2 8 2 8 1/11/2.5 2 5 1.0 :; 11111 . 1.0 :!t 11111 . 11111 . I~ ~~13.2. .2 I~ l~p2 2.2 L:.l I" •.:.: 136 .::: I~ u:: """ \:,; &:. :f J~ u I~ 2.0 " .... " 1.1 ...t:lf,.:.1.I. 1.1 La,:,,," --- -- 111111.8

1 4 1 6 111111.25 111111.4 111111.6 111111.25 11111 . 11111 .

MICROCOPY RESOLUTION TEST CHART MICROCOPY RESOLUTION TEST CHARl NATIONAL BUREAU OF STANDARDS·1963·A NATIONAL BUREAU Of STANDARDS·1963·A The Plusiinae (Lepidoptera: Noctuidae) of America North of Mexico, Empllasizing Genitalic and Larval Morpholog,ry

By THOMAS D. EICHLIN and HUGH B. CUNNINGHAM

Technical Bulletin No. 1567

Agricultural Research Service UNITED STATES DEPARTMENT OF AGRICULTURE ABSTRACT

T. D. Eichlin 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 Plusiinae 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 knovm, with setal maps and illustrations of other taxonomic characters, and keys to t:he a.dults based on the external morphology and on the male and female gemtalia, with photographs and illustrations ofthe structures. Two species are described, Syngrapha abstrusa n. sp. and S. cryptica n. sp., which form a species complex with S. alias (Ottolengui). S. viridisigma (Grote) 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 Hubner. Biological data, distributional information, and bibliographical notations are included. A phylog'eny 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, Vl1itefield, 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, Vernon, 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 Gniversity, 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 1. 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 0; Zoology-Entomology, Auburn University, Auburn, Ala. This De­ partment 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. lV The photographic work <\\raS done by the Smithsonian Photographic Laboratory. Although the senior author takes full responsibility for the drawings, he wishes to thank C'1arles S. Papp, illustrator, California Depa7tment 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.

v CONTENTS

Page Introduction ______1 Taxonomy ______1 Biology______3 Systematic treatment ______.______4 Methods and materials ______4 Diagnostic characters ______5 Key to genera ofplusiine adults ______7 Key to known species ofplusiine larvae ______8 Tribe Abrostolini, new tribe ______9 Abrostola Ochsenheimer ______9 Mouralia Walker ______11 Tribe Argyrogrammini, new tribe ______12 Argyrogramma Hubner ______12 Triclwplusia McDunnough ______14 Pseudoplusia McDunnough ______16 Tribe Autographini, new tribe ______17 Autoplusia McDunnough ______18 Rachiplusia Hampson ______20 Plusia Ochsenheimer ______.... ______21 Allagrapha FranclemonL______21 Polychrysia Hubner ______23 Adeva McDunnough ______24 Pseud.eva Hampson ______25 Chrysanympha Grote ______26

~osphoropteryx Dyar ______26 Autographa Hubner ______. ______27 Syngrapha Hubner ______35 Chrysaspidia Hubner ______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 LarvallVlorphology By THOMAS D. EICHLIN and HUGH B. CUNNINGHAM!

INTRODUCTION

The noctuid subfamily Plusiinae is distrib­ al'e 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 Hubner 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).2 The remainder of the species the late-instal' larvae of 35 species.

TAXONOMY

The early catalogers Walker (1857, 1858) and subfamily is based for the most part on the Smith (1893) included the majority of Plusiinae revision by McDunnough (1944). His checklist species in the genus Plusia Ochsenheimer; (1938) 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, Phytometm Haworth and on the work of Richards (1932) on skeletal Syngrapha Hubner, 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, Divil1ion 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 (1944) of Zoology-Entomology, Auburn University, Auburn, Ala., respectively. used these structures as the primary basis for 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 1 2 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, Triclwplusia, and Ch?"y­ subfamily. sodeixis. Group B, with one exception, is char­ Forbes (1954) placed the majority of the spe­ acteristically Nearctic-Neotropical, and in our cies in the genus Plusia, retaining the generic fauna he includes Antoplusia and Rachiplusia. categories employed by McDunnough as sub­ Group C is exclusively Holarctic, but for one genera. species in the Alpine Zone of the Andes Moun­ Crumb (1956), in his work on the noctuid tains. This group includes Caloplusia and Syn­ larvae, included 24 species in 9 genera of Plusi­ grapha. Group D is for all other genera, which inae. Based on larval characters, he combined are mostly Holarctic and particularly in the some of the genera as applied by McDunnough Palearctic Region. but did not specify any synonymy. For example, In studies much like this current undert2.k­ Crumb treats Trichoplusia McDunnough, Ar­ ing, Ichinose (1962 and 1973) revised the Japa­ gyrogramma Hilbner, and Pseudopl7.l.,Sia McD. nese Plusiinae. His classification is the result of in the same sense but does not indicate that the an integrated approach, using adult and larval genera should be synonymized with Argyro­ morphology. His work with the immatures, as gramma. Some of his diagnostic characters was the case with Crumb (1956), led him to have been shown to be too variable for identifi­ adopt a somewhat more conservative system cation. than those studies based mainly on adult geni­ Brown and McGuffin (1942), Forbes (1954), talia. McGuffin (1954), Ross and Evans (1957), and Wood and Neilson (1960) described the larvae of Recently, another plusiine fauna was de·· certain species or made short keys to the larval tailed by Dufay (1970). Working with the Mada­ forms of small groups of species. Their descrip­ gascar species, he pointed out disagreements he tions in many cases apply equally well to larvae had with some of Kostrowicki's and Ichinose's of other species, due to a reliance on color conclusions, mainly on the generic level. Dufay patterns. The use of color pattern for identifica­ defends his system on the basis that structures tion is not possible from preserved specimens, of the genitalia, particularly on the males, and which lose the normal color and become a the form of sternite 8 of males are the best set uniform cream color. In nature, the pattern is of characters upon which to determine the most usually a pale-green ground color with various correct natural relationships. longitudinal white or yellow striping. The classification that follows evolved from Unfortunately, Saunders (1863) and Nordman the integration of information obtained from (1942) used color exclusively in their descrip­ tions of the larvae. the study of larvae and adults. The classificf1­ tion is not as conservative a system as Ichi­ Gardner (1947), utilizing many morphological nose's and not as liberally divided up as Mc­ features, wrote on the larval stages of certain Dunnough, Kostrowicki, or Dufay have viewed Palearctic species, none of which are included the subfamily. The phylogeny derived from this in the Nearctic fauna. study of the Nearctic has been compared with Kostrowicki (1961), in his extensive study of those arrived at by Kostrowicki (1961) and Ichi­ the adult Palearctic Plusiinae, derived a classi­ nose (196£) for the Palearctic region~. A study fication along the lines of McDunnough's North of the Pantropical Plusiinae is essential to get American revision. In the former's paper, he the total worldwide phylogenetic picture of the suggests some changes, which are discussed subfamily. The authors hope that studies such later in the appropriate categories, and divides as this one and those mentioned above will the genera into zoogeographical groups. Group stimulate others to take up the challenge of A includes those genera that are characteristi­ clarifying the Plusiinae of the Tropics, and with cally tropical and subtropical in distribution the use of larval characters as well as adult and only poorly represented in the Holarctic characters, set up a classification that reflects fauna, occurring there only in the southern por­ the subfamily worldwide. PLUSIINAE OF AMERICA NORTH OF MEXICO 3 BIOLOGY

The information that follows is based on spec­ of marked and released specimens. Moths were imens reared in the laboratory. Average devel­ observed at night feeding on flowers of Cephal­ opment time from egg to adult was 30 days. The anthus occidentalis L. (button bush) not more species collected in the Southeastern Uni.ted than 15 feet from a black light trap. At least States required only 3 days from deposition of half of the flowers had moths feeding on them the egg to time of hatching. The duration of at a given time, yet only two moths came to the this developmental stage was about 8 days for light during the entire evening, which repre­ species taken in the Northern United States sented a small percentage of the moths in the and Canada. Most species pass through five area. The effectiveness of the traps depends instars with each stage requiring 3 to 5 days. upon many variables, such as the species of Pseudoplusia includens has six larval instars. moth, temperature, humidity, wind, intensity of The prepupal stage lasts from 1 to 2 days. the light, stages of the moon, and probably The adults emerge about 8 days following pupa­ others. One of the most productive evenings for tion and usually mate the second day following collecting at a black light was during a steady emergence. It was not uncommon to find 10 to rain, which is probably contrary to popular 12 spermatophores in the corpus bursae. These belief. numbers indicate that in nature, multiple mat­ Some of the species of plusiines flew, fed, and ings are the rule, not the exception. oviposited during daylight. Trichoplusia ni and The adults are active, strong fliers and feed Pseudoplusia includens were observed to be on nectar from flowers. The females produce a active in large numbers in fields of cotton and sex pheromone, which attracts males to the soybeans in the afternoon in Alabama. Syngra­ vicinity, but apparently the chemical is not pfut microgamma was observed actively moving specific (Berger, 1968). When close to the fe­ about in open bogs in northern Michigan in males, the males apparently rely on visual and bright sunlight as early as 11:00 a.m. Specimens other stimuli to locate the females. of Syngrapha u-aureum were most readily taken from the flowers of a small species of Intricate behavioral rituals and possibly me­ Solidago during daylight above the tree line on chanical structures supposedly constitute iso­ Mount Washington, N.H. This diurnal activity lating factors. The male emits a highly volatile is the characteristic behavior of many of the chemical for a short time when in close proxim­ species, particularly of those in the high alti­ ity to the female. This chemical is dispersed tudes and northern latitudes. from minute droplets on long hair pencils lo­ The females deposit ova singly on the larval cated in a pocket on the venter of the abdomen. host plant, placing them far apart, in most The hairs project caudad and are ventral to the cases on different parts of one or more plants vinculum of the male genitalia. When the during the same ovipositing session. One female pocket is everted, the hairs spread forming a usually deposits 200 to 300 eggs during her wide fan. This permits the maximum surface lifetime. area to be exposed, enabling the volatile chemi­ The species of Abrostola overwinter in the cal to be dispersed very rapidly into the air. The pupal stage. The remainder of the plusiines, so pheromone probably serves either to tranquil­ far as is known, overwinter as early instar ize or excite the female until copulation has larvae, probably third instar. Larvae reared been accomplished. The pheromone may also from females of species that are northern in serv,e as a recognition device, letting the female distribution would stop feeding in the labora­ "know" that the proper male of the same spe­ tory after reaching the third instar. Some cies has arrived (Birch, 1970). moulting was observed later but without in­ Light traps used for attracting moths are crease in size. Attempts to interrupt this dia­ probably inefficient, at least for species of the pause in the laboratory were unsuccessful. For­ subfamily Plusiinae, particularly as a means for tunately, a few specimens in each culture evaluating population densities or for recapture continued to develop. 4 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE SYSTEMATIC TREATMENT

The subfamily Plusiinae is worldwide in dis­ developed prolegs, one on each of segments 5, 6, tribution. The species are quite uniform in gen­ and 10. The caterpillars loop as they move, due eral appeara.nce and morphology except for spe­ to the reduced number of abdominal prulegs. cies of Ab?'ostola Ochsenheimer and Moumlia The crochets are arranged on the prolegs in a Walker. Paraphrasing Forbes (1954), the Plusi­ longitudinal, biordinal series, except for the inae have a prominent head, strong tongue, uniordinal arrangement on Chrysaspidia spe­ and large ocelli. Labial palpi are well developed cies. A small, corrugated sclerite on the hypo­ with segment 3 long; eyes are naked and heav­ pharynx, the raduloid, is characteristic of most ily lashed. The tympanum js the usual for plusiines but has been observed on a few noctuids, but with tympanic membrane deep in, nonplusiine species in closely related groups. not easily visible without dissection, and the wing venation is quadrifid. Male genitalia lack The pupae, according to Nakamura (1974), corona for the most part, at least for those insofar as the Japanese fauna is concerned at species in the limited fauna under discussion. least, can be distinguished from other noctuid Most species are stout bodied, with deep vesti­ subfamilies by the abnormally high position of ture and strong thoracic and abdominal tufting; the labrum, as in most Pyraustinae pupae, and palpi are upturned; antennae are simple with by the rounded projection mesally extending on very short, fine cilia in most species; a membra­ the abdominal segments, formed by the apex of nous lobe like a second hood is present behind the wings and maxillae, a condition common to the tympanic hood; and tympanic pocket 4 is some Cucullinae. much enlarged, producing a large swelling on We divide the subfamily into three tribes, the outside of the body. Abrostolini, n. tribe; Argyrogrammini, n. tribe; The larvae of all species, other than Abrostola and Autographini, n. tribe; on the basis of adult and Moumlia, have only three pairs of fully and larval characters.

