (Dyar) (Turtlehead Borer)
Gortyna nepheleptena Dyar, 1908, The Canadian Entomologist, 40: 77.
Type locality: “New York” [USA]. [USNM]
note—Described from the unique female holotype, with its exact locality unknown. The name is derived from the Greek nephele, cloudlike spot, and ptena, winged, likely referring to the appearance of the conspicuous white reniform stigma on the forewing.
Papaipema moeseri Bird, 1911, The Canadian Entomologist, 43: 42.
Type locality: Staten Island, [Richmond Co.], [New York City], New York, [USA]. [AMNH]
note—Described from eighteen specimens from the vicinity of Buffalo, N.Y. (West Seneca), Staten Island, N.Y., and Montréal, Québec. The male “type” is from Staten Island, N.Y.
Despite exhibiting little similarity of appearance in the coloration and maculation of the adult forewings, this species appears to be closely related to P.sciata and P.cerussata, based on larval characters, genitalia, and molecular data. This charming little species features a laterally compressed, figure 8-shaped reniform stigma which is disproportionately large in relation to the overall surface area of its forewings, and which is among its most easily recognizable diagnostic features. The only species with which P.nepheleptena is liable to be confused is the normally larger P.birdi. If uncertain as to identity, one can brush the scales off the tip of the male abdomen which will reveal the trigonate cucullus with its bird’s-head shape and relatively short ventral arm characteristic of P.nepheleptena, P.sciata, and P.cerussata. In contrast, the cucullus of P.birdi is strongly bifurcate with widely divergent subequal arms. DNA barcodes are available for six specimens of P.nepheleptena from Ontario, Québec, and North Carolina, intercalated with haplotypes of P.sciata, with at least one shared haplotype. Both are close to eupatorii, as discussed under that species.
The larva is among those with the first four heavily pigmented abdominal segments uninterrupted by the usual longitudinal white lines. The honey-colored prothoracic shield bears the usual dark lateral margins which are not continued as dark lines onto the head capsule. It is visually indistinguishable from the larva of the related P.sciata.
The only larval host recorded thus far from the eastern portions of the range of P.nepheleptena is white turtlehead (Chelone glabra). JRW (pers. comm.) reports that in the Midwest, the larva is commonly found boring in swamp verbena (Verbena hastata). Whether this reflects the only host used in the Midwest or simply collecting bias remains unknown. Once a person becomes familiar with the local occurrences of Papaipema larvae boring in their favorite hosts, additional searches tend to focus on the same host plants which have already proven productive, sometimes ignoring other possibilities. AEM has not found P.nepheleptena boring in verbena in the eastern United States despite repeated checks of this plant in localities where the larva bores in white turtlehead.
Larvae of P.nepheleptena initially bore a considerable length of the host plant’s stem before entering the root to complete their development. A detailed account of the early stages, under the name moeseri, is provided in
This species ranges from Nova Scotia to Minnesota and south to Nebraska, Kentucky, and North Carolina. Although widespread, it is not among the most commonly encountered Papaipema species.
Papaipema circumlucens GROUP
Despite dramatic differences in size, Papaipema circumlucens and P.blackorum are apparent sister species, based on nearly identical genitalic features in both sexes and distinctive larval habits as well as larval morphology. Both species are specialized borers of climbing woody vines and produce cylindrical galls on the vine stems. The forewing patterns have similar elements, and both species have unusually broad forewings relative to most other species in the genus.
Papaipema circumlucens (Smith) (Hop-gall Borer)
Hydroecia circumlucens Smith, 1899, Transactions of the American Entomological Society, 26: 43.
Type locality [lectotype]: “N. Jersey” [New Jersey, USA]. [USNM]
note—Confusion regarding the correct application of this taxon has reigned ever since Smith’s description more than 120 years ago. The initial problem was that Smith had two species mixed in his type series consisting of ten specimens. Seven of these syntypes are still extant, the other three are either lost or perhaps labelled in a manner as to be unrecognizable. Of these seven, five unambiguously agree in every detail with Smith’s description of both adult maculation and male genitalic characters, and the right valve from one male is illustrated on
Gortyna ochroptena Dyar, 1908, The Canadian Entomologist, 40: 77.
Type locality: Denver, Colorado, [USA]. [USNM]
note—Described from a single male specimen.
Gortyna marginidens; Dyar, 1908, not Guenée, 1852. Papaipema humuli Bird, 1915, The Canadian Entomologist, 47: 113.
Type locality: Rye, New York, [USA]. [AMNH]
note—Bird unnecessarily described this taxon from 30 reared specimens, because of his mistaken idea that Smith’s name applied to the dogbane borer which Bird had previously named baptisiae.
Papaipema ochropenta; Bird, 1916, misspelling.
This species is unlikely to be confused with any other. All Papaipema with similar general coloration have a contrasting cluster of basal spots, which P.circumlucens lacks. The species is somewhat variable in ground color, with some individuals exhibiting a reddish hue. Four DNA barcode samples from Michigan, Saskatchewan, and Illinois group within the large barcode BIN that includes the complex genetic variation of P.harrisii-P.pterisii, although forming a distinct subgroup therein. The nearest-neighbor barcodes are some haplotypes of P.pterisii, differing by a minimum of 0.5%.
The larva is a typical Papaipema of the group which bears only a single longitudinal middorsal white stripe crossing the dark purplish-brown pigmented band encircling the first four abdominal segments.
The sole known larval host plant is wild hops (Humulus lupulus) (
Larval development is confined to a surprisingly small section of the hop vine, where a slender, cylindrical gall up to ten centimeters long eventually forms, increasing in length and diameter commensurate with larval growth. Galls are often located at points where vine stems twist around each other, or where a vine wraps around fence wire or twigs. A small hole near the base of the gall serves for discarding larval frass and prevents any accumulation of rainwater, and a larger hole is ultimately chewed at the opposite end of the gall to allow for the mature larva to exit the gall, whereupon it descends the vine and pupates under a shallow layer of soil or detritus. Larval development is rapid, and galls are generally vacant by mid to late July. Accordingly, the adult moth is among the earliest species of the genus to emerge. The flight period commences around mid-August and continues for about ten days to two weeks.
Papaipema circumlucens was formerly more abundant and widespread than it is today. It appears to be entirely extirpated from the Atlantic portions of its range. Today this species occurs from western Pennsylvania to Saskatchewan, and southwest as far as Arizona, along margins of woodlands and fence rows where wild hops may be found climbing upon other vegetation. Should the seemingly disjunct montane southwestern populations be found to be genetically distinct, the name ochroptena Dyar is available for them.
Papaipema blackorum Quinter, NEW SPECIES (Pipevine Borer)
Papaipema blackorum Quinter.
Type locality: Johnson creek, 37.3871, -88.7779, S of Rt. 146 on Reevesville Rd., Johnson Co., Illinois, USA. [CNC]
note—Named to honor our dear friend, the late Captain William R. Black, Jr. of Paducah, Kentucky, for his outstanding dedication and persistence in documenting the occurrences of this moth, and also in recognition of his wife Nancy for her steadfast support of his herculean efforts and her gracious hospitality to all who were involved in that endeavor.
This is the largest member of the genus, having very broad forewings in addition to a wide wingspan. Occasional very large individuals of Papaipema cerussata, P.marginidens, P.matheri, or P.speciosissima approach but never exceed the forewing length of the largest individuals of this moth, which may reach 28 mm. Papaipema blackorum is remarkably uniform in both size and appearance, exhibiting little phenotypic variability. It is unlikely that P.blackorum could be confused with anything else, on account of its size and limited distribution. Some individuals of Papaipema unimoda, especially those from western portions of its distribution, although much smaller, are tawny brown similar to P.blackorum, but distributions of these two species barely overlap and genitalia are totally dissimilar.
Head: antennae of both sexes ciliate, laminate, less deeply in female; dorsum scaled; ventral lamination obscured by densely pubescent sensilla; eye smooth, rounded; haustellum unreduced; labial palpus laterally flattened, upcurved; first segment swollen basally, arching slightly upward, slightly more than half as long as second segment; second segment straight, about as long as eye width, scales broad; third segment ⅓ × length of second segment, and projecting slightly anteriorly, scales narrow; frons unmodified, smooth, convex, with unspecialized conical vestiture consisting of spatulate scales mixed with fine, hairlike scales. Thorax: vestiture of prothoracic collar, patagia, mesothorax, and metathorax concolorous, a mixture of coarse spatulate, thin apically spatulate, and hairlike scales; transverse, wedge-shaped, prothoracic crest prominent, mesoscutellar tuft moderate, metascutellar tuft subequal. Forewing: length similar in both sexes = 20–28 mm; dorsal surface ground in both sexes uniform wood brown; basal spots obscure; antemedial and postmedial lines concolorous with ground; antemedial line double, nearly straight until strongly bowed outward just before meeting inner margin; postmedial line double, scalloped, excurved around reniform then nearly straight, meeting inner margin diagonally; medial shade darker fuscous, touching lower loop of reniform, then angled below reniform and meeting inner margin nearly perpendicularly; subterminal line preceded by series of darker, charcoal chevrons filling spaces between veins, immediately followed by series of lighter, yellowish-brown spots, more pronounced apically; subterminal space filled with gray scales; terminal space fuscous; terminal line obscure; fringe dark fuscous; reniform very narrow and laterally compressed, with small, white, satellite spots surrounding thin, yellow, central lunule; orbicular and suborbicular stigmata very small, incorporating white scales but tending toward obsolescence in some. Hindwing: dorsal surface uniform dull fuscous; discal spot and medial shade obscure in most; subterminal area, terminal line, and fringe undistinguished. Abdomen: basal coremata and pockets absent in male; dorsal tufts on A1–A3 variably prominent, that on A3 the largest. Male genitalia: uncus narrowly spatulate, apically acuminate with sparse, dorsal, cristate patch of recumbent setae; tegumen rounded at base of uncus, flaring laterally and forming rounded peniculi, largely devoid of hairs, before narrowing at s-shaped pleural sclerite and articulating with v-shaped vinculum; juxta + anellus elongated, height 3 × width, with divergent, paired, dorsal projections; ventral margin of valve evenly convex at base, then nearly straight until angled dorsally into notch at anteroventral edge of cucullus; base of sacculus rounded, swollen, with dense patch of sensory hairs on bluntly rounded process on costal lobe; dorsal margin of valve concave basally, then nearly straight to broad neck of cucullus; cucullus bifurcate with dorsal arm broadly rounded, 3 × width of narrow, acute, spine-like ventral arm, together resembling a duck’s head in profile, the entire inner face of cucullus covered with dense patch of proximally oriented, spine-like setae; corona absent; clasper an appressed, sclerotized, crescent-shaped sclerite; ampulla absent; digitus heavily sclerotized, serrate or somewhat dentate on dorsal edge, sinuous, acuminate, extending almost to ventral margin of valve; phallus nearly straight, with terminal carina consisting of two opposing, subequal, sclerotized, ribbon-like, spined bars extending onto subbasal part of vesica; vesica with single, prominent, blunt, rose-thorn cornutus on a dorso-mesial diverticulum. Female genitalia: papilla analis dorsoventrally flattened, lyriform, with lateral margin prominently constricted toward anterior end, narrowly convex toward posterior end, acute apices divergent, with long setae set in tiny membranous circles on both dorsal and ventral surfaces; segment A8 2 × as wide as long, anterior apophysis 1.5 × length of segment A8 and posterior apophysis 2.5 × length of segment A8; lamella antevaginalis heavily sclerotized, laterally flared, hemi-cordate, with cleft on convex posterior rim, anterior portion of plate extended into a long, acute point, overall shape of lamella funnel-like; ostium moderately sclerotized; ductus bursae narrow, about as long as appendix bursae, heavily sclerotized; appendix bursae membranous with light rugosity, about as long as and half the width of corpus bursae, viewed ventrally, oriented left, ductus seminalis at posterior apex; corpus bursae membranous, round, with two signa.
DNA barcodes are available for four specimens of P.blackorum from Kentucky and Missouri, which form a discrete group and unique BIN at a minimum nearest-neighbor difference of 2.55%.
The larva belongs to the largest subgroup within the genus; i.e., those having only the middorsal white line continuous across segments A1–A4 which are otherwise completely encircled by dark purplish-brown pigmentation. The lateral margins of the honey-colored prothoracic shield bear the usual dark, nearly black pigmentation stripes which do not extend onto the reddish, chestnut head capsule in this species, as is often the case in others. The dorsal pinacula on A1–A7 are small, about average size for a Papaipema larva; those on A8 are very large, heavily sclerotized, and almost black. The D1 and D2 pinacula on A8 are ovoid and well separated. Those on A9 are fused into a single, heavily sclerotized bar, almost black, and of a distinctive shape reminiscent of a moustache, which covers most of the dorsal surface of the segment. The anal shield is brown, rugose, and heavily sclerotized.
TYPES. Holotype: ♀. Forewing length 23.5 mm [alar expanse 51.5 mm]. South of Rt. 146 on Reevesville Rd., Johnson Creek, C. Sampson property, N37.387106, -W088.777966, Johnson Co., Illinois; larva collected 12 August 2019 by J. R. Wiker / Larva gall forming in vine of Aristolochia tomentosa; pupated late August 2019, emerged 30 September 2019, #19-010. [CNC] Paratypes: 160 ♂, 19 ♀. Illinois. Johnson Co., Grantsburg, east of Reevesville Rd. + Johnson Creek, 37.3869, -88.7777 388´ [118 m]; 21 Aug 2017 ex larva in Aristolochia tomentosa reared in carrot, eclosed 5 Oct 2017; A. E., S., & L. McBride, J. Wiker (1 ♂). Same locality, 22 Jul 2020 ex larva in Aristolochia tomentosa reared in carrot, eclosed 5–10 Oct 2020; A. E. McBride w/ J. R. Wiker (4 ♂). Johnson Co., south of Rt. 146 on Reevesville Rd., Johnson Ck., C. Sampson property, N37.3871, -W088.7779; larva collected 12 Aug 2019 by J. R. Wiker / Larva gall forming in vine of Aristolochia tomentosa; pupated late Aug 2019, emerged 4 Oct 2019, #19-025 (1 ♂). Indiana. Posey Co., Hovey Lake FWA, Kron Tract boat ramp area, Ohio River, N37.7865,-W087.9415; 3 Oct 2019 UVT BLB; J. R. Wiker (1 ♂). Posey Co., Hovey Lake FWA, Kron Tract east of Union Ferry Road, N37.7843, -W087.9475; 3 Oct 2019 UVT BLB; J. R. Wiker (1 ♂). Kentucky. (Unless noted otherwise, all Kentucky records are of specimens collected by William R. Black, Jr. in UV light traps.) Ballard Co., Mayfield Creek, Hays’ Woods, 0.6 mi. W U.S. 62 bridge, cane site #9; 11 Oct 2005 (2 ♂). Calloway Co., Blood River, ca. Grubbs Rd. bridge, cane site #3; 17 Oct 2002 (1 ♂). Calloway Co., Blood River, cane sites #4 & #8; 15 Oct 2003 (7 ♂). Same locality; cane sites #6 & #9; 21 Oct 2003 (2 ♂). Same locality; cane site #4; 11 Oct 2006 (1 ♂). Same locality; cane site #4A; 6 Oct 2010 (10 ♂, 1 ♀). Carlisle Co., Sandy Branch, cane site #4; 12 Oct 1999 (1 ♂). Same locality; cane sites #3 & #4; 15 Oct 1999 (3 ♂). Carlisle Co., unnamed creek 1 mi. N Burkley; canebrake UV trap, 17–18 Oct 2001; M. C. Nielsen (2 ♂). Same locality; cane site #4; 18 Oct 2001 (2 ♂). Same locality; cane site #1; 11 Oct 2002 (3 ♂). Same locality; cane sites #4 & #5; 8 Oct 2004 (5 ♂). Same locality; cane site #2; 20 Oct 2004 (1 ♂). Carlisle Co., Mayfield Creek, cane site #1; 19 Oct 2003 (2 ♂). Fulton Co., Running Slough at Hwy 94 mile 6.7; 12 Oct 2001 (1 ♂). Fulton Co., Kentucky Bend, Stepp Rd., 90 m. from cane at creek mouth; 13 Oct 2001 (1 ♀). Graves Co., 2.8 mi. NE Wingo, Obion Creek canebrake, 36.6592, -88.6914 428´ [130 m]; 1 Jul 2016 ex larva in Aristolochia tomentosa, reared in carrot, eclosed 6 Oct 2016; Jim Wiker and Bill Black, Jr. (1 ♀). McCracken Co., old Mayfield Creek, 1 mi. W Clinton Rd., cane site #7; 13 Oct 1999 (1 ♀). Same locality; cane site #3; 19 Oct 1999 (1 ♂). McCracken Co., old Mayfield Creek, 1.5 mi. W Hwy 339, cane site #26; 16–23 Oct 2000 (3 ♂, 1 ♀). Mississippi. Lee Co., Natchez Trace Parkway, Chiwapa Creek, mile marker 253.1; 27 Sep 2013; R. Patterson (1 ♂). Warren Co., vicinity Bovina; coll. at MV light 11 Oct 1972; Bryant Mather (1 ♀). Missouri. Butler Co., Mark Twain National Forest, 360´ [110 m] jct. Rt. 67 & Black River 0.6 mi. WNW Hendrickson; MV light, 14–16 Oct 2000; E. Quinter & J. Wiker (14 ♂). Butler Co., Hendrickson, canebrake north of Black River, 36.9041, -90.4777 378´ [115 m]; 175w MV light, 15 Oct 2011; A. E. McBride w/ Jim Wiker (8 ♂). Same locality; reared ex ova 2012 in potato from parent female collected 15 Oct 2011, eclosed Jun 2012; reared by A. E. McBride (1 ♂, 2 ♀). Same locality and collector; 175w MV light, 12–13 Oct 2015 (4 ♂). Same locality; reared ex ova 2016 in potato from parent female collected 12 Oct 2015 at 8w BLT, eclosed Jul 2016; reared by A. E. McBride (5 ♂, 8 ♀). Wayne Co., Mark Twain National Forest, 400´ [122 m] jct. Rt. 49 & Black River, 2.7 mi. W Williamsville, Browns Crossing; ex larva found boring in emerging culm of Arundinaria gigantea 25 May 1994, pupated 22 Aug, emerged 18 Sep 1994; E. L. Quinter (1 ♀). Same locality; wandering larva found in canebrake 19 April 1995, reared on artificial diet, emerged: 6 Sep 1995; E. L. Quinter (1 ♂). Same locality, T27N-R4E, Sec. 23; coll. with UV light in canebrake 12 Oct 1999; J. Wiker & V. LaGesse (13 ♂, 1 ♀). Same locality; coll. with UV light in canebrake 12 Oct 1999; J. R. Wiker (11 ♂). Same locality; at UV/MV lights 14–17 Oct 2000; E. Quinter & J. Wiker (5 ♂). Butler Co., Mark Twain National Forest, 360´ [110 m] jct. Rt. 67 & Black River 0.6 mi. WNW Hendrickson; MV light, 14–16 Oct 2000; E. Quinter & J. Wiker (14 ♂). Tennessee. Henry Co., Blood River, cane sites #P2, #P3, & #P4; UV light traps, 12 Oct 2003; W. R. Black, Jr. (10 ♂, 1 ♀). Same locality and collector, cane sites #P5 & #P6; UV light traps, 13 Oct 2003 (3 ♂). Same locality and collector, cane site #P15; UV light trap, 21 Oct 2003 (5 ♂). Same locality and collector, cane site #P2; UV light trap, 17 Oct 2004 (1 ♂). Same locality and collector, cane site #P2; UV light trap, 20 Oct 2005 (1 ♂). Same locality and collector, cane site #P2; UV light trap, 11 Oct 2006 (1 ♂). Same locality and collector, cane site #17; UV light trap, 18 Oct 2006 (3 ♂). Same locality and collector, 50 m. upstream from cane site #P2; UV light trap (15W), 6 Oct 2010 (4 ♂). AMNH, CMNH, CNC, INHS, LACM, MCLB, MEM, NHML, NYSM, UCD, ULK, USNM, VADNH, YPM, AEM, ELQ, JRW, RPC, VAB.
This species was not known to exist until it came to ELQ’s attention with a single, large female from Mississippi (
In the interim, ELQ’s field research was focused on the noctuid fauna of canebrakes throughout the southern United States, where he had already discovered a number of unknown species. Two unknown Papaipema larvae detailed below ultimately resulted in the third and fourth known specimens of P.blackorum. Subsequent efforts to locate more larvae of this species boring in cane all met with failure. Alerted to the presence of this species in Wayne County, Missouri, JRW set up blacklight sheets in a large canebrake there in October 1999 and succeeded in collecting a large series of adult moths. This reinforced the notion that the species was an apparent canebrake specialist, which was further frustrated by the collective inability to locate additional larvae. Meanwhile Bill Black and others were intensively targeting canebrake habitats in western Kentucky and adjacent portions of neighboring states, collecting numbers of adult moths at black light traps placed in some canebrake sites, and relatively few or none in other seemingly suitable canebrakes. Years passed, and frustration grew, but finally came to closure with the discovery by AEM and JRW in 2016 that the larval host plant is actually woolly dutchman’s pipe (Aristolochia tomentosa), also known as woolly pipevine. Inasmuch as this plant occurs abundantly in the same general habitats as giant cane, but is not necessarily present at many canebrakes, accounts for the many years of inability to find the larvae. This example, and others like it such as the case with Papaipema aweme, serves to illustrate just how abundant a species thought to be extremely rare may actually prove to be, once the details of its biology have been elucidated.
The first encounter with the larva of this species occurred when ELQ found an individual of unknown identity, apparently nearly half grown, boring at ground level in a tender, emerging culm of giant cane on 25 May 1994 while examining a canebrake in Wayne County, Missouri. The canebrake was surrounded by extensive patches of woolly pipevine. The larva pupated 22 August and emerged 18 September 1994. Another wandering larva he collected 19 April 1995 at the same site appeared to be in the second instar. It molted seven times between that date and 6 July, pupating 13 August and emerging 6 September 1995. Both these larvae were reared to adults on artificial diet. Not until 29 Jun 2016, when AEM and JRW, accompanied in the field by William R. Black, Jr., discovered P.blackorum larvae boring in woolly pipevine along Obion Creek in Graves Co., Kentucky, was the true host plant correctly ascertained.
This is yet another of many examples of Papaipema species with specialized adaptations to boring in highly toxic host plants. Infested plants are indicated by large, cylindrical, woody galls 9–12 cm long, located from ground level to at least two meters high, which wrap with the contours of the vines that can climb at least ten meters up the supporting substrate. AEM found that the galls have a single orifice for ejection of frass at one end, and upon maturity the larva chews a second hole at the opposite end which serves as an exit. Galls of P.blackorum are similar in shape to those formed by Papaipema circumlucens in hops vines (Humulus lupulus), and as with that species, larvae of P.blackorum mature on relatively little plant material. Unlike hops, which grows a new complement of vines from its roots each season, the vines of woolly pipevine are perennial. These vines must reach a great age and a large diameter before they can support the larva of P.blackorum and should be conserved when one is searching for larvae of this species. The gall should be carefully split open on the long axis when extracting a P.blackorum larva; the vine itself should not be completely severed with cutting implements and thereby killed.
JRW has observed young larvae exhibiting ballooning behavior which may be characteristic of the few other arboreal Papaipema species as well, suggesting eggs may be deposited in large numbers well up into the canopy. Such behavior is an effective larval dispersal mechanism in groups such as lymantriines, which deposit eggs in large masses. Since many Papaipema larvae exhibit strong cannibalistic tendencies, it clearly is advantageous for young larvae to avoid inhabiting the same vine in numbers. A female collected by AEM in Butler Co., Missouri deposited 1,430 eggs on dried foliage before expiring. Pupation presumably occurs in the ground, among detritus or in a shallow cell in soil, as galls have not been found containing pupae. For further information regarding the life history of P.blackorum, see
Papaipema blackorum appears to have a rather limited distribution in floodplain forests in the southern parts of the midwestern United States, especially in the lower Ohio and Mississippi River drainages, in intimate association with local abundance of its host plant. This species is probably more widespread than records indicate, due to the fact that collecting efforts in the region this species was found were focused almost entirely upon canebrake habitats, and not upon populations of woolly pipevine, which are more generally distributed.
Papaipema impecuniosa GROUP
Papaipema impecuniosa and two very closely related species, P.nelita and P.sablensis, comprise this group. Papaipema sablensis is of recent origin, a derivation of the ancestral P.nelita lineage resulting from isolation, presumably during or slightly before the Pleistocene.
Papaipema impecuniosa (Grote) (Aster Borer)
Gortyna impecuniosa Grote, 1881, Bulletin of the United States Geological and Geographical Survey of the Territories, 6(2): 267.
Type locality: Massachusetts, [USA]. [NHML]
note—Described from a single male, 25 mm in expanse. Grote mentioned having seen larger specimens in other collections besides his own. This likely accounts for the fact that there is another specimen in the New York State Museum bearing one of Grote’s original, red-bordered type labels, identical to the one on the NHML specimen, with the inscription “ Gortyna impecuniosa Grote Type” in his handwriting. This male bears printed labels from the Wm. W. Hill collection and is labelled as having been collected at Centre, New York on September 25, 1877. However, this is actually a specimen of Papaipema unimoda. To preclude the possibility of future confusion and to stabilize the name impecuniosa, the male type in the NHML is hereby designated lectotype.
Papaipema inquaesita; Holland, 1903, not Grote and Robinson, 1868.
Grote’s choice of name for this species was never explained, but it seems likely it was derived from the Latin root pecuniosus, meaning rich, and in this instance impecunious, having little or no money. The name probably reflects the fact that this species never exhibits the silvery-white stigmata on the adult forewings which are so characteristic of most Papaipema species. This common, widespread species is among the smaller members of the genus. Adults resulting from larvae that were confined to feeding in stems of especially slender host plants can be quite tiny, since it does not appear that larvae will abandon their galleries in search of additional food sources. This species is sufficiently distinctive such that it is unlikely to be confused with any other.
DNA barcodes are available for 18 specimens of P.impecuniosa. Theses separate into two clusters, one unique to P.impecuniosa (North Carolina and eastern Canada) and the other within the P.nelita barcode cluster, with only 0.3% minimum divergence between the two. However, total variation among both species combined is low, and there is no indication that there is a taxonomic issue in P.impecuniosa as currently defined.
The larva is another of those with all longitudinal lines interrupted by the pigmented bands of segments A1–A4. The dark-pigmented lateral margins of the prothoracic shield do not extend onto the honey-colored head capsule. Additional details concerning the immature stages and larval detection are documented by
Papaipema impecuniosa is most often found boring in various species of asters, commonly flat-topped aster (Doellingeria umbellata) and purplestem aster (Symphyotrichum puniceum), but likely numerous others. In the Midwest, it has also been repeatedly reported boring in common sneezeweed (Helenium autumnale). Larvae of P.impecuniosa bore in the lower stems but mainly in the roots of their host plants. The presence of the larva in its host may be detected by a slight swelling or gall at the base of the stem. Small amounts of whitish frass may be noted on the soil beneath. Pupation invariably takes place in the basal gallery, rendering it unnecessary for one to rear larvae of this species because gathering pupae is easier. This species occurs in wet meadows throughout its range but also in drier prairie habitats. Adults are on the wing mainly from mid-September through the end of October.
The species is generally distributed throughout the northeastern states and upper Midwest, from Nova Scotia to Wisconsin and south to Missouri, Kentucky, and farther south at higher elevations along the Appalachian Mountains.
Papaipema nelita (Strecker) (Coneflower Borer)
Hydroecia nelita Strecker, 1898, Lepidoptera, Rhopaloceres and Heteroceres, Indigenous and Exotic. Supplement 1,.
Type locality: Chicago, [Cook Co.], Illinois, [USA]. [FMNH]
note—Described from two males.
Papaipema errans Barnes and McDunnough, 1912, The Canadian Entomologist, 44: 56.
Type locality: White Mts., Ariz. [Arizona, USA]. [USNM]
note—Described from a unique male holotype.
† Papaipema nelita ab. orbicularis Strand, 1915 [1916], Archiv für Naturgeschichte, 81A (11): 163.
Type locality: “Vereinigte Staaten” [no depository]
note—unavailable infrasubspecific name; described as an aberration of P.nelita from an illustration in
† Papaipema nelita linda Bird, 1916, The Canadian Entomologist, 48(1): 13-14.
Type locality: Wilmington, Del. [Delaware, USA]. [USNM]
note — Unavailable infrasubspecific name. Described from at least three specimens, based on the white-spotted form.
Papaipema melita; Tietz, 1936, misspelling. Papaipema engelhardti Bird, 1939, The Canadian Entomologist, 71: 74.
Type locality: Oak Creek Canyon, Coconino County, Arizona, [USA]. [AMNH]
note—Described from a male and female reared by George P. Engelhardt from larvae found boring in Rudbeckia ampla.
Papaipema unimoda; Forbes, 1954, part, not Smith, 1892.
The name is apparently derived from the Greek nelites, innocent. This common species, widespread throughout eastern North America, has a seemingly disjunct southwestern population generally distributed across mountainous areas of Colorado, New Mexico, and Arizona. This apparent disjunction may be only an artifact reflecting insufficient sampling in several midwestern states, but it may also reflect reality. If so, one might expect to find distinguishing features between these two populations. Distinguishing features have not been found, either morphological or genetic, between eastern and southwestern populations, and the host plants are essentially the same in both instances. The adult forewing in southwestern populations is uniformly dark charcoal gray, identical to that of northeastern and midwestern populations. The populations from the approximate latitude of Pennsylvania and south along the Appalachians are distinctive in the golden coloration of the medial forewing area. Accordingly, all populations are being treated as a single species with the caveat that the name errans, which was first applied to the southwestern populations, could be resurrected should future genetic studies reveal sufficient distinctions.
A species very commonly confused with P.nelita is P.unimoda. Both species occur together and can be similar in size and coloration. Both occur with the reniform, orbicular, and suborbicular spots either suppressed or strongly expressed. In the spotted form, P.unimoda may be recognized by the presence of yellowish basal spots, which are absent in P.nelita. In P.nelita, the course of the postmedial line is straight for nearly all its length which is diagnostic, whereas that of P.unimoda is widely excurved around the reniform. If one is still in doubt, the male genitalia are totally dissimilar and may be quickly examined by simply brushing the scales off the tip of the abdomen.
DNA barcodes are available for 20 P.nelita specimens from across the species’ range. Although taxonomically quite distinct, the minimum barcode divergence of P.nelita relative to P.duplicata and P.silphii is very low (0.3%), and partially intercalated with that of P.impecuniosa, as described under that species. Less surprising is the similarity to P.sablensis (0.46% minimum).
Cutleaf coneflower (Rudbeckia laciniata) is the most frequently reported larval host. ELQ and AEM have repeatedly found P.nelita larvae boring in stems of the non-native common burdock in upstate New York. The larva initially enters the stalk above ground but soon moves to the root where it completes most of its development. A slight gall may be formed near the soil line, and expelled whitish frass, together with signs of wilted foliage, may provide clues to larval presence. In burdock (Arctium), the larva typically bores low in the main stalk, though several larvae may inhabit the main stalk at different points in large plants. Pupation occurs in a cell in soil near the base of the bored stalk. This is one of the earliest species to be on the wing at any locality. Specimens have been collected in the last week of July, with the majority during August and September. Occasional stragglers are reported from early October.
In the eastern portions of its range, P.nelita ranges from the Maritime Provinces to Manitoba, south to Missouri and farther south at higher elevations in the Appalachian Mountains. The species is also widespread in the southwestern mountains of the United States.
Papaipema sablensis Quinter, NEW SPECIES (Sable Island Borer)
Papaipema sablensis Quinter.
Type locality: Sable Island, Nova Scotia, Canada. [CNC]
note—Named for its unique occurrence and fragile existence on the last remaining vestige of its Pleistocene refugium, Sable Island, Nova Scotia.
Although similar in appearance to P.nelita, there is no possibility of confusion with any other species due to the unique and remote occurrence of P.sablensis. The forewing coloration of adults of P.sablensis is straw yellow, much paler than those of typical mainland populations of the very closely related P.nelita. There appears to be little or no possibility for gene flow between these two entities, given the extremely sedentary nature of Papaipema adults in general and the separation of Sable Island from the closest point of the mainland by about 200 km of the Atlantic Ocean. There are no discernible differences in the genitalia of the two species, although there exists a slight difference in details of the postmedial line on the forewings. In P.sablensis, the inner of the parallel, double postmedial lines is strongly scalloped between the veins; in P.nelita, much less so. Also, the terminal area of P.sablensis is the same color as the medial area and contrasts with the darker subterminal area. In P.nelita, both subterminal and terminal areas of the forewing are about equally dark in comparison with the lighter medial area.
