The Plusiinae (Lepidoptera: Noctuidae) of America North of Mexico, Emphasizing Genitalic and Larval Morphology
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i The Plusiinae (Lepidoptera: Noctuidae) of America North of Mexico, Emphasizing Genitalic and Larval Morphology Technical Bulletin No. 1567 •:_.C^ ;-5C2 :—I m Agricultural Research Service UNITED STATES DEPARTMENT OF AGRICULTURE •^iyt$.i: The Plusiinae (Lépidoptera: Noctuidae) of America North of Mexico, Emphasizing Genitalic and Larval Morphology By THOMAS D. EICHLIN and HUGH B. CUNNINGHAM Technical Bulletin No. 1567 Agricultural Research Service UNITED STATES DEPARTMENT OF AGRICULTURE ABSTRACT T D. Eichlm and H. B. Cunningham. The Plusiinae (Lepidoptera: Noctui- dae) of America North of Mexico, Emphasizing Genitalic and Larval Morphology. U.S. Department of Agriculture Technical Bulletin 1567, 122 pp., 1978. Adults of 71 species and larvae of 35 species of the noctuid subfamily flusimae of America north of Mexico were studied. Included are keys to the genera based on adult structures, a key to the species for which larvae are known, with setal maps and illustrations of other taxonomic characters, and keys to the adults based on the external morphology and on the male and témale genitalia, with photographs and illustrations of the structures Two species are described, Syngrapha abstrusa n. sp. and S. cryptica n sp., which form a species complex with S. alias (Ottolengui). S. viridisiffma (Cxrote) IS elevated to species level, having previously been considered a variation of S. selecta (Walker). Caloplusia Smith and Anagrapha Mc- Dunnough are synonymized with Syngrapha Hübner. Biological data distributional information, and bibliographical notations are included A phylogeny for the species comprising the subfamily in North America is proposed. This phylogeny has been deduced by correlating characters of both adults and larvae and comparing it with similar studies of Palearctic species. Washington, D.C. Issued March 1978 iii ACKNOWLEDGMENTS This investigation would not have been possible without the cooperation of many individuals and institutions throughout North America and at the British Museum (Natural History). The senior author thanks the following people for showing me how and when to collect in their areas and later continued to send invaluable living and preserved material: L. P. Grey, Lincoln, Maine; Warren Kiel, VTiitefield, N.H.; Dox Lennox, Jefferson, N.H.; W. R. Martyn, Easton, Pa.; R. Michaels, Georgetown, Ontario; M. C. Nielsen, Lansing Mich.; D. H. Habeck, University of Florida, Gainesville, Fla.; G. W. Wood, Canada Department of Agriculture, Forestry Division, Fredricton, New Brunswick; B. A. Sugden and D. A. Ross, Canada Department of Agriculture, Forestry Division, Vemon, British Columbia, also forwarded living and preserved material. Those who contributed distributional and other biological data include: Bryant Mather, Clinton, Miss., and Christopher Henne, Pearblossom, Calif. The following individuals allowed us to examine specimens from their private or institutional collections: L. D. Anderson and S. Frommer, Univer- sity of California, Riverside; J. A. Powell, University of California, Berkeley; R. L. Schuster, University of California, Davis; P. Arnaud,. California Academy of Sciences, San Francisco; H. Clench, Carnegie Museum, Pitts- burgh, Pa.; J. P. Donahue, C. L. Hogue, and L. Martin, Los Angeles County Museum of Natural History, Los Angeles, Calif.; R. P. Holdsworth, Ohio State University, Columbus, Ohio; R. H. Leuschner, Manhattan Beach, Calif.; M. MacKay, Canadian National Collection, Ottawa, Ontario; N. McFarland, Adelaide, South Australia; A. H. Rose, Canada Department of Agriculture, Forestry Division, Sault Sainte Marie, Ontario; R. L. Fischer, Michigan State University, East Lansing; G. T. Okumura, Laboratory Services, California Department of Food and Agriculture, Sacramento; and D. L. Bauer, South Lake Tahoe, Calif. D. C. Ferguson, E. L. Todd, and D. M. Weisman, Systematic Entomology Laboratory, U.S. Department of Agriculture; W. D. Duckworth, National Museum of Natural History, Smithsonian Institution; and F. H. Ringe, American Museum of Natural History, deserve special thanks from the senior author for their instruction and assistance. We are indebted also to A. Watson and L W. B. Nye, British Museum (Natural History), for sending photographs of certain type material and genitalia of the types and for making comparisons with type material. This paper resulted from studies conducted by the senior author in partial fulfillment of the requirements for the Ph.D. degree for the Depart- ment of Zoology-Entomology, Auburn University, Auburn, Ala. This De- partmient furnished materials and funds for travel to collect and examine specimens in other collections. Members of the Auburn University staff