:Methods and Materials

The adults utilized in this study were either mens were collected by beating vegetation and borrowed from museums and private collec­ examining host plants. These methods pro­ tions, reared, captured at light traps, or caught duced no larvae in Michigan, New York, New by aerial net day or night. Moths were also Hampshire, Maine, and Quebec, the northern taken while feeding on flowers by day or by regions where collecting was done. using a headlamp to see them on flowers at night (Nielsen, personal correspondence). Reared larvae associated with adults were Eggs for rearing were obtained from gravid scarce in collections. Except for a few species, females. When the eggs hatched, one or two preserved plusiine adults were well represented first-instar larvae were placed in a l--oz plastic in the larger collections in the United States cup, containing 15 ml of an artificial medium and Canada. developed by Shorey (l96S). Larvae of each Selected specimens were sent to the British instal' and the pupae and the adults were killed Museum for comparison with type material. and preserved for further study. Immatures Type specimens as well as series of specimens were killed by dropping them in gently boiling were examined at the National Museum of water for 1 to 3 minutes, depending on their Natural History, Washington, D.C., and the size. The water was allowed to cool before the American Museum of Natural History, New larvae were transferred into vials containing 70 York. Identifications of adults were made pri­ percent ethyl alcohol for preservation. Adults marily by comparing genitalic preparations were killed and pinned. with descriptions, keys, and illustrations by In Alabama and Florida, some plusiine speci­ McDunnough (1944) and Forbes (1954). Many of PLTTSIINAE OF AMERICA NORTH OF MEXICO 5 these original identifications were later verified mies, have been made. The first entry in each by comparing them with type material. case is to the original description of the taxa. The genitalia of males and females of all species were removed, cleared, and preserved Due to the circumstances under which this either in glycerine in plastic microvials or study was conducted, we could not study all the mounted in balsam on slides. type material pertaining to the included spe­ Distribution and host plant data were com­ cies. Whenever possible, this material has been piled from several literature sources, included examined and lectotypes designated if neces­ Forbes (1954), Crumb (1956), Prentice (1962), sary. In those instances where syntypes are and Tietz (1972), and personal correspondence. deposited at more than one institution and Other data were taken directly from labels of insufficient syntypic material was examined, specimens examined and host files maintained lectotype designations have been deferred, by California Department of Food and Agricul­ pending review of the unexamined material. ture lepidopterists. In the latter cases, under The information provided concerning location the "Host Plant" category of each species, these of types is based on personal examination of the records are listed last and 'without a citation type material when possible, or has been ex­ following. Larval host records of the Canada tracted from various sources in the literature Forest Insect Survey (Prentice, 196:2) supplied and through personal correspondence. Where much data on the conifer feeders. In the latter questionable, the data are preceded by a ques­ case, the host records and insect species were tion mark. Type locality data are essentially as verified by rearing on the host from which the founJ in the original descriptions with an occa­ larvae were taken. An index to host plants (p. sional addition (State, district, Province, and so 116) is provided for quick reference. forth) for clarity. A list of references is provided under each The keys to species are mostly artificial. The generic and species heading. This is by no genera are arranged according to their phylo­ means a complete bibliography but incl udes genetic relationships as interpreted by us. Tax­ papers in which changes in nomenclature, re­ onomic changes instituted here are discussed in garding their history and pertinent synony­ the appropriate generic or specific category. Diagnostjc Characters

The male and female genitalia were described houses the inverted, membranous vesica. Any generally for each genus and for each species. sclerotized, spinel ike, or rodlike structure on For the species of Syngrapha, few generaliza­ the vesica is referred to as a cornutus. The tions could be made; thus, the species descrip­ terms "basal" and "apical cornuti," as used tions of the genitalia are presented in detail. here, need a note of explanation. Since Mc­ The species of Autogmplw are more homoge­ Dunnough's 1944 work, a new technique has nous than those of Syngmpha; therefore, the been developed that allows one to evert the species descriptions in the latter genus are vesica. When everted, what McDunnough re­ usually short, and references to structures as fers to as the basal cornutus becomes apical in "typical" indicate that these structures conform position and the apical cornutus is basal in to the characterization in the generalized de­ position. This everting technique has the ad­ scription of the genus. vantage of displaying the proper placement of The male genitalia (fig. 18)3 have many struc­ structures and their relation~hips on the vesica tures of diagnostic significance. Those most in addition to showing the size and conforma­ often used in this report are as follows: The tion of the vesica. aedeagus as used hel'e is the whole tubular The ~alves are two lateral, somewhat flat­ structure, often bulbous at the base, which tened, elongated structures, projecting caudad. The thickly sderotized section along the dorsal side is called the costa, and the dorsal edge is J Illustrations begin on p. 64. referred to as the costal margin. The ventral 6 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE side, and base of valve, is termed the "sacculus" on the e.nds. The apex may be shagreened, and is often expanded. The clavus is defined sclerotized, pigmented, strigate, or curved right here as a filiform or knoblike projection origi­ or left. The ductus seminalis is a very thin duct nating on the mesodorsal margin of the saccu­ arising from the apex in most species, but lus near the base of the valve. The clavus is arises from the fundus in a few species. rounded and setose on the apex. The term Measurements of the length of the ductus "ampulla" is used here to indicate the entire bursae were made in a straight line from the structure arising from the center near the mid­ base of the ostium bursae to the point of en­ point of the valve, more often basad than cen­ trance into the corpus bursae, without stretch­ trad. The ampulla may be only a small knob, ing or altering the normal shape of the ductus produced apically into a sharp spine, or long bursae. Reference to the width of the ostium and slender, rounded and setose at the apex. bursae means at its greatest width, which is The length of the valve was measured from the usually the caudal end and from a dorsal or most anterior basal portion to the caudal end. ventral aspect. The length of the corpus bursae The valves articulate basad on their mesal was determined from the tip of the apex to the surfaces with a platelike or shiald-shaped scler­ tip of the fundus in a straight line, without ite called the juxta. It is most often produced to disturbing its normal shape. a pointed apex, sometimes with a distinct and A portion of this study is concerned with the sclerotized projection on the apex referred to as differentiation of preserved larval forms of plu­ the spine of the juxta. siines. Chaetotaxy and other characters of the The tegumen was measured from the point of integument (fig. 1) were used along with struc­ articulation with the uncus to the basal end tures on the mouth parts, since these are not where it becomes narrow, not from the point of structures that can become altered during fIXa­ articulation with the valve. tion and preservation. A character of the hypo­ The cornutus often has a sclerotized plate pharynx first described by Eichlin and Cun­ extending from its base. When measuring the ningham (1969), referred to as the raduloid length of the cornutus to its tip, the plate was (Eichlin, 1975), made possible the differentiation not included. Measurements of the length of of some species of larval Plusiinae, which were the aedeagus did not include the membranous, previously inseparable. The presence of the rad­ apical extension, which is attached to the geni­ uloid can be determined without removing the tal capsule. hypopharynx and can be observed by deflecting The female genital structures most often the maxillary pal pus. The processes or ridges used (fig. 19) are the ostium bursae, ductus on the raduloid were counted with the aid of a bursae, corpus bursae, and ductus seminalis. compound microscope by observing the men­ The ostium bursae is that differentiated sec­ tum, which was removed and mounted in glyc­ tion from the external opening or ostium to the erine in a microculture slide. Many specimens point where the ductus bursae originates. In of the same species were examined from the great majority of species, the ostium bursae throughout its range with the result that the is well defined, sclerotizea, and pigmented. number of ridges on the raduloid apparently The ductus bursae is most often elongate, does not vary. For this reason, no range of the sinuous, tubular, partially sclerotized, and stri­ number of ridges was given under those spe­ gose. It most often enters anterior to the caudal cies' descriptions where the number of ridges end of the corpus bursae. on the raduloid was used as a specific attribute. The corpL.S bursae is usually a membranous The stereoscopic microscope provided sufficient sac but may have areas of pigmentation and magnification to study the other larval struc­ sclerotization. The cephalic end is called the tures. The setal nomenclature used in this fundus, and the caudal end is the apex, either study is that of Hinton (1946). or both of which may be somewhat defined by a An analysis of the species of Plusiinae fol­ constriction or elongation of the corpus bursae lows, presented according to the following for­ PLUSIINAE OF AMERICA NORTH OF MEXICO 7 mat: Keys to the genera of the three tribes of known larvae within each genus. A discussion plusiinae adults; a key to the species of known of deduced phylogenetic relationships follows plusiine larvae, which includes species from 13 the section on the analysis of the genera and of the 17 plusiine genera; keys to the species of species. The adult wing patterns are not de­ each genus based on male and female genitalia; scribed, but photographs of every species are descriptions of higher taxa and adults and provided for comparison.

Key to Genera of Plusiine Adults

1. Fore wing dorsally with transverse ridges of raised Males with ampullae and vinc:;ulum different from scales ...... 2 those descri~d above; females with ductus sem­ Fore wing smoothly scaled ...... 3 inalis arising from short apex of corpus bursae 2. Wing length more than 15 mm Moumlia Wlk. (p. 11) (fig. 124) ...... 9 Wing length less than 15 mm Abrostola Och. (p. 9) 3. All tibiae spined, or hind tibiae spined on terminal 9. Labial palpi long and curved outward at tip portion or with a few weak spines between spurs ...... •...... Polychr-ysia Hiib. (p. 23) •••••••••••••••••••••••••••••••••••••••••••• ..j Labial palpi not curved outward at tip. directed No tibiae spined ...... 6 dorsad ...... 10 4. Outer margin of fore wings straight or slightly con­ 10. Segment 3 of labial palpi long, acuminate ..... 11 cave ...... Autoplu8ia McD. (p. 18) Segment 3 of labial palpi ~horter, rounded apically Outer margIn of fore wings convex ...... 5 ...... 13 5. Males with clavus long and slender (fig. 26); females with ductus bursae entering corpus bursae ven­ 11. Fore wings with outer margin excised behind apex; trad: (Syngrapha ottolen{/uii males will key here palpal segment 3 hairy ventrad also, but are spined only on hind tibiae, while ...... Pse'udeva Hamp. (p. 25) Rachiplusia is spined on all tibiae) Fore wings with outer margin evenly rounded; pal­ ...... Rachiplusia Hamp. (p. 20) pal segment 3 not hairy ventrad ...... :l:~ Males with clavus short with wide base (fig. 88); 12. Fore wings short, apex rounded, and outer margin females with ductus bursae either not entering convex ...... Chrysanympha Grt. (p. 26) corpus bursae ventrad or if entering ventrad Fore wings long and narrow, apex pointed, and outer then ductus bursae flattened and L-shaped in margin oblique ... Eosphoro7lteryx Dyar (p. 26) ventral aspect ...... Syngrapha Hub. (p. 35) 6. Abdomen of males with lateral tufts of long scales on 13. Dorsal abdominal tufting lacking or greatly reduced segments 4 and 5; females with ductus seminalis ...... Chrysaspidia Hub. (p. 52) arising from the fundus (fig. 115) or if from the Dor;;al abdominal tufting well developed, especially apex, the ductus bursae is longer than the abdo­ on segment 3 ...... 14 men (fig. 126) ...... _. . . . 7 14. Males with clavus undeveloped (fig. 33), antennae Abdomen of males without lateral tufts of long moniliform; females with ductus bursae entering scales on segments 4 and 5; females with ductus corpus bursae dorsad, ductas bursae with no seminalis arising from the apex of corpus bursae excessory structure near entrance to corpus bur­ where ductus bursae is shorter than abdomen sae...... Adeva McD. (p. 24) (fig. 132) or from the end of a long, tubelike apex Males with clavus long and thin (fig. 47). antennae (fib. 123) ...... 8 finely ciliate; females with ductus bursa not 7. Caudal end of valves in males expanded, broadly entering corpus bursae dor~ad or if dorsad, duc­ rounded (fig. 31); females with ductus seminalis tus with a large, oval spiculate sac near en­ arising from the apex of corpus bursae, ductus trance to corpus bursae (fig. 119) ...... 15 bursae longer than abdomen (fig. 129) ...... TrichopluBia McD. (p. 14) 15. Fore wing with w