Head: antennae of both sexes ciliate, laminate, less deeply in female; dorsum scaled; ventral lamination obscured by densely pubescent sensilla; eye smooth, rounded; haustellum unreduced; labial palpus laterally flattened, upcurved; first segment swollen basally, arching slightly upward, slightly more than half as long as second segment; second segment straight, about as long as eye width, scales broad; third segment ⅓ × length of second segment, and projecting slightly anteriorly, scales narrow; frons unmodified, smooth, convex, with unspecialized conical vestiture consisting of spatulate scales mixed with fine, hairlike scales. Thorax: vestiture of prothoracic collar, patagia, mesothorax, and metathorax concolorous, a mixture of coarse spatulate, thin apically spatulate, and hairlike scales; transverse, wedge-shaped, prothoracic crest prominent, mesoscutellar tuft moderate, metascutellar tuft subequal. Forewing: length similar in both sexes = 13.5–15 mm; dorsal surface ground in both sexes uniform straw yellow; basal spots concolorous; filling of antemedial and postmedial lines concolorous with ground; antemedial line double, brown, nearly straight until strongly bowed outward just before meeting inner margin; postmedial line brown, double, the inner prominently scalloped, nearly straight until bending inward immediately before meeting costa and meeting inner margin slightly diagonally; medial shade darker fuscous, touching lower inner loop of reniform, then angled sharply below reniform and meeting inner margin diagonally; subterminal line preceded by series of darker, charcoal chevrons filling the spaces between veins; the subterminal space filled with gray scales; terminal space straw yellow, concolorous with ground, veins crossing terminal space prominent, contrasting dark fuscous; terminal line and fringe fuscous; reniform with either small, white, satellite spots or fuscous brown spots, in either phenotype prominently contrasting with paler ground; orbicular and suborbicular stigmata similar. Hindwing: dorsal surface uniform pale fuscous, paler than forewings and lacking the yellowish coloration; discal spot and medial shade visible but relatively obscure in most; subterminal area slightly paler; terminal line and fringe slightly darker than terminal area. Abdomen: basal coremata and pockets absent in male; dorsal tufts on A1–A3 not prominent, that on A3 the largest. Male genitalia: uncus narrowly spatulate, apically acuminate; tegumen rounded at base of uncus, flaring laterally and forming rounded peniculi, largely devoid of hairs, before narrowing at s-shaped pleural sclerite and articulating with v-shaped vinculum; juxta + anellus elongated, height 3 × width, with divergent, paired, dorsal projections; ventral margin of valve evenly convex at base, then nearly straight until angled dorsally at anteroventral edge of cucullus; base of sacculus rounded, swollen, with dense patch of sensory hairs on basal triangular process of costal lobe; dorsal margin of valve concave basally, then nearly straight to broad neck of cucullus; cucullus bifurcate with triangular dorsal arm broadly rounded, 3 × width of short, acutely pointed ventral arm; the entire inner face of cucullus covered with a dense patch of proximally oriented, spine-like setae; corona absent; clasper an appressed, sclerotized, crescent-shaped sclerite; ampulla absent; digitus heavily sclerotized, serrate on basal half of dorsal edge, sinuous, acuminate, extending more than halfway to ventral margin of valve; phallus slightly curved, with terminal carina consisting of two opposing, subequal, sclerotized, ribbon-like, spined bars extending onto subbasal part of vesica; vesica with single, prominent, blunt, rose-thorn cornutus on dorso-mesial diverticulum. Female genitalia: papilla analis dorsoventrally flattened, lyriform, with lateral margin prominently constricted toward anterior end, narrowly convex toward posterior end, acute apices divergent, with long setae set in tiny membranous circles on both dorsal and ventral surfaces; segment A8 2.8 × as wide as long, anterior apophysis 2 × length of segment A8 and posterior apophysis 2.8 × length of segment A8; lamella antevaginalis heavily sclerotized, laterally flared, triangular in outline with deep cleft on posterior rim, anterior portion of plate extended into a long, acute point, overall shape of lamella funnel-like; ostium moderately sclerotized; ductus bursae narrow, about as long as appendix bursae, heavily sclerotized; appendix bursae membranous with light rugosity, about as long as and half the width of corpus bursae, viewed ventrally, oriented left, ductus seminalis at posterior apex; corpus bursae membranous, round, with two signa.
DNA barcodes are available for two specimens of P.sablensis. Although these nest within the total variation exhibited by P.nelita, the presumed nearest relative, these two P.sablensis samples represent a unique haplotype differing by a minimum of 0.46%.
The larva is unknown; however, based on the close genetic relatedness of this species to Papaipema nelita, it is expected that the pigmented bands encircling the first four abdominal segments will be uninterrupted by the pale longitudinal lines.
TYPES. Holotype: ♂. Forewing length 13 mm [alar expanse 30 mm]. Sable Island, Nova Scotia, 11 Aug, 1978 D. C. Ferguson [CNC] Paratypes: 13 ♂, 3 ♀. Nova Scotia. Sable Island, Nova Scotia, 15 Sep 1967, J. E. H. Martin (1 ♂, 1 ♀). Same locality, Met Station, 10 Sep 1977, Barry Wright (1 ♂). Same locality, 28 Aug 1980, Barry Wright (1 ♂). Same locality, 31 Aug 1980, Barry Wright (1 ♂, 1 ♀). Same locality, NS Camp, 43.9366, - 60.0645, 25 Aug 2008, Zoe Lucas (4 ♂, 1 ♀). Same locality, Met Station, 3 Sep 1985, Agric. Can. (4 ♂). Same locality, Met Station, 15 Sep 1985, Agric. Can. (1 ♂). CNC, NSM, YPM.
Intense efforts by ELQ to identify the larval host on Sable Island in 1980 were unsuccessful. It still remains unknown. Due to the paucity of plant diversity on the island, the possibilities seem very limited. Only the abundant seaside goldenrod (Solidago sempervirens) and blueflag iris (Iris versicolor) were deemed suitable hosts. No Papaipema larvae were found in either plant during ELQ’s twelve-day exploration of the island beginning 2 July of that year, although Amphipoea americana larvae were generally found boring in the latter. The relatively few adult specimens of sablensis collected over four decades of sporadic visits to the island by others have been taken between 11 August and 15 September.
Papaipema sablensis represents another of several species of Lepidoptera endemic to Sable Island. It presents an opportunity, unique among Papaipema, to directly observe how much (or how little) evolution has occurred in an insect which has been isolated from its ancestral populations and nearest relative for perhaps 20,000 years. During the most recent of the Pleistocene glaciations, the ancestral populations which gave rise to P.nelita and P.sablensis likely were displaced westward, southward, and to the East, surviving in refugia there. In the case of P.sablensis, that likely occurred on or near the Grand Banks off the coast of Newfoundland (
Papaipema insulidens GROUP
This group consists of Papaipema insulidens, P.birdi, P.albimontana, and P.dribi, which are apparently more recent descendants of an ancestral clade. They also appear to reflect an ancestral preference for apiaceous plants. These four are poorly distinguished from one another by morphology or by barcodes but have nonetheless clearly evolved in allopatric isolation.
Papaipema insulidens (Bird) (Northwestern Umbel Borer)
Distribution of material examined of Papaipema albimontana, P.birdi, P.dribi, and P.insulidens.
Hydroecia insulidens Bird, 1902, The Canadian Entomologist, 34(5): 112.
Type locality: Vancouver Island, [British Columbia, Canada]. [AMNH]
note—Described from three examples including both sexes. The name refers to its insular type locality.
Papaipema pertincta Dyar, 1920, Insecutor Inscitiae Menstruus, 8(7–9): 146.
Type locality: Forest Grove, [Washington Co.], Oregon, [USA]. [USNM]
note—This taxon was described from a single male reared from lupine by L. P. Rockwood. It was synonymized with P.insulidens by
Some P.insulidens individuals cannot be reliably distinguished from those of P.birdi, other than by their geographic occurrence. Those populations of P.insulidens which occur along the northern and eastern boundaries of the Great Basin (
The eastern and western populations of these two taxa are broadly isolated from one another by the Rocky Mountains, and it appears all but certain that they represent sibling species, albeit not as distinct as might be expected. Otherwise, P.insulidens could only be confused with P.albimontana which occurs in the White Mountains of Arizona, well south of the range of P.insulidens. The reniform of P.insulidens is of the typical figure-8 shape; that of P.albimontana is smaller and more compressed laterally. DNA barcodes are available for 14 samples from Oregon, British Columbia, and Washington. As discussed under P.birdi, some barcodes are shared between multiple species of the P.insulidens group.
The larva is unusual in that it exhibits two different morphs, with some individuals showing an intergradation between morphs. Many larvae can be assigned to the group with only the dorsal white stripe continuous across the pigmented bands on the first four abdominal segments, while others have both the dorsal and subdorsal white stripes continuous across these same segments. The prothoracic shield shows the typical dark lateral margins with the dark lines extending onto the head capsule. The pinacula are of moderate size.
This species shows a strong preference for boring in stems of apiaceous plants such as cowparsnip (Heracleum), Angelica, water hemlock (Cicuta), and licorice-root (Ligusticum) but also has been recorded boring in burdock (Arctium), thistle (Cirsium), teasel (Dipsacus), burnweed (Erechtites), sweet coltsfoot (Petasites frigidus) (
Papaipema insulidens ranges south from Vancouver Island, British Columbia to Oregon along the coast with an interior distribution east to the more arid regions bordering the northern Great Basin.
Papaipema birdi (Dyar), REVISED STATUS (Water-hemlock Borer)
Gortyna birdi Dyar, 1908, The Canadian Entomologist, 40(3): 78.
Type locality: Rye, [Westchester Co.], New York, [USA]. [USNM]
note—Described from eight specimens, the male type specimen having been reared by Bird at Rye, New York. Papaipema birdi was synonymized with P.insulidens by
Hydroecia marginidens; Smith, 1899, in part, not Guenée, 1852.
Papaipema birdi has been widely called P.marginidens throughout the literature and is still commonly found misindetified as such in collections. The designation of a neotype for P.marginidens now stabilizes that name and distinguishes it from P.birdi. Populations of P.birdi appear to be largely monophagous in the austral portions of the range but blend imperceptibly into polyphagous populations northward. Individuals from these boreal populations tend to be smaller and were thought to possibly represent a cryptic sibling species. Genitalic characters, especially the lamella antevaginalis in the female, show greater variability than usual, suggesting possible introgression between two divergent entities, one essentially monophagous on water hemlock (Cicuta) and the other polyphagous. However, no consistent means of distinguishing individuals of both types which commonly co-occur could be found, so all eastern North American populations are being treated as the single species P.birdi, with the caveat that a second biological entity may be included.
Individuals of P.birdi are indistinguishable from those of widely allopatric northwestern populations of P.insulidens. Individuals of P.insulidens from more arid areas bordering the northern Great Basin are consistently paler than those of the eastern P.birdi. Papaipema birdi is broadly sympatric with several other eastern species with which it can be confused. See discussions under P.nepheleptena, P.eryngii, and P.marginidens for diagnostics. Members of the more distantly related P.rutila group are occasionally confused with both P.insulidens and P.birdi but can be recognized most easily by the color of the basal spots of the forewing. These spots are contrasting yellowish cream in P.birdi and P.insulidens but dull orange in P.rutila group species, none of which are readily distinguishable from P.birdi or P.insulidens by genitalic characters. All members of the P.insulidens group have very similar or even identical DNA barcodes. Seventeen samples of P.birdi from Michigan, Ontario, and North Carolina were compared to the remainder of the species-group, and included examples of barcode haplotypes shared with P.insulidens, P.albimontana, and P.dribi.
The early stages of P.birdi were first detailed by
Spotted water hemlock (Cicuta maculata) appears to be the preferred host for P.birdi throughout austral elements of its distribution. Rattlesnake-master (Eryngium) and water parsnip (Sium) have occasionally been reported as additional apiaceous hosts. Populations in more boreal areas have been recorded boring in a variety of hosts, commonly Angelica, water hemlock, and cowparsnip (Heracleum), but including non-apiaceous hosts; e.g., composites such as burdock (Arctium), Indian plantain (Arnoglossum), and thistle (Cirsium), as well as others such as Verbena and Iris. Larvae exit the nearly consumed root and pupate under ground nearby. Larval presence is readily noted by wilting, yellowing, or even blackening of the terminal foliage of water-hemlock as early in the season as mid-May. Lateral branches may also be bent at the point where internal girdling of the hollow stems has weakened them. Larvae boring in Cicuta typically spend most of their development feeding in the highly toxic roots which may be completely consumed by the time the larva is mature. That larvae deliberately consume the most toxic portion of the plant suggests the possibility they may be sequestering the cicutoxin responsible for the plant’s notorious toxicity, perhaps acquiring chemical protection for the adult stage, as suspected for other Papaipema species with similar preferences for host plants collectively bearing a vast array of phytotoxins. Recorded captures of adult P.birdi span from early August to the third week of October, but most records are from September.
Papaipema birdi is typically associated with wet meadows and other riparian environments and is broadly distributed throughout the upper austral, transition, and boreal zones of eastern North America west to Alberta. It follows the Appalachian Mountains well into the South but is otherwise absent throughout most of the southeastern United States.
Papaipema dribi Barnes and Benjamin (Sacramento Mountains Borer)
Papaipema dribi Barnes & Benjamin, 1926, Pan-Pacific Entomologist, 2(3): 112.
Type locality: High Rolls, [Otero Co.], New Mexico, [USA]. [USNM]
note—Described from a unique male holotype, date of collection given only as “August” without mention of year on its label. It could have been collected in any year prior to 1926. According to the original description, the collector was said to be J. G. Boniwell. Other labels indicate a slide was made of the genitalia by Foster Benjamin: “♂ gen. 436 4 Oct. 32 FHB” and “Genitalia Slide By USNM 39409”. Subsequently, a second male was located elsewhere among unidentified material, and it bore a crumbling, nearly illegible label, perhaps even older, which simply read “Bent, N.M.” This locality is about six miles west of Mescalero, and like the type locality High Rolls about 13 miles south, is also situated in Otero County.
This remained an enigmatic species known from only two males, both very old, prior to its rediscovery in 2021. The moth has a different appearance from other members of its group. Papaipema dribi is treated herein as a distinct species for this reason, and also because of its isolation from populations of its nearest relatives.
Papaipema dribi has a forewing length to 18 mm and is dark brownish with a pattern most nearly resembling that of P.albimontana. Both species have nearly identical forewing maculation which features a reniform stigma that is more laterally compressed and distinctly smaller than that of either P.insulidens or P.birdi. Brown scales are heavily dusted over a lighter yellowish medial area of the forewing of P.dribi which lacks the orange tones of P.albimontana, a paler species of about the same coloration as P.insulidens from the Great Basin. Some recent collections of P.depictata also have been misidentified as P.dribi even though they are yellowish-brown moths with a different habitus. Perhaps the most diagnostic feature of the forewing maculation of P.dribi is the near or actual obsolescence of the suborbicular group stigma (
The larva is unknown; however, based on the close genetic relatedness of this species to the other members of the group and especially to P.albimontana, it is expected that the larval phenotype exhibits pigmented bands encircling the first four abdominal segments, interrupted by only the middorsal pale longitudinal line.
The host plant remains unknown, but very likely is one which requires a mesic or riparian habitat as is typical of hosts for virtually all Papaipema species. As P.dribi appears to be a member of the Apiaceae-feeding clade, any suitable umbellifer growing at the appropriate elevation in the Sacramento Mountains would be high on the list of probable candidates. Based upon
Eric Metzler (pers. comm.) reported that “Bent and Mescalero are at the same elevation as High Rolls, and Bent and Mescalero are also at the exact place where the trees end and treeless desert begins. The topography is less steep at Bent.” In 2021, Metzler rediscovered the species at an undisclosed location in Otero County. For additional details, see
At present, P.dribi is known only from the Sacramento Mountains in Otero County, New Mexico at or above elevations of approximately 1,800 meters.
Papaipema albimontana Quinter, NEW SPECIES (White Mountains Borer)
Papaipema albimontana Quinter.
Type locality: 1 mi. s. Greer, Little Colorado River floodplain, 33.9928, -109.4653 8,500´ [2,591 m], White Mountains, Apache Co., Arizona, USA. [CNC]
note—The name albimontana, herein treated as a noun in apposition, refers to its type locality in the White Mountains of Arizona.
Papaipema albimontana appears to be a locally endemic species genetically isolated from its closest relatives which occupy habitats in isolated southwestern mountain ranges at relatively high elevations. First collected in 1947 by the Sperrys, this cryptic species has subsequently been taken by others in the same general area.
This species is most likely to be confused with P.insulidens or perhaps even P.dribi. Although perhaps less reddish, the coloration of the forewings of P.albimontana is similar to that of the rather pale P.insulidens characteristic of northern perimeter Great Basin populations, and the reniform is somewhat more compressed laterally. Papaipema albimontana is a smaller species than P.insulidens, seldom exceeding 16.5 mm in forewing length. These distinctions are subtle however; the best distinguishing feature being that the ranges of these two species do not appear to overlap. The forewing maculation of P.albimontana is nearly identical to that of P.dribi, which is much darker brown and displays a greater tendency for the suborbicular spots to trend toward obsolescence. Genitalic differences between all three species are not definitive. It is more likely that apparent subtle differences (e.g., in the female lamella) reflect intraspecific variation rather than actual species-level distinguishing features.
Head: antennae of both sexes ciliate, laminate, less deeply in female; dorsum scaled; ventral lamination obscured by densely pubescent sensilla; eye smooth, rounded; haustellum unreduced; labial palpus laterally flattened, upcurved; first segment swollen basally, arching slightly upward, slightly more than half as long as second segment; second segment straight, about as long as eye width, scales broad; third segment ⅓ × length of second segment, and projecting slightly anteriorly, scales narrow; frons unmodified, smooth, convex, with unspecialized conical vestiture consisting of spatulate scales mixed with fine, hairlike scales. Thorax: vestiture of prothoracic collar, patagia, mesothorax, and metathorax concolorous brown with admixed yellowish scales, collar with fringe of paler, contrasting, yellowish scales; vestiture a mixture of coarse spatulate, thin apically spatulate, and hairlike scales; transverse, wedge-shaped, prothoracic crest prominent, mesoscutellar tuft moderate, divided, metascutellar tuft darker. Forewing: length = 14.2–16.5 mm; dorsal surface ground in both sexes uniformly dull orange-brown; basal spots yellowish cream, highly contrasting with darker antemedial band; antemedial lines double, concolorous with medial area, sinuous until strongly bowed outward just before meeting inner margin; postmedial lines double, the outer heavier and darker, excurved around reniform and meeting inner margin diagonally; medial shade defined by darker scales, touching lower loop of reniform, then angled below reniform and meeting inner margin diagonally; subterminal line scalloped, double, outer line darker; subterminal space filled with plumbeous-gray scales defined outwardly by series of darker, charcoal chevrons filling the spaces between veins; space between subterminal area and outer margin of same lighter color as medial area, brightest apically; terminal line obscure; fringe dark fuscous; reniform relatively small, narrow, and laterally somewhat compressed, with small, white satellite spots surrounding thin, yellow, central lunule; orbicular and suborbicular stigmata white. Hindwing: dorsal surface dull fuscous, darker beyond disc; discal spot and medial shade obscure in most; subterminal area lighter as in disc, terminal line dark, and fringe light like subterminal area. Abdomen: basal coremata and pockets absent in male; dorsal tufts on A1–A3 variably prominent, that on A3 largest. Male genitalia: uncus narrowly spatulate, apically acuminate with sparse, dorsal, cristate patch of recumbent setae; tegumen rounded at base of uncus, flaring laterally and forming rounded peniculi, largely devoid of hairs, before narrowing at s-shaped pleural sclerite and articulating with v-shaped vinculum; juxta + anellus elongated, height 3 × width, with divergent anellar arms; ventral margin of valve evenly convex at base, then slightly sinuous until angled dorsally into notch at anteroventral edge of cucullus; base of sacculus rounded, swollen, with dense patch of sensory hairs on triangular process at base of costal lobe, hairs continuing along dorsal margin of sacculus; dorsal margin of valve concave basally, then nearly straight to moderately broad neck of cucullus; cucullus bifurcate with dorsal arm broadly rounded, 2.5 × width of narrow ventral arm terminating in acutely-pointed, recurved tip; entire inner face of cucullus covered with dense patch of proximally-oriented, spine-like setae; corona absent; clasper an appressed, sclerotized, crescent-shaped sclerite; ampulla absent; digitus heavily sclerotized, serrate or somewhat dentate on dorsal edge, sinuous, acuminate, extending halfway to ventral margin of valve; phallus slightly curved, with terminal carina consisting of two opposing, subequal, sclerotized, ribbon-like, spined bars extending onto subbasal part of vesica; vesica with single, prominent, blunt, rose-thorn cornutus on dorso-mesial diverticulum. Female genitalia: papilla analis dorsoventrally flattened, lyriform, with lateral margin prominently constricted toward anterior end, narrowly convex toward posterior end, acute apices slightly divergent, with long setae set in tiny membranous circles on both dorsal and ventral surfaces; segment A8 2.5 × as wide as long, anterior apophysis 2 × length of segment A8 and posterior apophysis 2.8 × length of segment A8; lamella antevaginalis heavily sclerotized, laterally flared, hemi-cordate, with deep cleft on convex posterior rim, anterior portion of lamellar plate extended into a long, acute point, overall shape of lamella funnel-like; ostium moderately sclerotized; ductus bursae narrow, about as long as appendix bursae, heavily sclerotized; appendix bursae membranous with slight rugosity, about as long as and half the width of corpus bursae, viewed ventrally, oriented left, ductus seminalis at posterior apex; corpus bursae membranous, ovate, with two signa.
Only one specimen of P.albimontana has been DNA-barcoded to date, yielding a haplotype shared with both P.birdi and P.insulidens.
The larva is a member of the most common group having only its middorsal longitudinal white line crossing the dark pigmented bands encircling segments A1–A4, the subdorsal lines being interrupted on these segments. It is indistinguishable from other larvae in the P.insulidens group. The lateral margins of the honey-colored prothoracic shield bear the usual dark, nearly black, pigmentation stripes which may extend onto the darker, chestnut-brown head capsule. The dorsal pinacula on A1–A7 are about average in size for a Papaipema larva; those on A8 are much larger, heavily sclerotized, and have dark pigmentation. The D1 and D2 pinacula on A8 are semi-ovoid and appressed to the white, middorsal line; the pinacula on A9 are dark brown to almost black and are slightly larger than those on A8. The chestnut anal shield has dark lateral margins.
TYPES. Holotype: ♂. Forewing length 16.5 mm [alar expanse 36.0 mm]. 1 mi. s. Greer, Little Colorado River floodplain, 33.99279, -109.4653 8,500 ft., Apache Co., Arizona; ex larva June 27, 2023; A. E. McBride, collector. Collected from Cicuta douglasii and reared in carrot; eclosed August 3, 2023. [CNC] Paratypes: 15 ♂, 15 ♀. Arizona. Apache Co., White Mts., Diamond Rock; 30 Aug–3 Sep 1947; Grace H. and John L. Sperry (3 ♂, 2 ♀). Apache Co., White Mts., Greer, 8,300´ [2,530 m]; 11 Sep 1966; Ron Leuschner (3 ♂). Apache Co., Greer; 6 October 2005; J. Vargo (3 ♂). Apache Co., elev. 6930´ [2,112 m], N34°8.689´, -W109°17.649´, Becker WMA off Hwy 60; to BL 11 Aug 2012; D. L. Wickle, DNA barcode voucher # CNCLEP 00120015 (1 ♂). Apache Co., 1 mi. s. Greer, edge of Little Colorado River, 33.9945, -109.4647 8,481´ [2,585 m]; ex larva 24 Jun 2022, A. E. McBride, collector; collected from Cicuta douglasii and reared in carrot, eclosed 30 Jul–2 Aug 2022 (4 ♂, 2 ♀). Apache Co., 1 mi. s. Greer, Little Colorado River floodplain, 33.9928, -109.4653 8,500´ [2,591 m]; ex larva 27 Jun 2023, A. E. McBride, collector; collected from Cicuta douglasii and reared in carrot, eclosed 27 Jul–15 Aug 2023 (1 ♂, 11 ♀). AMNH, CMNH, CNC, MCLB, NHML, UCD, USNM, YPM, AEM, ELQ, JVC.
In 2022 with JRW and again in 2023, AEM found P.albimontana larvae at Greer, Arizona in the White Mountains area of Apache County at 2,600 meters boring in western water hemlock (Cicuta douglasii) that was growing in large clumps right out of the water on the edges of the Little Colorado River. The larva was also encountered using Delphinium spp., which is often found in the same habitats. Larvae are readily detected by the presence of wilting, yellowing, or flagging foliage in western water hemlock. Entire stems may be bent over at ground level if larval boring is concentrated at that point. Lateral branches may also be bent at the point where internal girdling of the hollow stems has weakened them. In late June and early July, the larva may be found boring at any point along the main stem of the plant, and frass is easily visible just below the frass orifice. Eventually the larvae likely concentrate their efforts in the toxic roots. Based on the rather limited sample, adults have been recorded from mid-August through early October, when they still appeared to be in fresh condition.
Like P.dribi which thus far is known only from a highly isolated metapopulation in the Sacramento Mountains of New Mexico, P.albimontana appears to be another isolated entity that is restricted to the White Mountains of Arizona.
Papaipema sauzalitae GROUP
This group appears to represent another relatively recent derivation within Papaipema and may be most closely related to the P.insulidens group. Both the P.insulidens and P.sauzalitae groups retain an ancestral larval preference for Apiaceae. Included taxa are: Papaipema sauzalitae, P.angelica, P.harrisii, P.rubiginosa, and P.astuta. The two West Coast species, P.sauzalitae and P.angelica, have a modified frons and have lost the spines on their digiti. Their ancestral lineage could possibly be of East Asian origin via Beringia. These two species appear to represent an older lineage than the remaining three members of the group and may not be as closely related to the eastern subgroup as their appearance suggests.
The Papaipema harrisii subgroup appears to represent a complex of three sibling species, two previously masquerading under one name, which probably diverged from one another during the Pleistocene glaciations. The three species of this subgroup are Papaipema harrisii, P.rubiginosa, and P.astuta. Nominate P.harrisii is a boreal species. Its native host plant, Heracleum maximum (Apiaceae), occurs in a variety of mesic habitats including woodlands, forest openings, and grasslands, as well as in riparian areas such as wet meadows, stream terraces, alluvial benches, floodplains, and stream and lake margins. The distribution of nominate P.harrisii suggests its immediate ancestral lineage survived the Pleistocene glaciations isolated on coastal (now offshore) refugia to the east, along the Gulf of St. Lawrence or off the Grand Banks of Newfoundland.
A sibling species, P.rubiginosa, occupies a more southerly distribution and is associated with open, wetland marshes and calcareous fens where its primary host plant purplestem angelica (Angelica atropurpurea), also Apiaceae, frequently grows in dense colonies. It is a hollow-stem borer, and the larva never enters the root. AEM discovered a highly disjunct population in two fen complexes in northwestern New Jersey, and the historic capture by Otto Buchholz in 1909 in Hudson County, New Jersey suggests it was formerly more widespread in that state; otherwise, populations are scattered across the Great Lakes region, from the western environs of Lake Ontario and the historic occurrence at Buffalo, New York on Lake Erie, to areas farther west immediately encompassing the remaining Great Lakes south of Lake Superior. These Angelica-feeding populations apparently reach their northern limits along an ecotone wherever temperate deciduous forests contact lower boreal forests dominated by the nominate Heracleum-feeding populations. Some adults from these neighboring populations can be nearly indistinguishable. The biogeographical insights that may be gleaned from these observations suggest that, during maximum glacial periods, these Angelica-feeding populations probably shared a habitat and distribution very similar to those of the few remaining present-day populations of the southern wetland-associated satyrine butterfly Neonympha mitchellii French, isolated in a chain of widely scattered, relict pockets from Mississippi and Alabama to North Carolina. Elements of this familiar pattern are repeated in distributions of other wetland-associated satyrines such as Satyrodes eurydice (Linnaeus) and S. appalachia (Chermock), and more frequently with a number of skipper species which inhabit similar environments. Poanes viator (Edwards), P. massasoit (Scudder), Euphyes bimacula (Grote & Robinson), E. conspicua (Edwards), E.dukesi, E. dion (Edwards), E. alabamae (Lindsey), as well as some other Papaipema species (e.g., P.appassionata) exemplify this phenomenon. What all these taxa appear to have in common is a biogeographic history that is shared to varying degrees. Post-Wisconsin Glaciation recolonization of northeastern North America and the Great Lakes region from southern glacial refugia via the Hudson-Mohawk corridor provides the mechanism to account for today’s observed patterns. The familiar Atlantic Coastal Plain – Great Lakes disjunction pattern has been well documented (
The third sibling species in this complex is Papaipema astuta Bird. Its ecological requirements are dramatically different. It is a denizen of rich, damp woodlands where its host plant, Collinsonia canadensis (Lamiaceae), grows. Its larva bores in the lower portion of the slender stem and then enters the woody root where it completes its development, in stark contrast to the larval behavior of the two previous entities. Papaipema astuta populations show remarkable uniformity among individuals, and lack the dark phenotypes commonly expressed in populations of the Angelica-boring sibling, that has larger adults, on average. The present-day distribution of P.astuta suggests its ancestral lineage survived the maximum Pleistocene glaciations somewhere in the interior, perhaps well to the south of the Appalachians, but far removed from the Atlantic Coast. Upon recolonizing its former range in the post-glacial period primarily west of the mountains, an isolated population apparently became displaced to the northeast.
All three species are morphologically indistinguishable by their genitalia, which is to be expected among recently diverged species. This is analogous to a number of other species in the genus which are not reliably diagnosed by genitalia. It was hoped that barcodes might shed light in such instances but thus far, data derived from CO1 analyses have not proven useful in this regard. However, as
Papaipema sauzalitae (Grote) (Pacific Coast Stalk Borer)
Ochria sauzalitae Grote, 1874, Preliminary Catalogue of the Noctuidae of California, Part II, The Canadian Entomologist, 6(11): 216-217.
Type locality: “Sauzalito, Cal.” [Sausalito, California, USA]. [NHML]
note—Grote very briefly described this species in part II of his preliminary list of the Noctuidae of California, apparently based upon two available specimens, citing “Sauzalito, September 17th, Mr. Behrens, No. 161; Hy. Edwards, Esq. No. 135.” No sex of the syntypes is mentioned. A female in the NHML bears Grote’s original, red-bordered label with the words “ Gortyna sauzalitae Grote Type” in his own handwriting which must be considered the authentic holotype. Additional labels on the specimen were added subsequently, likely by Hampson’s assistant, in the same manner and handwriting as those on some other spurious types at NHML. Curiously, one of the added labels reads “California Sauzalito. 17.IX.77.” However, a date of 1877 is clearly impossible. It is also perplexing why Grote labelled the holotype as a Gortyna but described it as an Ochria.
Gortyna purpurifascia; Grote, 1873, in part, not Grote and Robinson, 1868. Gortyna sanzalitae; anonymous, 1882, misspelling. Papaipema sauzaelitae; authors, misspelling. Papaipema erubescens Bird, 1911, The Canadian Entomologist, 43: 39.
Type locality: San Francisco, California, [USA]. [AMNH]
note—Described from eleven males reared by F. X. Williams from larvae boring in Cirsium occidentale.
† Emboloecia sauzalitae form papaipemoides Barnes and Benjamin, 1929, Bulletin of the Brooklyn Entomological Society, 24: 180.
Type locality: San Francisco, California, [USA]. [USNM]
note—Described from two males and one female. This is merely typical P.sauzalitae with yellow instead of white filling of the reniform and orbicular stigmata, as indicated in the original description, and is therfore an unavailable name.
This is primarily a coastal species which one may encounter within sight of the Pacific Ocean, although the inland populations of southern California extend eastward nearly to Nevada. Adults from these inland populations tend to be much paler than their coastal counterparts, reflecting the more xeric environments inland. Papaipema sauzalitae bears a strong superficial resemblance to P.harrisii, although their distributions are completely allopatric. The forewings of P.sauzalitae are more acutely pointed, nearly falcate in comparison. The smooth digitus in the male genitalia and the ridge-like protuberance on the frons of both sexes of P.sauzalitae will definitively separate it from P.harrisii; for distinctions from P.angelica, see the discussion there. DNA barcodes are available for two specimens of P.sauzalitae. These form a discrete group and unique BIN closest to P.angelica, with 1.7% minimum divergence.
The larvae are typical of the group with only the pale middorsal line remaining continuous across A1–A4. Additional larval characters were provided by
The first reported host plant for this species was cobweb thistle (Cirsium occidentale), under the name P.erubescens, by
This species has been recorded in northern Washington, almost to the Canadian border, south to the Los Angeles area, and east as far as Inyo County, California.
Papaipema angelica (Smith) (Leather-root Borer)
Hydroecia angelica Smith, 1899, Transactions of the American Entomological Society, 26: 40.
Type locality: Los Angeles Co., California, [USA]. [USNM]
note—Described from a single male reared by D. W. Coquillett.