who were particularly helpful with suggestions, equipment and technical in- struction were: J. D. Harper, E. C. Mora, and G. W. Folkerts. IV The photographic work was done by the Smithsonian Photographic Laboratory. Although the senior author takes full responsibility for the drawings, he wishes to thank Charles S. Papp, illustrator, California Department of Food and Agriculture, for final preparation of line-drawing plates. The study is published as part of the ongoing research of the Agricultural Research Service, United States Department of Agriculture. CONTENTS Page Introduction 1 Taxonomy 1 Biology 3 Systematic treatment 4 Methods and materials 4 Diagnostic characters 5 Key to genera of piusiine adults 7 Key to known species of plusiine larvae 8 Tribe Abrostolini, new tribe 9 Abrostola Ochsenheimer 9 Mouralia Walker 11 Tribe Argyrogrammini, new tribe 12 Argyrogramma Hübner 12 Trichoplusia McDunnough 14 Pseudoplusia McDunnough 16 Tribe Autographini, new tribe 17 Autoplusia McDunnough 18 Rachipliisia Hampson 20 Pliisia Ochsenheimer 21 Allagrapha Franclemont 21 Polychrysia Hühner 23 Adeva McDunnough 24 Pseudeva Hampson 25 Chrysanympha Grote 26 Eosphoropteryx Dyar 26 Autographa Hübner 27 Syngrapha Hübner 35 Chrysaspidia Hübner 52 Phylogeny 54 Checklist of North American Plusiinae 58 Literature cited 59 Illustrations 64 Index to genera, species, and lower categories of Plusiinae 113 Index to host plants 116 VI The Plusiinae (Lepidoptera: Noctuidae) of America North of Mexico, Emphasizing Genitalic and Larval Morphology By THOMAS D. EICHLIN and HUGH B. CUNNINGHAM» INTRODUCTION The noctuid subfamily Plusiinae is distrib- are leaf feeders on a great array of plants. uted throughout the tropical, temperate, and Larvae of species in the genus Chrysaspidia arctic regions of the world. The North Ameri- Hübner feed on various species of grasses. can fauna includes representatives of each of This revision recognizes 71 species and places these climatic regions. This paper treats the them in 17 genera. We have endeavored to North American fauna, but the need to inte- demonstrate the relationships among the spe- grate our fauna with that of other faunal re- cies as interpreted from our data. Based on gions is great. larval studies, the species are closely related, Certain plusiines are frequently mentioned in except for the Abrostola- group. The species are the literature because their larval forms (loop- grouped according to their deduced phyloge- ers) are leaf feeders, which damage economi- netic relationships based on the adult and lar- cally important crops, garden vegetables, val characters of the extant species. Our classi- greenhouse plants, and ornamental herbs. The fication is explained under "Phylogeny." boreal species cause some injury to conifers This publication includes data and keys to (Prentice, 1962)? The remainder of the species the late-instar larvae of 35 species. TAXONOMY The early catalogers Walker (1857, 1858) and subfamily is based for the most part on the Smith {189S) included the majority of Plusiinae revision by McDunnough (19H\ His checklist species in the genus Plusia Ochsenheimer; (1988) was based on Dyar's generic designations Walker applied the name Plusidae for the (1902). The arrangement of McDunnough's list group. Hampson (1913) placed most of the spe- was based largely on genitalic characters and cies in two genera, Phytometra Haworth and on the work of Richards (1932) on skeletal Syngrapha Hübner, and included all plusiines morphology of the noctuid tympanum. Inter- in the subfamily Phytometrinae. pretations of the tympanal studies are dis- The North American classification of the cussed later in the present paper, since they pertain to phylogenetic associations of the subfamilies of the Noctuidae. McDunnough and 1 Systematic entomologist, Division of Plant Industry, his contemporaries were among the first to California Department of Food and Agriculture, Sacra- fully recognize the value of genitalic structures mento, Calif., and systematic entomologist. Department in taxonomic treatments. McDunnough (19U) of Zoology-Entomology, Auburn University, Auburn, Ala., used these structures as the primary basis for respectively. 2 The year in italic after the author's name refers to his classification and the delineation of species Literature Cited, p. 59. in the genus Syngrapha. The principal short- TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE coming of his revision was his failure to de- tions. From the Nearctic, he includes in this scribe adequately the remaining species in the group Argyrogramma, Trichoplusia, and Chry- subfamily. sodeixis. Group B, with one exception, is char- Forbes (195JÍ) placed the majority of the spe- acteristically Nearctic-Neotropical, and in our cies in the genus Plusia, retaining the generic fauna he includes Autoplusia and Rachiplusia,