1. Prolegs present on abdominal segments 3 and 4. 2 Crochets biordinal; edge of mandible not as above Prolegs absent or vestigial on abdominal segments 3 13 and 4 (fig. 1) ...... 4 13. Head spotted; middorsal band formed of dark micros­ 2. Mandible with a wide gap between dentes 2 and 3 pines ...... Syngrapha viridisigma (fig. 5) ...... Mouralia tinctoides S.orophila Mandible without wide gap between dentes 2 and 3 Head immaculate; microspines minute, not forming 3 distinct dorsal band ..... Syngraphafalcifera 3. Abdominal segments 1 and 2 with seta D-2 at poste­ 14. Vestigial prolegs present on abdominal segments 3 rior end of narrow pale area Abrostola urentis and 4 (fig 1) ...... 15 Abdominal segments 1 and 2 with a conspicuous Vestigial p:'vl,,;;;g absent on abdominal segments 3 white or pale spot including seta D-2 anrl 4 ...... 18 ...... Abrostola ovalis 15. Ribs,;! and 3 of mandible terminating in processes 4. Seta SV-2 present on abdominal segment 1 (fig. 9) before reaching cutting margin (fig. 2) ..... 16 5 Ribs 2 and 3 of mandible not terminating in proc­ Seta SV-2 absent on abdominal segment 1 (fig. 8) esses but continuing to cutting margin (fig. 7) 14 ...... Trichoplusia ni 5. Setae SV-l and SV-2 present on meso- and meta­ 16. Abdominal segment 2 with pinaculum of seta SV-1 thorax...... 6 closer to SV-2 than to SV-3 Seta SV-2 absent on meso- and metathorax .... 8 · ...... Argyrogramma basigera 6. Raduloid present on hypopharynx (fig. 12) Abdomina! segment 2 with pina!!ula of setae SV-3, ...... Syngrapha u-aureum SV-l and SV-2 approximately equidistant from Raduloid absent on hypopharynx ...... 7 one another ...... 17 7. Pinacula and areas surrounding bases of setae dark; 17. 10 ridges on raduloid ..... Pseudopl1lsia includens interval between pinacula of setae V-Ion ab­ 13 ridges on raduloid ..... Argyrogramma verruca dominal segment 4 more than one-half to sube­ 23 ridges on raduloid ... Trichoplusia oxygramma qual to interval between pinacula of seta V-I 18. At least one rib of mandible terminating in process and seta SV-1 (fig. 15) . Syngrapha rectangula before reaching cutting margin ...... 19 S. alias None of the ribs of mandible terminating in proc­ S. celsa esses but continuing to cutting margin .... 27 S. parilis 19. Pinacula of setae SV-1 and SV-2 at least partially Pinacula and areas surrounding bases of setae not fused on abdominal segment 2 (fig. 10) .... 20 dark; interval between pinacula of setae V-Ion Pinacula of setae SV-1 and SV-2 separated on ab­ abdominal segment 4 one-half the interval be­ dominal segment 2 (fig. 11) ...... 23 tween the pinacula of seta V-I and seta SV-l 20. Ribs 2 and 3 of mandible terminating in processes (fig. 16) ...... Syngrapha microgamma before reaching cutting margin (fig. 2) ..... 22 8. Ribs 2 and 3 of mandible terminating in processes Only rib 2 of mandible terminating in process before before reaching cutting margin (fig. 2) . . . . . 9 reaching cutting margin (fig. 6) ...... 21 Ribs 2 and 3 of mandible not terminating in proc­ 21. Raduloid of hypopharynx present (fig. 12) esses but continuing to cutting margin (fig. 7) ...... Autographa rubida 10 Raduloid of hypopharynx absent 9. Bases of setae on head and venter of thorax with · ...... Polychrysia morigera dark annuli ...... Autographa mappa 22. Head with a black, lateral stripe including ocelli 1-6 Bases of setae on head and venter of thorax not · ...... Autographa precation'is darkened ...... Autographa ampla Head without black, lateral stripe 10. Raduloid of hypopharynx present (fig. 12) ..... 11 ...... Autographa bimaculata Raduloid of hypopharynx absent ...... 12 23. Ribs 2 and 3 of mandible terminating in processes 11. Interval between pinacula of the two V-I setae on before reaching cutting margin (fig. 2) ..... 25 abdominal segment 4 one-half the interval be­ Only rib 2 of mandible terminating in process before tween pinacula of V-I and SV-l (fig. 16) reaching cutting margin, occasionally also on rib ...... 8yngrapha selecta 3, but then with setal pinacula of head ringed Interval between pinacula of the two V-I setae on with black ...... 24 abdominal segment 4 more than one-half to sub­ 24. Raduloid with 23 minute ridges; dorsal abdominal equal to the interval between pinacula of V-I setae on protruding conical pinacula and SV-1 (fig. 15) ...... Syngrapha epigaea · ...... Autoplu.sia olivacea 12. Crochets uniordinal; ventral edge of mandible Raduloid with 12 minute ridges; dorsal abdominal curved inward (fig. 3) .. Chrysaspidia contexta setae without strongly prominent pinacula C. putnami · ...... Autoplu.sia egena PLUSIINAE OF AMERICA NORTH OF MEXICO 9 25. Abdominal segment 2 with pinaculum of seta SV-2 27. Pinacula of setae SV-1 and SV-2 at least partially closer to SV-1 than to V-1 (fig. 14) fused on abdominal segment 2 (fig. 10) .... 28 ...... Allagrapha aerea Pinacula of setae SV-1 and SV-2 separated on ab­ Abdominal segment 2 with pinaculu of seta SV-1, dominal segment 2 (fig. 11) . . . Rachiplusia ou SV-2 and V-1 approximately equidistant from 28. Integument with microspines .. Autographa biloba one another (fig. 13) ...... 26 Integument without microspines, but with minute 26. Raduloid of hypopharynx present (fig. 12) granules ...... Autographa californica ...... Allagmpha ae1·oides Raduloid of hypopharynx absent .. Plusia balluca

Tribe ABROSTOLINI, New Tribe

The Abrostolini, the most primitive tribe in 2. Corpus bursae with sclerotized, pigmented plate dor­ our interpretation, is characterized in the adult sad (figs. 89 and 92) ...... parvula form by having areas of raised scales on the micrcvalis fore wings; tibiae without spines; male genitalia Corpus bursae without sclerotized plate (fig. 93) with a clavus in the form of a sclerotized, ...... ,...... urentis [For separation of parvula and microvalis, refer to adult platelike projection from the costa margin of figures (fig. 159 and 160).] the sacculus; and ampulla uniquely spatulate. The larvae have prolegs on abdominal seg­ Abrostola urentis Guenee ments 3 and 4, though slightly reduced on the third; and a raduloid on the hypopharynx. This Figure 158 tribe is represented in the North American Abrostola urentis Guenee, 1852, p. 322. fauna by two genera, Abrostola and Mouralia, Male genitalia (fig. 21).-Uncus long, thin, and with five species. pointed but unsplned apically; vinculum broadly rounded; valve with wide, sclerotized clavus, round and distal end, located near base Abrostola Ochsenheimer of sacculus; valve with costa produced into a large hump near middle,causing valve, to be Generotype.-Noctua triplasia Linnaeus, 1758, twice as wide at this point than elongate caudal p. 517. Designated by Stephens, 1829, p. 96. one-half; ampulla from a ·thick, inward project­ Abrostola Ochsenheimer, 1816, p. 88. ing base, spatulate, strongly incurved; aedea­ Adult and larval features generally as de­ gus with patch of spines external'on dorsum of scribed for the Abrostolini. apex, internally near apex with a ventral, bifur­ cate, sclerotized plate with two sharp projec­ KEY TO THE SPECIES BASED ON MALE GENITALIA tions on right edge, and dorsally with a cluster of 4 small cornuti on vesica, with another clus­ 1. Ampulla spatulate, incurved, extending beyond costa; clav<.ls wide, about same width throughout (fig. ter of about 12 cornuti just cephalad of first 21a) ...... urentis cluster. Ampulla narrow, pointed, outcurved at least apically, Female genitalia (fig. 93).-Two outward pro­ not extending to costa; clavus narrow at base, jecting, sclerotized, pigmented, granulose expanded, ovate apically (fig. 22) ..... , ...... 2 pouches laterad on cephalic end of abdominal 2. Vesica with at least two distinct patches of many small cornuti; ampulla much outcurved with apex segment 8; ostium bursae, sclerotized, granu­ directed somewhat ventrad (fig. 22). ".microvali,~ lose, wider than long, expanding laterad and Vesica with one continuous patch of cornuti; ampulla somewhat constricted at base; ductus bursae outcurved only slightly at apex (fig. 20) parvula initially narrow, granulate and sporadically [Male of ovalis not keyed out, no material examined.] sclerotized and membranous, making right an­ gle bend to left and another bend cephalad, KEY TO THE SPECIES BASED ON FEMALE gradually widening to about twice its initial GENITALIA width as it enters corpus bursae on left side; 1. Corpus bursae large and elongate (fig. 88) '" ovalis corpus bursae obovate, strigate caudad. Corpus bursae small and oval (fig. 93) ...... 2 Larvae.--Prolegs present on abdominal seg­ 10 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE ments 3-6, though slightly thinner on abdomi­ arises from poorly defined apex on the left side nal segment 3; abdominal segment 8 gibbose of corpus bursae, cephalad of entrance of duc­ subdorsad; head reticulate; mandible with tus bursae. (Described from USNM Type mesal processes on rib 2; raduloid on hypophar­ #41021.) ynx with 16 ridges; crochets average 20 to 28 in Types.--A male syntype of parvula in the number; abdominal segment 2 with pinacula of National Museum of Natural History has been setae SV-l and SV-2 widely separated;' abdomi­ selected and labeled and is presently desig­ nal segment ~. with SV-2 absent; and SV-2 nated as the lectotype; "Abrostola parvula B & absent on me"'o- and metathorax; no micros­ McD, Type o/So. Arizona, Poling/o gen. 353, 13 pines on integument; overall, very similar to M. Aug 32, FH!3." The holotype male_ ofmariana is tinctoides. also in the National Museum of Natural His­ Type.--British Museum (Natural History). tory. Type locality.--New York. TyP2 localities.--Southern Arizona (par· Host plant.·-Urtica lyalli Tietz (1972), Ur­ vula). Tempe, Ariz. (mariana). tica diotica. Host plant.-Unknown. Range (map IG).-Nova Scotia to British Col­ Range (map 8).-Arizona, California. umbia; United States except for southeast (no verified records from California, Eichlin, 1975): Abrostola microvalis Ottolengui Comments.--Larval specimens and the pa­ rental female were borrowed from the Cana­ Figure 160 dian National Collection, Ottawa, Ontario. Abrostola microvalis Ottolengui, 1919, p. 124. Male genitalia (fig. 22).-Description of male Abrostola parvula Barnes and McDunnough genitalia the same as for parvula with the following exceptions: Valve with ampulla Figure 159 strongly outcurved and projected caudad; ae­ Abrostola parvula Barnes and McDunnough, deagus with an apical and median patch of 1916, p. 225. many small, spinelike cornuti on vesica. Abrostola mariana Walter, 1928, p. 139.-Benja­ Female genitalia (fig. 92}.-From the speci­ min, 1932, p. 7. mens available, no consistent differences from Male genitalia (fig. 20).-Uncus pointed but A. parvula in female genitalia could be ascer­ not terminated in a spine; valve with ampulla tained. Both species are very rare in collections. slightly outcurved near apex and not, or Type.-Holotype male in the American Mu­ slightly, directed caudad; valve very narrow seum of Natural History. but expanded near the middle on costal edge to Type locality.--Texas. a high, rounded hump, basad of which is a Host plant.--Unknown. mushroom-shaped, flat, sclerotized clavus; ae­ Range (map 9).-Texas, Arizona. deagus with an apical patch of many small spinelike cornuti on vesica. Abrostola ovalis Guenee Female genitalia (fig. 89).-Ostium bursae a short but very wide membranous pouch with Figure 161 lateral membranous extensions caudad; lat­ Abrostola ovalis Guenee, 1852, p. 322. erally attached to these extensions are slightly Male genitalia.--Not examined. sclerotized, cup-shaped pockets open caudad; Female genitalia (fig. 88).-Ostium bursae ductus bursae, short, membranous, about one­ very shallow, sclerotized, shagreened, caudo­ third as wide as ostium bursae and curved left ventral edge folded down, with a median cleft, as it enters corpus bursae on the right side; sides folded in deeply subdorsad, causing «or­ corpus bursae a small oval sac with an evenly soreson to be completely open; ductus bursae, sclerotized and pigmented plate (signum) dor­ narrow and membranous for initialy short sec­ sad of ductus bursae entrance, rounded ce­ tion, abruptly expanding, becoming sclerotized, phalad and pointed ('~.udad; ductus seminalis darkly pigmented, much folded, shagreened, PLUSIINAE OF AMERICA NORTH OF MEXICO 11 with a fingerlike projection ventrad, and enters tapering to spoon-shaped apex, covered ven­ corpus bursae on left side, the entrance area as trally with hairlike scales and dorsally with long as one-third of total length of corpus bur­ long, spatUlate scales; ventrolateral margin of sae; corpus bursae elongate, mostly strigate, juxta with slender, sclerotized furca, just cau­ shagreened and lightly pigmented, caudal one­ dad of which is a sclerotized, lobed clavus; in half sharply angled toward the right, below center of valve are three projections, the center entrance of ductus bursae, produced ventrad as one being here considered as the actual am­ a broad, toughened ridge to apex; ductus semin­ pulla, which is narrow, rounded at apex, in­ alis arises from apex. curved, and projected toward sacculus; slightly Type.--British Museum eNatural History). above ampulla and toward base of valve is a Type locality.--New York. thick, short editum; below and caudad of am­ Host plant.--Urtica sp., Crumb (1956). pulla is a spatUlate clasper, wider than and Range (map 9).-Maine to North Carolina and subequal in length to ampulla. Aedeagus rela­ Ohio. tively small, about as long as tegumen, and Comments.--The larvae of Abrostola ovalis apically with a small cornutus on a broad, were not seen, but this species w~s placed in rounded base, and a club-shaped projection off the larval key on the basis of information ob­ vesica covered with small, broad spines. tained from Crumb (1956). Female genitalia (fig. 90).-No ductus. bursae as such, but with ostium bursae broad and short, slightly longer than wide, sclerotized, M ouralia Walker and partially strigate, leading directly into cor­ pus bursae mesoventrad; corpus bursae thinly Generotype.-Abrostola tinctoides Guenee, 1852, membranous, right side of corpus bursae pro­ p. 323. Designated by Hampson, 1913, p. 591. duced cephalad into a narrow, elongated fun­ Mouralia Walker, 1858, p. 1803 dus, left side produced cephalad into an elon­ Although the larvae of this species are much gate but shorter apex, about one-third as long like a larger version of the larvae of Abrostola as fundus, and slightly spiculate; ductus semin­ species, the genitalia of the adults indicate that aUs arises near tip of apex. they should not be considered as congeneric. Type.--Two male syntypes in Guenee Col­ The following description of M. tinctoides is an lection, [7] British Mu~um (Natural History). adequate description for the genus as well. Type locality.--Brazil. Host plants.--Tradescantia jZuminensis Mouralia tinctoides (Guenee) (wandering-Jew or spiderwort in part); Zebrina pendula (wandering-Jew in part). Figure 162 Range (map lO}.-Florida, Texas, and Califor­ Abrostola tinctoides Guenee, 1852, p. 323. nia; south to Brazil. Mouralia tinctoides: Hampson, 1913, p. 402. Cornrnents.--This species is considered to be Male genitalia (fig. 23).-Uncus C-shaped, closely related to Abrostola mainly because the broad at base, pointed at apex with a minute larval form is very similar to the larvae of curved spine at tip, distal one-fourth round, urentis and ovalis. The setal arrangement is basal three-fourths flat on top and much wider the same. A slight difference from the mandible in the middle, from which point arises a cluster of tinctoides and a difference from urentis in of hairlike scales; rectum near base of uncus the number of ridges on the raduloid of the with two subdorsal, sclerotized, somewhat tri­ hypopharynx were the only differentiating angular projections; juxta simple, shieldlike; characters found. Though the male genitalia vinculum not extending much beyond base of are highly modified, they still have characters valves, broadly rounde~ at apex; valves thickly peculiar to the Abrostolini, such as the odd, covered on inner surface with hairlike scales, platelike clavus on the costal margin of the particularly on sacculus all along ventral edge sacculus. Larvae were from the National Mu­ of valve to apex, valve widest near middle, seum of Natural History. 12 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE Tribe ARGYROGR..M'IMINI, New Tribe