This species was named for the locality it was first discovered, Los Angeles County, California, and not for a host plant (Angelica) often utilized by members of this group. The moth shares two unique, apomorphic features with its very close relative P.sauzalitae. It is possible that P.angelica merely represents the southernmost population of a P.sauzalitae cline, but barcode data suggest otherwise. Both P.angelica and P.sauzalitae have a smooth digitus on the male valve, lacking the dorsal tooth-like serrations typical of and unique to Papaipema species. Additionally, both exhibit a vertical ridge-like projection on the frons, a feature which was given so much weight by
Papaipema angelica is superficially indistinguishable from P.sauzalitae, except that the forewing stigmata lack the white filling which most P.sauzalitae specimens exhibit. Specimens of P.angelica average slightly larger than most P.sauzalitae, but having fed as larvae on a much more robust host could easily account for this difference. Indeed, P.sauzalitae is known to feed upon a range of unrelated hosts, and leather-root could simply represent a local choice. Papaipema angelica and P.sauzalitae are not synonymized herein due to the possibility that they are indeed genetically distinct, but until fresh specimens are found for additional molecular analysis, the matter will remain unresolved. Since only historical specimens are available of P.angelica, DNA barcode samples are degraded, with two sequences of incomplete length (540 base-pairs). However, these differ significantly from those of P.sauzalitae, the nearest-neighbor, at 1.8% divergence.
Details of larval morphology have not been fully documented, other than Bird’s statement that “mature larvae apparently are in full accord with the major group;” i.e., those having only the middorsal line crossing the first four abdominal segments. He also noted that the mature larva was reminiscent of P.furcata, and that the pinacula were blackish and unusually well-defined for the stage.
The only documented host plant utilized by larvae of P.angelica is large leather-root (Hoita macrostachya). The plant perhaps attained its best development in brackish coastal marshes in the greater Los Angeles area where the moth was first discovered by Coquillett in 1882. Numerous additional specimens were subsequently secured by local collectors who found the pupae in the large, hollow stems of the plants, which grew in large clumps with individual plants up to two meters tall. No records of specimens collected since the 1930s are known, and it seems very likely that such coastal wetlands have long since been lost to development. If P.angelica still survives anywhere, it would likely be in any remaining isolated interior wetlands or perhaps in montane riparian situations in southern California.
Papaipema angelica has only been recorded from Los Angeles, Orange, and San Bernardino Counties, California.
Papaipema harrisii (Grote) (Harris’s Stalk Borer)
Gortyna harrisii Grote, 1881, United States Geological and Geographical Survey of the Territories, 6(2): 268.
Type locality: Kittery Point, [York Co.], Maine, [USA]. [NHML]
note—Apparently described from three specimens, although this is unclear in Grote’s original description as he lists simply “♂♀.” He further states: “Expanse, 36 mil. Massachusetts, reared by Mr. Roland Thaxter.” Grote also indicated that “this species is represented in the Harris collection by two specimens under the number 361.” It may have been that Grote had a male in his possession and that he had seen two additional specimens in the Harris collection by the time of description, for there are three apparent syntypes, one male and two females, in the NHML. These three specimens all agree in measuring 36 mm and indeed appear to have been reared, as they show little wear despite their age. It must be noted however that all three specimens have been relabeled as having originated from Kittery Point, Maine. No Massachusetts specimens of that age have been found in either the NHML, the MCZ where the Harris collection is deposited, or elsewhere. Among his other writings, Grote mentioned having seen a variety of Papaipema species taken by Thaxter in Massachusetts, and it is known that Thaxter’s material originated from both Massachusetts and Kittery Point, Maine. For this reason, it must be assumed that Grote erred when he wrote that Massachusetts was the source. It is further apparent that when the specimens were relabeled prior to Hampson’s 1910 publication of his catalog, this error was corrected, perhaps based on some new information that was available at that time. As discussed elsewhere in this fascicle, Hampson’s assistant routinely relabeled specimens, frequently creating spurious types. In this instance the three specimens deposited in the NHML appear authentic, as the male additionally bears Grote’s original label. Inasmuch as Grote’s apparent error remains an assumption, and the unlikely possibility that other syntype material from Massachusetts might still exist somewhere, it is necessary to select a lectotype from the three NHML specimens. The male with Grote’s original label is hereby designated lectotype. It bears the following labels: “Maine Kittery Pt. 20.VIII” / “Grote Coll. 81–116” / “22” / “Type” / “ Gortyna Harrisii ♂ Type Grote” [The last label has a red border and is in Grote’s handwriting].
Hydroecia verona Smith, 1899, Transactions of the American Entomological Society, 26: 32.
Type locality: Winnipeg, Manitoba, [Canada]. [USNM]
note—Described from its unique male holotype.
Gortyna anargyrea Dyar, 1908, The Canadian Entomologist, 40: 79.
Type locality: Colorado, [USA]. [USNM]
note—Described from a single male. The exact locality was unknown to Dyar (collection of Wm. Schaus).
Papaipema impecuniosa; Hampson, 1910 part, not Grote, 1881. † Papaipema harrisi ab. mulieris Strand, 1915 [1916], Archiv für Naturgeschichte, 81A (11): 163.
Type locality: “Vereinigte Staaten” [Kittery Point, York Co., Maine, USA]. [AMNH]
note — Unavailable infrasubspecific name. Described as an aberration of P.harrisii from a figure in
Papaipema anargyria; Bird, 1944, misspelling. Papaipema harrisi; authors, incorrect subsequent spelling; unavailable, per ICZN Article 33.4.
Papaipema harrisii has the most boreal distribution of this group and is in fact among the most boreal species of the entire genus. As discussed above, the long-standing concept of P.harrisii has heretofore included another taxon, Papaipema rubiginosa. Distinguishing P.harrisii from P.rubiginosa is sometimes challenging as discussed under that species. The completely allopatric distributions of P.harrisii and P.astuta precludes confusion of these two otherwise very similar species. It is also possible that P.harrisii could be confused with P.arctivorens in parts of northeastern North America where their distributions overlap. The postmedial line of P.harrisii is nearly straight until sharply angled inwards just before meeting the costa; in P.arctivorens, the postmedial line is broadly rounded and evenly excurved beyond the reniform before it angles inward to the costa. Currently included in this concept of P.harrisii are populations in New Brunswick and eastern Québec which appear distinctive (
The DNA barcodes of P.harrisii are part of a large, complex cluster of haplotypes that involves not only members of the P.harrisii group, but also the P.pterisii group. All species in these two groups exhibit haplotype complements that are partially shared by at least one other species in either group, although species are intercalated among barcodes that range between 0 – 1.6% divergence. The haplotype complement of P.harrisii is partially shared by P.hesseli and P.rubiginosa. From a strictly taxonomic perspective, barcodes thus serve only to obscure the boundaries between species established as ecologically and biologically distinct. This high degree of mitogenomic similarity does however underscore the exceptional evolutionary plasticity of this section of Papaipema, where speciation appears to be tracking a recent dichotomy of specialization on two vastly different plant groups, ferns and umbellifers.
The larva is visually indistinguishable from those of the other members of the P.harrisii subgroup, all having the black pigmentation of the lateral margins of the prothoracic shield extending onto the head capsule as dark-pigmented lateral stripes, varying from prominent to obscure among individuals. Only the middorsal white line is continuous across the pigmented bands encircling the first four abdominal segments.
The preferred native host plant of P.harrisii is cowparsnip (Heracleum maximum). It has been reared from wild parsnip (Pastinaca sativa), an introduced invasive species, on Mt. Katahdin, Maine. ELQ found P.harrisii larvae in abundance boring in the introduced common hogweed (Heracleum sphondylium) on Cape Breton Island, Nova Scotia. Evidence of larval activity within the stems and thick petioles of the host plant is of the usual nature, including wilting of the foliage and presence of whitish frass. Pupation can occur either within the larval gallery or externally in a cell formed in soil. Adults have been recorded flying from early August through the third week of October.
Nominate P.harrisii is a boreal species found from Newfoundland and coastal maritime areas to its terminal southern historic occurrence on Long Island, New York. Westward in the interior, it ranges across Canada through the prairie provinces to northeastern British Columbia and south in the Rocky Mountains through Montana to central Colorado. Western populations were given the name verona but do not differ in any way from maritime populations of P.harrisii.
Papaipema rubiginosa Bird, REVISED STATUS (Angelica Borer)
Papaipema rubiginosa Bird, 1911, The Canadian Entomologist, 43(2): 40.
Type locality: Buffalo, [Erie Co.], New York, [USA]. [AMNH]
note—Described from 32 reared specimens. Although
This is the largest member of the sibling complex to which it belongs, with a forewing length ranging from 15.3–19.9 mm and averaging larger than either P.astuta or P.harrisii. Individuals of P.rubiginosa exhibit a much wider range of phenotypic variability compared to that of both P.harrisii and P.astuta, as these two species are consistently uniform in appearance (excepting the New Brunswick and eastern Québec populations mentioned above under P.harrisii). Even so, Papaipema rubiginosa is easily confused with both P.harrisii and P.astuta. Its populations lie sandwiched between the distributions of both siblings and can overlap with either. None can be distinguished by genitalic characters. The coloration of the forewing medial area of rubiginosa tends to be more brown, relative to the orange-yellow of both the smaller P.harrisii and P.astuta. The darkest P.rubiginosa are readily distinguishable from the consistently lighter P.astuta and P.harrisii. The palest, yellowish individuals of P.rubiginosa are problematic, and are most reliably identified by habitat and host association. The spots comprising the orbicular-reniform complex may be either white or yellow in all three species but are occasionally replaced by dark stigmata in P.rubiginosa.
The variability observed in P.rubiginosa is both most extreme and dramatic in those populations farthest isolated from its sibling P.harrisii; i.e., the New Jersey populations, the historic Buffalo, New York population, and the northern Illinois and southern Wisconsin populations, which all appear much alike. Individuals in P.rubiginosa populations from the southern counties of Michigan as well as the historic populations from the Chicago area tend to be more uniform in appearance, paler, and much more difficult to distinguish from P.harrisii, suggesting that genetic drift may be responsible there. Many specimens of P.rubiginosa reared by collectors in the general environs of Chicago, Illinois in the early twentieth century are widely distributed in institutional collections today under the name P.harrisii. Dark phenotypes from the Buffalo area represent nominate P.rubiginosa. Similar dark individuals with either light or dark stigmata (plate A-7, figures 2 and 4) regularly occur in populations of P.rubiginosa from New Jersey to Wisconsin but have not been observed in the boreal populations of nominate P.harrisii or in populations of P.astuta.
DNA barcodes are available for ten samples from southern Ontario and Michigan. Some haplotypes are shared with P.harrisii, and the complex barcode variation of this entire group is discussed in the P.harrisii account.
The larva belongs to the group with only the middorsal longitudinal white line crossing the dark pigmented bands encircling segments A1–A4, the subdorsal lines being interrupted on these segments. The lateral margins of the honey-colored prothoracic shield bear the usual dark, nearly black, pigmentation stripes which commonly extend onto the darker, chestnut-brown head capsule. The small, nearly black, dorsal pinacula on A1–A7 are about average size for a Papaipema larva; those on A8 are much larger, heavily sclerotized, with pigmentation varying from honey-colored to almost black. The D1 and D2 pinacula on A8 are round and well separated; those on A9 are larger, more elliptical, but also well separated, honey-colored to almost black. The honey-colored anal shield bears dark lateral margins.
The predominant host plant is purplestem angelica (Angelica atropurpurea), but also recorded are other apiaceous plants such as cowparsnip (Heracleum maximum) and the introduced wild parsnip (Pastinaca sativa), as well as occasional composites such as burdock (Arctium) and Indian plantain (Arnoglossum) in situations where such plants coexist with purplestem angelica in wetland fens. Henry
Papaipema rubiginosa is associated with calcareous fens distributed across a relatively narrow latitudinal zone from northern New Jersey west to southern Ontario, northern Iowa, and eastern Minnesota. Its northern limits are unclear, due to the difficulty of distinguishing it from neighboring Heracleum-feeding populations of P.harrisii, with which it may hybridize. The distribution pattern of P.rubiginosa at least partially mirrors those of several satyrine butterflies and a number of rare or declining eastern North American Hesperiidae. The Great Lakes fen populations of Hemileuca maia complex are also replicated at the New Jersey P.rubiginosa sites. The distribution of the nominate subspecies of the federally endangered Neonympha mitchellii nearly exactly matches that of P.rubiginosa, as it also occurred historically in northern New Jersey fens. A thriving colony of P.rubiginosa boring in purplestem angelica occurs within sight of a protected N.mitchellii locality in Cass County, Michigan, and a few hundred meters from this purplestem angelica population in the marsh, a small population of the sibling species Papaipema astuta occurs in the nearby woods, boring in stoneroot (Collinsonia). These two populations could potentially interbreed.
Papaipema astuta Bird (Stoneroot Stem Borer)
Papaipema astuta Bird, 1907, The Canadian Entomologist, 39: 272.
Type locality: [Rye, Westchester Co., New York, USA]. [AMNH]
note—The type locality was not given in the original description, other than Bird’s mention that he discovered the species on his “home preserves,” which were at Rye, New York. Accompanied by his son Junius, ELQ walked in a small wood on the Bird estate, and indeed found the host plants still growing there in the 1970s. Described from a single female, expanse 31 mm.
Some individuals of P.astuta are nearly indistinguishable from P.rubiginosa, especially in those limited areas where distributions of these two species overlap. In such instances, adults need to be reared from known host plants for confirmation of identity, and hybridization remains a possible factor as well. Papaipema astuta has a more southerly, largely allopatric distribution, and its average size is normally smaller than P.rubiginosa. The general appearance of P.astuta is consistently uniform, lacking the significant phenotypic variability seen in P.rubiginosa. There is also a tendency for the subterminal area of the forewing of P.astuta to be more solidly dark than that of P.rubiginosa, contrasting strongly with the yellow medial area. The forewing stigmata are most commonly yellow, and occasionally white. Papaipema harrisii is also extremely similar in appearance to P.astuta, but its forewing stigmata are never yellow, except for certain individuals from western populations, and these species are completely allopatric. Four DNA barcode samples from Michigan, Kentucky, and New Jersey, although very similar to the P.harrisii-P.pterisii barcode cluster (0.8% divergence) discussed in the P.harrisii account, nevertheless form a discrete haplotype group.
The larva is typical of the largest group of Papaipema which have only the single middorsal white line continuous across the dark-pigmented bands encircling the first four abdominal segments. Papaipema astuta is a specialist borer of stoneroot (Collinsonia canadensis), throughout its range. AEM, Loran Gibson, and Ellis Laudermilk found P.astuta boring in the small, woodland-dwelling filmy angelica (Angelica triquinata) in Harlan County, Kentucky, which was found growing in the same habitat as stoneroot plants infested with P.astuta larvae. This likely represents an instance of reversal to an ancestral adaptation to, and preference for, apiaceous hosts. The host plant is bored from a point several centimeters above the ground into the hard, flattened, woody root where some of the larval development occurs after having consumed the available resources above ground, but boring in the hard root does not occur to the extent seen in P.duplicata. Infested plants exhibit wilting, discoloration, and drying; whitish frass may be seen accumulated on soil beneath the bore hole in the plant’s stem. Pupation occurs in a shallow cell in soil or detritus near the base of the plant. The adult flight period extends from late August through September into early October.
This generally uncommon species occurs from Connecticut to Illinois and south in the mountains to western North Carolina. It tends to be overlooked unless specifically sought in its preferred habitat.
Papaipema leucostigma GROUP
This group consists of Papaipema leucostigma, P.lysimachiae, P.pterisii, and P.hesseli. It is possible that the group may share a distant phylogenetic relationship with the Papaipema speciosissima group inasmuch as the last two members have either retained or reverted to a fern-feeding specialization and show similarities in the peculiar nature of the adult forewing orbicular and suborbicular maculation. Perhaps more likely is a closer connection to some members of the Papaipema sauzalitae group, as supported by molecular data.
Papaipema leucostigma (Harris) (Columbine Borer)
PL. A-7, FIGS 21–23 (adult);
Gortyna leucostigma Harris, 1841, A Report on the Insects of Massachusetts, Injurious to Vegetation, 320.
Type locality: [Massachusetts, USA]. [MCZ]
note—Described from a single female.
Gortyna purpurifascia Grote and Robinson, 1868, Transactions of the American Entomological Society, 1: 341.
Type locality: “Atlantic District. (Mass.!)” [Massachusetts, USA]. [Type lost]
note—This taxon was described from one male and one female. The female syntype was described as having an alar expanse of 43 mm and a body length of 20 mm. The description of this specimen perfectly fits leucostigma in all details; however, this syntype is lost. The female syntype was illustrated on
Papaipema pterisii; Hampson, 1910, not Bird, 1907. † Papaipema pterisii ab. luteipicta Strand, 1916, Neue Aberrationen der Noctuiden-Subfamilie Acronyctinae, 163.
Type locality: Newtonville, Massachusetts [USA]. [NHML]
note — Unavailable infrasubspecific name. The male holotype specimen is missing its abdomen and is actually a specimen of P.leucostigma.
In the sense that it was the first named, this colorful species has the distinction of being the oldest member of the genus. It is perhaps fitting that the holotype resides in America’s oldest university, the Museum of Comparative Zoology at Harvard. It is also very likely that this holotype may also be the oldest surviving Papaipema specimen in existence. The type specimen emerged September 24, 1834 from a pupa formed the 19th of August. The larva had been found boring in a columbine root by Thaddeus W. Harris earlier in July of that year. This is documented in his 1841 publication, making that the earliest life history report for a Papaipema species.
As evident from Grote and Robinsons’ conflation of two species under the name P.purpurifascia, P.leucostigma and P.lysimachiae have also been confused by others. Aside from the significant difference in size, these two species share a particularly bright and similar pattern of forewing coloration that renders them unlikely to be confused with other Papaipema species, but more likely to be confused with each other. Papaipema lysimachiae rarely if ever approaches the size of even the smallest P.leucostigma. A consistent difference in forewing maculation readily distinguishes the two. In P.leucostigma, the lower white spot of the suborbicular is invariably the larger of the two, often ovoid, and also larger than the orbicular spot. In some specimens, including the holotype, the lower suborbicular is the only white spot on the entire forewing. In P.lysimachiae, the white orbicular spot is normally about the same size as the lower suborbicular spot, with a somewhat smaller but often obsolescent white spot in between. Perhaps less likely to be confused, the forewing coloration of P.pterisii is less bright orange, and its white suborbicular spots tend to be fused into a vertical bar, which is never seen in P.leucostigma. DNA barcodes are available for 18 specimens from Ontario and Minnesota. These form a discrete cluster with little variation, although with only 0.8% minimum divergence from the P.pterisii- P.harrisii barcode cluster.
The larva embodies all the features characteristic of the genus, and groups with those having only the middorsal white line unbroken across the first four abdominal segments. This line in P.leucostigma is thread-like and much narrower than usual, and in combination with the strong, uniformly pink pigmentation of all segments, gives the larva an appearance unlike that of any other.
This species is a monophagous specialist on columbine (Aquilegia). Larvae abandon boring activity in the stem early and proceed to the root where they complete their development by consuming the entire root except for a thin outer skin. Above-ground portions of the plant quickly wither and die, and ultimately the entire plant is usually killed. In wild columbine plants, which are often rather small and slender, this often necessitates moving to an additional nearby plant to secure adequate food. Cultivated garden varieties of columbine are sufficiently robust to produce adults larger than those that have fed upon wild plants. Evidence of boring activity is sometimes indicated only by dried frass on the ground where the plant had been, and the larva responsible for this damage may be located nearby boring in a new plant. Pupation occurs in a shallow cell in soil or detritus near the base of the plant bored last. The adult flight period begins in mid-August and peaks in September, when the majority of captures have been recorded.
The species is widespread in northeastern North America. It ranges as far west as Manitoba and as far south as Missouri, Kentucky, and West Virginia. It is generally uncommon but can be abundant in those situations where columbine growth is prolific.
Papaipema lysimachiae Bird (Loosestrife Borer)
PL. A-7, FIGS 19–20 (adult);
Papaipema lysimachiae Bird, 1914, The Canadian Entomologist, 46: 70.
Type locality: Harrison, Westchester Co., New York, USA. [AMNH]
note—Described from 20 specimens representing equal numbers of the sexes. See
Gortyna purpurifascia Grote and Robinson, 1868, Transactions of the American Entomological Society, 1: 342, in part, male.
Type locality: “Atlantic District. (Mass.!)” [Massachusetts, USA]. [ANSP]
note—One of the two syntypes of Papaipema purpurifascia is a male P.lysimachiae. See discussion under P.leucostigma.
The species most likely to be confused with P.lysimachiae is P.leucostigma. For distinguishing features, refer to the discussion under P.leucostigma. Worn specimens of P.pterisii can also be confused with P.lysimachiae. The two suborbicular spots of P.lysimachiae are sometimes merged or even fused, creating an appearance similar to the same spots in P.pterisii, which are normally fused into an elongate, vertical bar. The white satellite spots of the reniform of P.pterisii, especially those basad of the central lunule, are always a prominent feature of its forewing maculation. In P.lysimachiae, these white markings are either greatly diminished or completely absent. Papaipema pterisii is almost always significantly larger than P.lysimachiae. DNA barcodes are available for four specimens of P.lysimachiae from Québec and Ontario, which form a discrete group and unique BIN at a minimum nearest-neighbor difference of 1.35%.
The pigmented portions of the integument and especially the first four abdominal segments of the larva are dark brown, and only the middorsal white line interrupts this characteristic Papaipema feature. The larva is very slender, as necessitated by its occupancy of a very thin root. For a detailed account of the early stages, including parasitoids, see
This smallest member of the genus is unusual in that it occurs in two quite different habitats. Like the other specialists in the genus, it is restricted to the habitat of the host plant. In this instance, the two common host plant species in the genus Lysimachia utilized by the borer grow in different situations. Whorled loosestrife (L.quadrifolia) occurs in poor, dry, upland soils, and swamp candles (L.terrestris) is a denizen of wetland habitats. Presence of the larvae is readily detected by wilted or browned plants which stand out among the healthy plants surrounding them in what is often a rather extensive colony of the host. The hole in the lower stem and the presence of frass on the soil confirms that a larva has bored the root and killed the plant. The roots are quite slender which accounts for the diminished size of this species.
The species is moderately common, ranging from Nova Scotia west to North Dakota and south in the Appalachians as far as Georgia.
Papaipema pterisii Bird (Bracken Borer)
PL. A-7, FIGS 13–18 (adult);
Papaipema pterisii Bird, 1907, The Canadian Entomologist, 39: 310.
Type locality: [Rye, Westchester Co., New York, USA.] [AMNH]
note—Described from three specimens, their sex not stated. No type locality was specifically mentioned, but it is clear from the accompanying text that the type specimens were collected within the immediate vicinity of the author’s home at Rye, New York.
Gortyna triorthia Dyar, 1908, The Canadian Entomologist, 40: 79.
Type locality: Holderness, [Grafton Co.], New Hampshire, [USA.] [USNM]
note—Described from a single female, although Dyar associated the holotype with additional specimens from various localities.
Papaipema pterissi; Moore, 1955, misspelling.
Papaipema pterisii can be difficult to distinguish from its close relative P.hesseli, which however occurs in very different habitats. See the discussion under that species for distinguishing features. Although less likely, in the northern portions of its range, P.pterisii might also be mistaken for P.harrisii. The most readily recognizable distinction lies in the forewing maculation, wherein both suborbicular spots are more or less fused into a narrow, nearly vertical bar which is almost confluent with the narrowed, vertical orbicular spot in P.pterisii. In P.harrisii, the suborbicular spots are better separated and rounded or ovoid, and the orbicular spot is round or sometimes subtriangular. DNA barcodes that are confidently placed as P.pterisii (based on reared specimens) include about 15 specimens, most from Ontario. Although overall variation is considerable, nearly all haplotypes are shared at least with P.hesseli, which in turn are intercalated among several species with very similar barcodes – this includes all species of the P.pterisii group and P.harrisii group. As such, the barcodes of P.pterisii are not diagnostic.
The larva belongs to the common group wherein only the middorsal longitudinal white line crosses the pigmented bands encircling the first four abdominal segments. The only known host is western brackenfern, Pteridium aquilinum (Dennstaedtiaceae). Larval boring remains confined to a relatively small section of the stipe base for most of the development period, suggesting that perhaps like certain other Papaipema species, P.pterisii derives most of its nutrition by consuming nutrient-rich fluids rather than by masticating an extensive amount of plant tissues.
This familiar species occurs almost anywhere within its range that its host fern grows in abundance on poor soils. A notable exception regards its rarity in areas of the New Jersey pine barrens where its host grows in vast, near monocultures in the sandy soil. These areas are subject to frequent burns which heavily impact survival of the immature stages. The adult flight period lasts about six weeks, from mid-August to the end of September. This species is widespread from Newfoundland to Manitoba and south as far as North Carolina in the mountains.
Papaipema hesseli Quinter, NEW SPECIES (Hessel’s Fern Borer)
PL. A-7, FIGS 6–12 (adult);
Papaipema hesseli Quinter.
Type locality: Normans Kill, 30 m., 42°40’40”, -73°54’23” Albany Co., New York, USA. [CNC]
note—Named in memory of Sidney Adolphus Hessel, a most enthusiastic proponent for this genus who compiled a useful guide to finding the early stages, and who discovered and first brought this cryptic new species to my attention.
Given that evolution is a dynamic process, it is not surprising that flying alongside congeners representing the living descendants of ancient lineages dating back millions of years are relatively modern Papaipema species which may have diverged as recently as the Pleistocene or post-glacial Holocene epochs. Papaipema hesseli appears to be one such recently derived species, better defined by ecological association than by morphology or molecular data.
Some individuals of P.hesseli are practically indistinguishable from its nearest relative P.pterisii, with which it is broadly sympatric throughout its more limited distribution. However, the two species inhabit very different niches and are usually not found in immediate association. Papaipema hesseli is perhaps most easily recognized by its restriction to wooded swamps where its host plants grow in abundance. Papaipema pterisii is an upland species inhabiting drier environments where bracken grows. Papaipema hesseli is a larger species than P.pterisii, which has an average forewing length of 15.2 mm (n = 69). The average forewing length of P.hesseli is 16.7 mm (n = 114), although some females have forewing lengths of 18.5 mm and wingspans to 40.5 mm. The coloration of the forewing of P.hesseli is yellow in many, with a dusting or even heavy suffusion of overlying cinnamon-brown scales. A minority of individuals in some populations occur as quite distinctive, dark reddish-brown morphs (plate A-7, figure 11), analogous with those forms also occurring in some populations of P.rubiginosa. However, in P.pterisii, these overlying scales tend to be brighter orange-red and never as extensively suffused as in dark individuals of P.hesseli. A comparable morph of P.pterisii is not known to exist, with many hundreds of specimens having been examined from across its range. The tendency of the orbicular and suborbicular stigmata to become fused into a vertical bar, characteristic of fern-feeding Papaipema, is somewhat less pronounced in P.hesseli than in P.pterisii. The genitalia of the two species are indistinguishable. Papaipema hesseli might also be confused with members of the P.harrisii complex, although these are not likely to be found together. Both P.harrisii and P.rubiginosa are similar in size and appearance to P.hesseli, but in addition to their exclusive larval association with Apiaceae, may be most readily distinguished from P.hesseli by the characteristic nature of the orbicular and suborbicular maculation of the adult forewings. In P.hesseli, the white orbicular spot is vertically elongated rather than round, and lacks the dark central spot seen in many individuals of the other two. Additionally, the two bright white suborbicular spots are fused into a single, vertically oriented bar, similar to the vertical bar characteristic of P.pterisii and the other fern feeders.
Head: antennae of both sexes ciliate, laminate, less deeply in female; dorsum scaled; ventral lamination obscured by densely pubescent sensilla; eye smooth, rounded; haustellum unreduced; labial palpus laterally flattened, upcurved; first segment swollen basally, arching slightly upward, slightly more than half as long as second segment; second segment straight, about as long as eye width, scales broad; third segment ⅓ × length of second segment, and projecting slightly anteriorly, scales narrow; frons unmodified, smooth, convex, with unspecialized conical vestiture consisting of spatulate scales mixed with fine, hairlike scales. Thorax: vestiture of prothoracic collar, patagia, mesothorax, and metathorax concolorous brown with admixed yellowish scales, collar with fringe of paler, contrasting, yellowish scales; vestiture a mixture of coarse spatulate, thin apically spatulate, and hairlike scales; transverse, wedge-shaped, prothoracic crest prominent, mesoscutellar tuft moderate, divided, metascutellar tuft subequal, less prominent. Forewing: length = 15.2–19.8 mm, females typically larger than males; dorsal surface ground in both sexes variable, ranging from yellowish-orange to uniform dark reddish-brown, medial area heavily peppered with darker scales which contrast with underlying yellowish-orange scales; basal spots yellowish-orange, contrasting slightly with darker antemedial band, about same color as medial area; antemedial lines double, concolorous with medial area, nearly straight until strongly bowed outward just before meeting inner margin; postmedial lines double, the outer comprising darkest element of forewing pattern, nearly straight until just before meeting costa and meeting inner margin diagonally; medial shade ranging from obscure to defined by darker scales, touching lower loop of reniform, then angled below reniform and meeting inner margin diagonally; subterminal line scalloped, sometimes preceded by series of darker, charcoal chevrons filling spaces between veins, immediately followed by series of lighter, yellowish-brown scales, more pronounced apically but often obscure; subterminal space filled with plumbeous-gray scales; terminal line obscure; fringe normally dark fuscous; reniform narrow, laterally compressed, with small, white satellite spots surrounding thin, yellow, central lunule; orbicular and suborbicular stigmata white, often more prominent than the reniform stigma. Hindwing: dorsal surface dull fuscous, often darker beyond disc; discal spot and medial shade obscure in most; subterminal area, terminal line, and fringe undistinguished. Abdomen: basal coremata and pockets absent in male; dorsal tufts on A1–A3 variably prominent, that on A3 the largest. Male genitalia: uncus narrowly spatulate, apically acuminate with sparse, dorsal, cristate patch of recumbent setae; tegumen rounded at base of uncus, flaring laterally and forming rounded peniculi, sparsely covered in hairs, before narrowing at s-shaped pleural sclerite and articulating with v-shaped vinculum; juxta + anellus elongated, height 3 × width, with divergent anellar arms; ventral margin of valve evenly convex at base, then slightly sinuous until angled dorsally into notch at anteroventral edge of cucullus; base of sacculus rounded, swollen, with dense patch of sensory hairs on triangular process at base of costal lobe, the hairs continuing along dorsal margin of sacculus; dorsal margin of valve concave basally, then nearly straight to moderately broad neck of cucullus; cucullus bifurcate with dorsal arm broadly rounded, 2.5 × width of narrow ventral arm; entire inner face of cucullus covered with dense patch of proximally-oriented, spine-like setae; corona absent; clasper an appressed, sclerotized, crescent-shaped sclerite; ampulla absent; digitus heavily sclerotized, serrate or somewhat dentate on dorsal edge, sinuous, acuminate, extending almost to ventral margin of valve; phallus curved, with terminal carina consisting of two opposing, subequal, sclerotized, ribbon-like, spined bars extending onto subbasal part of vesica; vesica with single, prominent, blunt, rose-thorn cornutus on dorso-mesial diverticulum. Female genitalia: papilla analis dorsoventrally flattened, lyriform, with lateral margin prominently constricted toward anterior end, narrowly convex toward posterior end, acute apices slightly divergent, with long setae set in tiny membranous circles on both dorsal and ventral surfaces; segment A8 2.8 × as wide as long, anterior apophysis 2 × length of segment A8 and posterior apophysis 3 × length of segment A8; lamella antevaginalis heavily sclerotized, laterally flared, hemi-cordate, with deep cleft on convex posterior rim, anterior portion of lamellar plate extended into a long, acute point, overall shape of lamella funnel-like; ostium moderately sclerotized; ductus bursae narrow, about as long as appendix bursae, heavily sclerotized; appendix bursae membranous with slight rugosity, about as long as and half the width of corpus bursae, viewed ventrally, oriented left, ductus seminalis at posterior apex; corpus bursae membranous, ovoid, with two signa.
Comparison of barcodes from over 20 confidently identified (reared) specimens of P.hesseli from Ontario and New York shows that there is extensive haplotype sharing with its sister species, P.pterisii. Nonetheless, there appear to be haplotypes that are unique to either species. Barcode variation of P.hesseli is further discussed under P.harrisii.
The larva is a member of the most common group having only its middorsal longitudinal white line crossing the dark pigmented bands encircling segments A1–A4, the subdorsal lines being interrupted on these segments. It is indistinguishable from the larva of P.pterisii. The lateral margins of the honey-colored prothoracic shield bear the usual dark, nearly black, pigmentation stripes which commonly extend onto the darker, chestnut-brown head capsule. The small, nearly black, dorsal pinacula on A1–A7 are about average in size for a Papaipema larva; those on A8 are much larger, heavily sclerotized, with pigmentation varying from honey-colored to almost black. The D1 and D2 pinacula on A8 are ovoid and well separated; those on A9 are honey-colored to almost black, larger, elliptical, and fused. The honey-colored anal shield bears dark lateral margins.