The Argyrogrammini is for the most part and states that Triehoplusia cannot be conge­ southern in distribution, ranging into the Trop­ neric with it. He also finds Kostrowicki's placing ics for Tl-iehoplusia ni and Pseudoplusia inelu­ of seven species in Argyrogramma to be an dens, which are generally distributed in the erroneous application of the limits of this ge­ north temperate zone of America as well. This nus. trIbe corresponds to Kostrowicki's Group A A more accurate interpretation of these ge­ (1961, p. 369). The adults have no tribal spines; nera probably lies somewhere in between the males have a well-developed, elongate clavus concepts based strictly on adults or mostly on and saccus; and, except for P.ineludens, males immatures. To add further to the problem, have strong lateral tufts of scales caudad on Nakamura (1974) states that based on pupal the abdomen. The larvae have a raduloid on the chaetotaxy, the generic concepts of both Dufay hypopharynx; and the prolegs on abdominal and Ichinose, as regards Chrysodeixis, Argyro­ segments 3 and 4 are vestigial, heing reduced to gramma, and Triehoplusia, are apparenl:ly minute, peglike structures having no crochets. somewhat in error, because species of each The Argyrogrammini in North America is com­ genus, as recognized by those authors, fit into posed of three genera, Argyrogramma, Pseudo­ more than one of Nakamura's categories. pL1tsia, and Tl-iehoplusia, which include six spe­ The limited representation of this tribe in the cies. Nearctic does not permit us to resolve the Kostrowicki (1961), Dufay (1970), and Ichinose controversy at this time. We believe a thorough (1962 and 1973) differ as to the generic concepts investigation of the Neotropical fauna in this within the Argyrogrammini. According to the complex will be necessary before more accurate larval key to genera in Ichinose (1962), the conclusions can be drawn. species of this tribe in North America are all congeneric with Chrysodeixis Hub. Ichinose Argyrogramma Hiibner (1973) synonymizes Tr-ichoplusia with Argyro­ gramma and Pse1ldoplusia with Chrysodeixis. Generotype.~Argyrogramma omega Hubner The larvae of the Nearctic species do not fit the [1821] (=verruca Fabricius, 1794). generic limits as defined by Ichinose. The lar­ Designated by McDunnough, 1944, p. 206. vae of the type species for the genus Argyro­ g}'amma, A.ven"1wa (Fab.), do not fit his Argyrogramma Hubner, [1821], p. 29. character whereby, "...DJ on Tz not so close to Phytometra Hampson (nee Haworth), 1913, p. 452 (partim). D2, situated on the imaginary line connecting D2, and SV~." As a matter of fact, the species is Autographa: McDunnough, 1938, p. 113. similar to the character state described for Argyrogramma: McDunnough, 1944, p. 205. Chrysodei:ris whereby, "... D, on T2 near to D2, The larvae and features of the males are situated just behind the imaginary line con­ generally as descri oed for the tribe, the differ­ necting Dz and SD2," but the larval form does ences being more useful in a specific sense. not agree on a second character state of the In North America, the females of the two latter genus. In like fashion, Pseudophlsia in­ Argyrogramma species are characterized by cludens larvae are similar to Chrysodei.ris on having the ductus seminalis arising from the one character but differ from this genus on a fundus on the corpus bursae, a rather unusual second. situation in the subfamily. One could perhaps synonymize all of the above genera of the Argyrogrammini under KEY TO THE SPECIES BASED ON MALE GENITALIA Ch rysodei:ris and redefine the genus. Dufay Aedeagus with one COrt1utus; apex of ampulla knobbed (1970), relying on genitalic differences and the (fig. 25) ••.....•••.....•...•..•.•.•...... verruca last abdominal sclerite of the males, does not Aedeagus with two cornuti; apex of ampulla hooked agree with Ichinose's concept of Chlysodei:J."is (fig. 24) •.•••...••••.••••.••..••..•..••• ba8igera PLUSIINAE OF AMERICA NORTH OF MEXICO 13 KEY TO THE SPECIES BASED ON FEMALE GENITALIA ment 2; crochets range from 17 to 20 in number; dark, lateral band on head if present, appar­ Ductus bursae thin, ropelike, somewhat sc1erotized and strigate for most of its length (fig. 96) ... verruca ently not including ocelli 1-4; ribs 2 and 3 of Ductus bursae straplike, sc1erotized throughout and mandible terminating in processes before strigate (fig. 91) ...... basigera reaching cutting margin; raduloid on hypophar­ ynx with 13 ridges. Argyrogramma verruca (Fabricius) Type.--Fabricius Collection, Kiel, Denmark, [Zimsen, 1964] (verruca). [?] Lost (omega). [?] Figure 163 Rijksmuseum van Natuurliijke Historie, Lei­ Noctua ve1"rUca Fabricius, 1794, p. 81. den, Netherlands (00). [?] Lost (omicron). Plusia olnega (Hubner), 1823, p. 29, figs. 373, Type locaIity.-South America (verruca). Sa­ 374; Plusia 00 (Cramer), 1782, pI. 311, fig. e vanna, Ga. (omega). [?] (00). [?] (omicron). (pm"tim); Plusia omicron (Hubner), 1821, p. Host plant.--Sagittaria sp., Forbes (1954); 251.-Guenee, 1852, p. 342. Gossypium hirsutum, Justicia americana, Nico­ Autographa verruca: Dyar, 1902, p. 81. tiana tabaccum, Vigna unguiculata; Crumb Phytometra verruca: Hampson, 1913, pp. 465, (1956); Calendula sp., Kimball (1965); Rumex 466. crispus, Eichlin and Cunningham (1969); Poly­ Autographa ve7lUCa: McDunnough, 191.6, p. 400. gonum punctatum, P. setaceum, Heppner and Habeck (1976); Hyptis sp., Mentha sp., Pelargon­ Argyrogramma verruca: McDunnough, 1944, pp. ium sp., Spathiphyllum sp., Tradescantia flumi­ 205,206. nensis, Trichosporum sp. Male genitalia (fig. 25).-Uncus widest near Range (map 4).-Ontario, Maine to Florida, apex, with large, curved, terminal spine; vincu­ west to Colorado, and south to tropical South lum broad, same width throughout, truncate at America. apex; valve elongate, rounded apically, with Comments.--Parental females of laboratory short, knobbed, heavily setose ampulla, and cultures were collected from a light trap at long, filiform clavus; aedeagus long, slender, Auburn, Ala. only slightly expanded at base, with flat, basal cornutus, pointed at both ends, and an S­ shaped, spinose plate near the middle, and a A rgyrogramma basigera (Walker) sclerotized, spinose plate near apex; apex of Figure 164 aedeagus externally, minutely spinose. Female genitalia (fig. 96).-Ostium bursae Plusia basigera Walker, 1865, p. 836. small, weakly sclerotized and strigate; ductus Plusia laticlavia Morrison, 1875, p. 98.-Smith, bursae slender, initial short portion S-shaped 1893, p. 225. when viewed from the left side, membranous, Autogmpha basigem: Dyar, 1902, p. 81. becoming somewhat sclerotized, granulate, and Phytometm basigera: Hampson, 1913, pp. 486, strigate, bending dorsally and laterad toward 487. the left, entering the corpus bursae dorsally Autographa basigera: McDunnough, 1916, p. toward the right of meson; corpus bursae finely 400. spiculate, tapering to a blunt fundus from Argyrogramma basigera: McDunnough, 1944, which arises the ductus seminalis. pp. 205, 206. Larvae.--Prolegs vestigial on abdominal Male genitalia (fig. 24).-Uncus thin through­ segments 3 and 4, reduced to small peglike out, with terminal spine; vinculum narrow, structures (fig. 1): No microspines on integu­ elongate, bluntly pointed at apex; valve elon­ ment; mesothorax with pinaculum of D-2 closer gate and rounded apically; with relatively long to D-l than to SD-2; seta SV-2 absent on meso­ ampulla, strongly hooked at apex, directed ce­ and metathorax; seta SV-2 absent on abdomi­ phalad, and long filiform clavus; aedeagus long nal segment 1; pinaculum of seta SV-2 equidis­ and slender, expanded at base, with flat, basal tant from SV-3 and SV-l on abdominal seg­ cornutus rounded at both ends and a sharply 14 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE pointed median cornutus, apex of aedeagus ex­ to study the foreign material under question ternally, minutely spinose; juxta relatively nar­ first hand. At this time, we prefer to view row, strongly keeled medially. Trichoplusia as interpreted by McDunnough Female genitalia (fig. 91).-Ostium bursae (1944), recognizing that this interpretation is small; ductus bursae initially narrow but ab­ based on the very limited North American ruptly expands to as wide as widest part of Argyrogrammini fauna. ostium bursae, the caudal one-half of ductus Though modified, the male genitalia in gen­ bursae flat and granulose, becoming more eral form are as described for the tribe. The heavily sclerotized, strigate and twisted, enter­ characters of the genitalia are most useful for ing the corpus bursae dorsally toward the right; specific determinations. corpus bursae narrow about four times longer The female genitalia are characterized by the than wide, blunt at apex and tapering to a unique development of the ductus bursae, blunt point cephalad, with ductus seminalis which is extremely long and tapelike. The duc­ arising from fundus. tus seminalis arises from an elongate, narrowed Larvae.--Mesothorax with pinaculum of apex of the corpus bursae. seta D-2 equidistant from D-l and SD-2; pina­ culum of SV-2 closer to SV-1 than to SV-3 on KEY TO THE SPECIES BASED ON MALE GENITALIA abdominal segment 2; crochets range in num­ Ampulla produced caudad, then making a right angle ber from 20 to 23; dark, lateral band on head, if bend dorsad, sickle shaped (fig. 34) .,...... ni present, including ocelli I-B in all specimens Ampulla long and thin, protruding well above valve (fig. examined; raduloid on hypopharynx with 17 30) ...... •...... •...... oxygramma ridges; otherwise very similar to verruca. KEY TO THE SPECIES BASED ON FEMALE Type.--Female, Carter's collection, British GENITALIA Museum (Natural History) (basigem). Tepper Collection, Michigan State University (laticla­ Ductus bursae does not enter through a pouch on corpus bursae (fig. 102) ...... •...... ni via). Ductus bursae enters corpus bursae through a long Type locality.--North America (basigera). membranous pouch on left side of corpus bursae New York (laticlavia). (fig. 105) ...... oxygramma Host plant.--Unknown. Range (map l).-Maine south to Florida, west Trichoplusia ni (Hiibner) to Texas and Mexico. Comments.--Parental females of laboratory Figure 165 cultures were collected at a black light in Noctua ni Hiibner, [1800-1803], pI. 58, fig. 284. Gainesville, Fla., and from a light trap at Au­ Autogmpha ni: Hiibner, 1821, p. 25l. burn, Ala. Plw;ia ni: Treitschke, 1823, p. 189. Plusia ni: Grote, 1873, p. 147 [=bmssicae]. Trichoplusia IVlcDunnough Plusia echinocystidis Strecker, 1874, p. 94.- Smith, 1893, p. 253. Generotype.-Plusia ni var. bmssicae Riley, Phytometm ni: Hampson, 1913, pp. 468-470. 1870, p. 110. Original designation. Autogmpha ni: McDunnough, 1916, p. 400. Trichoplusia McDunnough, 1944, p. 204. Autogmpha bmssicae (Riley), 1870, p. 110.- Chrysode'in.s: Ichinose, 1962, p. 95. McDunnough 1938, p. 113. Trichoplusia: Dufay, 1970, p. 15. Trichopltulia ni: McDunnough, 1944, p. 204. Argyrogmmma: Ichinose, 1973, p. 138. Male genitalia (fig. 34).