TYPES. Holotype: ♀. Forewing length 19 mm [alar expanse 39.5 mm]. USA, New York, Albany Co., Normans Kill, 30 m., 42.40.40-73.54.23; 21 Sept. 1989, T. L. McCabe. Specimen ID CNCLEP 00116420. Barcode of Life Project, Leg removed DNA extracted. [CNC] Paratypes: 77 ♂, 34 ♀. Connecticut. Fairfield Co., Boardman Bridge; emerged 2–20 Sep 1955 ex larva in Matteuccia struthiopteris, S. A. Hessel, collector (2 ♂, 2 ♀). Same locality and collector, emerged 2 October 1956 ex larva in Matteuccia struthiopteris (1 ♀). Litchfield Co., Rt. 128 1.6 mi. E West Cornwall; ex larva in Matteuccia struthiopteris; emerged 29 Aug–11 Sep 1975, E. L. Quinter, collector (1 ♂ 2 ♀). Same locality and collector, emerged 2 Sep 1977 ex larva in Matteuccia struthiopteris (1 ♀). Massachusetts. Berkshire Co., Pittsfield; larvae collected 26 Jul 1958, emerged 2 Sep–5 October 1958 ex larva in Matteuccia struthiopteris, S. A. Hessel, collector (2 ♂, 5 ♀). Berkshire Co., Sheffield; pupa collected in soil 1–2 inches [2.5–5 cm] deep, ca. 5 inches [12 cm] from ostrich fern roots 15 Aug 1957; emerged 29 Aug–1 Sep 1957, S. A. Hessel, collector (2 ♀). Same locality and collector, ex larva collected 26 Jul 1958 in Matteuccia struthiopteris; emerged 7–23 Sep 1958, S. A. Hessel, collector (2 ♂ 1 ♀). New York. Albany Co., Normans Kill, 30 m., 42°40’40”, -73°54’23”; 1–21 Sep 1989, T. L. McCabe (44 ♂, 2 ♀). Same locality and collector, 27 Sep 1992 (2 ♂). Albany Co., Normans Kill; ex larva in Matteuccia struthiopteris, emerged 28 Aug–16 Sep 1994, E. L. Quinter, collector (3 ♂, 4 ♀). Albany Co., Guilderland, along Normans Kill w. Rt. 155, 42.6799, -73.9063 202´ [62 m]; 21 Jul 2013 ex larva in Matteuccia struthiopteris, reared in potato, eclosed 25 Aug–15 Sep 2013, A. E. McBride, collector (7 ♂, 7 ♀). Same locality and collector, 20 Jul 2014 ex larva in Matteuccia struthiopteris, reared in potato, eclosed 29 Aug–16 Sep 2014 (5 ♂, 6 ♀). Same locality and collector, 3 Jul 2016 ex larva in Matteuccia struthiopteris, reared in potato, eclosed 17–20 Aug 2016 (2 ♂). Columbia Co., New Lebanon, Shaker Road; emerged 30 Aug–30 Sep 1957 ex larva in Matteuccia struthiopteris, J. Bird, collector (5 ♂, 1 ♀). Pennsylvania. Forest Co., King Island, Allegheny River, 41.5567N 79.4259W, ex Matteuccia struthiopteris, collected 19 Jul 2023, emerged 8 Sep 2023, P. Woods (2 ♂). AMNH, CMNH, CNC, CUIC, INHS, LACM, MCLB, NHML, NYSM, UCD, USNM, VADNH, YPM, AEM, ELQ, JRW.
The larva is a borer in the crown of ostrich fern (Matteuccia struthiopteris), although there is strong evidence suggesting P.hesseli may at least occasionally also utilize regal ferns (Osmunda spp.) often found in close association with ostrich fern. The moths have been taken in habitats where ostrich fern was not observed but Osmunda species were in abundance. It is likely this species is much more habitat dependent than specific host plant dependent. Larval presence in the host plant may often be detected by a small, stunted, dead stipe at the base of the crown which was the initial entry point of the first instar early in the season. The subsequent growth of healthy, luxuriant fronds tends to obscure this evidence, so very careful examination is necessary to find larvae early in the season. Occasionally a small amount of orange frass may have accumulated on soil around the base of the bored plant, but the pervasive flooding in this habitat often washes away this evidence. In late July and early August, mature larvae are easier to find since their boring in the enlarged stipe bases of mature fronds causes the entire frond to wilt or turn yellow and then brown (
Due to the difficulty of distinguishing some individuals of P.hesseli from those of P.pterisii, the type series was restricted to specimens from those localities where both the habitat and host utilization are known to us. However, P.hesseli has a much broader distribution based upon specimens examined from central Wisconsin to New Brunswick, occurring between latitudes 41.5° and 47°. The recent discovery of what is likely this species in eastern Saskatchewan, where ostrich fern is abundant in river floodplains (but bracken fern is absent), indicates that P.hesseli probably has a much larger range than previously thought. Still, the species exhibits a much narrower distribution than that of P.pterisii, extending neither as far south nor as far north.
Papaipema limpida GROUP
This group consists of three species, Papaipema limpida, P.maritima, and P.beeriana. Papaipema limpida and P.maritima share some features with each other but not with P.beeriana. All three species have dark forewings reflecting their association with host plants in the Asteraceae, which have scabrous foliage that turns black after frost.
Papaipema limpida (Guenée) (Ironweed Root Borer)
Gortyna limpida Guenée, 1852, in Boisduval and Guenée, Histoire Naturelle des Insectes. Species Général des Lépidoptѐres, 5: 124.
Type locality: “État des Illinois” [Illinois, USA]. [Types lost]
note—Described from one male (26 mm) and one female (40 mm). Although Guenée’s original types were apparently destroyed, it is fortunate that the identity of limpida can be ascertained from his description. Guenée named both limpida and nebris in the same publication, and it is clear to which species his names apply. Although not authentic, the NHML “types” of limpida are correct for the species.
Papaipema limpida is another of Guenée’s “lost” species, the correct identity of which has long been debated. This confusion led
Specimens of P.limpida have been misidentified in collections as the much more common P.nebris. The forewing of P.limpida is darker brown than that of P.nebris, and the subterminal area is the same color as the medial area; in P.nebris, the subterminal area is lighter and contrasts to varying degrees with the medial area. The postmedial line of P.limpida is broadly sinuous; that of P.nebris is straight from the inner margin to near the costa, where it bends inward to meet the costa. The reniform, orbicular, and suborbicular stigmata of P.limpida are larger than those of P.nebris and are bright white. Only the central lunule of the reniform is cream. Occasional specimens of P.limpida lack the white stigmata on the forewings (
The DNA barcodes of the seven P.limpida specimens sampled separate into two different BINs that are near each other but are not sister groups, possibly indicating a genetic artefact such as hybridizing introgression. This should be investigated further. One group with specimens from Missouri and Pennsylvania places closer to P.sciata- P.nepheleptena barcodes, the second (Texas, Nebraska, Illinois, Kansas) is more isolated with a minimum divergence of 1.03%.
The larval description provided here is based on nearly mature larvae. The penultimate and final instars are grub-like, whitish, and largely devoid of pattern, in stark contrast to those of P.cerussata which are quite typical Papaipema larvae and utilize the same host plant. The head capsule is light reddish-brown and the prothoracic shield of T1 is a paler honey color, lacking dark pigmentation on both anterior and lateral margins. This is unique among late instar Papaipema species; all others have dark, almost black pigmentation on either the anterior margin in a few species or defining the lateral margins of the shield in most. The normal purplish brown cuticular pigmentation of A1–A4, characteristic of Papaipema species, is obscure or lacking in later instars of P.limpida, but apparently not in earlier ones. Subdorsal, supraspiracular, spiracular, and subspiracular pinacula are greatly reduced and not strongly pigmented, a typical condition of subterranean borers. The anal plate is reduced and is of the same color as the prothoracic shield. Aaron Brees (pers. comm.) has kindly provided images and details concerning his observations of the earlier instars of P.limpida. Based on these images, it is now clear that P.limpida belongs to the subgroup of Papaipema larvae without longitudinal white lines crossing the dark pigmented bands encircling segments A1–A4. It is also noteworthy that these longitudinal lines on A5-A8 are represented as white spots at the anterior margins of these segments but are not continuous white stripes across them, as in many other Papaipema species. However, most informative is the discovery that dark pigmentation on the lateral margins of the prothoracic shield is not a stable character across instars; it is strongly expressed in early instars of P.limpida and completely lost in later instars.
The host plant and larva of P.limpida remained unknown until 1977, when ELQ first found larvae feeding in Baldwin’s ironweed (Vernonia baldwinii) in Jasper County, Missouri. Earlier reports from other plants such as burdock (Arctium) were based upon misidentifications. This moth has since been reared from Missouri ironweed (Vernonia missurica), and it seems likely any ironweed species growing within the range of P.limpida may be utilized.
Both P.limpida and P.cerussata are specialists on ironweed. JRW has found multiple larvae of both species boring in the same clump of ironweed having as many as twenty separate stems. Both their larvae and their feeding habits differ dramatically, and their identity is immediately apparent upon inspection of the affected plants. Larvae of P.limpida enter the stem at or near ground level, rarely more than 2.5 centimeters above ground. Larval mining is confined to the base of the stem where a slight swelling and hole with ejected brown frass might be seen, and then proceeds to the root where boring is extensive and most larval development occurs. The mature larva exits the plant and pupates in soil nearby. In the case of P.cerussata, the larva enters the stem as much as one third of a meter above ground and bores in the stem extensively before eventually entering the root, as described in the discussion of that species. Adults are on the wing throughout September and into October.
Papaipema limpida is essentially a prairie inhabitant with a comparatively restricted distribution. It is unusual in that it occurs neither as far north, nor as far south, as other midwestern Papaipema species. It also occurs in old fields with abundant ironweed and sparse successional growth of trees. It ranges from western Pennsylvania to western Nebraska.
Papaipema maritima Bird (Giant Sunflower Borer)
Papaipema maritima Bird, 1909, The Canadian Entomologist, 41: 59.
Type locality: [Rye, Westchester Co., New York, USA.] [AMNH]
note—No specific type locality other than “Atlantic seaboard in the neighboring latitude of New York City” was mentioned in the original description, but Bird’s male type and female cotype are both labelled “Rye, N.Y.” The species was described from 108 specimens.
Papaipema necopina; authors, not Grote, 1876.
Papaipema maritima is easily confused with several other Papaipema species, especially P.necopina, but also P.silphii, P.beeriana, and perhaps P.duplicata, all of which share dark forewing color adapted for a strong cryptic resemblance to the frosted, scabrous, dead foliage of their host plants, upon which the moths deposit their eggs and rest by day. From P.necopina, P.maritima is distinguished by its darker hindwings. The postmedial line on the forewing of P.maritima is often faint but still visible as a double, white-filled line for at least its posterior half. In P.necopina, this line is obscure but defined by the boundary it forms with the contrasting bluish subterminal area. The unique thoracic crest of P.maritima (
Neither the longitudinal mid-dorsal nor subdorsal white lines cross the pigmented bands of the first four abdominal segments in this species. The prothoracic shield has prominent, black, lateral margins, but these dark lines do not extend onto the head capsule.
The maritime Atlantic population exclusively utilizes giant sunflower (Helianthus giganteus) as its host. Elsewhere, other thick-stemmed sunflowers are commonly bored, including Jerusalem artichoke (H.tuberosus), cheerful sunflower (H. laetiflorus), ashy sunflower (H.mollis), and sawtooth sunflower (H. grosseserratus). Isolated records of larvae in Indian plantains (Arnoglossum spp.) clearly reflect overflow in prairie habitats where sunflowers remain the dominant hosts. The larva spends its entire development period within a rather large, ovoid gall which forms near the base of the stem of its host plant and serves as a clear indication of infestation. Upon maturity, pupation occurs within this protective structure after an exit hole for the adult, covered with a thin, parchment-like, easily breakable layer of the gall’s outer skin, is prepared by the larva. The pupa of P.maritima has a feature unique among Papaipema species. At the cephalic end of the pupa, a heavily sclerotized, keel-like ridge protrudes from the prothoracic area which is not homologous with the protrusion seen on the pupae of P.sauzalitae and P.angelica, which corresponds to a conical projection of the frons in the adult. Adult P.maritima have no vestige of any similar structure. The distinctive, round gall formed near the base of the sunflower stem serves to distinguish P.maritima from P.necopina, which has larvae that form a long, cylindrical gall. Both species may be encountered in some midwestern populations occupying the same patch of sunflowers. For a highly detailed account of the immature stages of P.maritima, see
The present-day distribution of this species reveals a significant disjunction which likely reflects the effects of prior glaciations on ancestral populations. The Atlantic Coastal Plain maritime populations, now imperiled, feed exclusively as larvae on giant sunflower. The widespread, abundant, midwestern populations utilize a variety of other native sunflowers. This species was formerly common along the Atlantic Coast from Connecticut to Delaware, and small remnants of this disjunct population are still extant. Otherwise, it is widespread throughout the Midwest from southeastern Ontario to Kansas, northwest to Alberta, and south to Missouri, Tennessee, and northern Mississippi, as well as at higher elevations in the southern Appalachians. The Payson, Utah records have been verified by ELQ, but could represent an introduction breeding in sunflowers transplanted from some eastern population.
Papaipema beeriana Bird (Blazing Star Borer)
Papaipema beeriana Bird, 1923, The Canadian Entomologist, 55: 106.
Type locality: Chicago, [Cook Co.], Illinois, [USA]. [AMNH]
note—Described from forty specimens from the same locality.
† Papaipema beeriana lacinariae Bird, 1923, The Canadian Entomologist, 55: 107.
note—Described from eight specimens from the same locality as above, based on the white-spotted form.
Papaipema lacinaria; Hessel, 1954, misspelling.
This species is similar in appearance to several other species. For distinguishing features, refer to the discussion of this matter in the treatment of P.maritima. Throughout most of its range, P.beeriana is consistent in appearance, with both white-spotted and dark-spotted phenotypes (plate A9, figures 21 & 23), though the white-spotted phenotype is generally less common. The ground color of the forewings is mouse gray, which is mixed with a warmer brown in the medial area, and the wings are heavily flecked in light gray. Possibly due to genetic drift, a distinctive phenotype prevails in the northernmost known population of P.beeriana, isolated in Otsego County in the northern part of Michigan’s lower peninsula. These individuals (plate A9, figure 24) exhibit a prominent rusty coloration of the medial area of the forewing, which is typically mixed with gray and less reddish elsewhere throughout the range of P.beeriana. Additionally, the reniform and orbicular stigmata are typically white in these populations but are most commonly suppressed in populations elsewhere. This population occurs in a jack pine dominant prairie environment but, aside from the different coloration of the individuals, they are morphologically and genetically indistinguishable from those occurring elsewhere, in both the larval stage and the adult.
The phylogenetic affinities of P.beeriana remain unclear. The larvae of P.beeriana retain pigmentation features unique among all other Papaipema species and reminiscent of those seen in more basal genera such as Fibigeria and Hydraecia. The male genitalia exhibit the most extreme bifurcation of the genus. Molecular data suggest it is a basal outlier that may be part of a lineage that includes P.eupatorii, but morphological features of both the larva and adult do not support placement either there or anywhere else. Seven DNA barcode samples from Michigan and Illinois represent an isolated barcode cluster differing by at least 3.04% from other species. The larvae of P.beeriana exhibit a peculiar pigmentation pattern unique among all other Papaipema species. In addition to the usual dark, reddish-brown pigmented bands encircling the first four abdominal segments, this pigmentation continues on the remaining segments, creating a banded effect reminiscent of some more primitive apameine genera. Otherwise, the larval morphology does not differ significantly from that of its congeners. For a much more detailed discussion of the immature stages, see
Various species of blazing stars in the genus Liatris serve as hosts for this prairie specialist. The larva remains within the subterranean bulb of its host for nearly its entire development, likely an adaptation for surviving frequent prairie fires as well as drought.
Papaipema beeriana can be a challenging species to locate in its larval stages, especially in very dry years, as there may be little remaining above-ground evidence of its presence in the bulbs. Occasionally, some dried vestige of the initial entry stalk may provide the only clue. The mature larva commonly pupates in the hollowed bulb which had served as its refuge, but also in soil nearby. Adults appear in late August and fly throughout September into early October.
This species occupies both moist meadows and fens as well as dry, sandy, prairie environments throughout the central and upper Midwest from southeastern Ontario to North Dakota and south to Virginia and Arkansas.
LITERATURE
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Monochrome Plates: 1–11
WING PATTERN AND THORACIC VESTITURE 1. Forewing and hindwing of Papaipema limata. Thoracic vestiture: 2. Papaipema cerussata, lateral view. 3. Papaipema maritima, lateral view. 4. Papaipema silphii, lateral view. 5. Papaipema cerussata, frontal view. Figures 2–4 illustrated by Henry Bird. Figure 5 illustrated by A. E. McBride.
Genitalia of Acrapex and Sarita 1. Acrapex prisca (Walker) ♀. 2. Sarita relicta (Ferguson) ♀. 3. Sarita relicta (Ferguson) ♂.
Larvae of Arundinaria-Feeding Genera 1. Arundinariaphila hayesi Quinter, dorsal view. 2. Arundinariaphila hayesi Quinter, lateral view. 3. Cherokeea slotteni Quinter, dorsal view. 4. Cherokeea slotteni Quinter, lateral view. 5. Cherokeea gabrielleae Quinter, dorsal view. 6. Cherokeea gabrielleae Quinter, lateral view.
Larvae of Arundinaria-Feeding Genera 1. Cherokeea trigona (Smith), dorsal view. 2. Cherokeea trigona (Smith), lateral view. 3. Arundinariaphila hayesi Quinter, head. 4. Cherokeea slotteni Quinter, head. 5. Cherokeea gabrielleae Quinter, head. 6. Cherokeea trigona (Smith), head.
Larvae of Franclemontia and Sarita 1. Franclemontia interrogans Ferguson, dorsal view. 2. Franclemontia interrogans Ferguson, lateral view. 3. Sarita relicta (Ferguson), dorsal view. 4. Sarita relicta (Ferguson), lateral view. 5. Franclemontia interrogans Ferguson, head. 6. Sarita relicta (Ferguson), head.
Larvae of Papaipema 1. Papaipema sullivani Quinter, dorsal view. 2. Papaipema sullivani Quinter, lateral view. 3. Papaipema matheri Quinter, dorsal view. 4. Papaipema matheri Quinter, lateral view. 5. Papaipema sullivani Quinter, head. 6. Papaipema matheri Quinter, head.
Larval Damage The blackened, withered, drooping top on Cicuta maculata bored by an early instar Papaipema birdi larva in mid-May. Illustration by Henry Bird.
Larval Damage 1. Early boring by the larva of Papaipema stenocelis causing wilting in the frond tip of Woodwardia virginica. 2. The withered and browned frond of Onoclea sensibilis caused by the boring of Papaipema inquaesita. Illustrations by Henry Bird.
Larval Damage Larva of Papaipema unimoda boring in the rootstock of Thalictrum polygamum. Illustration by Henry Bird.
Adults: Color Plates A1–A11
: Figs 1–43; SCALE 1.25:1 Arundinariaphila hayesi Quinter, ♀. Paratype. Rt. 121 SE of Wickliffe, Stovall Creek, Ballard Co., Kentucky, with head lamps in canebrake, 15 Jun. 1999, E. L. Quinter and J. R. Wiker, ELQ. Arundinariaphila hayesi Quinter, ♂. Paratype. St. Catherines Island, Liberty Co., Georgia, bred ex larva in Arundinaria tecta, emerged 18 May 1994, E. L. Quinter, ELQ. Arundinariaphila hayesi Quinter, ♀. Paratype. Sandy Branch, E of old G.M.&O. right of way, Westvaco WMA, 5 mi SW of Bardwell, Carlisle Co., Kentucky, 14 Jun. 1999, E. Quinter, W. Black, and J. Wiker, ELQ. Arundinariaphila hayesi Quinter, ♀. Paratype. Mark Twain National Forest, Jct. Rt. 49 and Black River, 2.7 mi W of Williamsville, Browns Crossing, 400´ [123 m], Wayne Co., Missouri, 21–22 Jun. 1993, ex larva in Arundinaria gigantea, emerged 18 Jul. 1993, E. L. Quinter, ELQ. Arundinariaphila hayesi Quinter, ♂. Paratype. St. Catherines Island, Liberty Co., Georgia, bred ex larva in Arundinaria tecta, emerged 30 May 1993, E. L. Quinter, ELQ. Arundinariaphila hayesi Quinter, ♀. Paratype. Mark Twain National Forest, Jct. Rt. 49 and Black River, 2.7 mi W of Williamsville, Browns Crossing, Wayne Co., Missouri, ex larva collected 25 May 1994 in Arundinaria gigantea, reared on artificial diet, pupated 2 Jun. 1994, emerged 19 Jun. 1994, E. L. Quinter, ELQ. Arundinariaphila hayesi Quinter, ♂. Holotype. St. Catherines Island, Liberty Co., Georgia, bred ex larva in Arundinaria tecta, emerged 19 May 1994, E. L. Quinter, CNC. Cherokeea attakullakulla Sullivan & Quinter, ♀. Fontana Lake Estates, vicinity Trails End Drive, 35.3873, -83.5515, 1,782´ [543 m], Swain Co., North Carolina, 25 Apr. 2023 ex larva, collected on Arundinaria appalachiana and reared on ornamental bamboo, pupated 3 May 2023, eclosed 28 May 2023, A. E. McBride, AEM. Cherokeea attakullakulla Sullivan & Quinter, ♂. Fontana Lake Estates, vicinity Trails End Drive, 35.3873, -83.5515, 1,782´ [543 m], Swain Co., North Carolina, 25 Apr. 2023 ex larva, collected on Arundinaria appalachiana and reared on ornamental bamboo, pupated 6 May 2023, eclosed 29 May 2023, A. E. McBride, AEM. Cherokeea attakullakulla Sullivan & Quinter, ♂. Fontana Lake Estates, vicinity Trails End Drive, 35.3873, -83.5515, 1,782´ [543 m], Swain Co., North Carolina, 25 Apr. 2023 ex larva, collected on Arundinaria appalachiana and reared on ornamental bamboo, pupated 2 May 2023, eclosed 24 May 2023, A. E. McBride, AEM. Cherokeea attakullakulla Sullivan & Quinter, ♀. Fontana Lake Estates, 35.3844N, 83.5579W, 2000´ [610 m], Swain Co., North Carolina, mesic mixed hardwoods, pine, 15w UV trap, 13–14 Jun. 2004, J. Bolling Sullivan, CNC (CNCLEP # 113485). Cherokeea slotteni Quinter, ♀. Paratype. St. Catherines Island, Liberty Co., Georgia, bred ex larva in Arundinaria tecta, pupated 26 Apr. 1994, emerged 14 May 1994, E. L. Quinter, ELQ. Cherokeea slotteni Quinter, ♂. Paratype. St. Catherines Island, Liberty Co., Georgia, bred ex larva in Arundinaria tecta, pupated 12 Apr. 1994, emerged 4 May 1994, E. L. Quinter, ELQ. Cherokeea slotteni Quinter, ♀. Paratype. St. Catherines Island, Liberty Co., Georgia, bred ex larva in Arundinaria tecta, pupated 28 Apr. 1994, emerged 19 May 1994, E. L. Quinter, ELQ. Cherokeea slotteni Quinter, ♂. Holotype. St. Catherines Island, Liberty Co., Georgia, bred ex larva in Arundinaria tecta, pupated 12 Apr. 1994, emerged 1 May 1994, E. L. Quinter, CNC. Cherokeea slotteni Quinter, ♂. Paratype. Great Dismal Swamp National Wildlife Refuge, City of Suffolk, Virginia, bred ex larva in Arundinaria tecta, emerged 14 May 1998, E. L. Quinter, ELQ. Cherokeea gabrielleae Quinter, ♂. Holotype. Great Dismal Swamp National Wildlife Refuge, City of Suffolk, Virginia, bred ex larva in Arundinaria tecta, pupated 19 Jun. 1998, emerged 4 Jul. 1998, Eric Quinter and Steven Roble, CNC. Cherokeea gabrielleae Quinter, ♂. Paratype. Great Dismal Swamp National Wildlife Refuge, City of Suffolk, Virginia, bred ex larva in Arundinaria tecta, pupated 19 Jun. 1998, emerged 3 Jul. 1998, Eric Quinter and Steven Roble, ELQ. Cherokeea gabrielleae Quinter, ♀. Paratype. Great Dismal Swamp National Wildlife Refuge, City of Suffolk, Virginia, ex larva collected 23–24 Apr. 2000 in Arundinaria tecta, reared on artificial diet, emerged 14 Jun. 2000, Eric Quinter and Steven Roble, ELQ. Cherokeea gabrielleae Quinter, ♀. Paratype. Great Dismal Swamp National Wildlife Refuge, City of Suffolk, Virginia, bred ex larva in Arundinaria tecta, emerged Jul. 1999, Eric Quinter and Zhiwei Liu, ELQ. Cherokeea gabrielleae Quinter, ♀. Paratype. Great Dismal Swamp National Wildlife Refuge, City of Suffolk, Virginia, bred ex larva in Arundinaria tecta, pupated Jun. 1998, emerged 30 Jun. 1998, Eric Quinter and Steven Roble, ELQ. Cherokeea gabrielleae Quinter, ♀. Paratype. Great Dismal Swamp National Wildlife Refuge, City of Suffolk, Virginia, bred ex larva in Arundinaria tecta, pupated 19 Jun. 1998, emerged 4 Jul. 1998, Eric Quinter and Steven Roble, ELQ. Cherokeea odonnellae Quinter, ♀. Paratype. Fontana Lake Estates, vicinity Trails End Drive, 35.3873, -83.5515, 1,782´ [543 m], Swain Co., North Carolina, 8 May 2022 ex larva, collected from new leaf petiole on older culm of Arundinaria appalachiana, eclosed 22 Jun. 2022, A. E. McBride, AEM. Cherokeea odonnellae Quinter, ♀. Paratype. Great Smoky Mountains National Park, Tom Branch Rd., 0.8 mi NE of Deep Creek Campground, 2,080´ [634 m], Swain Co., North Carolina, bred ex larva in Arundinaria appalachiana first year culm, pupated 4 Jun. 2001, emerged 18 Jun. 2001, E. L. Quinter and J. O’Donnell, ELQ. (p.27). Cherokeea odonnellae Quinter, ♀. Holotype. Little Tennessee River, Lower Needmore Road, 35.3285, -83.522, 1,787´ [545 m], Swain Co., North Carolina, 4 May 2021 ex larva, collected on one year old culm of Arundinaria appalachiana, pupated 27 May 2021, eclosed 15 Jun. 2021, larva photographed (#1), A. E. McBride with Eric Quinter, CNC. Cherokeea odonnellae Quinter, ♀. Paratype. Great Smoky Mountains National Park, Tom Branch Rd., 0.8 mi NE of Deep Creek Campground, 2,080´ [634 m], Swain Co., North Carolina, bred ex larva in Arundinaria appalachiana first year culm, pupated 4 Jun. 2001, emerged 19 Jun. 2001, E. L. Quinter and J. O’Donnell, ELQ. Cherokeea odonnellae Quinter, ♂. Paratype. Little Tennessee River, Lower Needmore Road, 35.3285, -83.522, 1,787´ [545 m], Swain Co., North Carolina, 4 May 2021 ex larva, collected on one year old culm of Arundinaria appalachiana, pupated 26 May 2021, eclosed 16 Jun. 2021, A. E. McBride with Eric Quinter, AEM. Cherokeea odonnellae Quinter, ♂. Paratype. Little Tennessee River, Lower Needmore Road, 35.3285, -83.522, 1,787´ [545 m], Swain Co., North Carolina, 25 Apr. 2023 ex larva, collected on Arundinaria appalachiana and reared on ornamental bamboo, eclosed 13 Jun. 2023, A. E. McBride, AEM. Cherokeea trigona (Smith), ♀. Mark Twain National Forest, Jct. Rt. 49 and Black River, 2.7 mi W of Williamsville, Browns Crossing, Wayne Co., Missouri, ex larva collected 25 May 1994 in Arundinaria gigantea, reared on artificial diet, pupated 14 Jul. 1994, emerged 30 Jul. 1994, E. L. Quinter, ELQ. Cherokeea trigona (Smith), ♂. Cordell Hull Lake, Big Bottom, Jackson Co., Tennessee, ex larva in Arundinaria gigantea, pupated 12 Aug. 1994, emerged 30 Aug. 1994, E. L. Quinter, ELQ. Cherokeea trigona (Smith), ♀. Cordell Hull Lake, Big Bottom, Jackson Co., Tennessee, ex larva in Arundinaria gigantea, pupated 12 Aug. 1994, emerged 29 Aug. 1994, E. L. Quinter, ELQ. Cherokeea trigona (Smith), ♂. War Bluff Sanctuary, 5 mi N of Golconda, Pope Co., Illinois, mature larva collected 25 Jul. 1996 ex Arundinaria gigantea, Quinter, Wiker, Day, ELQ. Cherokeea trigona (Smith), ♂. Mark Twain National Forest, Jct. Rt. 49 and Black River, 2.7 mi W of Williamsville, Browns Crossing, Wayne Co., Missouri, ex larva collected 25 May 1994 in Arundinaria gigantea, reared on artificial diet, pupated 11–12 Jul. 1994, emerged 28 Jul. 1994, E. L. Quinter, ELQ. Cherokeea trigona (Smith), ♂. War Bluff Sanctuary, 5 mi N of Golconda, Pope Co., Illinois, mature larva collected 25 Jul. 1996 ex Arundinaria gigantea, Quinter, Wiker, Day, ELQ. Cherokeea trigona (Smith), ♀. Cave Valley, 1 mi N of Pomona, Old G.M.&O. ROW, Jackson Co., Illinois, mature larva collected 28 Jul. 1996 ex Arundinaria gigantea, emerged 19 Aug. 1996, Quinter, Wiker, Day, ELQ. Franclemontia interrogans (Walker), ♀. St. Catherines Island, Liberty Co., Georgia, pupa collected 8 Apr. 1995, emerged 12 Apr. 1995, E. L. Quinter, ELQ. Franclemontia interrogans (Walker), ♀. Great Dismal Swamp National Wildlife Refuge, City of Suffolk, Virginia, ex pupa, emerged 15 Apr. 1998, E. L. Quinter, ELQ. Franclemontia interrogans (Walker), ♂. St. Catherines Island, Liberty Co., Georgia, pupa collected 2 Apr. 1993, emerged 5 Apr. 1993, E. L. Quinter, ELQ. Franclemontia interrogans (Walker), ♂. St. Catherines Island, Liberty Co., Georgia, pupa collected 4 Apr. 1993, emerged 12 Apr. 1993, E. L. Quinter, ELQ. Franclemontia interrogans (Walker), ♀. St. Catherines Island, Liberty Co., Georgia, ex larva in Arundinaria tecta, emerged 6 Apr. 1994, E. L. Quinter, ELQ. Sarita relicta (Ferguson), ♂. Revell’s Landing, Liberty Co., Florida, emerged 10 May 2001, Sarah and Eric Quinter, ELQ. Sarita relicta (Ferguson), ♀. St. Catherines Island, Liberty Co., Georgia, ex larva in Arundinaria tecta, pupated 8–9 May 1993, emerged 25 May 1993, E. L. Quinter, ELQ. Sarita relicta (Ferguson), ♀. Great Dismal Swamp National Wildlife Refuge, City of Suffolk, Virginia, ex larva, emerged 16 Jun. 1999, E. L. Quinter, ELQ.