-Tegumen long and Chrysodei~:is: Nakamura, 1974, p. 6. slender; vinculum long, tapering to a point; As previously mentioned in the discussion of valve widest at broadly rounded apex with the Argyrogrammini, we will not attempt here sickle-shaped ampulla produced dorsad as a to resolve the controversy surrounding the ge­ thin point, passing just beyond costal margin of neric limits of Triciwplusia, Chrysodeixis, and valve, and clavus slightly shorter than width of Argyrogmmma worldwide, having been unable valve at this point; aedeagus about as long as PLUSIINAE OF AMERICA NORTH OF MEXICO 15 valve, slender for most of its length, bulbous at sp., Mentha sp., Nicotiana glauca, N. tabaccum, base, with no cornuti. N. tngonophylla, Pectis papposa, Phaseolus vul­ Female genitalia (fig. 102).-Ostium bursae garis, Pisum sati1lUm, Plantago sp., Raphanus small, almost spherical, dorsal surface membra­ sativus, Reseda odorata, Rumex sp., Senecio nous, vertically strigate laterad and smoothly cruentis, S. mikanioides, Smilax rotundifolia, sclerotized ventrad; ductus bursae initial1y one­ Solanum tuberosum, Spinacea oleracea, Tarax­ half the width of ostium bursae and membra­ acum officinale, Trifolium sp., T?·opaeolum ma­ nous, but quickly and abruptly expanding to as jus, Vigna sinensis, (Tietz, 1972); Acacia greggii, wide as ostium bursae and continuing to left Asclepias sp., Be?llenia sp., B1·assica nap?ts, Ci­ side of corpus bursae as a strigate, sclerotized, neraria sp., Citrus sp., C. sinensis, Coleus sp., narrow ribbonlike tube, extremely long, being Crotalaria 1YJtundifolia, Cucurbita pepo, Daucus about 4 to 5 times longer than length of corpus caTata, Euphorbia pulcher'rima, F1'agana sp., bursae; corpus bursae saclike, narrowing to a Helianthus sp., Hibiscus e.c;culentus, Ipomoea blunt point at apex from which arises the duc­ purpurea, Lativa sp., Lupinus sp., Lycopersicon tus seminalis. esculentum, Medicago sativa, Papaver sp., Petu­ Larvae.--Prolegs on abdominal segments 3 nia sp., Phaseolus sp., P. limensis, P. lunatus, and 4 vestigial, reduced to tiny peglike struc­ Philodendron sp., Rumex C?-1.spUS, Salvia sp., tures around the bases of which are clustered Tribulus sp., Zea mays. setae SV-2, SV-3, and V-I; seta SV-2 absent on Range (map 6).-United States and Southern abdominal segment 1; seta SV-2 absent on Canada; south to Tropics. meso- and metathorax, on mesothorax D--2 equi­ Comments.--This species is commonly distant from D-l and SD-2; seta SV-l and SV-2 termed the cabbage looper. Parental females of on abdominal segment 2 well separated, ap­ laboratory cultures were collected in a light proximately equidistant from each other and trap at Auburn, Ala., and near Pittsburgh, Pa. seta SV-3; crochets on ventral prolegs range in number from 19 to 22; no microspines on inte­ Trichoplusia oxygramma (Geyer) gument; ribs of mandible not terminating in processes before reaching cutting margin; radu­ Figure 166 loid on hypopharynx with 10 ridges. Autogmpha oxygmmnza Geyer, 1832, p. 37, figs. Type.--Not known to us (ni). A male syn­ 769, 770. type of bmssicae in the National Museum of Natural History has been selected and labeled Plu.c;ia oxygmmnw: Guenee, 1852, p. 353. and is presently designated as the lectotype: Autographa o:rygmmma: Dyar, 1902, p. 81. "294,ibs. Nov. 12/75iUSNM Type #275." [?] Autographa indigna (Walker) 1857, pp. 909, Strecker Collection, Chicago Field Museum 910-0ttolengui, 1902, p. 67. (echinocystidis). Phytometra oxygramma: Hampson, 1913, pp. Type locality.--Not known to us (n.i). Mis­ 494, 495. souri (bmssicae). California (echinocystidis). Autographa oxygramnw.: McDunnough, 1916, p. Host plant.--Pastinaca sati'ua, Crumb 400. (19,56); Ageratum conyzoides, Althaea ?"Osea, An­ Trichoplusia oxygmmma: McDunnough, 1944, t·irrhinum sp., Apium gmveolens, Argemone pla­ p.204. tyceras var. hispidia, Asparagus officinalis, Male genitalia (fig. 30).-Tegumen shortel' Beta mtigans, BmssicCL hirta, B. campestris, B. and wider than T. ni: vinculum much the same nigra., B. olemcea, B. rapa, Calendula officin­ as in ni; valve similar in shape to ni but very nlis, Chaenactis stevioides, Chaenomeles jnpon­ narrow before apex with ampulla straight, slen­ ica, Chenopodium album, Chrysnnthem1tm sp., der, rounded apically, two-thirds of its length Citrullus llulgnns, C?·epiB sp., Cucurbita ma:rima projected beyond costal margin, arising more C. mela, C. .'IntivLlS, Dianthus ca17Jophyllus, Er­ basad midway between base and middle of odiu,m sp., E. circ-utarium, Eschscholzia caUfor­ valve, and clavus similar to ni; aedeagus longer nica, Gemnium sp., Gossypium sp., Lactuca than valve, slender and slightly expanded bas­ canadens1:s, L. sativa, Lathyrus odoratus, Maf1,a ally with a long, needlelike, slightly curved, 16 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE basal cornutus, and a sclerotized, elongate F), 1782 = includens Walker, 1857, p. 94. Origi­ patch in apical one-third of aedeagus curving nal designation. from dorsal side ventrad to apex. Pseudoplusia McDunnough, 1944, p. 206. Female genitalia (fig. 105).-Ostium bursae Chrysodeixis: Ichinose, 1973, p. 139. entirely membranous, short, about three times The genus Psettdoplusia contains a single wider than long; ductus bursae similar to ni, species, includens. P. includens is closely re­ shorter, about three to four times longer than lated to other species of the Argyrogrammini, length of corpus bursae and enters corpus bur­ but due to differences in adult and larval mor­ sae into a thickened, membranous pouch; cor­ phology and our lack of a comprehensive treat­ pus bursae essentially the same as ni. ment of this tribe in the Neotropics, we have Larvae.--Very similar to ni with the follow­ chosen not to remove includens from Pseudo­ ing exceptions: On mesothorax, D-2 closer to D­ plusia, pending further study. 1 than to SD-2; number of crochets on ventral The males of this genus lack the lateral scale prolegs range from 22 to 25; mandible with ribs tufts on the abdomen, which characterize other 2 and 3 terminating in processes before reach­ Argyrogrammini species. ing cutting margin; raduloid on hypopharynx with 23 ridges; dark lateral stripe on head if present, not including ocelli 1-4 on all speci­ Pseudoplusia includens (Walker) mens examined. Type.--Not known to us (oxygramma). Syn­ Figure 167 types not seen by us, Milne's collection, British Plusia includens Walker, 1857, p. 94. Museum (Natural History) (indigna). Type locality.--Georgia (oxygmmma). Geor­ Plusia hamifera Walker, 1857, p. 917; P. dyaus gia {indigna). Grote, 1875c, pp. 203, 204.-Smith, 1893, p. 250. Host plant.--Aster sp., Erigemn canadensis, Autogmpha culta (Litner), 1885, pp. 94-97.­ Nicotiana tabaccum, Solidago sp., Crumb (1956). Ottolengui, 1902, p. 66. Range (map 12).-Eastern one-half and Autogmpha includens: Ottolengui, 1902, p. 66. Southwest United States, south to Tropics. [=mgationis of authors, nee Guenee] Comments.--Eichlin and Cunningham Autographa 00: McDunnough (nee Cramer), (1969) reported finding numerous larvae of this 1938, p. 133. species feeding on a narrow-leafed goldenrod. Pseudoplusia 00: McDunnough (nee Cramer), Though some pupae were found on this plant, 1944, pp. 206, 207. where two or three leaves had been held to­ Plusia inclttdens Walker, Forbes, 1954, p. 303. gether with silk, many more pupae were found Male genitalia (fig. 28).-Uncus long and slen­ attached to the underside of a broad-leafed der; vinculum extremely narrow, tapering to a species of ragweed (Iva annua L.) nearby in the point; aedeagus swollen in apical one-half, con­ same field upon which no larval feeding had stricted, then expanded again basad, base of taken place. In subsequent observations near aedeagus with patch of small cornuti; with the Tall Timbers Research Station, Tallahas­ cornutus near entrance of ejaculatory duct, and see, Fla., pupae were found on Chenopodi'um apex of aedeagus spinose; juxta narrow, pro­ ambrosiodes L. and on leaves of Acer negundo duced apically to a strong, sclerotized spine; L. (box elder) with no evidence of feeding dam­ valve with basal one-half narrow, expanding age. Apparently, for this species at least, many gradually in apical one-half to broadly rounded of the last-instar larvae leave their host to apex, costa near caudal edge of tegumen pro­ pupate on broad-leafed plants upon which they duced inward as an elongate spine, curved ce­ do not feed. phalad near distal end, clavus very long and relatively thick, ampulla extremely long and filiform, longer than one-half the length of Pseudoplusia ~lcDunnough valve, arising from sacculus near base of valve. Female genitalia (fig. 99).-Ostium bursae Generotype.-Phalaena 00 Cramer (fig. E, not shallow, but flared out laterad, sclerotized, stri­ PLUSIINAE OF AMERICA NORTH OF MEXICO 17 gate cephalad, and heavily shagreened; ductus Albany (culta). [?] Syntypes, British Museum bursae with initial one-half membranous, nar­ (Natural History) (dyaus). A male syntype of row at first but widening, cephalic one-half rogationis in the National Museum of Natural begins where ductus bursae abruptly narrows History has been selected and labeled and is to about one-third its width, then becoming presently designated as the lectotype: "Pl. roga­ somewhat sclerotized, strigate and widening tionis Gn., Spec. 116gJType rogationis, alc Hofer again to entrance into corpus bursae on right Austral/Ex. Musaeo, Ach. Guenee/Oberthur side; corpus bursae an elongate, shagreened, Collectionl& genitalia slide: USNM 51, J. G. membranous sac with an extremely elongate, Franclemont." [?] Rijksmuseum van Natuur­ narrow, tubelike apex, projecting from the right lijke Historie, Leiden, Netherlands (00). of ductus bursae entrance cephalad, following Type locality.--St. Domingo (includens). the side of the main body of ostium bursae Santarem and Venezuela (hamifera). West around fundus, and then caudad along left side Troy, N. Y. (culta). Texas, Jamaica (dyaus). almost to caudal end, where it terminates in a North America (mgationis). [?] (00). point, from which arises the ductus seminalis. Larvae.--Prolegs vestigial on abdominal Host plant.---Commelina sp.. Cmton capita­ segments 3 and 4, reduced to very small, peg­ tus, Lactuca sativa, Medicago sativa, Nicotiana like structures; no microspines on integument; tabaccum, Phaseolus sp., Crumb (1956); Glycine mesothorax with pinaculum of D-2 closer to D­ max, Gossypium hirsutum, Solidago sp., Eichlin 1 than to SD-2; seta SV-2 absent on meso- and and Cunningham (1969); Brassica oieracea, Eu­ metathorax; seta SV-2 absent on abdominal patorium sp., Geranium sp., Hibiscus esculen­ segment 1; crochets range in number from 22 to tus, Pela?'gonium sp., Zebrina pendula, Tietz 25; dark, lateral band if present on head not (1972); Chrysanthemum sp., Coleus sp., Lantana including ocelli 1-4 on all specimens examined; sp., Lycopersicon esculentum, Pe'rsea ameri­ cana. ribs 2 and 3 of mandible terminating in proc­ esses before reaching cutting margin; raduloid Range (map 4).-Quebec to West Indies; Flor­ on hypopharynx with 10 ridges. ida to California; south to South America. Type.--Syntypes not seen by us, British Comments.--This species is commonly Museum (Natural History) (includens). Syn­ called the soybean looper. Parental females of types not seen by us, British Museum (Natural laboratory cultures were collected from a light History) (hamifera). New York State Museum, trap at Auburn, Ala.

Tribe AUTOGRAPHINI, New Tribe

The Autographini is the largest tribe in clavus is long, filiform, and setose on the apex. North America and is composed of the following In Adeva and Pseudeva, only a setose bump 12 genera with 61 species: Autoplusia, Plusia, remains of the clavus and in Syngrapha the Allagrapha, Rachiplusia, Polychrysia, Pseu­ clavus is short and aris~R from a broad base. deva, Adeva, Chrysanympha, Eosphoropteryx, The larvae have no vestiges of prolegs on ab­ Autographa, Syngrapha, and Chrysaspidia. dominal segments 3 and 4. Most species have a The Autographini could be divided into two raduloid on the hypopharynx. The exceptions or three tribes based on adult characters, but are species comprising the genera Polychrysia has not been so treated because of the similari­ and Chrysaspidia, and a group of species we ties of the larval forms. The tibiae are unspined include in the genus Syngrapha. The larvae of in all genera with the following exceptions: In Chrysaspidia are unique in the subfamily in all but one species of' Syngrapha and Autoplu­ that they have uninordinal crochets. Based on sia, only the hind tibiae are spined. Syngrapha studies of larval forms, several species appear ignea, however, and the genus Rachiplu.sia transitional between the two largest genera, have all tibiae spined. In most genera, the male Autogmpha and Syngrapha. The basis for the 18 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE interpretation of the relationships and defini­ ward as a long, slender process and ventral tions of these genera is presented in more detail margin produced inward as a longer sclerotized, in the section on "Phylogeny." sharply pointed spine; sacculus greatly ex­ panded and extended caudad on ventral margin A utoplllsia McDunnough to an elongate point, the points of both valves coming close together mesad; aedeagus longer Generotype--Pl1lsia egena Guenee, 1852, p. 328. than valve, basal one-half with many small Original designation. cornuti, a long, thick, apical cornutus, and apex of aedeagus spinose. A1doplusia McDunnough, 1944, p. 203. This is essentially a Neotropical genus con­ Female genitalia (fig. I07).-Ostium bursae taining three species, one of which, A. olivacea broad and about twice as long as wide, evenly (Skinner), may be endemic to the southern sclerotized ventrally and entirely membranous coastal areas of California. 'The other two spe­ dorsally; ductus bursae just slightly wider than cies are clearly Tropical, extending into sub­ ostium bursae and slightly longer, dorsoven­ tropical and warmest, temperate regions in the trally flattened, evenly sclerotized throughout, Southern United States. entering corpus bursae ventrad; corpus bursae The genitalia are unique in the subfamily. elongate, narrow, S-shaped from dorsal aspect, The larvae are typical for species of Autogra­ with elongate, rounded apex, having rugose phini. strigae; ductus seminalis arises from apex of ostium bursae. KEY TO THE SPECIES BASED ON MALE GENITALIA Larvae.--No vestige of prolegs on abdomi­ 1. Sacculus produced caudally to a long, fine. ventral nal segments 3 and 4; microspines on integu­ point (fig. 39) ...... olivacea ment; all setal pinacula of head ringed with Sacculus not produced caudally to a long, fine, ventra! black; crochets range from 19 to 25 in number; point (fig. 38) ...... •...... 2 seta SV-2 absent on meso- and metathorax; 2. Valve with curved, inward projecting, spinelike proc­ seta SV-2 absent on abdominal segment 1; ess on ventral margin near apex (fig. 38) egena Valve without ventral process near apex (fig. 59) abdominal segment 2 with pinaculum of SV-1 ...... • ...... ill1t.~t1·ata approximately equidistant from SV-2 and V-1; abdominal segment 4 with V-1 pinacula as close KEY TO THE SPEClES BASED ON FEMALE to each other as V-1 is to SV-2, or slightly GE:-I ITA LIA closer to SV-2 than to each other; on meso­ 1. Ductus seminalis arises from apex of ostium bursae thorax pinaculum of seta D-2 equidistant from 2 D-1 and SD-2; rib 2 and often rib 3 of mandible Ductus seminalis arises from fundus of corpus bursae terminate in a process before reaching cutting (fig. 95) ...... ill1tstraia margin; raduloid on hypopharynx with 23 2. Apex of corpus bursae bent dorsad over ductus bursae (fig. 107) ...... olivacea ridges. Dorsal setae on protruding conical pina­ Apex of corpus bursae twisted ventrad of ductus cula. fun nel (fig. 109) ...... egell(t Type.--Academy of Natural Sciences, Phila­ delphia. Type locality.--Los Angeles, Calif. Autoplusia olivacea (Skinner) Host plant.--Heliotropium sp., Lantana sp., Mimulus cardinalis, Pha.<;eolus vulgaris, P. li­ FiGure 168 mensis, Senecio sp., S. jCLCobaea. Autogmpha olivacea Skinner, 1917, p. 329. Range (map 7).-Southern coastal California. AutopluBia olivacea: McDunnough, 1944, p. 203. Male genitalia (fig. 39).-Uncus slender, no Alltoplusia illustrata (Guenee) apical spine; tegumen slender; juxta narrow; Figure 169 elongate, clavuslike structures arise from sides of juxta; vinculum tapered to round apex; Plusia itlustrata Guenee, 1852, p. 328. valves narrow for caudal two-thirds, apex Syngraph{L illustmta: Hampson, 1913, p. 437. round, before apex dorsal margin produced in­ Autoplusia illu.stmt(L: IGmball, 1965, p. 119. PL USIINAE OF AMERICA NORTH OF MEXICO 19 Male genitalia (fig. 59).-Uncus slender with on ventral edge as a sharply pointed process; strong, apical spine; valve with dorsal and ven­ aedeagus more expanded basally, patch of tral margins parallel, slightly rounded apically, small cornuti more extensive, occupying basal clothed mesally with setaceous scales, extended two-thirds of aedeagus, and apex not spinose. from base ventrally, at apex of which are at­ Female genitalia (fig. l09).-Ventral posterior tached scale tufts; ampulla lacking; clavus elon­ margin of ostium bursae greatly elongated ven­ gate, slender, setose apically; apex of vinculum trad, ostium bursae broad, evenly sclerotized, short, narrowly rounded; juxta elongate, some­ and punctate, bending ventrally to where duc­ what keeled, but without apical spine; aedeagus tus bursae arises; ductus bursae as wide as thick, slightly expanded apically; vesica with ostium bursae and ribbonlike, sclerotized, and short, pointed, apical cornutus, short, stout, with strigations only on right side and on into pointed, medial cornutus, and an elongate row corpus bursae which it enters far down on the of approximately 25 slender cornuti medially. ventral side; apex of corpus bursae elongate, Female genitalia (fig. 95).-Ostium bursae flattened, about as wide as ductus bursae, small, cone shaped, strigate, weakly sclerotized, mostly membranous and strigate, bending to not clearly distinct from ductus bursae; ductus the left, twisting, and protruding ventrad of bursae mostly membranous, threadlike, enter­ ostium bursae; the ductus seminalis arises from ing corpus bursae dorsad on right, very near the truncate margin of the apex; corpus bursae apex; corpus bursae shagreened, with apex small, tapering cephalad and granulose. thickened internally, and with a heavily sclero­ Larvae.--No vestige of prolegs on abdomi­ tized, bilobed plate near entrance of ductus nal segments 3 and 4; minute microspines on b'lrsae; ductus seminalis arises from tip of fun­ integument; all setal pinacula of head ringed dus. with black; crochets range from 24 to 26 in Type.--Female, British Museum (Natural number; seta SV-2 absent on meso- and meta­ History). thorax; seta SV-2 absent on abdominal seg­ Type locality.--Raiti. ment 1; pinaculum of SV-l approximately equi­ Host plant.--Unknown. distant from SV-2 and V-Ion abdominal Range (map l).-Florida; West Indies; South segment 2; abdominal segment 4 with pinacu­ America. lum of V-I closer to SV-2 than setae V-I are to Comments.--The ductus seminalis, which each other; on mesothorax pinaculum of seta arises from the fundus of the corpus bursae, is D-2 equidistant from D-l and SD-2; rib 2 and, an unusual condition in the subfamily, occur­ occasionally, rib 3 of mandible terminates in a ring also in females of species of Argyro­ process before reaching cutting margin; radu­ gramma. Kimball (1965) placed illw;tmtn in Au­ loid on hypopharynx with 12 ridges. toplu,,<;in tentatively. The male structures tend to support this placement. Type.--A male syntype of egena in the Na­ tional Museum of Natural History has been selected and labeled and is presently desig­ A utoplusia egena (Cuenee) nated as the lectotype: "Type, P. egena Gn., alc Hofer/egena, Bresil/Ex Musaeo, Ach. Guenee/ Figure 170 Oberthur CollectiOn/Barnes Collection." Plusia egena Guenee, 1852, p. 328. Type locality.--Brazil. Autographa egena: Dyar, 1902, p. 81. Host plant.--Agapanthus sp., Althaea ro­ Syngrapita egena: Hampson, 1913, pp. 435, 436. sea, Apium gmveolens, Bmssica olemcea vari­ eties, Chrys(Lnthemum sp., C. fndescens, D(l'/(­ Autogmpita egena: McDunnough, 1916, p. 400. CllS camta, Delphinium sp., Glycine ma;r, McLiva Autoplu.sia egena: McDunnough, 1944, p. 203. sp., Mentha sp., M. spicata, Phaseol1J~S sp., P. Male genitalia (fig. :J8).-Similar to olh}(lcea limensis, P. VUlgaris, Senecio sp., Symphytum with the following exceptions: Valve without sp., Tagetes e?·ecta. dorsal margin produced inward as slender proc­ Range (map 3).-Florida, New Mexico, and ess before apex, simply humped; sacculus California; south to Brazil. though greatly expanded, not produced caudad Comments.--This species is called the bean 20 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE leaf skeletonizer. Larvae examined were bor­ about one-third length of valve; aedeagus rowed from laboratory cultures maintained at rather thick throughout, slightly expanded at the University of California, Riverside. base, with three groups of small cornuti; one group basad with about three cornuti, two groups mesad with three to six cornuti in each Rachiplusia Hampson group. Female genitalia (fig. 97).-Ostium bursae Generotype.-Plusia n'/.l Guenee, 1852, p. 347. broad, dorsoventrally compressed, sclerotized, Original designation. with longitudinal grooves dorsally and ven­ RachipluS'ia Hampson, 1913, p. 410. trally, bilobed caudoventrad, tapered slightly to Autographa: McDunnough, 1916, p. 400. base; initial short section of ductus bursae Rachiplusia: McDunnough, 1944, p. 207. membranous, narrow, leading to a short, well­ This is another basically Neotropical genus. defined sclerotized section, becoming membra­ The generotype occurs in South America, nous until it widens and becomes strigate and whereas R. ou (Guenee) occurs from Venezuela partially sclerotized, entering the corpus bur­ northward and has successfully invaded all but sae on the right side; corpus bursae constricted the northernmost areas of the United States. cephalad, producing a small, well-defined fun­ Its success in North America is due mainly to dus; cephalic one-half shagreened and membra­ its ability to feed on several common, wide­ nous, caudal one-half strigate and partially spread host plants. sclerotized with apical region bent to the right The presence of spines on all tibiae character­ dorsad of ductus bursae; ductas seminalis izes this genus. On the larvae, the setal pina­ arises from sclerotized bulge on cephalic side of cula of SV-1 and SV-2 are separated, which is a apical region ventrad. character state different from that of Autogra­ Larvae.--No vestige of prolegs present on pha species. abdominal segments 3 and 4; integument with dark microspines particularly on venter and Rachiplusia ou (Guenee) around base of pinaculum of seta SD-2, the amount of spination is highly variable; crochets range from 20 to 22 in number; seta SV-2 Figure 171 absent on meso- and metathorax; seta SV-2 Plw;ia ou Guenee, 1852, p. 348. absent on abdominal segment 1; pinacula of Plusia fratella Grote, 1875a, p. 161.-Morrison, seta SV-l and SV-2 separated on abdominal 1875, p. 219. segment 2; pinacula of the two V-I setae about Plusia ou val'. russea Edwards, 1886, p. 170. the same distance from each other as V-I is PluS'ia ou val'. californica Speyer, 1875, p. 164; from SV-2 on abdominal segment 4; dark lat­ Smith, 1893, p. 252. eral band on head if present, not including Autographa ou: Dyar, 1902, p. 81. ocelli 1-4 on all specimens examined; ribs of mandibles without processes, continuing to cut­ Autographa ou: Dyar, 1903, p .. 200. [=race of ting margin; raduloid on hypopharynx with 10 gamma] ridges. Phytometra ou: Hampson, 1913, pp. 542, 543. Autographa ou: McDunnough, 1916, p. 400. Type.-.-Syntypes, not seen by us, British Museum (Natural History) (ou). British Mu­ Autographa pedalis (Grote), 1875c, p. 204.- seum (Natural History) (pedalis). Syntypes, not Barnes and McDunnough, 1917, p. 84. seen by us, British Museum (Natural History) Rachiplusia ou: McDunnough, 1944, p. 207. (fratella). Male genitalia (fig. 26).-Vinculum of even Type locality.--Boreal America (ou). Kansas width throughout, truncate at apex; juxta pro­ (pedalis). Texas (fratella). duced to a point at apex; valve narrow at base, expanding gradually to truncate apex, with no Host plant.--Chenopodium ambrosioides, ampulla; costa and sacculus well defined from Cosmos sp., Nicotiana tabaccum, Trifolium sp., base to caudal margin; clavus elongate, filiform, Crumb (1956); Mentha sp., Tietz (1972); Triticum PLUSIINAE OF AMERICA NORTH OF MEXICO 21 aestivum, T7'Opaeolum sp., Eichlin and Cun­ Lanrae.--No vestige of prolegs on abdomi­ ningham (1969). nal segments 3 and 4; integument without mi­ R~nge (map 3).-United States except north­ crospines; crochets range from 18 to 20 in num­ ernmost States. ber; seta SV-2 absent on meso- and Comments.--Parental fema.les of laboratory metathorax; seta SV-2 absent on abdominal cultures were collected from a light trap at segment 1; abdominal segment 2 with pinacula Auburn, Ala., and, in addition, numerous speci­ of setae SV-l, SV-2, and V-I approximately mens were examined from field collections. equidistant from each other, widely separated; abdominal segment 4 l,vith pinacula of two V-I Plusia Ochsenheimer setae about one-half the distance from each other as V-I is from SV-2; on mesothorax Generoiype.-Noetua ehrysitis Linnaeus, 1758. pinaculum of D-2 closer to D-l than to SD-2; Designated by Duponchel, 1829, p. 71. ribs 2 and 3 of mandible terminating in proc., Plv,sia Ochsenheimer, 1816, p. 89. esses before reaching cutting margin; raduloid Agmpha Hubner, [1821], p. 250 (partim). absent on hypopharynx. Plusia: Dyar, 1902, p. 81. Type.--British Museum (Natural History). Type localitY.--Georgia. Phytometm: Hampson (nee Haworth), 1913, p. 452. Host plant.--Humul'us lupulus, Tietz (1972); Populus tremuloides. Plusia: McDunnough, 1916, p. 400. Range (map 7}.-Nova Scotia to Manitoba; One species represents this genus in North south to Pennsylvania and Minnesota (no speci­ America. There is no stigma or other silvering mens seen from Southeastern United States). on the forewing as with most plusiinl~s. Plusia is a genus of northern temperate regions, and is essentially a Palearctic genus. Allagrapha Franclemont