: Figs 1–21; SCALE 1.25:1 Hydraecia medialis Smith, ♂. Leeds Canyon, Washington Co., Utah, 2 Sep. 1988, R. C. Mower, ELQ. Hydraecia medialis Smith, ♀. Fort Collins, Larimer Co., Colorado, at light, 21 Sep. 1930, CNC. Hydraecia medialis Smith, ♀. Laramie, 7,500 feet [2,286 m], 41° 17’N 105° 31’W, Albany Co., Wyoming, 15 Sep. 2000, Cliff Ferris, CNC. Hydraecia medialis Smith, ♀. Elkwater, Cypress Hills, Alberta, 2 Sep. 1990, G. Anweiler, CNC. Hydraecia medialis Smith, ♂. Weberg Rd., 1 mile N of Weberg, Crook Co., Oregon, 5 Sep. 1997, J. Troubridge, ELQ. Hydraecia medialis Smith, ♂. Hayes Creek, 49° 32’N 102° 25’W, Princeton, British Columbia, 7 Oct. 2000, J. Troubridge, CNC. Hydraecia medialis Smith, ♂. Hayes Creek, 49° 32’N 102° 25’W, Princeton, British Columbia, 7 Oct. 2000, J. Troubridge, CNC. Hydraecia medialis Smith, ♀. Hayes Creek, 49° 32’N 102° 25’W, Princeton, British Columbia, 7 Oct. 2000, J. Troubridge, ELQ. Hydraecia obliqua (Harvey), ♂. Nelson Creek, Plumas Co., California, 17 Sep. 1940, W. R. Bauer, ELQ. Hydraecia obliqua (Harvey), ♂. Deception Pass, Island Co., Washington, 31 Aug. 1996, J. Troubridge and L. Crabo, CNC. Hydraecia obliqua (Harvey), ♂. Bodega Bay Dunes, Sonoma Co., California, 25 Jul. 1992, R. Robertson, AEM. Hydraecia intermedia (Barnes & Benjamin), ♂. Holotype. “Fort Calgary, British Columbia” [Crowsnest Pass region, Alberta], Aug. 1916, USNM (USNMENT 01897614). Hydraecia perobliqua Hampson, ♀. Sandy Point Campground, 50° 43’N 110° 04’W, Alberta, 13 Aug. 2001, J. Troubridge and G. Anweiler, CNC. Hydraecia perobliqua Hampson, ♂. 17 km N of Winnemucca, 41° 06’N 118° 35’W, 1,300 m, Humboldt Co., Nevada, 21 Jul. 2001, D. Lafontaine and J. Troubridge, CNC. Hydraecia stramentosa Guenée, ♂. 13 mi SE of Cloudcroft, Otero Co., New Mexico, 9 Sep. 1975, Lafontaine and Bowen, CNC. Hydraecia stramentosa Guenée, ♂. Warm Springs Mountain N of Ingall’s Field, 37.9858, -79.8151, 3,532´ [1,077 m], Valley Springs, Bath Co., Virginia, 4 Jul. 2015 ex larva, ex burrow in root of Scrophularia lanceolata, eclosed 2 Aug. 2015, A. E. McBride, AEM. Hydraecia stramentosa Guenée, ♂. Tinytown, S. Turkey Creek Rd. across from train park, 39°36.17’N 105°13.33’W, 6,870´ [2,094 m], Jefferson Co., Colorado, UV trap, 18 Aug. 2008, C. Harp, ELQ. Hydraecia immanis Guenée, ♀. Carroll Co., Illinois, bred ex ovo on artificial diet, parent stock ex larva on seedling corn, Apr. 1982, Eli Levine, ELQ. Hydraecia immanis Guenée, ♂. Carroll Co., Illinois, bred ex ovo on artificial diet, parent stock ex larva on seedling corn, Apr. 1982, Eli Levine, ELQ. Hydraecia micacea (Esper), ♂. Hanwell, York Co., New Brunswick, 11 Sep. 2017, G. Hensel, CNC. Hydraecia micacea (Esper), ♀. Aylesford, Nova Scotia, 26 Sep. 1958, B. Wright, ELQ.
: Figs 1–33; SCALE 1.25:1 Hydraecia micacea (Esper), ♂. Orono, Penobscot Co., Maine, Aug. 1933, ELQ. Hydraecia micacea (Esper), ♂. 0.6 mi ESE of Round Top, 1,660´ [506 m], Richfield Twp., Otsego Co., New York, bred ex larva in Phalaris, emerged 18 Jul. 1982, E. L. Quinter, ELQ. Fibigeria erepta (Grote), ♂. Orchard Beach meadows, Bronx, Bronx Co., New York, bred ex larva in Tripsacum dactyloides, collected 12 Jun. 1994, pupated 17 Jun. 1994, emerged 29 Jun. 1994, E. L. Quinter, ELQ. Amphipoea interoceanica (Smith), ♂. Clarks Corner, 0.6 mi ESE of BM 577 power line cut, 630´ [192 m], Hampton, Windham Co., Connecticut, 19 Jul. 2018, UV/MV, E. L. Quinter, ELQ. Amphipoea interoceanica (Smith), ♂. Clarks Corner, 0.6 mi ESE of BM 577 power line cut, 630´ [192 m], Hampton, Windham Co., Connecticut, 20–21 Jun. 2012, BLT, E. L. Quinter, ELQ. Amphipoea interoceanica (Smith), ♂. prairie S of Pike Lake, Manitoulin Isl., Ontario, bred ex larva in crown of Geum triflorum, collected 13 Jun. 2011, emerged 30 Jul. 2011, E. L. Quinter and A. E. McBride, ELQ. Amphipoea interoceanica (Smith), ♂. Summer Hill, 1.9 mi W of Auburn, mixed woods, 860´ [262 m], Schuylkill Co., Pennsylvania, 15 Jul. 1973, E. L. Quinter, ELQ. Amphipoea interoceanica (Smith), ♀. prairie S of Pike Lake, Manitoulin Isl., Ontario, bred ex larva in crown of Geum triflorum, collected 13 Jun. 2011, emerged 30 Jul. 2011, E. L. Quinter and A. E. McBride, ELQ. Amphipoea americana (Speyer), ♂. Clarks Corner, 0.6 mi ESE of BM 577 power line cut, 630´ [192 m], Hampton, Windham Co., Connecticut, 20–21 Jun. 2012, BLT, E. L. Quinter, ELQ. Amphipoea americana (Speyer), ♂. Mt. Mitchell State Park, 6,400´ [1,951 m], Yancey Co., North Carolina, 31 Jul. 1974, J. Bolling Sullivan, ELQ. Amphipoea americana (Speyer), ♂. Mt. Mitchell State Park, 6,400´ [1,951 m], Yancey Co., North Carolina, 30 Jul. 1974, J. Bolling Sullivan, ELQ. Amphipoea pacifica (Smith), ♀. Hat Creek R. S., Shasta Co., California, 20 Jul. 1947, Fred H. Rindge, ELQ. Amphipoea pacifica (Smith), ♀. Mill Valley, Marin Co., California, 30 Aug. 1937, E. C. Johnston, CNC. Amphipoea pacifica (Smith), ♂. Frissell Pt., 44° 13’N 122° 06’W, 4,900´[1,494], Lane Co., Oregon, 28 Aug. 2002, J. Troubridge, CNC. Amphipoea pacifica (Smith), ♂. 8 Dollar Rd., 42° 14’N 123° 40’W, 1,500´ [457 m], Josephine Co., Oregon, 27 Jul. 2001, J. Troubridge, CNC. Amphipoea cottlei (McDunnough), ♀. Camp O-Ongo near Running Springs, 6,300´ [1,920 m], San Bernardino Co., California, 27 Aug. 1971, C. L. Hogue, ELQ. Amphipoea keiferi (Benjamin), ♂. Rt. 140 W of Adel, 42° 12’N 120° 00’W, 5,200´ [1,585 m], Lake Co., Oregon, 21 Sep. 2001, J. Troubridge and L. Crabo, ELQ. Amphipoea keiferi (Benjamin), ♂. Cornet Creek, 44° 29’N 117° 51’W, 3,600´ [1,097 m], Baker Co., Oregon, 31 Aug. 2000, J. Troubridge, ELQ. Amphipoea keiferi (Benjamin), ♂. Satus Creek, Yakima Co., Washington, 17 Sep. 1949, E. C. Johnston, CNC. Amphipoea keiferi (Benjamin), ♂. Jordan Valley, Malheur Co., Oregon, 13–16 Sep. 1963, W. C. Cook, CNC (CNCLEP # 116075). Amphipoea senilis (Smith), ♀. along Hwy. 285 just N of jct. of Hwy. 9, gas station MV lights, 39°13’02”N 105°59’34”W, 9,869´ [3,008 m], Fairplay, Park Co., Colorado, 31 Aug. 2007, 11 p.m., Chuck Harp, ELQ. Amphipoea senilis (Smith), ♀. White Mountains, 8,500´ [2,591 m], Greer, Apache Co., Arizona, 9 Aug. 1962, black light, E. and I. Munroe, CNC. Amphipoea senilis (Smith), ♂. 5 mi N of Mancos, 7,300´ [2,225 m], Montezuma Co., Colorado, 22 Aug. 1971, D. F. Hardwick, CNC. Amphipoea senilis (Smith), ♂. Headquarters Area, Florissant Fossil Beds, T13S, R71W, Sec. 13, 8,400–8,500´ [2,560–2,591 m], Teller Co., Colorado, 24 Sep. 1976, UV light, F. M. Brown, ELQ. Amphipoea senilis (Smith), ♀. Rt. 244, 1.15 mi. NE of Cloudcroft, 32.9967, -105.6658, 8,023´ [2,445 m], Otero Co., New Mexico, ex larva 25 Jun. 2023, collected from and reared in Iris missouriensis, eclosed 4 Aug. 2023, A. E. McBride, AEM. Amphipoea senilis (Smith), ♂. N32.808 W105.649, 8,100´ [2,469 m], Otero Co., New Mexico, 9 Aug. 2021, J. Vargo, AEM. Selicanis cinereola Smith, ♂. T18N, R3W, Sec. 35, Monroe Co., Wisconsin, 1 Oct. 1996, L. A. Ferge, ELQ. Parapamea buffaloensis (Grote), ♀. Green Village, Morris Co., New Jersey, 26 Aug. 1949, J. Muller, ELQ. Parapamea buffaloensis (Grote), ♀. Eldora, Cumberland Co., New Jersey, sheet on shed, 5 Oct. 1982, C. B. Worth, ELQ. Papaipema unimoda (Smith), ♂. Hook and Bullet, 44°48’21”, -96°23’38”, 330 m, Clay Co., Minnesota, 10 Sep. 1983, T. McCabe, ELQ. Papaipema unimoda (Smith), ♂. Rt. 28, 1 mi W of West Lebanon, Warren Co., Indiana, bred ex larva in Thalictrum, emerged 1 Oct. 1974, E. L. Quinter, ELQ. Papaipema unimoda (Smith), ♂. 0.3 mi S of Auburn, 500´ [152 m], Schuylkill Co., Pennsylvania, 16 Oct. 1972, E. L. Quinter, ELQ. Papaipema unimoda (Smith), ♂. Thompson Rd. Prairie, T7S, R16W, Sec. 5, Cass Co., Michigan, 14 Oct. 1989, G. Balogh, ELQ.
: Figs 1–27; SCALE 1.25:1 Papaipema furcata (Smith), ♂. Trout Brook Wildlife Management Area, 41.1149, -74.8614, 938´ [286 m], Stillwater Twp., Sussex Co., New Jersey, 20w BLS, 18 Sep. 2017, J. R. Garris, Sr., AEM. Papaipema furcata (Smith), ♀. Woodcock management area on Red Rose Rd., 40.6752, -77.8998, 901´ [275 m], Petersburg, Huntingdon Co., Pennsylvania, ex ovo 2018 from parent female collected 26 Sep. 2017 by S. Smith and A. E. McBride at 275w MVS, reared in potato and then in Fraxinus americana after third instar, eclosed Sep. 2018, A. E. McBride, AEM. Papaipema furcata (Smith), ♂. Hunterdon Co., New Jersey, 24 Sep. 1962, Joseph Muller, ELQ. Papaipema furcata (Smith), ♀. Chicago, Cook Co., Illinois, 28 Aug. 1921, Emil Beer, ELQ. Papaipema appassionata (Harvey), ♀. calcareous fen, 41.0367, -74.7485, 599´ [183 m], Andover Twp., Sussex Co., New Jersey, 3 Jul. 2016 ex larva, collected from and reared in Sarracenia purpurea, eclosed 1 Aug. 2016, A. E. McBride, AEM. Papaipema appassionata (Harvey), ♂. Hallsboro, Columbus Co., North Carolina, bred ex ovo in Sarracenia flava, emerge 3 Oct. 1927, Emil Beer, AMNH. Papaipema appassionata (Harvey), ♂. Cheboygan Co., Michigan, bred ex larva in Sarracenia purpurea, collected 14 Jul. 1979, emerged 16 Aug. 1979, E. L. Quinter, ELQ. Papaipema amianthii Quinter, ♂. Holotype. Appalachian Mts., Kittatinny Ridge, Blue Mountain, acid bog, 4.2 mi. SW of Auburn, 40.5494, -76.1459, 1,604´ [489 m], Schuylkill Co., Pennsylvania, bred ex larva in bulb of Amianthium muscitoxicum, emerged 7 Sep. 1972, E. L. Quinter, CNC. Papaipema amianthii Quinter, ♀. Paratype. Hickory Run State Park, Carbon Co., Pennsylvania, collected at building light, 9 Sep. 1977, E. L. Quinter, ELQ. Papaipema amianthii Quinter, ♀. Paratype. Hickory Run State Park, Carbon Co., Pennsylvania, bred ex pupa in Amianthium muscitoxicum, collected 29 Aug. 1976, emerged 2–7 Sep. 1976, E. L. Quinter, ELQ. Papaipema marginidens (Guenée), ♂. Blood River, Calloway Co., Kentucky, UV light trap at cane #6, 21 Oct. 2003. W. R. Black, Jr., ELQ. Papaipema marginidens (Guenée), ♀. Woodcock management area on Red Rose Rd., 40.6772, -77.8939, 922´ [281 m], Petersburg, Huntingdon Co., Pennsylvania, ex ovo 2018 from parent female collected 26 Sep. 2017 by A. E. McBride in 15w BLT, reared in potato, eclosed Jul. 2018, A. E. McBride, AEM. Papaipema marginidens (Guenée), ♀. New Lisbon, Burlington Co., New Jersey, 2 Oct. 1948, John W. Cadbury, ELQ. Papaipema marginidens (Guenée), ♀. Neotype. “New Yorck?”, USNM (USNMENT 01979913). Papaipema eryngii Bird, ♂. Holly Shelter Game Land, Pender Co., North Carolina, 19 Oct. 1995, J. B. Sullivan, JBS. Papaipema eryngii Bird, ♀. Cicero, Cook Co., Illinois, 16 Sep. 1925, Emil Beer, ELQ. Papaipema eryngii Bird, ♀. Wah-Kon-Tah Prairie, Foust Tract, Speedwell Twp., St. Clair Co., Missouri, ex larva in Eryngium yuccifolium 9 Jul. 2015, reared in carrot, eclosed 1 Oct. 2015, James R. Wiker, AEM. Papaipema dribi Barnes & Benjamin, ♂. Near Mescalero on Mescalero Apache Tribal Land, floodplain habitat, Otero Co., New Mexico, 14 Aug. 2021, E. H. Metzler, CNC. Papaipema dribi Barnes & Benjamin, ♂. Near Mescalero on Mescalero Apache Tribal Land, floodplain habitat, Otero Co., New Mexico, 13 Aug. 2021, E. H. Metzler, CNC. Papaipema albimontana Quinter, ♀. Paratype. Little Colorado River floodplain, 1 mi. S of Greer, 33.9928, -109.4653, 8,500´ [2,591 m], Apache Co., Arizona, 27 Jun. 2023 ex larva, collected in Cicuta douglasii and reared in carrot, eclosed 10 Aug. 2023, A. E. McBride, AEM. Papaipema albimontana Quinter, ♂. Holotype. Little Colorado River floodplain, 1 mi. S of Greer, 33.9928, -109.4653, 8,500´ [2,591 m], Apache Co., Arizona, 27 Jun. 2023 ex larva, collected in Cicuta douglasii and reared in carrot, eclosed 3 Aug. 2023, A. E. McBride, CNC. Papaipema insulidens (Bird), ♂. Pullman, Whitman Co., Washington, ex Senecio, emerged 4 Aug. 1935, S. E. Crumb, ELQ. Papaipema insulidens (Bird), ♀. Steptoe Butte, Whitman Co., Washington, 2,500–3,000´ [762–914 m], bred ex larva in Heracleum, emerged 7 Sep. 1995, E. L. Quinter, ELQ. Papaipema insulidens (Bird), ♂. Sissons, Siskiyou Co., California, 22 Aug. 1918, F. M. Jones, ELQ. Papaipema insulidens (Bird), ♂. Rt. 101, 12 mi. S of junction with Rt. 22, Tillamook Co., Oregon, bred ex larva in Heracleum, emerged 8 Aug. 1995, E. Quinter and P. Goldstein, ELQ. Papaipema insulidens (Bird), ♀. Rt. 101, 1–2 mi. N of Manzanita, Tillamook Co., Oregon, bred ex larva in Ligusticum apiifolium, emerged 9 Aug. 1995, E. Quinter and P. Goldstein, ELQ. Papaipema insulidens (Bird), ♀. Rt. 101, 1–2 mi. N of Manzanita, Tillamook Co., Oregon, bred ex larva in Ligusticum apiifolium, emerged 12 Aug. 1995, E. Quinter and P. Goldstein, ELQ.
: Figs 1–29; SCALE 1.25:1 Papaipema insulidens (Bird), ♂. Rt. 101, 8 mi. S of Astoria, Clatsop Co., Oregon, bred ex larva in Cicuta douglasii, emerged 3 Aug. 1995, E. Quinter and P. Goldstein, ELQ. Papaipema insulidens (Bird), ♀. Three Rocks Road, 1.5 mi. W of Rt. 101, Lincoln Co., Oregon, bred ex larva in Cirsium, emerged 11 Aug. 1995, E. Quinter and P. Goldstein, ELQ. Papaipema birdi (Dyar), ♀. 0.6 mi. ESE of Round Top, 1,600´ [488 m], Richfield Twp., Otsego Co., New York, bred ex larva in Angelica, pupated 3–9 Aug. 1995, emerged 31 Aug. 1995, E. Quinter and P. Goldstein, ELQ. Papaipema birdi (Dyar), ♀. Goshen TNC Preserve, Litchfield Co., Connecticut, ex larva in Cicuta maculata, emerged 8 Sep. 2007, E. Quinter and D. Wagner, ELQ. Papaipema birdi (Dyar), ♂. 0.6 mi. ESE of Round Top, 1,600´ [488 m], Richfield Twp., Otsego Co., New York, bred ex larva in Iris versicolor, emerged 14 Sep. 1982, E. L. Quinter, ELQ. Papaipema birdi (Dyar), ♀. 0.6 mi. ESE of Round Top, 1,600´ [488 m], Richfield Twp., Otsego Co., New York, bred ex larva in Angelica, emerged 31 Aug. 1995, E. Quinter and P. Goldstein, ELQ. Papaipema birdi (Dyar), ♂. 0.6 mi. ESE of Round Top, 1,600´ [488 m], Richfield Twp., Otsego Co., New York, bred ex larva in Angelica, pupated 4–7 Aug. 1995, emerged 29 Aug. 1995, E. Quinter and P. Goldstein, ELQ. Papaipema birdi (Dyar), ♀. T7N, R1E, Sec. 3, Ingham Co., Michigan, ex larva in Cicuta maculata, emerged 15 Sep. 1973, M. C. Nielsen, ELQ. Papaipema birdi (Dyar), ♂. 0.6 mi. ESE of Round Top, 1,600´ [488 m], Richfield Twp., Otsego Co., New York, bred ex larva in Angelica, pupated 4–7 Aug. 1995, emerged 29 Aug. 1995, E. Quinter and P. Goldstein, ELQ. Papaipema nepheleptena (Dyar), ♂. Junction of Follett Rd. and Potters Brook, 240´ [73 m], 0.7 mi. ESE of Windham, Windham Co., Connecticut, UV trap, 16 Sep. 2005, E. L. Quinter, ELQ. Papaipema nepheleptena (Dyar), ♂. T5S, R14W, Sec. 30, Cass Co., Michigan, UV trap, 19 Sep. 1993, Mogens C. Nielsen, ELQ. Papaipema nepheleptena (Dyar), ♀. Rt. 79, 0.5 mi. E of Slaterville Springs, Tompkins Co., New York, bred ex larva in Chelone glabra, emerged 29 Aug. 1978, E. L. Quinter, ELQ. Papaipema nepheleptena (Dyar), ♀. Richardson Wildlife Foundation, Muskrat Pond, T20N, R11E, Sec. 13, Lee Co., Illinois, bred ex larva in Verbena hastata, emerged 26 Sep. 2007, James R. Wiker, ELQ. Papaipema sciata Bird, ♀. 1.5 mi. SE of Lee’s Summit, Jackson Co., Missouri, bred ex larva in Veronicastrum virginicum, emerged 28 Sep. 1975, E. L. Quinter, ELQ. Papaipema sciata Bird, ♂. Pecan Run, South Kilbourne, Rt. 97 and IMRR, T19N, R8W, Sec. 3, Mason Co., Illinois, ex larva in Veronicastrum virginicum, emerged 1 Oct. 2007, James R. Wiker, ELQ. Papaipema cerussata (Grote), ♀. Dyer Hill Creek, Rt. 60 SW of Buma, E of Birdsville, Livingston Co., Kentucky, bred ex larva in Cirsium, emerged 28 Oct. 1999, E. L. Quinter, ELQ. Papaipema cerussata (Grote), ♀. Rt. 28, 1 mi. W of West Lebanon, Warren Co., Indiana, bred ex larva in Vernonia sp., emerged 25 Sep. 1974, E. L. Quinter, ELQ. Papaipema cerussata (Grote), ♀. Rt. 28, 1 mi. W of West Lebanon, Warren Co., Indiana, bred ex larva in Vernonia sp., emerged 17 Sep. 1974, E. L. Quinter, ELQ. Papaipema blackorum Quinter, ♀. Paratype. About 8 miles NNW of Poplar Bluff, E of US Highway 67 and just E of Old Bridge over the Black River, T26N, R5E, Sec. 13, Butler Co., Missouri, collected with MH light in canebrake, 12 Oct. 2002, James R. Wiker and Mogens C. Nielsen, JRW. Papaipema blackorum Quinter, ♀. Holotype. S of Rt. 146 on Reevesville Rd., Johnson Creek, C. Sampson property, N37.3871, W088.7779, Johnson Co., Illinois, 12 Aug. 2019 ex larva, larva gall forming in vine of Aristolochia tomentosa, pupated late Aug. 2019, emerged 30 Sep. 2019, James R. Wiker, CNC. Papaipema blackorum Quinter, ♀. Paratype. Blood River, cane site #P3, UV light trap, Henry Co., Tennessee, 12 Oct. 2003, William R. Black, Jr., ELQ. Papaipema blackorum Quinter, ♀. Paratype. Blood River, cane site #4A, UV light trap, Calloway Co., Kentucky, 6 Oct. 2010, William R. Black, Jr., ELQ. Papaipema blackorum Quinter, ♀. Paratype. vicinity of Bovina, Warren Co., Mississippi, collected at MV light, 11 Oct. 1972, Bryant Mather, genitalic preparation #349 female by E. Quinter, 1976, ELQ. Papaipema circumlucens (Smith), ♀. 3 mi. E of Good Hope, Short Fork Marsh, T7N, R2W, Sec. 27, McDonough Co., Illinois, ex larva in Humulus lupulus 25 Jul. 2003, emerged 12 Sep. 2003, James R. Wiker, JRW. Papaipema circumlucens (Smith), ♀. Three Bridges, Hunterdon Co., New Jersey, bred ex larva in Humulus lupulus, emerged 28 Aug. 1980, E. Quinter and J. Muller, ELQ. Papaipema circumlucens (Smith), ♂. 2.5 mi. SW of West Lebanon, Warren Co., Indiana, bred ex larva in Humulus lupulus, emerged 11 Aug. 1978, E. L. Quinter, ELQ. Papaipema circumlucens (Smith), ♂. Lectotype. New Jersey, USNM (USNMENT 01979912). Papaipema impecuniosa (Grote), ♂. Snyder Heights, Ithaca, Tompkins Co., New York, ex pupa in Aster, emerged 10 Sep. 1977, E. L. Quinter, ELQ. Papaipema impecuniosa (Grote), ♀. 2.6 mi. NE of New Ringgold, Schuylkill Co., Pennsylvania, ex pupa in Arctium minus, emerged 18 Sep. 1972, E. L. Quinter, ELQ.
: Figs 1–36; SCALE 1.25:1 Papaipema impecuniosa (Grote), ♂. Brook Hollow Farm meadows, 40.9617, -75.0541, 333´ [101 m], Knowlton Twp., Warren Co., New Jersey, ex pupa 11 Sep. 2015, ex burrow in lower stem of Aster puniceus, eclosed 16 Sep. 2015, A. E. McBride. AEM. Papaipema nelita (Strecker), ♂. 0.3 mi. S of Auburn, Schuylkill Co., Pennsylvania, bred ex larva in Rudbeckia laciniata, emerged 31 Aug. 1974, E. L. Quinter, ELQ. Papaipema nelita (Strecker), ♀. Diamond Rock, White Mountains, Apache Co., Arizona, 6 Sep. 1947, Grace H. and John L. Sperry, ELQ. Papaipema nelita (Strecker), ♀. 0.6 mi. ESE of Round Top, 1,660´ [488 m], Richfield Twp., Otsego Co., New York, bred ex larva in Arctium minus, pupated 21 Jul. 1995, emerged 7 Aug. 1995, E. L. Quinter, ELQ. Papaipema nelita (Strecker), ♀. Little Colorado River floodplain, 1 mi. S of Greer, 33.9945, -109.4646, 8,487´ [2,587 m], Apache Co., Arizona, 24 Jun. 2022 ex larva, collected in Delphinium and reared in carrot, eclosed 25 Jul. 2022, A. E. McBride, AEM. Papaipema nelita (Strecker), ♀. Forest Park, Cook Co., Illinois, ex Rudbeckia, 27 Aug. 1922, Emil Beer, ELQ. Papaipema nelita (Strecker), ♂. Palos Park, Cook Co., Illinois, ex Rudbeckia, 21 Aug. 1917, Emil Beer, ELQ. Papaipema nelita (Strecker), ♀. 0.6 mi. ESE of Round Top, 1,660´ [488 m], Richfield Twp., Otsego Co., New York, bred ex larva in Arctium minus, pupated 21 Jul. 1995, emerged 8 Aug. 1995, E. L. Quinter, ELQ. Papaipema nelita (Strecker), ♂. State Rd. 128, 0.6 mi. off Blue Ridge Parkway, 5,500´ [1,676 m], Yancey Co., North Carolina, 21 Aug. 1974, J. B. Sullivan, ELQ. Papaipema nelita (Strecker), ♂. L’Assomption, Quebec, 21 Aug. 1979, CNC. Papaipema sablensis Quinter, ♂. Holotype. Sable Island, Nova Scotia, 11 Aug. 1978, D. C. Ferguson, CNC. Papaipema sablensis Quinter, ♂. Paratype. Meteorological Station, Sable Island, Nova Scotia, 10 Sep. 1977, Barry Wright, CNC. Papaipema sauzalitae (Grote), ♀. 4 mi. W of Sixes, Curry Co., Oregon, 15 Jul. 2017, ex larva in Angelica hendersonii, reared in carrot, eclosed 29 Aug. 2017, A. McBride and J. Wiker, AEM. Papaipema sauzalitae (Grote), ♂. Petaluma, Sonoma Co., California, 30 Sep. 1937, E. C. Johnston, ELQ. Papaipema sauzalitae (Grote), ♀. Inverness, Marin Co., California, 10 Oct. 1950, W. R. Bauer, ELQ. Papaipema sauzalitae (Grote), ♀. Anderson Springs, Lake Co., California, 24 Sep. 1949, W. R. Bauer, ELQ. Papaipema sauzalitae (Grote), ♀. Oso Flaco Lake, 5 mi. S of Oceano, San Luis Obispo Co., California, 3 Oct. 1972, at bl. and wh. Lights, J. Powell, AEM. Papaipema angelica (Smith), ♂. San Bernardino, San Bernardino Co., California, C. M. Dammers, 27 Jul. 1935, ELQ. Papaipema angelica (Smith), ♀. San Bernardino, San Bernardino Co., California, C. M. Dammers, 19 Jul. 1935, ELQ. Papaipema angelica (Smith), ♀. Huntington Beach, Orange Co., California, ex Psoralea macrostachya, emerged 24 Aug. 1930, ELQ. Papaipema astuta Bird, ♂. Johnsonburg, Warren Co., New Jersey, bred ex larva in Collinsonia canadensis, emerged 29 Aug. 1994, E. L. Quinter, ELQ. Papaipema astuta Bird, ♀. Fredon-Springdale Rd., 1 mi. W of Springdale, Sussex Co., New Jersey, bred ex larva in Collinsonia canadensis, emerged 6 Sep. 1976, E. L. Quinter, ELQ. Papaipema astuta Bird, ♂. T5S, R13W, Sec. 30, Cass Co., Michigan, UV trap, 17 Sep. 1991, M. C. Nielsen, ELQ. Papaipema astuta Bird, ♂. White Lake Wildlife Management Area, Hardwick Twp., Warren Co., New Jersey, 26 Jul. 2017, ex larva in Collinsonia canadensis, reared in potato, eclosed 29 Aug. 2017, A. E. McBride, AEM. Papaipema astuta Bird, ♀. Black Mountain Ridge Rd., 4 mi. S of Lynch, 36.9159, -82.8939, 3,881´ [1,183 m], Harlan Co., Kentucky, 1 Jul. 2016, ex larva in Angelica triquinata, reared in potato, eclosed 3 Sep. 2016, A. McBride, L. Gibson, E. Laudermilk, AEM. Papaipema harrisii (Grote), ♂. Beinn Breigh, Cape Breton Island, Nova Scotia, bred ex larva in Heracleum sphondylium, emerged 31 Aug. 1980, E. L. Quinter, ELQ. Papaipema harrisii (Grote), ♂. Beinn Breigh, Cape Breton Island, Nova Scotia, bred ex larva in Heracleum sphondylium, emerged 24 Aug. 1980, E. L. Quinter, ELQ. Papaipema harrisii (Grote), ♂. Beinn Breigh, Cape Breton Island, Nova Scotia, bred ex larva in Heracleum sphondylium, emerged 29 Aug. 1980, E. L. Quinter, ELQ. Papaipema harrisii (Grote), ♂. South Turkey Creek Rd. across from train park, Tinytown, 39°36.17’N, 105°13.33’W, 6,870´ [2,094 m], Jefferson Co., Colorado, MV trap in parking lot, 20 Aug. 2007, 11:00 pm, Chuck Harp, ELQ. Papaipema harrisii (Grote), ♂. Edmundston, New Brunswick, 24 Aug. 1992, Henry Hensel, CNC. Papaipema harrisii (Grote), ♀. Edmundston, New Brunswick, 7 Sep. 1993, Henry Hensel, CNC. Papaipema rubiginosa Bird, ♀. Giant Fen, N of Old Prospect School Rd., Hardyston Twp., Sussex Co., New Jersey, bred ex larva in Angelica atropurpurea, emerged 31 Aug. 2008, A. E. McBride, ELQ. Papaipema rubiginosa Bird, ♀. Rose Lake State Game Area, Clinton Co., Michigan, bred ex larva in Angelica atropurpurea, emerged 5 Sep. 1974, M. C. Nielsen and E. L. Quinter, ELQ. Papaipema rubiginosa Bird, ♂. Thompson Rd. Prairie, T7S, R16W, Sec. 5, Cass Co., Michigan, ex larva in Angelica atropurpurea, emerged 16 Aug. 1999, M. C. Nielsen, ELQ. Papaipema rubiginosa Bird, ♀. Bakertown Fen, Berrien Co., Michigan, bred ex larva in Angelica atropurpurea, emerged 21 Aug. 1979, M. C. Nielsen and E. L. Quinter, ELQ. Papaipema rubiginosa Bird, ♀. Stewart Park, Dane Co., Wisconsin, ex Angelica, emerged 13 Aug. 1999, E. L. Quinter, ELQ.
PLATE A-7: Figs 1–29; SCALE 1.25:1 Papaipema rubiginosa Bird, ♂. Elgin, Cook Co., Illinois, 27 Aug. 1930, M. Bristol, ELQ. (p.147). Papaipema rubiginosa Bird, ♂. Giant Fen, N of Old Prospect School Rd., 41.1137, -74.6274, 534´ [163 m], Hardyston Twp., Sussex Co., New Jersey, 25 Jun. 2010 ex larva in Angelica atropurpurea, reared in carrot, emerged 9 Aug. 2010, A. E. McBride, AEM. Papaipema rubiginosa Bird, ♀. Giant Fen, N of Old Prospect School Rd., 41.1137, -74.6274, 534´ [163 m], Hardyston Twp., Sussex Co., New Jersey, bred ex larva in Angelica atropurpurea, emerged 26 Aug. 2008, A. E. McBride, ELQ. Papaipema rubiginosa Bird, ♀. Giant Fen, N of Old Prospect School Rd., 41.1137, -74.6274, 534´ [163 m], Hardyston Twp., Sussex Co., New Jersey, 20 Jun. 2023 ex larva in Angelica atropurpurea, reared in carrot, emerged 14 Aug. 2023, A. E. McBride, AEM. Papaipema rubiginosa Bird, ♀. Giant Fen, N of Old Prospect School Rd., 41.1137, -74.6274, 534´ [163 m], Hardyston Twp., Sussex Co., New Jersey, 28 Jun. 2020 ex larva in Angelica atropurpurea, reared in carrot, emerged 30 Aug. 2020, A. E. McBride, CNC. Papaipema hesseli Quinter, ♀. Paratype. Pittsfield, Berkshire Co., Massachusetts, ex larva in Matteuccia struthiopteris, collected 26 Jul. 1958, emerged 17 Sep. 1958, S. A. Hessel, YPM. Papaipema hesseli Quinter, ♀. Paratype. Normans Kill, 42.6799, -73.9063, 202´ [62 m], Albany Co., New York, 5 Sep. 1994, bred ex larva in Matteuccia struthiopteris, E. L. Quinter, ELQ. Papaipema hesseli Quinter, ♂. Paratype. Jock River, Moodie Drive bridge, 45.2493, -75.7916, Ottawa-Carleton, Ontario, floodplain/Matteuccia, 11 Sep. 2019, B. C. Schmidt, UV, CNC (CNCLEP # 231913). Papaipema hesseli Quinter, ♂. Paratype. Rt. 128, 1.6 mi. E of West Cornwall, Litchfield Co., Connecticut, bred ex larva in Matteuccia struthiopteris, emerged 3 Sep. 1975, E. L. Quinter, ELQ. Papaipema hesseli Quinter, ♀. Paratype. Normans Kill, 42.6799, -73.9063, 202´ [62 m], Albany Co., New York, 18 Sep. 1989, T. L. McCabe, ELQ. Papaipema hesseli Quinter, ♀. Paratype. Rt. 128, 1.6 mi. E of West Cornwall, Litchfield Co., Connecticut, bred ex larva in Matteuccia struthiopteris, emerged 11 Sep. 1975, E. L. Quinter, ELQ. Papaipema hesseli Quinter, ♀. Holotype. Normans Kill, 42°40’40” 73°54’23”, 30 m., Albany Co., New York, 21 Sep. 1989, T. L. McCabe, CNC (CNCLEP # 116420). Papaipema pterisii Bird, ♀. Summer Hill, 1.9 mi. W of Auburn, 860´ [262 m], Schuylkill Co., Pennsylvania, bred ex larva in Pteridium aquilinum, emerged 29 Aug. 1976, E. L. Quinter, ELQ. Papaipema pterisii Bird, ♀. Summer Hill, 1.9 mi. W of Auburn, 860´ [262 m], Schuylkill Co., Pennsylvania, bred ex larva in Pteridium aquilinum, emerged 4 Sep. 1982, E. L. Quinter, ELQ. Papaipema pterisii Bird, ♂. Clarks Corner, 0.6 mi ESE of BM 577 power line cut, 630´ [192 m], Hampton, Windham Co., Connecticut, 24–25 Aug. 2011, 15w BLT, E. L. Quinter, ELQ. Papaipema pterisii Bird, ♂. Enfield, Penobscot Co., Maine, 3 Sep. 1981, L. P. Grey, ELQ. Papaipema pterisii Bird, ♀. Triune, Monongalia Co., West Virginia, bred ex larva in Pteridium aquilinum, emerged 10 Aug. 1983, E. L. Quinter, ELQ. Papaipema pterisii Bird, ♂. Clarks Corner, 0.6 mi ESE of BM 577 power line cut, 630´ [192 m], Hampton, Windham Co., Connecticut, 10–12 Sep. 2011, 15w BLT, E. L. Quinter, ELQ. Papaipema lysimachiae Bird, ♀. Appalachian Trail, 3.1 mi. ESE of Summit Station, 1,600´ [488 m], Schuylkill Co., Pennsylvania, bred ex larva in Lysimachia quadrifolia, 11 Sep. 1972, E. L. Quinter, ELQ. Papaipema lysimachiae Bird, ♂. Whiting, Ocean Co., New Jersey, bred ex larva in Lysimachia terrestris, larva collected 21 Jun. 1994, emerged 28 Aug. 1994, E. L. Quinter, ELQ. Papaipema leucostigma (Harris), ♂. Sand Road Quarry, Canaan, 800´ [244 m], Litchfield Co., Connecticut, bred ex larva in Aquilegia canadensis, emerged 16 Sep. 2007, E. Quinter and D. Wagner, ELQ. Papaipema leucostigma (Harris), ♀. Wausau, Marathon Co., Wisconsin, 22 Aug. 1952, H. M. Bower, ELQ. Papaipema leucostigma (Harris), ♀. Wausau, Marathon Co., Wisconsin, ex pupa, 2 Sep. 1962, H. M. Bower, ELQ. Papaipema limata Bird, ♂. Payson, Utah Co., Utah, 20 Sep. 1993, Joel Johnson, CNC. Papaipema limata Bird, ♂. High Creek Canyon Rd., Cove, 41.9581, -111.7857, 4,754´ [1,149 m], Cache Co., Utah, 29 Jun. 2023 ex larva in Apocynum cannabinum, reared in carrot, eclosed 16 Aug. 2023, A. E. McBride, AEM. Papaipema limata Bird, ♀. High Creek Canyon Rd., Cove, 41.9581, -111.7857, 4,754´ [1,149 m], Cache Co., Utah, 29 Jun. 2023 ex larva in Apocynum cannabinum, reared in carrot, eclosed 16 Aug. 2023, A. E. McBride, AEM. Papaipema limata Bird, ♀. Dayton, 1,620´ [494 m], Columbia Co., Washington, 27 Sep. 1957, R. E. Miller, ELQ. Papaipema depictata Benjamin, ♀. 42D Forest Rd., 0.4 mi. SE of Rustler Park, 31.8987, -109.2759, 8,650´ [2,637 m], Cochise Co., Arizona, 23 Jun. 2022 ex larva in Hymenoxys hoopesii, reared in carrot, eclosed 26 Jul. 2022, A. E. McBride and J. Wiker, AEM. Papaipema depictata Benjamin, ♂. 42D Forest Rd., vicinity of Long Park sign, 31.8882, -109.2792, 9,010´ [2,746 m], Cochise Co., Arizona, 23 Jun. 2022 ex larva in Hymenoxys hoopesii, reared in carrot, eclosed 30 Jul. 2022, A. E. McBride and J. Wiker, AEM.
: Figs 1–24; SCALE 1.25:1 Papaipema rutila (Guenée), ♀. Moth Ridge, 4 km. NE of Ruff Creek, 39°59´N, 80°09´W, 240 m., Greene Co., Pennsylvania, 24 Sep. 1994, J. E. Rawlins, ELQ. Papaipema rutila (Guenée), ♀. Valley Station, Jefferson Co., Kentucky, 9 Sep. 1974, ELQ. Papaipema rutila (Guenée), ♀. 5 mi. W of Greenview, T19N, R7W, Sec. 12, Menard Co., Illinois, ex larva in Podophyllum peltatum, emerged 29 Sep. 2007, J. R. Wiker, ELQ. Papaipema rutila (Guenée), ♂. Neotype. Area of Beadles Barrens N.P.A. About 4 miles SW of Albion, N38.351218-W088.126076, Edwards Co., Illinois, 20 Jun. 2019, larva boring in stem/root of Podophyllum peltatum, pupated 16 Aug. 2019, emerged 17 Sep. 2019, James R. Wiker, CNC. Papaipema rutila (Guenée), ♂. Black Farm, 6 mi. S of Paducah, 485´ [148 m], McCracken Co., Kentucky, 4 Oct. 1974, at light, William R. Black, Jr., ELQ. Papaipema baptisiae (Bird), ♂. Summer Hill, 1.9 mi. W of Auburn, 860´ [262 m], Schuylkill Co., Pennsylvania, 15 Sep. 1972, UV trap, E. L. Quinter, ELQ. Papaipema baptisiae (Bird), ♂. Summer Hill, 1.9 mi. W of Auburn, 860´ [262 m], Schuylkill Co., Pennsylvania, 3 Sep. 1972, UV trap, E. L. Quinter, ELQ. Papaipema baptisiae (Bird), ♀. Summer Hill, 1.9 mi. W of Auburn, 860´ [262 m], Schuylkill Co., Pennsylvania, bred ex larva in Apocynum, 23 Aug. 1994, E. L. Quinter, ELQ. Papaipema baptisiae (Bird), ♀. Summer Hill, 1.9 mi. W of Auburn, 860´ [262 m], Schuylkill Co., Pennsylvania, bred ex larva in Apocynum, 31 Aug. 1994, E. L. Quinter, ELQ. Papaipema baptisiae (Bird), ♀. Lodi Marsh, Hawk Hill, Dane Co., Wisconsin, ex Apocynum, emerged 7 Sep. 1999, E. L. Quinter, ELQ. Papaipema baptisiae (Bird), ♂. Rt. 168, 6 mi. SW of Darlington, Beaver Co., Pennsylvania, 10 Sep. 1967, at UV light, M. A. Carter, ELQ. Hydroeciodes serrata (Grote), ♂. 72 Beaver Flat Trail, Absarokee, Stillwater Co., Montana, 22–25 Jul. 2016, MV and UV lights, Ben D. Williams, ELQ. Papaipema arctivorens Hampson, ♂. Washington, Litchfield Co., Connecticut, 10 Sep. 1965, S. A. Hessel, ELQ. Papaipema arctivorens Hampson, ♂. WMA, Rt. 3, 1 mi. S of Southwick Beach State Park, Jefferson Co., New York, pupa collected in Arctium, emerged 21 Aug. 1977, E. L. Quinter, ELQ. Papaipema arctivorens Hampson, ♂. Richardson Wildlife Foundation remnant prairie, T20N, R11E, Sec. 13, Lee Co., Illinois, bred ex larva in Cacalia tuberosa, emerged 24 Sep. 2007, J. R. Wiker, ELQ. Papaipema araliae Bird & Jones, ♀. Cretaceous Hills Area, 3 mi. SW of Bay City, Pope Co., Illinois, bred ex larva in Aralia spinosa, 26 Jul. 1996, emerged 3 Sep. 1996, E. L. Quinter, ELQ. Papaipema araliae Bird & Jones, ♀. Great Dismal Swamp NWR, City of Suffolk, Virginia, bred ex larva in Aralia spinosa, emerged 25 Sep. 1997, E. L. Quinter, ELQ. Papaipema araliae Bird & Jones, ♀. Waverly Park, 1.5 mi. ENE of Waverly Hills, Jefferson Co., Kentucky, bred ex larva in Aralia spinosa, emerged 16 Aug. 1978, E. L. Quinter, ELQ. Papaipema araliae Bird & Jones, ♀. Croatan National Forest, Junction of Forest Rts. 167 and 132, Craven Co., North Carolina, bred ex larva in Aralia spinosa, emerged 12 Oct. 1978, E. L. Quinter, ELQ. Papaipema polymniae Bird, ♂. ca. 5 mi. SE of Fairmount, Junction Salacoa Rd. and Salacoa Creek, E of rd., N/S sides of creek, Bartow Co., Georgia, light trap, 28 Sep. 2001, James K. Adams, ELQ. Papaipema polymniae Bird, ♀. Pine Hills Campground, T11S, R3W, Sec. 34, Union Co., Illinois, bred ex larva in Smallanthus uvedalia, emerged 8 Oct. 2007, James R. Wiker, ELQ. Papaipema polymniae Bird, ♂. Rt. 649, 2.3 mi. NW of Glen Lyn, along New River, Giles Co., Virginia, bred ex mature larva in Smallanthus uvedalia, collected 28 Jul. 1974, emerged 25 Sep. 1974, E. L. Quinter, ELQ. Papaipema polymniae Bird, ♂. River Rd. along French Broad River, 2.7 mi. NW of Hot Springs, 35.9239, -82.8598, 1,290´ [393 m], Madison Co., North Carolina, ex larva 23 Jul. 2023 in Smallanthus uvedalia, reared in carrot, eclosed 6 Oct. 2023, A. E. McBride, AEM. Papaipema polymniae Bird, ♀. River Rd. along French Broad River, 2.7 mi. NW of Hot Springs, 35.9239, -82.8598, 1,290´ [393 m], Madison Co., North Carolina, ex larva 23 Jul. 2023 in Smallanthus uvedalia, reared in carrot, eclosed 9 Oct. 2023, A. E. McBride, AEM.
: Figs 1–27; SCALE 1.25:1 Papaipema nebris (Guenée), ♂. 2 mi. NE of Greenview, T19N, R5W, Sec. 6, Menard Co., Illinois, ex larva in Ambrosia trifida, emerged 22 Sep. 2007, J. R. Wiker, ELQ. Papaipema nebris (Guenée), ♀. Berwyn, Cook Co., Illinois, ex Ambrosia, 17 Sep. 1928, E. Beer, ELQ. Papaipema nebris (Guenée), ♀. Junction of Follett Rd. and Potters Brook, 240´ [73 m], 0.7 mi. ESE of Windham, Windham Co., Connecticut, bred ex larva in Bidens, emerged 15 Sep. 2005, E. L. Quinter, ELQ. Papaipema nebris (Guenée), ♂. Edgard, Saint John the Baptist Parish, Louisiana, UV light trap, 22 Oct. 1980, V. A. Brou, ELQ. Papaipema nebris (Guenée), ♀. Massac Creek at Hinkleville Rd. (Old US 60), McCracken Co., Kentucky, UV, 3–4 Oct. 1999, M. C. Nielsen, ELQ. Papaipema limpida (Guenée), ♂. Sarcoxie, Jasper Co., Missouri, bred ex larva in Vernonia baldwinii, emerged 12 Sep. 1977, Rae Letsinger and Eric Quinter, ELQ. Papaipema limpida (Guenée), ♀. Short Fork Marsh, 3 mi. E of Good Hope, T7N, R2W, Sec. 27, McDonough Co., Illinois, ex larva in Vernonia 25 Jul. 2003, emerged 4 Sep. 2003, J. R. Wiker, JRW. Papaipema limpida (Guenée), ♀. Short Fork Marsh, 3 mi. E of Good Hope, T7N, R2W, Sec. 27, McDonough Co., Illinois, ex larva in Vernonia, emerged 19 Sep. 2007, J. R. Wiker, ELQ. Papaipema limpida (Guenée), ♂. Rt. 168, 6 mi. SW of Darlington, Beaver Co., Pennsylvania, 22 Sep. 1966, M. A. Carter, ELQ. Papaipema limpida (Guenée), ♀. Madison Twp., Polk Co., Iowa, ex larva in Vernonia missurica 31 May 2018, reared in potato, eclosed 9 Sep. 2018, A. Brees, AEM. Papaipema maritima Bird, ♀. Jones Island, 2.8 mi. SSW of Cedarville, Cumberland Co., New Jersey, ex pupa in Helianthus giganteus 23 Sep. 1979, emerged 11 Oct. 1979, E. L. Quinter, ELQ. Papaipema maritima Bird, ♀. Tucker Prairie, 3 mi. W of Kingdom City, Callaway Co., Missouri, ex larva in Helianthus mollis, emerged 9 Oct. 1975, E. L. Quinter, ELQ. Papaipema maritima Bird, ♀. W of Greenview, T19N, R7W, Sec. 10, Menard Co., Illinois, ex larva in Helianthus, emerged 28 Sep. 2007, J. R. Wiker, ELQ. Papaipema necopina (Grote), ♀. Rt. 28, 1 mi. W of West Lebanon, Warren Co., Indiana, ex larva in Helianthus tuberosus 3 Aug. 1974, emerged 19 Sep. 1974, E. L. Quinter, ELQ. Papaipema necopina (Grote), ♀. Knightsville Dam, Hampshire Co., Massachusetts, ex larva in Helianthus tuberosus, emerged 10 Sep. 2010, E. L. Quinter, ELQ. Papaipema duplicata Bird, ♂. Thunder Hill, mile post 76, Blue Ridge Parkway, 3,920´ [1,195 m], Bedford Co., Virginia, MV light, 25 Sep. 1994, E. Quinter, P. Goldstein, S. Roble, ELQ. Papaipema duplicata Bird, ♀. Black Mountain Ridge Road, 4 mi. S of Lynch, 36.9159, -82.8939, 3,881´ [1,183 m], Harlan Co., Kentucky, 1 Jul. 2016 ex larva in Collinsonia canadensis, reared in potato, eclosed 9 Oct. 2016, A. E. McBride, AEM. Papaipema duplicata Bird, ♀. Thunder Hill, mile post 76, Blue Ridge Parkway, 3,920´ [1,195 m], Bedford Co., Virginia, ex larva in Collinsonia canadensis, emerged 7 Oct. 2008, A. E. McBride, ELQ. Papaipema silphii Bird, ♂. 1.5 mi. SE of Lee’s Summit, Jackson Co., Missouri, ex larva in Silphium laciniatum, emerged 13 Sep. 1975, E. L. Quinter, ELQ. Papaipema silphii Bird, ♀. Cline property along Gudgel Rd., 1 mi. SW of Athens, Menard Co., Illinois, ex larva in Silphium terebinthinaceum 30 Jul. 2004, emerged 28 Sep. 2004, James R. Wiker, JRW. Papaipema beeriana Bird, ♀. Richardson Wildlife Foundation, prairie S of Woody Pond, T20N, R11E, Sec. 13, NW quarter, Lee Co., Illinois, ex larva in Liatris, emerged 26 Sep. 2007, J. R. Wiker, ELQ. Papaipema silphii Bird, ♀. Chicago, Cook Co., Illinois, 28 Sep. 1917, A. K. Wyatt, ELQ. Papaipema beeriana Bird, ♂. Chicago, Cook Co., Illinois, H. H. Newcomb, ELQ. Papaipema beeriana Bird, ♂. barrens S of Lovells Rd., Chester, Otsego Co., Michigan, 44.8999, -84.5342, 1,230´ [375 m], ex larva in Liatris aspera 14 Jun. 2011, emerged 2 Sep. 2011, A. McBride, M. Nielsen, E. Quinter, and T. Herig, ELQ. Papaipema eupatorii (Lyman), ♀. Hawley Bog, 4 mi. S of Charlemont, 1,780´ [543 m], Franklin Co., Massachusetts, bred ex larva in Eutrochium purpureum, emerged 21 Sep. 1982, E. L. Quinter, ELQ. Papaipema eupatorii (Lyman), ♂. Edgebrook, Cook Co., Illinois, ex Eutrochium, 22 Sep. 1919, E. Beer, ELQ. Papaipema eupatorii (Lyman), ♀. 1 mi. SE of Schuylkill Haven, Schuylkill Co., Pennsylvania, bred ex larva in Eutrochium purpureum, emerged 2 Oct. 1972, E. L. Quinter, ELQ.
: Figs 1–29; SCALE 1.25:1 Papaipema rigida (Grote), ♀. Maiden Creek Rd., 2.6 mi. E of Molltown, Berks Co., Pennsylvania, ex larva in Helianthus decapetalus, emerged 7 Oct. 1972, E. L. Quinter, ELQ. Papaipema rigida (Grote), ♂. Richardson Wildlife Foundation, prairie E of Fell Rd., T20N, R11E, Sec. 14, SE quarter, Lee Co., Illinois, ex larva in Zizia aurea, emerged 1 Oct. 2007, J. R. Wiker, ELQ. Papaipema rigida (Grote), ♀. Maiden Creek Rd., 2.6 mi. E of Molltown, Berks Co., Pennsylvania, ex larva in Helianthus decapetalus, emerged 7 Oct. 1972, E. L. Quinter, ELQ. Papaipema cataphracta (Grote), ♀. Massac Creek, Highway 60, Paducah, McCracken Co., Kentucky, ex larva in Verbesina alternifolia, emerged 28 Sep.–6 Oct. 1999, E. L. Quinter, ELQ. Papaipema cataphracta (Grote), ♂. Rt. 92, 1 mi. S of Evansville, Preston Co., West Virginia, bred ex larva in Vernonia, emerged 12 Oct. 1983, E. L. Quinter, ELQ. Papaipema cataphracta (Grote), ♂. vicinity of Cary, McHenry Co., Illinois, 10 Oct. 1976, Irwin Leeuw, ELQ. Papaipema cataphracta (Grote), ♂. 7 mi. W of Reading, Berks Co., Pennsylvania, ex pupa in Thalictrum, emerged 21 Sep. 1972, E. L. Quinter, ELQ. Papaipema cataphracta (Grote), ♂. Abbott Rd., Buffalo, Erie Co., New York, ex Arctium, 28 Sep. 1909, F. E. Moeser, Genitalia Slide #23, J. Bauer, CMNH. Papaipema cataphracta (Grote), ♀. 6 mi. N of Wolf Lake, T11N, R3W, Sec. 4, Union Co., Illinois, ex larva in Smallanthus uvedalia 3 Aug. 2003, emerged 6 Oct. 2003, James R. Wiker, JRW. Papaipema sulphurata Bird, ♀. Martha’s Vineyard, Dukes Co., Massachusetts, bred ex larva in Decodon verticillatus, emerged 1996, P. Z. Goldstein, ELQ. Papaipema sulphurata Bird, ♀. Barnstable, Barnstable Co., Massachusetts, 28 Sep. 1950, C. P. Kimball, ELQ. Papaipema sulphurata Bird, ♀. Rt. 151, W of Old Barnstable Rd., vicinity of Mashpee, Barnstable Co., Massachusetts, ex pupa in Decodon verticillatus, emerged 21 Sep. 1994, P. Z. Goldstein, ELQ. Papaipema sulphurata Bird, ♂. Martha’s Vineyard, Dukes Co., Massachusetts, bred ex larva in Decodon verticillatus, emerged 1996, P. Z. Goldstein, ELQ. Papaipema duovata (Bird), ♂. Jones Island, 2.8 mi. SSW of Cedarville, Cumberland Co., New Jersey, emerged ex pupa in Solidago sempervirens 26 Sep. 1979, E. L. Quinter, ELQ. Papaipema duovata (Bird), ♀. Port Norris, Cumberland Co., New Jersey, bred ex larva in Solidago sempervirens 10 Jul. 1994, pupated 29 Aug. 1994, emerged 27 Sep. 1994, E. L. Quinter, ELQ. Papaipema aweme (Lyman), ♂. Bark Bay Slough State Natural Area, 3 mi. NE of Herbster, Bayfield Co., Wisconsin, 26 Jun. 2017 ex larva in Menyanthes trifoliata, eclosed 12 Aug. 2017, A. McBride and L. Ferge, AEM. Papaipema aweme (Lyman), ♀. Wanamaker Lake Peatland E of Rt. 421, 46.3636, -85.7672, 725´ [221 m], McMillan, Luce Co., Michigan, 24 Jun. 2017 ex larva in Menyanthes trifoliata, eclosed 26 Jul. 2017, A. E. McBride, AEM. Papaipema cerina (Grote), ♀. Richard Bong State Recreation Area, Kansasville, 42.6321, -88.1625, 792´ [241 m], Kenosha Co., Wisconsin, reared by A. E. McBride in 2019 ex ovo from parent female caught 17 Sep. 2018 by Steve Bransky, AEM. Papaipema cerina (Grote), ♀. Swamp Lovers Preserve, 43.1332, -89.6652, 1,023´ [312 m], Cross Plains, Dane Co., Wisconsin, 2 Jul. 2014 ex larva in Elymus riparius, reared in carrot, eclosed 14 Sep. 2014, A. E. McBride, AEM. Papaipema sullivani Quinter, ♀. Paratype. Great Dismal Swamp National Wildlife Refuge, City of Suffolk, Virginia, bred ex larva in Arundinaria tecta on artificial diet, emerged 25 Sep. 2000, E. Quinter, P. Goldstein, and S. Roble, ELQ. Papaipema sullivani Quinter, ♂. Paratype. Great Dismal Swamp National Wildlife Refuge, City of Suffolk, Virginia, bred ex larva in Arundinaria tecta on artificial diet, emerged 23 Sep. 2000, E. Quinter, P. Goldstein, and S. Roble, ELQ. Papaipema sullivani Quinter, ♀. Paratype. 0.2 mi. S of junction of Rts. 101 and 1154, Carteret Co., North Carolina, bred ex larva in Arundinaria tecta, emerged 13 Oct. 1980, E. L. Quinter, ELQ. Papaipema sullivani Quinter, ♂. Paratype. 0.2 mi. S of junction of Rts. 101 and 1154, Carteret Co., North Carolina, bred ex larva in Arundinaria tecta 1 Aug. 1978, emerged 6 November 1978, E. L. Quinter, ELQ. Papaipema sullivani Quinter, ♀. Holotype. Great Dismal Swamp National Wildlife Refuge, City of Suffolk, Virginia, bred ex larva in Arundinaria tecta, emerged 30 Sep. 1999, E. L. Quinter, CNC. Papaipema matheri Quinter, ♀. Paratype. USAE Waterways Experiment Station, Vicksburg, Warren Co., Mississippi, bred ex larva in Arundinaria gigantea, emerged 26 Oct. 1980, E. L. Quinter, ELQ. Papaipema matheri Quinter, ♀. Holotype. McCrite Farm, N of cemetery, 37.2418, -89.3258, 387´ [118 m], Diswood, Alexander Co., Illinois, 30 Jun. 2014 ex larva in Arundinaria gigantea, reared in potato/carrot, eclosed 2 Oct. 2014, A. E. McBride, CNC. Papaipema matheri Quinter, ♂. Paratype. Cordell Hull Lake, Big Bottom, Jackson Co., Tennessee, ex larva in Arundinaria gigantea 23 Jun. 1995, pupated 27 Aug. 1995, emerged 25 Sep. 1995, E. L. Quinter, ELQ. Papaipema matheri Quinter, ♀. Paratype. unnamed creek at Berkley, cane #5, Carlisle Co., Kentucky, UV light trap, 8 Oct. 2004, William R. Black, Jr., ELQ. Papaipema matheri Quinter, ♂. Paratype. 5 mi. SW of Bardwell, E of old G.M.&O. R.O.W., in canebrake along Sandy Branch, Carlisle Co., Kentucky, bred ex larva in Arundinaria gigantea 14 Jun. 1999, pupated 19 Aug. 1999, emerged 16 Sep. 1999, James R. Wiker, ELQ.
Figs 1–24; SCALE 1.25:1 Papaipema matheri Quinter, ♂. Paratype. 0.25 mi. W of Kincaid Mounds, cane site #2, in wooded slough, Massac Co., Illinois, UV light trap, 18 Oct. 2000, William R. Black, Jr., ELQ. Papaipema matheri Quinter, ♂. Paratype. Massac Creek, McCracken Co., Kentucky, ex pupa in Arundinaria gigantea 7 Sep. 2002, emerged 1 Oct. 2002, W. R. Black, Jr., CNC (CNCLEP # 116422). Papaipema matheri Quinter, ♀. Paratype. Terrapin Creek State Nature Preserve, Area #4, 0.5 mi. SW of Bell City, Graves Co., Kentucky, 13 Oct. 2000, W. R. Black, Jr., CNC (CNCLEP # 116423). Papaipema speciosissima (Grote & Robinson), ♂. Lakehurst, Ocean Co., New Jersey, bred ex larva in Osmunda, emerged 30 Sep. 1973, E. L. Quinter, ELQ. Papaipema speciosissima (Grote & Robinson), ♂. Hessville, Lake Co., Indiana, bred ex larva in Osmunda regalis, emerged 25 Aug. 1929, A. Kwiat, ELQ. Papaipema speciosissima (Grote & Robinson), ♀. Lakehurst, Ocean Co., New Jersey, bred ex larva in Osmunda, emerged 20 Sep. 1973, E. L. Quinter, ELQ. Papaipema speciosissima (Grote & Robinson), ♀. Everglades National Park, Florida, 12 May 1972, Charles V. Covell, Jr., ELQ. Papaipema inquaesita (Grote & Robinson), ♀. Junction of Follett Rd. and Potters Brook, 240´ [73 m], 0.7 mi. ESE of Windham, Windham Co., Connecticut, UV/MV lights, 28 Sep. 2008, E. L. Quinter, ELQ. Papaipema inquaesita (Grote & Robinson), ♂. Junction of Follett Rd. and Potters Brook, 240´ [73 m], 0.7 mi. ESE of Windham, Windham Co., Connecticut, bait trap, 25 Sep. 2007, E. L. Quinter, ELQ. Papaipema stenocelis (Dyar), ♀. Rt. 563, 3 mi. S of Chatsworth, Burlington Co., New Jersey, bred ex larva in Woodwardia virginica, emerged 3 Sep. 1973, E. L. Quinter, ELQ. Papaipema stenocelis (Dyar), ♀. Howardsville cranberry bogs, 39.8141, -74.3665, 135´ [41 m], Ocean Co., New Jersey, 30 Jul. 2019 ex larva, collected from and reared in Woodwardia virginica, eclosed 21 Sep. 2019, A. E. McBride, AEM. Papaipema stenocelis (Dyar), ♂. Rt. 563, 3 mi. S of Chatsworth, Burlington Co., New Jersey, bred ex larva in Woodwardia virginica, emerged 8 Sep. 1973, E. L. Quinter, ELQ. Papaipema stenocelis (Dyar), ♀. Alligator Tram, Forest Rt. 124, 2 mi. N of Mill Creek Rd., Croatan National Forest, Carteret Co., North Carolina, bred ex larva in Woodwardia virginica, emerged 28 Sep. 1978, E. L. Quinter, ELQ. Papaipema stenocelis (Dyar), ♀. Rt. 563, 3 mi. S of Chatsworth, Burlington Co., New Jersey, bred ex larva in Woodwardia virginica, emerged 11 Sep. 1973, E. L. Quinter, ELQ. Papaipema [probable hybrid impecuniosa? × cataphracta?], ♂. Goose Lake Prairie, Grundy Co., Illinois, 18 Sep. 1989, R. Panzer, ELQ. Papaipema [probable hybrid], ♀. Union Co., New Jersey, 17 Sep. 1912, O. Buchholz, AMNH. Papaipema [probable hybrid], ♂. Union Co., New Jersey, 12 Sep. 1912, O. Buchholz, AMNH. Papaipema aerata (Lyman) [probable hybrid necopina × cataphracta], ♂. Montreal, Quebec, bred ex Arctium, 12 Sep. 1909, H. Lyman, AMNH. Papaipema [probable hybrid rubiginosa? × cataphracta], ♂. Chicago, Cook Co., Illinois, ex Arctium, 17 Sep. 1921, E. Beer, ELQ. Papaipema [probable hybrid necopina × cataphracta], ♀. Elgin, Cook Co., Illinois, ex pupa in Arctium, 14 Sep. 1928, E. Beer, ELQ. Papaipema lab reared F1 necopina ♂ × cataphracta ♀. Hybrid ♀, ex ovo from cage mating by Ted Herig, 2017. Parent ♂ Papaipema necopina: Rt. 27 roadside, 2.2 mi. N of St. Johns, 42.9696, -84.5429, 740´ [226 m], Bingham Twp., Clinton Co., Michigan, ex larva in Helianthus, Jul. 2017. Parent ♀ Papaipema cataphracta: 0.1 mi. W of Stoll and Upton Rds., 42.7989, -84.4005, 839´ [256 m], Bath Twp., Clinton Co., Michigan, ex larva Jul. 2017. Ex ovo 2018 in potato, eclosed Jul. 2018, A. E. McBride, AEM. Papaipema [probable hybrid necopina × cataphracta], ♀. Abbott Rd., Buffalo, Erie Co., New York, ex Arctium, 23 Sep. 1909, F. E. Moeser, Genitalia Slide USNM #39451 and #1327 J. G. Franclemont, USNM. Papaipema [probable hybrid necopina × cataphracta], ♂. Buffalo, Erie Co., New York, 23 Sep. 1908, F. E. Moeser, CMNH. Papaipema unimoda (Smith), ♀. 5 mi. E of Eagle Harbor, adjacent to Lake Superior, T59N, R30W, Sec. 36, Keweenaw Co., Michigan, UV trap, 30 Sep. 2002, Mark Netznik, MSU (Michigan State Univ. Arthropod Res. Coll. Cat#: 218988).
Male Genitalia: Color Plates M1–M19
MALE GENITALIA 1. Arundinariaphila hayesi Quinter; Liberty Co., GA; CNC18090. 2. Cherokeea attakullakulla Sullivan & Quinter; Swain Co., NC; CNC18082. 3. C. slotteni Quinter; Jones Co., NC; CNC18084. 4. C. gabrielleae Quinter; Suffolk, VA; CNC18088. Scale bar = 1mm.
MALE GENITALIA 1. Cherokeea odonnellae Quinter; Swain Co., NC; CNC18285. 2. C. trigona (Sm.); Jackson Co., IL; CNC18086. 3. Franclemontia interrogans (Wlk.); Liberty Co., GA; CNC18274. 4. Sarita relicta (Ferguson); Craven Co., NC; CNC18276. Scale bar = 1mm.
MALE GENITALIA 1. Amphipoea interoceanica (Sm.); Ottawa Dist., ON; CNC18092. 2. A.americana (Speyer); St.-Basile, NB; CNC18095. 3. A.pacifica (Sm.); Jackson Co., OR; CNC18100. 4. A.cottlei (McD.); San Bernardino Co., CA; CNC18333. Scale bar = 1mm.
MALE GENITALIA 1. Amphipoea keiferi (Benj.); Whitman Co., WA; CNC18106. 2. A.senilis (Sm.); Grand Co., CO; CNC18104. 3. Fibigeria erepta (Grt.); Charleston Co., SC; CNC18113. 4. Hydraecia medialis Sm.; Princeton, BC; CNC18115. Scale bar = 1mm.
MALE GENITALIA 1. Hydraecia obliqua (Harv.); Island Co., WA; CNC18117. 2. H.intermedia (B. & Benj.); Holotype; Crowsnest Pass, AB; USNM 39390. 3. H.micacea (Esp.); Waterloo, ON; CNC18121. 4. H.perobliqua Hamp.; Spirit Dunes, MB; CNC18124.