Generotype.-Noetua aerea Hubner, [1800-1803], Plusia balluca (Geyer) pI. 56, fig. 27. Original designation. Figure 172 _4llagmpha Franclemont, 1964, p. 73. Dyaehrysia balluca Geyer, 1832, p. 2, figs. 681, Plusia: Ichinose, 1973, p. 139, (Agmpha Hub.). 682. Like the North American species of Plusia, the species of Allagmpha lack the metallic sil­ Plusia balluca: Guenee, 1852, p. 344. ver or gold signs on the forewings. The male Phytometra balluca: Halnpson, 913, p. '577. genitalia characteristically have a highly modi­ Plusia balluca: McDunnough, 1916, p. 400. fied sacculus "vith a heavily sclerotized and Male genitalia (fig. 29).-Uncus long and specialized apex. The larvae are basically like thick; tegumen wide; vinculum elongate, nar­ those of Plusia, but have a hypopharyngial rowed apically to a point; juxta with large, raduloid, whereas the latter does not. sclerotized projection ventrad; valve broad with McDunnough (1944) and Franclemont (1964) rounded apex, clavus filiform, ampulla nar­ agreed thatA. aerea (Hub.) differed significantly rowed abruptly at distal one-fourth, extended in characters of the male genitalia from the to costal margin; aedeagus with basal cornutus, generotype of Plusia, P. chrysitis L., to remove and median cluster of a few small cornuti. the former from Plusia. Recently, Crumb (1956), Female genitalia (fig. 94).-Lateral invagina­ Kostrowicki (1971), and Ichinose (1973) regarded tions of abdominal segment 8 forming deep Agrapha (=Allagrapha) as a synonym of Plusia pockets; ductus bursae thick, sclerotized, stri­ on the basis that the splitting of the genus gate, with a spiculate sac on right side on tended to obscure the relationships among the cephalic one-half, ductus bursae entering bursa species of this group. Ichinose argues that the copulatrix on right side; corpus bursae elon­ differences of the male genitalia are not signifi­ gate, membranous, spiculate at apex; ductus cant enough in themselves and cites the simi­ seminalis arises from apex of corpus bursae. larities in female genitalia and larval morphol­ 22 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE ogy as reasons for not splitting aerea from loop projecting dorsad before entering corpus Plusia. bursae on right side; corpus bursae membra­ Undoubtedly, the species involved are all nous, cephalic one-half saclike, bent to the left, closely related; however, A. aerea and A. caudal one-half more narrow with apex ventrad aeroides are similar to each other in adult and of ductus bursae; ductus seminalis arises from larval features, with a strong superficial ap­ apex of corpus bursae. pearance which cannot be ignored. In addition, Larvae.--N0 vestige of prolegs on abdomi­ the two differ from sp.ecies of Plusia in the nal segments 3 and 4; integument without mi­ modifications of the male genitalia and the crospines; crochets range from 17 ·to 20 in num­ presence of a raduloid on the larvae of Allagra­ber; seta SV-2 absent on meso- and pha, a raduloid being absent at least on the metathoraY; seta SV-2 absent on abdominal Nearctic Plusia species. For these reasons, the segment 1; abdominal segment 2 with pinacu­ authors feel it is useful to separate aerea and lum of seta SV-l approximately equidistant aeroidl:'s from Plusia to demonstrate better from SV-2 and V-I, widely separated; abdomi­ their dose relationship to each other and dis­ nal segment 4 with pinacula of two V-I setae tinctivenp3s from other Plusiinae. There are about one-half the distance from each other as apparently no comparable Palearctic species. V-I is from SV-2; on mesothorax pinaculum of D-2 closer to D-l than to SD-2; ribs 2 and 3 of KEY TO THE SPECIES BASED ON MALE GENITALIA mandible terminating in processes before Aedeagus with sclerotized apical plate; saccus longer reaching cutting margin; raduloid on hypophar­ than one-half length of valve (fig. 35) ..... aerea ynx with 12 ridges. Aedeagus without sclerotized apical platej saccus Type.--[?] British Museum (Natural His­ shorter than one-half length of valve (fig. 33) tory)...... aeroides Type locality.--Massachusetts, New York, KEY TO THE SPECIES BASED'ON FEMALE Pennsylvania. GENITALIA Host plant,--Mentha arvensis, Crumb (1956); Spiraea salicifolia, Tietz (1972); Aster sp., Ostium bursae large, mostly sclerotizedj corpus bursae with elongate apical region (fig. 104) ... aeroides Forbes (1954). Ostium bursae poorly defined, mostly membranousj Range (map 7).-Nova Scotia to British Col­ corpus bursae saclike, lacking definite apical re­ umbia; south to Pennsylvania in the east and gion (fig. 98) ...... aerea the Colorado and California in the west. Comments.--Larvae of this species were Allag'rapha aeroides (Grote), new combination reared on artificial medium. The parental fe­ males were collected in Groveton, N.H., at a Figure 173 black light. Plusia aeroides Grote, 1864, p. 83. Allagrapha aerea (Hubner) Phytometra aeroides: Hampson, 1913, p. 572. Plusia aeroides: McDunnough, 1916, p. 400. Male genitalia (fig. 33).-Vinculum pointed; Figure 174 valve approximately of same width throughout, Noctua aerea Hubner, [1800-1803], pI. 56, fig. 27. though slightly narrowed at round apex; clavus Agrapha aerea: Hubner, 1821, p. 251. filiform; ampulla broad, heavily sclerotized, Plusia aerea: Boisduval, 1829, p. 59. bladelike, extending dorsocaudad to costal mar­ Phytometra aerea: Hampson, 1913, p. 575. gin; aedeagus without cornuti, spiculate at Autographa aerea: McDunnough, 1916, p. 400. apex. Female genitalia (fig. 104).-Ostium bursae Agrapha aerea: McDunnough, 1944, p. 216. large, round, dorsoventrally flattened, sclero­ Allagrapha aerea: Franclemont, 1964, p. 73. tized ventrally and laterally, infolded subdor­ Male genitalia (fig. 35).-Uncus thin; tegumen sally, membranous dorsad; ductus bursae less very short; juxta small, somewhat circular with than one-half as wide as ostium bursae, entirely a thick, stubby spine ventrad; vinculum long, membranous, somewhat strigate, forming a tapered to a blunt point; valves highly modified, PLUSIINAE OF AMERICA NORTH OF MEXICO 23 sacculus heavily sclerotized, expanded on dor­ Pseudeva Hampson, but further verification is sal edge into a rounded plate, and extending needed through a comparison of the imma­ ventrocaudad, forming a sharp, upcurved pro­ tures, which are unknown or not available at jection near middle of valve; ampulla wide, this time. large, heavily sclerotized, curved caudad, with projection cephalad of midpoint; clavus ex­ KEY TO THE SPECIES BASED ON MALE GENITALIA panded at apex, setose; aedeagus with a sclero­ tized apical plate, usually having three strong Valves oval, broadly rounded at apices; small pointed process on valve near base of ampulla (fig. 37a) spines on its ventral edge and. one spine on ...... 7nOrigera caudal edge. Valves not oval, tapering somewhat toward apices; no Female genitalia (fig. 98).-Ostium bursae process on valve near base of ampUlla (fig. 36a) mostly membranous, poorly defined; ductus ...... 7nOneta bursae short, U-shaped from lateral aspect, par­ tially sclerotized, strigate where it enters apex KEY TO THE SPECIES BASED ON FEMALE GENITALIA of corpus bursae on left side very near origin of ductus seminalis; corpus bursae membranous, Corpus bursae saclike, apex broadly rounded (fig. 108) ...... morigera saclike with no well defined fundus or apical Corpus bursae narrow, tapering at both ends, some­ region. what S-shaped from dorsal aspect (fig. 110) Larvae.--Very similar to A. aeroides, see ...... 7nOneta following description of that species; crochets range from 19 to 21 in number; abdominal Polychrysia moneta (Fabricius) segment 2 with pinaculum of seta SV-1 closer to SV-2 than to V-1; and raduloid on hypophar­ Figure 175 ynx with 18 ridges. Type.---[?]. Noctua moneta Fabricius, 1787, p. 162. Type locality.-[?]. Phalaena-Noctua moneta: Gmelin, 1789, p. 2556. Host plant.--General feeder on low plants, Polychrysia moneta: Hubner, 1821, p. 251. Forbes (1954); Urtica sp., Crumb (1956); Aster Plusia moneta: Treitschke, 1823, p. 158. umbeUatus, Glycine max. Chrysoptera moneta: Latreille, 1825, p. 476. Range (map 2).-Nova Scotia to Alberta; Polychrysia trabea (Smith), 1895, p. 340, p. 15, south to New Mexico and Gulf of Mexico. fig. 16,-Dyar, J.903, p. 198. Comments.--Parental females of laboratory Polychrysia moneta: Dyar, 1902, p. 80. cultures were collected from a light trap at Chrysoptera moneta: Hampson, 1913, pp. 444, Auburn, Ala. Other material was reared from 445. adults collected near Pittsburgh, Pa. Polychrysia moneta: McDunnough, 1944, p. 212. Male genitalia (fig. 36).-Very similar to the following species, morigera, but with valve Hiibner Polychrysia slightly tapered toward apex, ampulla longer, arising more ventrad, and clavus more elon­ Generotype.-Noctua moneta Fabricius, 1787, p. gate; apex of aedeagus not spinose in those 162. Monobasic. specimens examined. Polychrysia Hubner, 1821, p. 25l. FemaJe genitalia (fig. llO).-Ostium bursae Chrysoptera Latreille, 1825, p. 476. short, broad, round, mostly membranous; duc­ Deva: Smith, 1893, p. 246, (partim). tus bursae strigate, sclerotized, expanded grad­ Polychrysia: Dyar, 1902, p. 80. ually to entrance into corpus bursae on left Chrysoptera: Hampson, 1913, p. 439. side; corpus bursae elongate, tapered at both ends and somewhat S-shaped from dorsal as­ Polychrysia: McDunnough, 1944, p. 212. pect; ductus seminalis arises from apex of cor­ Polychrysia is a Holarctic group with two pus bursae. species in the Nearctic region. This genus is Type.-Lost (Zimsen, 1964) (moneta). Male, apparently related to Adeva McDunnough and National Museum of Natural History (trabea). 24 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE Type locality.-Austrian mountains (rrwneta). Host plant.-Delphinium sp. Calgary, Alberta (trabea). Range (map 7).-Oregon, Wyoming, Montana, Host plant.-Delphinium sp., Crumb (1956). In California, and Colorado. Europe, it feeds on many species of Aconitum, Comments.-Larvae and associated parental Cucumis, Delphinium, He lianthis, and Trollius female were borrowed from the McFarland Col­ (Kostrowicki,1961). lection at the Los Angeles County Museum of Range (map 14).--Ontario to Alberta; north to Natural History, Calif. Adult specimens agreed Alaska; Europe. with type examined in the American Museum of Natural History. Polychrysia morigera (Edwards) Adeva McDunnough Figure 176 Generotype.-Autographa albavitta Ottolengui, Deva morigera Edwards, 1886, p. 169. 1902, p. 75. Original designation. Adeva Mc­ Panchrysia morigera: Dyar, 1902, p. 81. Dunnough, 1944, p. 213. Chrysoptera morigera: Hampson, 1913, pp. 445, Adeva is a monobasic genus closely allied to 446. Pseudeva, based on genitalic structures. The Polychrysia morigera: McDunnough, 1944, p. most obvious feature which the two genera 212. have in common is the very reduced clavus of Male genitalia (fig. 37).-Overall size small, the males. The larvae of both groups are as yet similar to Adeva albavitta; valve oblong, unknown. The only species in the genus occurs rounded at apex; clavus more developed than in in the high deserts and mountains of south­ albavitta; ampulla slender, elongate, projecting western United States. Its biology is unknown. just beyond costal margin; no point on juxta; aedeagus with apex spinose, and with a slender Adeva albavitta (Ottolengui) apical cornutu8. Female genitalia (fig. 108).-Ostium bursae Figure 177 mostly membranous; ductus bursae with initial Autographa albavitta Ottolengui, 1902, p. 75. one-third membranous, expanded dorsad ab­ ruptly to three times its initial width, sclero­ Phytometra hutsonii (Smith), 1904, p. 60- tized, strigate, doubling width on final one­ Hampson, 1913, p. 570. third, entering corpus bursae on the left side Autographa albavitta: McDunnough, 1916, p. very near apex of corpus bursae; corpus bursae 400. broad at apex and with a bluntly pointed fun­ Adeva albavitta: McDunnough, 1944, pp. 213, dus. 214. Larvae.-No vestige of prolegs on abdominal hlale genitalia (fig. 32).-Overall siZe small, segments 3 and 4; minute microspines on inte­ compact; uncus thick with apical spine; tegu­ gument; all pinacula of setae on head ringed men short and wide; juxta produced into a long with black; crochets range from 21 to 23 in point at apex; valves short, oblong; ampulla number; seta SV-2 absent on meso- and meta­ arises approximately in center of valve, elon­ thorax; pinacula of setae SV-1 and SV-2 at f;(,ate, slender, extending just beyond costal mar­ least partially fused on abdominal segment 2; gin; clavus reduced to a setose knob; vinculum abdominal segment 4 with pinacula on setae V­ short, tapered to a point apically; aedeagus I closer than to SV-2; on mesothorax pinacu­ J.onger than valve, with short basal cornutus, lum of seta D-2 equidistant from D-1 and SD-2; with a circular base. only rib 2 of mandible terminating in a process F'emale genitalia (fig. l06).-Ostium bursae before reaching cutting margin; raduloid of hy­ undifferentiated from ductus bursae, which is popharynx absent. membranous and very narrow on caudal one­ Type.-Female, American Museum of Natu­ half, becoming lightly sclerotized and strigate, ral History. and widening to about four times its initial Type locality.-Colorado. width as it enters the corpus bursae on the PLUSIINAE OF AMERICA NORTH OF MEXICO 25