MALE genitalia 1. Hydraecia stramentosa Gn.; Ottawa Dist., ON; CNC18134. 2. Hydraecia immanis Gn.; Wyandot Co., OH; CNC18119. 3. Selicanis cinereola Sm.; Washakie Co., WY; CNC18136. 4. Parapamea buffaloensis (Grt.); DeSoto Co., FL; CNC18138. Scale bar = 1mm.
MALE genitalia 1. Papaipema unimoda (Sm.); Edmundston, NB; CNC18140. 2. P.appassionata (Harv.); York Co., NB; CNC18143. 3. P.amianthii Quinter; Carbon Co., PA; CNC18163. 4. P.furcata (Sm.); Arner, ON; CNC18145. Scale bar = 1mm.
MALE genitalia 1. Papaipema cerina (Grt.); Norfolk Co., ON; CNC18142. 2. P. sullivani Quinter; Isle of Wight Co., VA; CNC18261. 3. P. matheri Quinter; Calloway Co., KY; CNC18256. 4. P. marginidens (Gn.); Clay Co., IL; CNC18153. Scale bar = 1mm.
MALE genitalia 1. Papaipema eryngii Bird; Grundy Co., IL; CNC18147. 2. P. cataphracta (Grt.); Vaudreuil-Soulanges Co., QC; CNC18228. 3. P.sulphurata Bird; Nantucket Co., MA; CNC18230. 4. P.aweme (Lyman); Aweme, MB; CNC18246. Scale bar = 1mm.
MALE genitalia 1. Papaipema necopina (Grt.); Haldimand Co., ON; CNC18252. 2. P.duovata (Bird); Carteret Co., NC; CNC18342. 3. P.speciosissima (G. & R.); DeSoto Co., FL; CNC18159. 4. P.inquaesita (G. & R.); Ottawa Dist., ON; CNC18161. Scale bar = 1mm.
MALE genitalia 1. Papaipema stenocelis (Dyar); Alachua Co., FL; CNC18157. 2. P.rigida (Grt.); Edmundston, NB; CNC18242. 3. P.eupatorii (Lyman); Manitoulin Dist., ON; CNC18236. 4. P.nebris (Gn.); Haldimand Co., ON; CNC18234. Scale bar = 1mm.
MALE genitalia 1. Papaipema arctivorens Hamp.; Ottawa Dist., ON; CNC18217. 2. P.araliae Bird & Jones; Alexander Co., IL; CNC18215. 3. P.polymniae Bird; Ashe Co., NC; CNC18213. 4. P. rutila (Gn.); Menard Co., IL; CNC18221. Scale bar = 1mm.
MALE genitalia 1. Papaipema baptisiae (Bird); Haldimand Co., ON; CNC18219. 2. P.limata Bird; Yakima Co., WA; CNC18280. 3. P.depictata Benj.; Taos Co., NM; CNC18224. 4. P.duplicata Bird; Westchester Co., NY; CNC18238. Scale bar = 1mm.
MALE genitalia 1. Papaipema silphii Bird; Cook Co., IL; CNC18254. 2. P.cerussata (Grt.); Essex Co., ON; CNC18149. 3. P.sciata Bird; Clay Co., IL; CNC18151. 4. P.nepheleptena (Dyar); Ottawa Dist., ON; CNC18342. Scale bar = 1mm.
MALE genitalia 1. Papaipema circumlucens (Sm.); Estevan, SK; CNC18165. 2. P.blackorum Quinter; Johnson Co., IL; CNC18258. 3. P.impecuniosa (Grt.); Hanwell, NB; CNC18232. 4. P.nelita (Stkr.); Edmundston, NB; CNC18244. Scale bar = 1mm.
MALE genitalia 1. Papaipema sablensis Quinter; Sable Island, NS; CNC18260. 2. P.insulidens (Bird); Harney Co., OR; CNC18168. 3. P.birdi (Dyar); Marion Co., IL; CNC18170. 4. P.dribi B. & Benj.; Otero Co., NM; CNC18279. Scale bar = 1mm.
MALE genitaia 1. Papaipema albimontana Quinter; Apache Co., AZ; CNC18334. 2. P.sauzalitae (Grt.); Curry Co., OR; CNC18180. 3. P.angelica (Sm.); Orange Co., CA; CNC18182. 4. P.harrisii (Grt.); Madawaska Co., NB; CNC18363. Scale bar = 1mm.
MALE genitalia 1. Papaipema rubiginosa Bird; McHenry Co., IL; CNC18175. 2. P.astuta Bird; Harlan Co., KY; CNC18179. 3. P.leucostigma (Harr.); Pakenham , ON; CNC18186. 4. P.lysimachiae Bird; Ottawa Dist., ON; CNC18184. Scale bar = 1mm.
Female Genitalia: Color Plates F1–F10
FEMALE GENITALIA 1. Arundinariaphila hayesi Quinter; Liberty Co., GA; CNC18091. 2. Cherokeea attakullakulla Sullivan & Quinter; Swain Co., NC; CNC18083. 3. C. slotteni Quinter; St. Tammany Parish, LA; CNC18085. 4. C. gabrielleae Quinter; Suffolk, VA; CNC18089. 5. C. odonnellae Quinter; Swain Co., NC; CNC18286. 6. C. trigona (Sm.); Alexander Co., IL; CNC18087. 7. Franclemontia interrogans (Wlk.); Liberty Co., GA; CNC18275. 8. Sarita relicta (Ferguson); Craven Co., NC; CNC18277. Scale bar = 1mm.
FEMALE GENITLIA 1. Amphipoea interoceanica (Sm.); Haldimand Co., ON; CNC18094. 2. A.americana (Speyer); Edmundston, NB; CNC18097. 3. A.pacifica (Sm.); Clackamas Co., OR; CNC18102. 4. A.cottlei (McD.); San Bernardino Co., CA; CNC18103. 5. A.keiferi (Benj.); Whitman Co., WA; CNC18107. 6. A.senilis (Sm.); San Juan Co., UT; CNC18105. 7. Fibigeria erepta (Grt.); Bronx Co., NY; CNC18114. 8. Hydraecia medialis Sm.; Fergus Co., MT; CNC18116. Scale bar = 1mm.
FEMALE GENITALIA 1. Hydraecia obliqua (Harv.); Fitsgerald, BC; CNC18118. 2. H.micacea (Esp.); Waterloo, ON; CNC18122. 3. H.perobliqua Hamp.; Tofield, AB; CNC18125. 4. H.stramentosa Gn.; Ottawa Dist., ON; CNC18135. 5. H.immanis Gn.; Waterloo, ON; CNC18120. 6. Selicanis cinereola Sm.; Albany Co., WY; CNC18137. 7. Parapamea buffaloensis (Grt.); Henry Co., TN; CNC18139. 8. P.unimoda (Sm.); Edmundston, NB; CNC18281. Scale bar = 1mm.
female genitalia 1. Papaipema appassionata (Harv.); Frontenac Co., ON; CNC18345. 2. P.amianthii Quinter; Carbon Co., PA; CNC18164. 3. P.furcata (Sm.); Cook Co., IL; CNC18283. 4. P.cerina (Grt.); Lenawee Co., MI; CNC18332. 5. P.sullivani Quinter; Carteret Co., NC; CNC18338. 6. P.matheri Quinter; McCracken Co., KY; CNC18257. 7. P.marginidens (Gn.); Swain Co., NC; CNC18154. 8. P.eryngii Bird; Cook Co., IL; CNC18148. Scale bar = 1mm.
female genitalia 1. Papaipema cataphracta (Grt.); Haldimand Co., ON; CNC18229. 2. P.sulphurata Bird; Plymouth Co., MA; CNC18231. 3. P.aweme (Lyman); Lambton Co., ON; CNC18247. 4. P.necopina (Grt.); Arner, ON; CNC18253. 5. P.duovata (Bird); Ocean Co., NJ; CNC18227. 6. P.speciosissima (G. & R.); DeSoto Co., FL; CNC18160. 7. P.inquaesita (G. & R.); Ottawa Dist., ON; CNC18162. 8. P.stenocelis (Dyar); Carteret Co., NC; CNC18158. Scale bar = 1mm.
FEMALE GENTALIA 1. Papaipema rigida (Grt.); Edmundston, NB; CNC18243. 2. P.eupatorii (Lyman); Manitoulin Dist., ON; CNC18237. 3. P.nebris (Gn.); Norfolk Co., ON; CNC18235. 4. P.arctivorens Hamp.; Haldimand Co., ON; CNC18218. 5. P.araliae Bird & Jones; Pope Co., IL; CNC18216. 6. P.polymniae Bird; Bartow Co., GA; CNC18214. 7. P.rutila (Gn.); Hunterdon Co., NJ; CNC18222. 8. P.baptisiae (Bird); Haldimand Co., ON; CNC18220. Scale bar = 1mm.
EMALE GENITALIA 1. Papaipema limata Bird; Cache Co., UT; CNC18331. 2. P.depictata Benj.; Taos Co., NM; CNC18225. 3. P.duplicata Bird; Westchester Co., NY; CNC18239. 4. P.silphii Bird; Cook Co., IL; CNC18255. 5. P.cerussata (Grt.); Union Co., NJ; CNC18150. 6. P.sciata Bird; Jackson Co., MO; CNC18152. 7. P.nepheleptena (Dyar); Ottawa Dist., ON; CNC18156. 8. P.circumlucens (Sm.); Westchester Co., NY; CNC18167. Scale bar = 1mm.
FEMAE GENITALIA 1. Papaipema blackorum Quinter; Carlisle Co., KY; CNC18259. 2. P.impecuniosa (Grt.); Hanwell, NB; CNC18233. 3. P.nelita (Stkr.); Edmundston, NB; CNC18245. 4. P.sablensis Quinter; Sable Island, NS; CNC18284. 5. P.insulidens (Bird); Harney Co., OR; CNC18169. 6. P.birdi (Dyar); Schuylkill Co., PA; CNC18171. 7. P.albimontana Quinter; Apache Co., AZ; CNC18344. 8. P.sauzalitae (Grt.); Curry Co., OR ; CNC18181. Scale bar = 1mm.
FEMALE GENITALIA 1. Papaipema angelica (Sm.); Orange Co., CA; CNC18183. 2. P.harrisii (Grt.); Montreal, QC; CNC18364. 3. P.rubiginosa Bird; Lenawee Co., MI; CNC18346. 4. P.astuta Bird; Harlan Co., KY; CNC18173. 5. P.leucostigma (Harr.); Ottawa Dist., ON; CNC18187. 6. P.lysimachiae Bird; Ottawa Dist., ON; CNC18185. 7. P.pterisii Bird; Lanark Co., ON; CNC18211. 8. P.hesseli Quinter; Albany Co., NY; CNC18212. Scale bar = 1mm.
Larvae: Color Plates L1–L7
Figs 1–12; VARIABLE SIZES Photographs by A. E. McBride. Arundinariaphila hayesi Quinter, larva in Arundinaria gigantea, Ela Public Access Area, Tuckasegee River, Swain Co., North Carolina, A. E. McBride and E. L. Quinter. Arundinariaphila hayesi Quinter, larva in Arundinaria gigantea, Ela Public Access Area, Tuckasegee River, Swain Co., North Carolina, A. E. McBride and E. L. Quinter. Cherokeea attakullakulla Sullivan & Quinter, larva in Arundinaria appalachiana, Fontana Lake Estates, vicinity Trails End Drive, Swain Co., North Carolina, A. E. McBride. Cherokeea slotteni Quinter, larva in Arundinaria appalachiana, Lower Needmore Rd., Little Tennessee River, Swain Co., North Carolina, A. E. McBride and E. L. Quinter. Cherokeea gabrielleae Quinter, larva in Arundinaria tecta, Railroad Ditch, Great Dismal Swamp National Wildlife Refuge, City of Suffolk, Virginia, A. E. McBride. Cherokeea gabrielleae Quinter, larva in Pseudosasa japonica, Port Norris, Cumberland Co., New Jersey, A. E. McBride and D. F. Schweitzer. Cherokeea odonnellae Quinter, larva in Arundinaria appalachiana, Lower Needmore Rd., Little Tennessee River, Swain Co., North Carolina, A. E. McBride and E. L. Quinter. Cherokeea odonnellae Quinter, larva in Arundinaria appalachiana, Lower Needmore Rd., Little Tennessee River, Swain Co., North Carolina, A. E. McBride and E. L. Quinter. Cherokeea trigona (Smith), larva in Arundinaria gigantea, Ela Public Access Area, Tuckasegee River, Swain Co., North Carolina, A. E. McBride and E. L. Quinter. Cherokeea trigona (Smith), larva in Arundinaria gigantea, Ela Public Access Area, Tuckasegee River, Swain Co., North Carolina, A. E. McBride and E. L.. Franclemontia interrogans (Walker), larva in Arundinaria tecta, Beaufort, Carteret Co., North Carolina, 3 March 2023, A. E. McBride and J. B. Sullivan. Franclemontia interrogans (Walker), larva in Arundinaria tecta, Beaufort, Carteret Co., North Carolina, 3 March 2023, A. E. McBride and J. B. Sullivan.
Figs 1–12; VARIABLE SIZES Photographs by A. E. McBride. Hydraecia immanis Guenée, larva in root of Silphium laciniatum, Richardson Wildlife Foundation, prairie S of Shaw Rd., Lee Co., Illinois, 27 June 2014, A. E. McBride and J. R. Wiker. Hydraecia stramentosa Guenée, larva in root of Scrophularia lanceolata, Warm Springs Mountain N of Ingall’s Field, Valley Springs, Bath Co., Virginia, 4 July 2015, A. E. McBride. Hydraecia micacea (Esper), larva in grass culm, Federal Hill Rd. 2, Delhi, Delaware Co., New York, A. E. McBride. Fibigeria erepta (Grote), larva in Tripsacum dactyloides, Pelham Bay, Bronx, Bronx Co., New York, 2 June 2020, A. E. McBride. Amphipoea senilis (Smith), larva in root of Iris missouriensis, Rt. 244, 1.15 mi NE of Cloudcroft, Otero Co., New Mexico, 25 June 2023, A. E. McBride. Amphipoea americana (Speyer), larva in root of Iris, Federal Hill Rd. 2, Delhi, Delaware Co., New York, 11 June 2023, A. E. McBride. Parapamea buffaloensis (Grote), larva in Saururus cernuus, Blackbird Creek, New Castle Co., Delaware, 16 August 2015, A. E. McBride. Papaipema eryngii Bird, larva in Eryngium yuccifolium, Washington Twp., Pettis Co., Missouri, 23 July 2020, A. E. McBride. Papaipema amianthii Quinter, larva in bulb of Amianthium muscitoxicum, State Game Lands #129, E of Rt. 903, Kidder Twp., Carbon Co., Pennsylvania, 25 July 2015, A. E. McBride. Papaipema appassionata (Harvey), larva in root of Sarracenia flava, Pender Co., North Carolina, A. E. McBride, J. B. Sullivan, and S. P. Hall. Papaipema cerussata (Grote), larva in Vernonia noveboracensis, Brook Hollow Farm, Knowlton Twp., Warren Co., New Jersey, 26 July 2014, A. E. McBride. Papaipema cerussata (Grote), mature larva in Vernonia noveboracensis, Brook Hollow Farm, Knowlton Twp., Warren Co., New Jersey, 10 August, 2014, A. E. McBride.
Figs 1–12; VARIABLE SIZES Photographs by A. E. McBride. Papaipema furcata (Smith), larva in Fraxinus pennsylvanica, Teal Rd., Petersburg State Game Area, Monroe Co., Michigan, A. E. McBride. Papaipema impecuniosa (Grote), larva in Symphyotrichum puniceum, Brook Hollow Farm, Knowlton Twp., Warren Co., New Jersey, 19 July 2014, A. E. McBride. Papaipema nepheleptena (Dyar), larva in Chelone glabra, wet meadow W of Emmons Lane, North Canaan, Litchfield Co., Connecticut, 21 July 2014, A. E. McBride. Papaipema sciata Bird, larva in Veronicastrum virginicum, Richardson Wildlife Foundation, prairie N of Shaw Rd., Lee Co., Illinois, A. E. McBride and J. R. Wiker. Papaipema rigida (Grote), mature larva in Helianthus decapetalus, Brook Hollow Farm, Knowlton Twp., Warren Co., New Jersey, 25 July 2014, A. E. McBride. Papaipema unimoda (Smith), larva in root of Thalictrum pubescens, Palmer River floodplain SW of Water Street, Rehoboth, Bristol Co., Massachusetts, 6 August 2014, A. E. McBride. Papaipema beeriana Bird, larva in corm of Liatris spicata, prairie S of Lulu Rd., Petersburg State Game Area, Monroe Co., Michigan, 26 June 2014, A. E. McBride. Papaipema nelita (Strecker), larva in Rudbeckia laciniata, New Columbia Rd., New Columbia, Union Co., Pennsylvania, 18 June 2014, A. E. McBride and S. Johnson. Papaipema maritima Bird, mature larva in Helianthus giganteus, Cape May Point, Cape May Co., New Jersey, 30 July 2014, A. E. McBride. Papaipema limpida (Guenée), larva in Vernonia, Matanzas Prairie, Bath, Mason Co., Illinois, 28 June 2014, A. E. McBride and J. R. Wiker. Papaipema marginidens (Guenée), ex ovo, reared in potato, parent female collected at bird management area N of Red Rose Rd., Petersburg, Huntingdon Co., Pennsylvania, A. E. McBride, S. Smith, and S. Johnson. Papaipema marginidens (Guenée), larva in underground storage stem of Dioscorea villosa, bird management area N of Red Rose Rd., Petersburg, Huntingdon Co., Pennsylvania, A. E. McBride.
Figs 1–12; VARIABLE SIZES Photographs by A. E. McBride. Papaipema eupatorii (Lyman), larva in Eutrochium, Riverside Park near Bean Creek, Morenci, Lenawee Co., Michigan, 26 June 2014, A. E. McBride. Papaipema inquaesita (Grote & Robinson), larva in rhizome of Onoclea sensibilis, fen N of Old Prospect School Rd., Hardyston Twp., Sussex Co., New Jersey, A. E. McBride. Papaipema speciosissima (Grote & Robinson), larva in rhizome of Osmunda regalis, Braidwood, Will Co., Illinois, A. E. McBride and J. R. Wiker. Papaipema stenocelis (Dyar), larva in rhizome of Woodwardia virginica, Howardsville cranberry bogs, Barnegat Twp., Ocean Co., New Jersey, A. E. McBride. Papaipema araliae Bird & Jones, larva in Aralia spinosa, Bean Ridge Rd., Tamms Precinct, Alexander Co., Illinois, 1 July 2014, A. E. McBride and J. R. Wiker. Papaipema araliae Bird & Jones, mature larva in Aralia spinosa, Bean Ridge Rd., Tamms Precinct, Alexander Co., Illinois, 1 July 2014, A. E. McBride and J. R. Wiker. Papaipema polymniae Bird, larva in Smallanthus uvedalia, Pine Hills area off Larue Rd., Union Precinct, Union Co., Illinois, 1 July 2014, A. E. McBride and J. R. Wiker. Papaipema nebris (Guenée), larva in Ambrosia trifida, Beaver Brook Wildlife Management Area, White Twp., Warren Co., New Jersey, 10 July 2014, A. E. McBride. Papaipema arctivorens Hampson, larva in Cirsium, White Lake Wildlife Management Area, Hardwick Twp., Warren Co., New Jersey, 6 June 2015, A. E. McBride. Papaipema arctivorens Hampson, mature larva in Cirsium, White Lake Wildlife Management Area, Hardwick Twp., Warren Co., New Jersey, 18 June 2014, A. E. McBride. Papaipema astuta Bird, larva in Collinsonia canadensis, Whittingham Wildlife Management Area, Fredon Twp., Sussex Co., New Jersey, 19 July 2014, A. E. McBride. Papaipema astuta Bird, mature larva in Collinsonia canadensis, White Lake Wildlife Management Area, Hardwick Twp., Warren Co., New Jersey, 19 July 2014, A. E. McBride.
Figs 1–12; VARIABLE SIZES Photographs by A. E. McBride. Papaipema rubiginosa Bird, larva in Angelica atropurpurea, fen N of Old Prospect School Rd., Hardyston Twp., Sussex Co., New Jersey, 21 June 2014, A. E. McBride. Papaipema harrisii (Grote), larva in Heracleum maximum, Indian Creek Campground, N of Rt. 67, Douglas Co., Colorado, 15 July 2014, J. R. Wiker. Papaipema sauzalitae (Grote), larva in Angelica hendersonii, 4 mi. W of Sixes, Curry Co., Oregon, 15 July 2017, A. E. McBride and J. R. Wiker. Papaipema birdi (Dyar), larva in Cicuta maculata, Teal Rd., Petersburg State Game Area, Monroe Co., Michigan, A. E. McBride and D. R. Badgero. Papaipema insulidens (Bird), larva in Heracleum maximum, US Rt. 101, 0.3 mi N of Hawk Creek, Neskowin, Tillamook Co., Oregon, 11 July 2017, A. E. McBride and J. R. Wiker. Papaipema insulidens (Bird), larva in Heracleum maximum, Steptoe Butte, Whitman Co., Washington, 12 July 2017, A. E. McBride and J. R. Wiker. Papaipema insulidens (Bird), larva in Heracleum maximum, Steptoe Butte, Whitman Co., Washington, 12 July 2017, A. E. McBride and J. R. Wiker. Papaipema albimontana Quinter, larva in root of Cicuta douglasii, Little Colorado River floodplain, 1 mi. S of Greer, Apache Co., Arizona, 27 June 2023, A. E. McBride. Papaipema depictata Benjamin, larva in Hymenoxys hoopesii, 42D Forest Rd., Long Park, Cochise Co., Arizona, 24 June 2023, A. E. McBride. Papaipema baptisiae (Bird), larva in Apocynum cannabinum, C.R. 539, Greenwood Forest Wildlife Management Area, Lacey Twp., Ocean Co., New Jersey, 26 July 2014, A. E. McBride. Papaipema limata Bird, larva in Apocynum cannabinum, U.S. Highway 20, 2.5 mi. NW of Juntura, Malheur Co., Oregon, 13 July 2017, A. E. McBride and J. R. Wiker. Papaipema rutila (Guenée), larva in root of Podophyllum peltatum, Riverside Park near Bean Creek, Morenci, Lenawee Co., Michigan, 26 June 2014, A. E. McBride.
Figs 1–12; VARIABLE SIZES Photographs by A. E. McBride. Papaipema necopina (Grote), larva in Helianthus, 0.1 mi. W of junction of Stoll and Upton Rds., Bath Twp., Clinton Co., Michigan, 15 July 2015, A. E. McBride and T. Herig. Papaipema duplicata Bird, larva in root of Collinsonia canadensis, State Game Lands #156, N of U.S. Rt. 322, South Lebanon Twp., Lebanon Co., Pennsylvania, 16 July 2014, A. E. McBride. Papaipema silphii Bird, larva in root of Silphium laciniatum, Richardson Wildlife Foundation, prairie S of Shaw Rd., Lee Co., Illinois, 28 June 2014, A. E. McBride and J. R. Wiker. Papaipema pterisii Bird, larva in Pteridium aquilinum, State Game Lands #129, E of Rt. 903, Kidder Twp., Carbon Co., Pennsylvania, 16 July 2014, A. E. McBride. Papaipema hesseli Quinter, larva in Matteuccia struthiopteris, Normans Kill floodplain W of Rt. 155, Guilderland, Albany Co., New York, 20 July 2014, A. E. McBride. Papaipema lysimachiae Bird, larva in Lysimachia quadrifolia, powerline ROW S of North Polktown Rd., Bethlehem Twp., Hunterdon Co., New Jersey, 22 June 2014, A. E. McBride. Papaipema leucostigma (Harris), larva in Aquilegia canadensis, Cross Plains, Dane Co., Wisconsin, 2 July 2014, A. E. McBride. Papaipema blackorum Quinter, larva in Aristolochia tomentosa, canebrake near Obion Creek, 2.8 mi. NE of Wingo, Graves Co., Kentucky, 29 June 2016, J. R. Wiker, A. E. McBride, and W. R. Black, Jr. Papaipema circumlucens (Smith), larva in Humulus lupulus, C.R. 3600E, Allens Grove, Mason Co., Illinois, 28 June 2014, A. E. McBride and J. R. Wiker. Papaipema circumlucens (Smith), larva in Humulus lupulus, C.R. 3600E, Allens Grove, Mason Co., Illinois, 28 June 2014, A. E. McBride and J. R. Wiker. Papaipema cataphracta (Grote), larva in Collinsonia canadensis, White Lake Wildlife Management Area, Hardwick Twp., Warren Co., New Jersey, 19 July 2014, A. E. McBride. Papaipema sulphurata Bird, larva in Decodon verticillatus, dike at north end of Hoxie Pond, Sandwich, Barnstable Co., Massachusetts, M. Mello, 4 August 2014, A. E. McBride, and E. L. Quinter.
Figs 1–8; VARIABLE SIZES Photographs by A. E. McBride. Papaipema aweme (Lyman), larva in Menyanthes trifoliata, Pinecreek Peatland State Natural Area, Roseau Co., Minnesota, 24 June 2016, A. E. McBride and K. E. Johnson. Papaipema aweme (Lyman), mature larva in Menyanthes trifoliata, Pinecreek Peatland State Natural Area, Roseau Co., Minnesota, A. E. McBride and K. E. Johnson. Papaipema cerina (Grote), larva in Podophyllum peltatum, Riverside Park near Bean Creek, Morenci, Lenawee Co., Michigan, A. E. McBride. Papaipema cerina (Grote), mature larva in Podophyllum peltatum, Riverside Park near Bean Creek, Morenci, Lenawee Co., Michigan, A. E. McBride. Papaipema sullivani Quinter, larva in Arundinaria tecta, Railroad Ditch, Great Dismal Swamp National Wildlife Refuge, City of Suffolk, Virginia, A. E. McBride. Papaipema matheri Quinter, larva in Arundinaria gigantea, McCrite Farm, Diswood, Alexander Co., Illinois, 30 June 2014, A. E. McBride and J. R. Wiker. Papaipema duovata (Bird), larva in Solidago sempervirens, Seven Bridges Rd., Great Bay Boulevard Wildlife Management Area, Little Egg Harbor Twp., Ocean Co., New Jersey, 27 July 2014, A. E. McBride. Papaipema duovata (Bird), larva in Solidago sempervirens, Seven Bridges Rd., Great Bay Boulevard Wildlife Management Area, Little Egg Harbor Twp., Ocean Co., New Jersey, 27 July 2014, A. E. McBride.
Larval Damage: Color Plates D1–D9
Figs 1–6 Photographs by A. E. McBride, except as noted. Cherokeea trigona (Smith), larva on mature culm of Arundinaria gigantea, Ela Public Access Area, Tuckasegee River, Swain Co., North Carolina, 2 May 2021, A. E. McBride and E. L. Quinter. The dead and dried leaf sheath on this mature culm has been pulled down to the right, exposing an early instar appressed to the culm alongside new axillary growth. Early instars of Cherokeea and Arundinariaphila commonly shelter underneath leaf sheaths. Cherokeea trigona (Smith), larva on mature culm of Arundinaria gigantea, Ela Public Access Area, Tuckasegee River, Swain Co., North Carolina, 2 May 2021, A. E. McBride and E. L. Quinter. In this close-up of the early instar in Figure 1, note the accumulation of frass and plant debris in this larval shelter, as well as the striped pattern of the larva. Cherokeea slotteni Quinter, larval damage of newly emergent culm of Arundinaria tecta after larva has vacated culm for pupation in the soil, St. Catherines Island, Liberty Co., Georgia, 20 May 1993, Eric L. Quinter. Note that the tender terminal node which was consumed by the larva is missing, but the exit hole at its base remains visible in the image at the tip of the second node. Photograph by Erik Neff. Cherokeea odonnellae Quinter, larval damage in new growth of Arundinaria appalachiana, Lower Needmore Rd., Little Tennessee River, Swain Co., North Carolina, 9 May 2022, A. E. McBride. Early instars utilize new terminal growth as well as fresh axillary growth similarly to C. trigona in Figures 1 and 2, above. Note the accumulated white frass outside the larval burrow. Fibigeria erepta (Grote), larval damage in Tripsacum dactyloides, Pelham Bay, Bronx, Bronx Co., New York, 13 Jun. 2023, A. E. McBride. Withered and dried grass blades among healthy shoots are evidence of larval damage in this species. Fibigeria erepta (Grote), larval damage in Tripsacum dactyloides, Pelham Bay, Bronx, Bronx Co., New York, 13 Jun. 2023, A. E. McBride. After larval feeding is completed in the large culm base, the larva chews a characteristic round exit hole and abandons the burrow to pupate in soil.
Figs 1–6 Photographs by A. E. McBride. Franclemontia interrogans (Walker), larval damage in mature culm of Arundinaria tecta, Beaufort, Carteret Co., North Carolina, 4 March 2023, A. E. McBride and J. B. Sullivan. The mature larva overwinters in the culm gallery, undergoes a molt in early Spring, and then prepares this exit aperture for the adult before pupating in the culm. While feeding, larvae girdle the inside of the culm similarly to Papaipema matheri in Figure 4, below. Amphipoea senilis (Smith), larval damage in Iris missouriensis, Rt. 244, 1.15 mi NE of Cloudcroft, Otero Co., New Mexico, 25 Jun. 2023, A. E. McBride. Larval boring in the rhizome causes the leaves to yellow and die. Parapamea buffaloensis (Grote), pupa in larval burrow in Saururus cernuus, Bernards Twp., Morris Co., New Jersey, 7 Sep. 2023, A. E. McBride. The larva prepares an exit opening for the adult, seen at the top of this exposed burrow, before pupation. Papaipema matheri Quinter, larval damage in mature culm of Arundinaria gigantea, McCrite Farm, Diswood, Alexander Co., Illinois, 30 Jun. 2014, A. E. McBride and J. R. Wiker. The larva girdles the inside of the culm at one point, which makes the culm vulnerable to breakage by wind (inset). The larva can be found feeding in the section of culm above the girdled point, and pupates within the culm below the girdled section after preparing an exit opening for the adult. Papaipema eupatorii (Lyman), larval damage in Eutrochium purpureum, New Columbia Rd., New Columbia, Union Co., Pennsylvania, 18 Sep. 2021, A. E. McBride and S. Johnson. After completing feeding in the root of the plant, the larva fashions a characteristic ovoid exit aperture at the base of the stem through which the adult will emerge and then pupates in the burrow. In this image, the original round orifice the larva used to expel frass from the burrow can be seen above the adult exit opening. Papaipema rubiginosa Bird, larval damage in Angelica atropurpurea, fen N of Old Prospect School Rd., Hardyston Twp., Sussex Co., New Jersey, 25 Jun. 2010, A. E. McBride. Larval burrowing in the plant stalk causes yellowing of the leaves. In Angelica, a biennial, larvae may be found in tall, blooming-size plants as well as in short, vegetative first-year plants, as in this image.
Figs 1–6 Photographs by A. E. McBride. Papaipema amianthii Quinter, larval damage in Amianthium muscitoxicum, State Game Lands #129, E of Rt. 903, Kidder Twp., Carbon Co., Pennsylvania, 30 Jul. 2020, A. E. McBride. As the larva feeds in the bulb, a plug of accumulated frass develops at the burrow entrance, and the leaves of the plant turn yellow and begin to senesce. Papaipema amianthii Quinter, larval damage in Amianthium muscitoxicum, Appalachian Mts., Kittatinny Ridge, Blue Mountain, acid bog, 4.2 mi. SW of Auburn, Schuylkill Co., Pennsylvania, 15 Aug. 2021, A. E. McBride. With feeding and growth completed, the larva uses frass, plant fragments, and silk to construct a protective plug over the burrow entrance and then pupates within the remnants of the bulb. In Aug., the dead leaves and plug can be lifted to reveal the pupa in the chamber below. Papaipema appassionata (Harvey), larval damage in Sarracenia purpurea, Glovers Pond, Frelinghuysen Twp., Warren Co., New Jersey, 28 Jul. 2014, A. E. McBride. As the larva feeds in the root, orange frass is deposited at the burrow entrance, and the pitchers turn yellow and eventually die. Papaipema cerussata (Grote), larval damage in Vernonia noveboracensis, Brook Hollow Farm, Knowlton Twp., Warren Co., New Jersey, 29 Aug. 2023, A. E. McBride. The plant on the right is stunted and shows the characteristic “witches’ broom” that develops after the larva bores in the terminal meristem during the early instars. The larva then exits the terminal burrow and completes its growth by boring in the lower stalk and root below. Papaipema cerussata (Grote), larval damage in Vernonia noveboracensis, Brook Hollow Farm, Knowlton Twp., Warren Co., New Jersey, 29 Aug. 2023, A. E. McBride. The base of the plant stalk in this image shows the characteristic vertical, elongate frass opening made by the larva. Papaipema eryngii Bird, larval damage in Eryngium yuccifolium, Washington Twp., Pettis Co., Missouri, 23 Jul. 2020, A. E. McBride, J. R. Wiker, and P. McKenzie. Larval feeding in the root causes the leaves to turn brown.
Figs 1–6 Photographs by A. E. McBride. Papaipema furcata (Smith), larval damage in Fraxinus pennsylvanica, Teal Rd., Petersburg State Game Area, Monroe Co., Michigan, 4 Jul. 2014, A. E. McBride. The early instars bore in the terminal growth, causing wilting and yellowing of the leaves and stem. Papaipema furcata (Smith), larval damage in Fraxinus pennsylvanica, Teal Rd., Petersburg State Game Area, Monroe Co., Michigan, 4 Jul. 2014, A. E. McBride. After reaching the third instar, the larva abandons the burrow in the terminal shoot, crawls down the branch, and bores into the woody growth from the prior year. This entrance opening is always made just below a branch node. Papaipema impecuniosa (Grote), larval damage in Symphyotrichum puniceum, Brook Hollow Farm, Knowlton Twp., Warren Co., New Jersey, 19 Sep. 2020, A. E. McBride. Before pupating in the root, the larva fashions this characteristic opening for the adult to exit the burrow. Papaipema sciata Bird, larval damage in Veronicastrum virginicum, Richardson Wildlife Foundation, prairie N of Shaw Rd., Lee Co., Illinois, 26 Jun. 2014, A. E. McBride and J. R. Wiker. Boring in the root by the larva of this species causes the entire plant to blacken and die. Papaipema rigida (Grote), larval damage in Helianthus decapetalus, Brook Hollow Farm, Knowlton Twp., Warren Co., New Jersey, 25 Jul. 2014, A. E. McBride. In Helianthus, this species typically leaves meager aboveground sign of its presence. Here, sawdust-like frass can be seen near the stem base. Papaipema beeriana Bird, larval damage in Liatris, Richardson Wildlife Foundation, prairie N of Shaw Rd., Lee Co., Illinois, 27 Jun. 2014, A. E. McBride and J. R. Wiker. Boring of the stem in early instars causes the plant’s leaves to blacken, though this can be difficult to detect in a large patch of Liatris. In this figure, frass can be seen at the burrow entrance. Most of the feeding in later instars takes place below ground in the corm.
Figs 1–6 Photographs by A. E. McBride. Papaipema maritima Bird, larval damage in Helianthus giganteus, Cape May Point, Cape May Co., New Jersey, 27 Sep. 2013, A. E. McBride. Larval boring causes the plant to produce a characteristic egg-shaped gall at the plant base. Before pupation in the gallery, the larva fashions an exit aperture for the eventual escape of the adult. The position of the pupa in the burrow can be seen in the right-hand inset, where the gall has been sliced open to reveal the hollow inside. Papaipema marginidens (Guenée), larval damage in Dioscorea villosa, bird management area N of Red Rose Rd., Petersburg, Huntingdon Co., Pennsylvania, 4 Jul. 2020, A. E. McBride. Boring in the rhizome of this plant causes the aboveground vine to shrivel and die. The dead vine must be carefully traced back to the ground to determine which underground rhizome supports the larva. Papaipema inquaesita (Grote & Robinson), larval damage in Onoclea sensibilis, Richardson Wildlife Foundation, wet prairie N of Shaw Rd., Lee Co., Illinois, 27 Jun. 2014, A. E. McBride and J. R. Wiker. Boring in the rhizome causes the fern to curl up and dry, displaying a characteristic rust-colored dead frond. Papaipema stenocelis (Dyar), larval damage in Woodwardia virginica, Howardsville cranberry bogs, Barnegat Twp., Ocean Co., New Jersey, 16 Jul. 2009, A. E. McBride. Boring in the rhizome causes the fern to yellow and then turn brown. Two of these deadened fronds can be seen in this image. Papaipema stenocelis (Dyar), larval damage in Woodwardia virginica, Howardsville cranberry bogs, Barnegat Twp., Ocean Co., New Jersey, 27 Jul. 2014, A. E. McBride. An accumulation of reddish-brown frass can be found at the base of bored stipes. Papaipema polymniae Bird, larval damage in Smallanthus uvedalia, River Rd. along French Broad River, 2.7 mi. NW of Hot Springs, Madison Co., North Carolina, 23 Jul. 2023, A. E. McBride, J. Petranka, and B. Elkin. Boring by the larva causes the plant to develop swellings of various widths, and frass may accumulate on the ground below. Otherwise, the plant may show little sign of larval presence.
Figs 1–6 Photographs by A. E. McBride. Papaipema nelita (Strecker), larval damage in Rudbeckia laciniata, New Columbia Rd., New Columbia, Union Co., Pennsylvania, 15 Jun. 2019, A. E. McBride and S. Johnson. In Rudbeckia, larval presence in the root causes a slight gall to develop at or just below the soil line, inside which the larva feeds. Some frass may be present on the surface. Papaipema araliae Bird & Jones, larval damage in Aralia spinosa, Bean Ridge Rd., Tamms Precinct, Alexander Co., Illinois, 30 Jun. 2014, A. E. McBride and J. R. Wiker. Yellowing followed by blackening of the large leaves of this thorny shrub are a sign of larval presence. Upon closer inspection, a bore opening with accumulated frass may be seen, often accompanied by a thick, brown, gummy exudate released by the plant. Papaipema arctivorens Hampson, larval damage in Cirsium, White Lake Wildlife Management Area, Hardwick Twp., Warren Co., New Jersey, 7 Jul. 2014, A. E. McBride. The larva bores the terminal section of the stalk, creating a characteristic concentration of browned leaves and frass confined to the top of the plant. Papaipema astuta Bird, larval damage in Collinsonia canadensis, White Lake Wildlife Management Area, Hardwick Twp., Warren Co., New Jersey, 19 Jul. 2014, A. E. McBride. Bored plants wither and dry and are easy to spot amongst healthy stems. Papaipema sauzalitae (Grote), larval damage in Angelica hendersonii, 4 mi. W of Sixes, Curry Co., Oregon, 15 Jul. 2017, A. E. McBride and J. R. Wiker. Various signs of larval infestation may be presented in this plant, including wilted stems, flagged stems, dried stalks, and bore openings with bright orange frass. Papaipema nebris (Guenée), larval damage in Ambrosia trifida, Beaver Brook Wildlife Management Area, White Twp., Warren Co., New Jersey, 21 Jul. 2014, A. E. McBride. Larval boring causes the plant stalk to swell. Several larvae can inhabit a tall plant, with frass accumulating outside the gallery openings.
Figs 1–6 Photographs by A. E. McBride. Papaipema albimontana Quinter, larval damage in Cicuta douglasii, Little Colorado River floodplain, 1 mi. S of Greer, Apache Co., Arizona, 27 Jun. 2023, A. E. McBride. Several presentations of damage can occur, including wilted stems, flagged stems, dried stalks, and bore openings at any point on the main stem, with accumulated frass. Papaipema depictata Benjamin, larval damage in Hymenoxys hoopesii, 42D Forest Rd., Long Park, Cochise Co., Arizona, 24 Jun. 2023, A. E. McBride. Bore openings can occur at various points along the main stalk, often resulting in wilted stems and leaves. Papaipema baptisiae (Bird), larval damage in Apocynum cannabinum, C.R. 539, Greenwood Forest Wildlife Management Area, Lacey Twp., Ocean Co., New Jersey, 27 Jul. 2014, A. E. McBride. In Apocynum, boring by the larva in the lower stem and root causes the leaves to turn a characteristic yellow color. The stems often bend and fall over at the point low on the stem where the frass opening was made, and copious light, brownish-red frass can be found on the soil surface. Papaipema necopina (Grote), larval damage in Helianthus, 0.1 mi. W of junction of Stoll and Upton Rds., Bath Twp., Clinton Co., Michigan, 15 Jul. 2015, A. E. McBride and T. Herig. Boring by the larva causes the plant to produce a characteristic gall at ground level. Papaipema duplicata Bird, larval damage in Collinsonia canadensis, State Game Lands #156, N of U.S. Rt. 322, South Lebanon Twp., Lebanon Co., Pennsylvania, 16 Jul. 2014, A. E. McBride. As with the plant in the right-hand side of this image, bored plants wither and dry and are easy to spot amongst healthy stems. The plants often bend and fall over at the point on the lower stem where the frass opening was created by the larva. Papaipema duplicata Bird, larval damage in Collinsonia canadensis, Thunder Hill, mile post 76, Blue Ridge Parkway, 3,920´ [1,195 m], Bedford Co., Virginia, 20 Jul. 2023, A. E. McBride. This is a typical location on the lower stem where the frass orifice is created. Note the browned coloration of the stem, which is normally green. Light brown frass can also be seen on the moss below.
Figs 1–6 Photographs by A. E. McBride. Papaipema silphii Bird, larval damage in Silphium laciniatum, Richardson Wildlife Foundation, prairie S of Shaw Rd., Lee Co., Illinois, 27 Jun. 2014, A. E. McBride and J. R. Wiker. The warm brown dead leaf that was initially bored stands out starkly in contrast with healthy leaves. In midsummer, the larva can be found boring in the extensive rootstock below. Papaipema pterisii Bird, larval damage in Pteridium aquilinum, State Game Lands #129, E of Rt. 903, Kidder Twp., Carbon Co., Pennsylvania, 16 Jul. 2014, A. E. McBride. Larvae of pterisii can frequently be found boring in plants in relatively moist environments, and the entire frond turns yellow and eventually brown. On drier sites, Pteridium tends to die or senesce for various reasons other than boring by larvae, so yellowed or browned fronds do not always indicate larval damage. Papaipema hesseli Quinter, larval damage in Matteuccia struthiopteris, Normans Kill floodplain W of Rt. 155, Guilderland, Albany Co., New York, 20 Jul. 2014, A. E. McBride. Bored fronds turn yellow and then brown. In mid to late Jul., larvae can be found either in the enlarged stipe base, or just below the stipe between the basal scales of the fern clump. Papaipema lysimachiae Bird, larval damage in Lysimachia quadrifolia, powerline ROW at west end of Pascoe Rd., Sparta Twp., Sussex Co., New Jersey, 3 Jul. 2020, A. E. McBride. Bored plants wilt and dry and stand out in contrast to healthy plants. Copious light, pinkish frass may be found on the ground below. If the leaves are wilted but still green, the larva can be found in the lower stem or root. If the leaves are completely brown and dead, the larva will have vacated the burrow. Papaipema circumlucens (Smith), larval damage in Humulus lupulus, C.R. 3600E, Allens Grove, Mason Co., Illinois, 28 Jun. 2014, A. E. McBride and J. R. Wiker. The swollen gall containing the larva will usually be found where the vine twists around another vine, plant stem, or fence wire. The swelling is usually yellowish in color and is paler than the adjacent vine. The burrow opening in the gall is usually found very close to the object the gall twists around, as in this image, affording the larva some additional shielding. Papaipema blackorum Quinter, larval damage in Aristolochia tomentosa, S of Rt. 146 on Reevesville Rd., Johnson Creek, C. Sampson property, Johnson Co., Illinois, 23 Jul. 2020, A. E. McBride and J. R. Wiker. In this image, the thick, pale, vertical vine section just to the left of center is the gall. At the lower end of the gall near the point where the vine twists around other vines, note the blackened and moist frass orifice.
Figs 1–6 Photographs by A. E. McBride. Papaipema sulphurata Bird, larval damage in Decodon verticillatus, dike at north end of Hoxie Pond, Sandwich, Barnstable Co., Massachusetts, 15 Sep. 2017, A. E. McBride. The larva pupates in the stem, which is slightly swollen in this image due to larval boring activity. Before pupation, the larva creates a large, round opening to facilitate adult emergence. A smaller frass opening previously used by the larva can be seen a short distance below the adult exit opening on the right. Papaipema duovata (Bird), larval damage in Solidago sempervirens, Seven Bridges Rd., Great Bay Boulevard Wildlife Management Area, Little Egg Harbor Twp., Ocean Co., New Jersey, 27 Jul. 2014, A. E. McBride. This is the typical appearance of the gallery opening at the base of a stem, with light brown frass on the soil below. The leaves above may or may not show signs of yellowing. Papaipema duovata (Bird), larval damage in Solidago sempervirens, Seven Bridges Rd., Great Bay Boulevard Wildlife Management Area, Little Egg Harbor Twp., Ocean Co., New Jersey, 22 Sep. 2016, A. E. McBride. Before pupating in the lower stem or root, the larva fashions an exit aperture through which the adult will emerge. The larva also plugs the hollow stem above this aperture with chewed plant fragments to prevent the adult from making the mistake of crawling up inside the stalk upon emergence. Papaipema duovata (Bird), larval damage in Solidago sempervirens, Seven Bridges Rd., Great Bay Boulevard Wildlife Management Area, Little Egg Harbor Twp., Ocean Co., New Jersey, 22 Sep. 2016, A. E. McBride. Cutaway view of the gallery revealing the pupa within. Papaipema cerina (Grote), larval damage in Elymus riparius, Cross Plains, Dane Co., Wisconsin, 2 Jul. 2014, A. E. McBride. In Jun. and very early Jul., the early instars may be found boring in the upper internodes of grass culms, and this activity causes the grass tops to turn brown. Larvae do not bore in the lower parts of the culm or in the roots. Once it grows too large to bore in the slender upper grass stem, the larva abandons the burrow and crawls off in search of plants with stems that are large enough for it to complete development, such as Podophyllum and Lilium. Papaipema sullivani Quinter, larva boring in Arundinaria tecta, Railroad Ditch, Great Dismal Swamp National Wildlife Refuge, City of Suffolk, Virginia, 8 Jun. 2015, A. E. McBride. The larva in this image was photographed entering a new cane culm after it had abandoned a thinner shoot. The process of entering a new culm can take a half hour or more, and this is a very vulnerable time for the larva.
Habitats: Color Plates H1–H5
Figs 1–6 Photographs by A. E. McBride, except as noted. Windmill Pond, St. Catherines Island, Liberty Co., Georgia. Type locality of Arundinariaphila hayesi Quinter and Cherokeea slotteni Quinter. Habitat for Franclemontia interrogans (Walker), Sarita relicta (Ferguson), and Papaipema sullivani Quinter. Photograph by Erik Neff. Canebrake on St. Catherines Island, Liberty Co., Georgia, at the type locality designated in Figure 1, above. Mesic loblolly pine (Pinus taeda) forest with switch cane (Arundinaria tecta) understory. Photograph by Erik Neff. Giant cane (Arundinaria gigantea) in Swain Co., North Carolina, showing dense microhabitat of a single culm. Habitat for Cherokeea trigona (Smith), Arundinariaphila hayesi Quinter, and Papaipema matheri Quinter. Photograph by Jane O’Donnell. Switch cane (Arundinaria tecta) at type locality of Cherokeea gabrielleae Quinter and Papaipema sullivani Quinter at Great Dismal Swamp National Wildlife Refuge, City of Suffolk, Virginia. Mesic bottomland forest. Habitat for Cherokeea slotteni Quinter, Franclemontia interrogans (Walker), and Sarita relicta (Ferguson). Photograph by Jane O’Donnell. Hill cane (Arundinaria appalachiana) at type locality of Cherokeea odonnellae Quinter in floodplain of the Little Tennessee River off Lower Needmore Rd. in Swain Co., North Carolina. Habitat for Cherokeea attakullakulla Sullivan & Quinter and Cherokeea slotteni Quinter. Meadow with gamagrass (Tripsacum dactyloides) at Orchard Beach, Bronx, Bronx Co., NY. Habitat for Fibigeria erepta (Grote).
Figs 1–6 Photographs by A. E. McBride. Bur oak prairie with prairie smoke (Geum triflorum) at the south end of Pike Lake on Manitoulin Island, Ontario. Habitat for Amphipoea interoceanica (Smith). Grassy meadow with Iris missouriensis and Rudbeckia laciniata along Rt. 244, 1.15 mi NE of Cloudcroft, Otero Co., New Mexico. Habitat for Amphipoea senilis (Smith) and Papaipema nelita (Strecker). Floodplain of the Passaic River with Saururus cernuus in Bernards Twp., Morris Co., New Jersey. Habitat for Parapamea buffaloensis (Grote). Rich, mesic, deciduous forest with Collinsonia canadensis on Warm Springs Mountain in Bath Co., Virginia. Habitat for Papaipema duplicata Bird, Papaipema rigida (Grote), Papaipema amianthii Quinter, Papaipema marginidens Guenée, Papaipema pterisii Bird, and Hydraecia stramentosa Guenée. Floodplain of the Little Delaware River with Matteuccia struthiopteris in Delhi, Delaware Co., New York. Habitat for Papaipema hesseli Quinter. Dry roadside prairie with Apocynum cannabinum in Cove, Cache Co., Utah. Habitat for Papaipema limata Bird.
Figs 1–6 Photographs by A. E. McBride. Wet meadow with Onoclea sensibilis, Vernonia noveboracensis, Eutrochium purpureum, and Symphyotrichum puniceum at Brook Hollow Farm in Knowlton Twp., Warren Co., New Jersey. Habitat for Papaipema inquaesita (Grote & Robinson), Papaipema cerussata (Grote), Papaipema eupatorii (Lyman), and Papaipema impecuniosa (Grote). Papaipema unimoda (Smith) can also be found in wet meadows. Old field habitat with Apocynum cannabinum and Cirsium sp. in Stillwater, Sussex Co., New Jersey. Habitat for Papaipema baptisiae (Bird), Papaipema arctivorens Hampson, Papaipema cataphracta (Grote), and Papaipema nebris (Guenée). Papaipema rigida (Grote) can also be found in old fields. Wet swale along Interstate 195 with Decodon verticillatus in Marion, Plymouth Co., Massachusetts. Habitat for Papaipema sulphurata Bird. Pacific Ocean bluffs with Angelica hendersonii at Cape Blanco State Park in Curry Co., Oregon. Habitat for Papaipema sauzalitae (Grote). Saltmarsh meadow with Solidago sempervirens off Cedar Run Dock Rd., Stafford Twp., Ocean Co., New Jersey. Habitat for Papaipema duovata (Bird). Floodplain of the Little Colorado River with Cicuta douglasii, 1 mi. S of Greer, Apache Co., Arizona. Habitat for Papaipema albimontana Quinter.
Figs 1–6 Photographs by A. E. McBride, except as noted. Grassy meadow in pine forest with Hymenoxys hoopesii, off the 42D Forest Rd. at Long Park, Cochise Co., Arizona. Habitat for Papaipema depictata Benjamin. Tallgrass prairie with Silphium terebinthinaceum at Killdeer Plains Wildlife Area, Marion Co., Ohio. Habitat for Papaipema silphii Bird and Hydraecia immanis Guenée. Photograph by John Peacock. Acidic woodland at Hickory Run State Park with Amianthium muscitoxicum in Carbon Co., Pennsylvania. Habitat for Papaipema amianthii Quinter and Papaipema pterisii Bird. Tallgrass prairie with Eryngium yuccifolium at Joseph A. Tauer Prairie State Natural Area, Brown Co., Minnesota. Habitat for Papaipema eryngii Bird. Photograph by Kyle E. Johnson. Mesic coastal meadow with Helianthus giganteus at Cape May Point State Park, Cape May Point, Cape May Co., New Jersey. Habitat for Papaipema maritima Bird. Eastern deciduous forest with Fraxinus americana, Collinsonia canadensis, and Podophyllum peltatum, White Lake Natural Resource Area, Hardwick Twp., Warren Co., New Jersey. Habitat for Papaipema furcata (Smith), Papaipema astuta Bird, and Papaipema rutila (Guenée).
Figs 1–5 Photographs by A. E. McBride, except as noted. Eric Quinter examining giant cane culm (Arundinaria gigantea) at the type locality of Cherokeea trigona (Smith) in Smith Co., Tennessee. Habitat for ten species of cane-specializing Lepidoptera. Photograph by Nancy Hanson. Anthony E. McBride on the United States – Canada border in a rich fen with Menyanthes trifoliata and Sarracenia purpurea at Pine Creek Peatland State Natural Area, Roseau Co., Minnesota. Habitat for Papaipema aweme (Lyman) and Papaipema appassionata (Harvey). Photograph by Kyle Johnson. Eric Quinter, Mogens “Mo” Nielsen, Jim Wiker, and Ted Herig on a sandy prairie with Liatris aspera, Chester, Otsego Co., Michigan. Habitat for Papaipema beeriana Bird. Photograph by Jane O’Donnell. J. Bolling Sullivan and Stephen P. Hall in a mesic prairie with Sarracenia flava at Croatan National Forest in Carteret Co., North Carolina. Habitat for Papaipema appassionata (Harvey). Jim Wiker holding a mature larva of Papaipema blackorum Quinter in a mesic bottomland forest along Johnson Creek with Aristolochia tomentosa off of Reevesville Rd., Johnson Co., Illinois.
NOTES
1. ABBREVIATIONS OF CITED INSTITUTIONAL AND PRIVATE COLLECTIONS
AEM Anthony E. McBride Collection, Blairstown, NJ, USA
AMNH American Museum of Natural History, New York, NY, USA
ANSP Academy of Natural Sciences, Philadelphia, PA, USA
CMNH Carnegie Museum of Natural History, Pittsburgh, PA, USA
CNC Canadian National Collection of Insects and Arachnids, Ottawa, ON, Canada
CUIC Cornell University Insect Collection, Ithaca, NY, USA
ELQ Eric L. Quinter Collection, Windham, CT, USA
FMNH Field Museum of Natural History, Chicago, IL, USA
GDW Glenn Darryl Willis Collection, Miramar Beach, FL, USA
INHS Illinois Natural History Survey Collection, Champaign, IL, USA
JBS J. Bolling Sullivan Collection, Beaufort, NC, USA
JKA James K. Adams Collection, Calhoun, GA, USA
JRS Jeffrey R. Slotten Collection, Gainesville, FL, USA
JRW James R. Wiker Collection, Greenview, IL, USA
JVC James Vargo Collection, Mishawaka, IN, USA
LACM Los Angeles County Museum of Natural History, Los Angeles, CA, USA
LAF Leslie A. Ferge Collection, Middleton, WI, USA
LDG Loran D. Gibson Collection, Florence, KY, USA
LEM Lyman Entomological Museum, Montréal, QC, Canada
MCLB McGuire Center for Lepidoptera and Biodiversity, University of Florida, Gainesville, FL, USA
MCN Mogens C. Nielsen Collection, now in MSU
MCZ Museum of Comparative Zoology, Harvard University, Cambridge, MA, USA
MEM Mississippi Entomological Museum, Mississippi State University, Starkville, MS, USA
MNHU Museum für Naturkunde der Humbolt-Universität, Berlin, Germany
MSU Michigan State University Collection, East Lansing, MI, USA
NHML The Natural History Museum, London, England (formerly British Museum of Natural History)
NSM Nova Scotia Museum, Halifax, Nova Scotia, Canada
NYSM New York State Museum, Albany, NY, USA
RPC Ricky L. Patterson Collection, Vicksburg, MS, USA
SJC Steven Johnson Collection, Sunbury, PA, USA
UCD University of California at Davis, Richard Bohart Museum, Davis, CA, USA
UCMS University of Connecticut Museum, Storrs, CT, USA
ULK University of Louisville, Louisville, KY, USA
USNM National Museum of Natural History – SI, Washington, DC, USA
VAB Vernon A. Brou Collection, Abita Springs, LA, USA
VADNH Virginia Division of Natural Heritage Collection, Virginia Department of Conservation and Recreation, Richmond, VA, USA
WPC Wedge Plantation Collection, Columbia, SC, USA
WVU West Virginia University, Morgantown, WV, USA
YPM Peabody Museum, Yale University, New Haven, CT, USA
2. ADDITIONAL COLLECTIONS EXAMINED
Auburn E. Brower Collection, Augusta, ME, USA
Bryant Mather Collection, now in MEM and AMNH
Bureau of Plant Industry Collection, Harrisburg, PA, USA
C. Brooke Worth Collection, Eldora, NJ, USA
Charles G. Merker Collection, now in CMNH
C. P. Gillette Museum of Arthropod Diversity, Colorado State University, Fort Collins, CO, USA
Dale F. Schweitzer Collection, now in YPM
Dwayne R. Badgero Collection, MI, USA
Eric H. Metzler Collection, Alamogordo, NM, USA
Graham Bell Fairchild Collection, Beinn Bhreagh, Nova Scotia, Canada
Great Smoky Mountains National Park Collection, Gatlinburg, TN, USA
H. Godwin Stevenson Collection, Baltimore, MD, USA
Iowa State University Collection, Ames, IA, USA
John G. Franclemont Collection, Ithaca, NY, USA
J. Richard Heitzman Collection, now in MCLB
Ken Stead Collection, Fort Myers, FL, USA
Lance A. Durden Collection, Statesboro, GA, USA
Lloyd Center for the Environment, South Dartmouth, MA, USA
Missouri Department of Agriculture Collection, Jefferson City, MO, USA
Nova Scotia Museum, Halifax, NS, Canada
Oregon State University Collection, Corvallis, OR, USA
Pennsylvania State University, Frost Entomological Museum, University Park, PA, USA
Rae Letsinger Collection, Sarcoxie, MO, USA
Richard Kergosien Collection, Biloxi, MS, USA
Ronald H. Leuschner Collection, now in MCLB
Ronald Panzer Collection, Chicago, IL, USA
Rutgers University Collection, New Brunswick, NJ, USA
Sidney A. Hessel Collection, Washington, CT, USA
Timothy L. McCabe Collection, Albany, NY, USA
University of California at Berkeley, Berkeley, CA, USA
University of Connecticut, Storrs, CT, USA
University of Massachusetts, Amherst, MA, USA
University of Michigan, Ann Arbor, MI, USA
University of Missouri, Columbia, MO, USA
University of Nebraska, Lincoln, NE, USA
Vermont Agricultural Experiment Station Collection, University of Vermont, Burlington, VT, USA
Virginia Polytechnic Institute, Blacksburg, VA, USA
Warren J. Kiel Collection, Groveton, NH, USA
Western Illinois University, Macomb, IL, USA
Wilfred S. Craig Collection, Columbia, MO, USA
William D. Winters Collection, Dedham, MA, USA
William E. Sieker Collection, Madison, WI, USA
William Lamond Collection, Brantford, ON, Canada
William T. Davis Collection, Staten Island Institute of Arts & Sciences, Staten Island, NY, USA
Yale Sedman Collection, Macomb, IL, USA
3. CITATIONS OF AUTHORS
Authors’ names without parentheses indicate that the specific name is associated with the genus in which it was described.
Authors’ names in parentheses indicate that the specific name has been transferred from the genus in which it was described to another genus.
4. WING MEASUREMENTS
Forewing length is the measurement in millimeters from the base of the wing to the apex.
Alar expanse is the maximum lateral measurement in millimeters of the forewings on spread specimens.
5. LOCATIONS OF PRIMARY TYPES
The current location of the primary type is given by the appropriate abbreviation in square brackets immediately following the type locality.
6. BOTANICAL NAMES
All botanical names used throughout the fascicle are in accord with the current standardized taxonomy of the PLANTS Database of the United States Department of Agriculture (https://plants.usda.gov/).
CHECK LIST
NOCTUINAE
Apameini Gn., 1841, tribe
Apameina Gn., 1841, subtribe
ARUNDINARIAPHILA Quinter, 2025
A.hayesi Quinter, 2025
CHEROKEEA Quinter & Sullivan, 2014
C. attakullakulla Sullivan & Quinter, 2014
C. slotteni Quinter, 2025
C. gabrielleae Quinter, 2025
C. odonnellae Quinter, 2025
C. trigona (Sm., 1902)
FRANCLEMONTIA Ferguson, 1992
F. interrogans (Wlk., 1856)
SARITA Quinter, 2025
S.relicta (Ferguson, 1991)
AMPHIPOEA Billberg, 1820
A.interoceanica (Sm., 1899)
A.americana (Speyer, 1875)
atlantica (Sm., 1899)
A.pacifica (Sm., 1899)
A.cottlei (McD., 1948)
A.keiferi (Benj., 1935)
A.senilis (Sm., 1892)
flavostigma (B. & Benj., 1924)
FIBIGERIA Quinter, 2025
F. erepta (Grote, 1881)
ryensis (Bird, 1913)
HYDRAECIA Gn., 1841
H.medialis Sm., 1892
pallescens Sm., 1899
H.obliqua Harv., 1876
columbia (B. & Benj., 1924)
ximena (B. & Benj., 1924)
H.intermedia (B. & Benj., 1924)
H.micacea (Esp., 1789)
cypriaca (Hübner, [1803])
H.perobliqua Hamp., 1910
H.stramentosa Gn., 1852
diplocyma Hampson, 1910
H.immanis Gn., 1852
SELICANIS Sm., 1900
S.cinereola Sm., 1900
PARAPAMEA Bird, 1927
P.buffaloensis (Grt., 1877)
latia (Strecker, 1899)
†simplicissima (Bird, 1916); (form)
PAPAIPEMA Sm., 1899
P.unimoda (Sm., 1892)
frigida (Sm., 1899)
thalictri (Lyman, 1905)
†perobsoleta (Lyman, 1905); (form)
imperspicua Bird, 1908
†terminalis Strand, 1915 [1916]; (form)
P.appassionata (Harv., 1876)
horni Strand, 1914
P.amianthii Quinter, 2025
P.furcata (Sm., 1899)
P.cerina (Grt., 1874)
P.sullivani Quinter, 2025
P.matheri Quinter, 2025
P.marginidens (Gn., 1852)
nephrasyntheta (Dyar, 1908)
P.eryngii Bird, 1917
P.cataphracta (Grt., 1864)
†aerata (Lyman, 1901); (hybrid)
†fluxa Bird, 1911; (form)
P.sulphurata Bird, 1926
P.aweme (Lyman, 1908)
P.necopina (Grt., 1876)
imperturbata Bird, 1907
mulleri Buchholz, 1956
P.duovata (Bird, 1902)
P.speciosissima (G. & R., 1868)
†regalis Wyatt & Beer, 1927; (form)
P.inquaesita (G. & R., 1868)
†wyatti B. & Benj., 1929; (form)
P.stenocelis (Dyar, 1907)
P.rigida (Grt., 1877)
P.eupatorii (Lyman, 1905)
P.nebris (Gn., 1852)
nitela (Gn., 1852)
P.arctivorens Hamp., 1910
P.araliae Bird & Jones, 1921
P.polymniae Bird, 1917
P.rutila (Gn., 1852)
merriccata Bird, 1907
placida Bird, 1921
P.baptisiae (Bird, 1902)
P.limata Bird, 1908
vaha Benj., 1933
†ochroptenoides Bird, 1944; (form)
P.depictata Benj., 1933
P.duplicata Bird, 1908
†obsolescens Strand, [1916]; (form)
P.silphii Bird, 1915
P.cerussata (Grt., 1864)
P.sciata Bird, 1908
P.nepheleptena (Dyar, 1908)
moeseri Bird, 1911
P.circumlucens (Sm., 1899)
ochroptena (Dyar, 1908)
humuli Bird, 1915
P.blackorum Quinter, 2025
P.impecuniosa (Grt., 1881)
P.nelita (Stkr., 1898)
errans B. & McD., 1912
†orbicularis Strand, [1916], (form)
†linda Bird, 1916 (form)
engelhardti Bird, 1939
P.sablensis Quinter, 2025
P.insulidens (Bird, 1902)
pertincta Dyar, 1920
P.birdi (Dyar, 1908)
P.dribi B. & Benj., 1926
P.albimontana Quinter, 2025
P.sauzalitae (Grt., 1874)
erubescens Bird, 1911
†papaipemoides (B. & Benj.), 1929, (form)
P.angelica (Sm., 1899)
P.harrisii (Grt., 1881)
verona (Smith, 1899)
anargyrea (Dyar, 1908)
†mulieris Strand, [1916], (form)
P . rubiginosa Bird, 1911
P.astuta Bird, 1907
P.leucostigma (Harr., 1841)
purpurifascia (G. & R., 1868)
†luteipicta Strand, [1916], (form)
P.lysimachiae Bird, 1914
P.pterisii Bird, 1907
triorthia (Dyar, 1908)
P.hesseli Quinter, 2025
P.limpida (Gn., 1852)
P.maritima Bird, 1909
P.beeriana Bird, 1923
†lacinariae Bird, 1923 (form)
EXCLUDED TAXA
“Amphipoea” lunata (Smith)
Apamea lunata (Smith), 1891, Transactions of the American Entomological Society, 18: 110.
Type Locality: Alameda Co., California, [USA]. [USNM]
NOTE—The systematic placement of this species within Apameini remains unknown; it is currently misplaced in Amphipoea.
“Hydraecia” arnymai (Dyar)
Hydroecia arnymai Dyar, 1913, Proceedings of the United States National Museum, 44: 296.
Type Locality: Zacualpan, [Veracruz], Mexico. [USNM]
NOTE—This species is currently misplaced in Hydraecia; its systematic placement is in or near Hydroeciodes Hampson (Noctuinae: Eriopygini).
“Papaipema” apicata Dyar
Papaipema apicata Dyar, 1912, Proceedings of the United States National Museum, 42: 65.
Type Locality: Zacualpan, [Veracruz], Mexico. [USNM]
NOTE—This species is currently misplaced in Papaipema; its systematic placement is in or near Hydroeciodes Hampson (Noctuinae: Eriopygini).