right side; corpus bursae somewhat laterally Very similar to above species, but apparently without compressed, at least when spermatophore is the additional lobe caudally on left side of corpus present, broadest just caudad of middle, ce­ bursae (fig. 101) ...... pUl-PlO-igera phalic one-half narrower and elongate; ductus seminalis arises from apex of corpus bursae. Pseudeva palligera (Grote) Type.-Female, National Museum of Natural History (albamtta). Lectotype male, American Figure 178 Museum of Natural History (hutsonii) desig­ Deva palligem Grote, 1881, p. 35. nated by Todd (1972). Panch1ysia palligem: Dyar, 1902, p. 80. Type.-Middle California (albavitta). Yuma Co., Ariz. (hutsonii). Pseudeva palligem: Hampson, 1913, p. 449. Host plant.-Unknown. Pseudeva Il.lbigem Hampson, 1913, p. 449 (new Range (map 3).-Oregon, California, Arizona, synonymy). and Nevada. Male genitalia (fig. 3l).-Overall size small, generally like males of Polych?ysia spp., vincu~ lum short and pointed; juxta simple, not Pseudeva Hampson pointed or spined; valves short, wide and rounded apically, with ampulla arising approxi­ Generotype.-Deva pl.l:rpurigem Walker, 1858, p. mately in center of valve midway from base to 1791. Original designation. Pseudeva Hampson, apex, and extending a little beyond costal mar­ 1913, p. 447. gin; clavus reduced to a mere setose knob' This genus is apparently restricted to tem­ aedeagus with median, small cornutus, and ~ perate North America. The adult morphology curved, sclerotized, spinose plate at apex. suggests a close relationship to Adeva and prob­ Female genitalia (fig. lOO).-Ostium protected ably Polychrysia. These three genera all have by folds of tissue from abdominal segment 8; relatively small and compact male genitalia ostium bursae membranous; ductus bursae similar in general form. In Polychrysia, the granulose, strigate, entering corpus bursae on clavus is short, but is reduced to a small swell­ the left side; corpus bursae granulose, with ing in Adeva and Polychrysia. The female geni­ pointed lobe caudad on left side on all speci­ talia differ sufficiently to distinguish the spe­ mens examined; ductus seminalis arises from cies, but also demonstrate the relative closeness granulose apex. of the above three genera in general features. Type.-A male syntype of palligera in the Once associated with their adult forms, the American Museum of Natural History has been larvae of species in the three genera when selected and labeled and is presently desig­ compared will probably verify the proposed re­ nated as the lectotype: "Deva palligera Gr., lationships. Type/35 11 , Sierra Nev., CalJNo. 11226, Collec­ tion Hy. Edwards." Male, British Museum (Nat­ KEY TO THE SPECIES BASED O:-..r MALE GENITALIA ural History) (nlbigeru). Aedeagus with a curved, sclerotized plate apically, to Type locaIity.-Sierra Nevada, Calif. (palli­ which are attached several small spines (fig. 31) gem). California (1I.lbigem)...... '" ...... palligera Host plant.-Unknown. Aedeagus with 20 to 25 stout, sharp, scIerotized spines Range (map 8).-California, Montana, Idaho, apically, joined together at their bases forming a and Colorado. row, longer than the spines of the above species, the longest being about one-third the width of Comments.-We could find no difference be­ aedeagus (fig. 27) ...... purpurigem tween palligem and mbigera in the genitalia and consider any small differences in wing KEY TO THE SPECIES BASED ON FEMALE pattern to be in the expected range of species GENITALIA variation. On specimens examined, corpus bursae slightly differ­ ent in shape from the other species, having not Pseudeva purpurigera (Walker) only a well-defined apical region, but an additional lobe to the left of apex (fig. 100) ...... palligem Figure 179 26 TECHN"ICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE Deva p1l?-p'u7-igera Walker, 1858, p. 1791. Chrysanympha formosa: McDunnough, 1938, p. Plus':-: purpm-igem: Grote, 1874a, p. 30. 114. Male genitalia (fig. 40).-Overall size small, Deva pW7J1H-igera: Grote, 1881, p. 35. uncus long and slender, juxta narrow, tapered Paneh1ysia purpul'igem: Dyar, 1902, p. 80. to a down-curved point; valve wide, slightly Pseudeva PW']J'l.u-igera: Hampson,. 1913, pp. 448, rounded apically, ampulla slender, just reach­ 449. ing costal margin, clavus slender and about as Male genitalia (fig. 27).-Very similar to palli­ long as one-half the median width of valve; gem except aedeagus with a patch of many vinculum narrowed apically to a blunt point; small cornuti apically. aedeagus with club-shaped, basal cornutus. Female genitalia (fig. lOl).-Ostium protected Female genitalia (fig. l11).-Ostium bursae above and below by folds of punctate, mostly short, broad, sclerotized and strigate; ductus membranous tissue of abdominal segment 8; bursae short, straight, initially membranous, ostium bursae entirely membranous; ductus then strigate, entering corpus bursae into an bursae granulose throughout, only sclerotized elongate, granulate pouch on left side; corpus initially, then broadening and membranous, el­ bursae elongate, narrow, fundus recurved to bowed to the left before entering corpus bursae the left, projecting caudad; ductus seminalis on the left side; corpus bursae granulose with arises from narrow apex. ductus seminalis arising from apex. Type.-[?] Museum of Comparative Zoology, Type.-British Museum (Natural History). Harvard University. Type locality.-Orilla [Ontario], Canada. Type locality.-Lawrence, Mass. Host plant.-Thalietrum sp., Tietz (1972). Host plant.--Gaylussaeia dumosa, V(leeinium Range (map 8).-Nova Scotia to Manitoba; sp., Tietz (197:2). south to Delaware, west to Montana, then Range (map 4).-Nova Scotia to Manitoba; south to New Mexico. south to Michigan and North Carolina.

Chrysanympha Grote Eosphoropteryx Dyar Generotype.-Plus'ict thyatyroides Guenee, 1852, Generotype.-Leptina formosa Grote, 1865, p. p. 337. Monobasic. 323. Monobasic. Chrysanympha Grote, 1895a, p. 417. EosphoroptelYx Dyar, 1902, p. 80. As with the preceding genus, EosphoroptelYx Abrostola, Hampson (nee Haworth), 1913, p. 582, contains a unique species. Its placement in (pmtim). relation to the other species is highly tenuous. Chrysanympha: McDunnough, 1944, p. 211. Hopefully, an examination of the immatures Chrysanympha is represented by a single will eventually clarify the problem. This genus species. It is unique in its superficial appe.ar­ and Chrysanympha are native to the Nearctic ance. The male genitalia are generally hke fauna. species of Autogmpha. Its phylogenetic position cannot be accurately determined, pending ex­ (Guenee) amination of the larva. Eosphoropteryx thyatyroides

Figure 181 Chrysanympha formosa (Grote) Plwlia thyatyroides Guenee, 1852, p. 337. Eosphoroptye1Yx thyatYl'oides: Dyar, 1902, p. 80. Male genitalia (fig. 41).-Uncus long and slen­ Figure 180 der; tegumen short and broad; vaJv~ narrow at Leptina formolw Grote, 1865, p. 323. apex, very wide medially, bulging ventrad; am­ Phu;ia fOTlnosa: Morrison, 1875, p. 98. pu lla small, short; clavus long, slender; vincu­ Polychrysia lOllnos(L' Dyar, 1902, p. 79. lum tapered to a point; aedeagus without cor­ Abrostola formosa: Hampson, 1913, pp. 583, 584. nuti, vesica somewhat spiculate. PLUSIINAE OF AMERICA NORTH OF MEXICO 27 Female genitalia (fig. I03).-Ostium bursae certainty on the basis of genitalic characters twice as long as wide, rectangular, flattened examined in this study. caudad, with venter evenly sclerotized and dor­ The female genitalia are also relatively ho­ sum membranous, ventrocaudal edge turned mogenous, but enough features permit differen­ downward; ductus bursae about as long as tiation. In general, the ostium bursae is short ostium bursae and narrower, somewhat sclero­ with no caudal extensions, thus leaving the tized and weakly strigate, bending to the right, ostium unprotected. The ductus bursae is most and entering corpus bursae on left side beyond often a slender, heavily strigate tube, having a apex, corpus bursae oblong, with apex straight, short lateral process near the base of the os­ rugose, irregularly pigmented; ductus seminalis tium bursae in several species. In all but three arises from apex of corpus bursae. of the species, the ductus bursae enters the Syntypes.-Barnes Collection in the National corpus bursae on the right side. The corpus Museum of Natural History and the British Museum (Natural History). bursae is large and saclike, but varies in shape among species. Most species have a rounded, Type locality.-New York. semilunate protuberance or otherwise tough, Host plant.-Thalictrum sp., Forbes (1954). thickened area on the corpus bursae cephalad Range (map 4).-Nova Scotia to British Col­ of the entrance of the ductus bursae. The duc­ umbia; south to Virginia, Minnesota, and Ore­ gon. tus seminalis arises from the apex of the corpus bursae in most species. The larvae of the genus Autographa have no vestige of prolegs on abdominal segments 3 and Autographa Hubner 4. Pinacula of SV-l and SV-2 are partially fused on abdominal segments 2--4. SV-2 is absent on Generotype.-Noctua gamma Linnaeus, 1758, p. abdominal segment 1, and SV-2 is absent on 513. Designatec. by Grote, 1895b, p. 61. the meso- and metathorax on all species. The Autographa Hiibner, 1821, p. 251. two V-I setae on abdominal segment 4 are Phytometm Hampson (nee Haworth), 1913, p. usually closer to each other on most species 452, (partim). than V-I is to SV-2. The presence of a raduloid Autographa: McDunnough, 1916, p. 400. is a constant feature, with the number of The species of this genus can be separated for ridges, though stable in number for individual the most part by cursory examination of the species, ranging from 8 on the generotype, wing patterns and colors (refer to Forbes, 1954, gamma, to 18 on ampla. pp. 298-308, and photographs provided here). The genitalia by contrast, particularly of the KEY TO THE SPECIES BASED ON MALE GENITALIA males, are similar to each other. The male genitalia are characterized by hav­ 1. Length of tegumen greater than one-half of the length of the valve...... 2 ing long, filiform clavi, setose on the apex. The Length of tegumen one-half or less than one-half the tegumen is lengthened and the valves are long length of valve ...... 7 and broad with only a few exceptions. The 2. Aedeagus with cornutus ...... 3 sacculus of the valve is expanded and some­ Aedeagus without cornutus (fig. 45) ..••... biloba what keeled ventrally. The ampulla is generally 3. Length of aedeagus about six times the length of cornutus (fig. 52) ...... •...... v. alba slender and extends to the costal margin at Length of aedeagus about five times the length of least and beyond in most species. The vinculum comutus (fig. 46)...... •...... Irpeciosa is relatively long and bluntly pointed. The ae­ (fig', 48) ....•...•.•...... ••••••....• eorusea, deagus is long, slender for most of its length, (fig. 50) ...••.•••.•...... •..•.••.•.• Labrona and bulbous at the base, which has a single Length of aedeagus about four times length of cor­ nutus ...... , ...... 4 cornutus with an extended, flat, sclerotized, 4. Ratio of length of tegumen to length of valve about basal plate in most species. The juxta is typi­ 3:5 (fig. 55) ...... ample cally produced to a blunt point apically, but not Ratio of le\igth of tegumen to length of valve slightly into a spinelike process. Males of eoru,$ea, la­more than 1:2 (fig. 42) ...... 5 brosa, and speeiosa. cannot be separated with 5. Caudal end of valve narrowed and strongly upturned (fig. 43) ...... 6 28 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE Caudal end of valve rounded and slightly upturned Caudal one-half of corpus bursae not strigate except (fig. 42) ...... mappa. perhaps on right side near entrance of ductus 6. Clavus about as long as wi<1th of valve from base of bursae (fig. ll8) ...... corusca clavus to ventral margin (fig. 43). bimaculata 8. Ductus bursae simple, strigate, approximately same Clavus longer than width of valve from base of width throughout ...... 9 clavus to ventral margin (fig. 47) ... flagellum Cephalic one-half of ductus bursae at least twice as 7. Length of valve about three times the length of wide as caudal one-half (fig. 119) . .. flagellum tegumen (fig. 49) ...... precationis 9. Corpus bursae strongly recurved to the right, either Length of valve about two times the length of tegu­ at the middle (fig. 117) or apical region (fig. 122) men (fig. 56) .....•...... 8 ...... 10 8. Aedeagus with small patch of about 8 to 10 tiny Corpus bursae not recurved to the right ...... 11 cornuti apically and one large cornutus basally 10. Apex of corpus bursae recurved to the right; main (fig. 56) ..•...... •...... rubida portion of corpus bursae disk shaped (fig. 122) Aedeagus with one basal cornutus only ...... 9 ...... rubida 9. Length of aedeagus at most four times length of Corpus bursae recurved at the middle, U-shaped (fig. cornutus ...... 10 117) ...... sansoni Length of aedeagus at least five times length of 11. Corpus bursae with pigmented protuberance ceo cornutus ...... 11 phalad of entrance of ductus bursae (fig. 123) 10. Length of aedeagus about three times length of 12 cornutus; costal margin of valve slightly up­ Corpus bursae without a protuberance but with a turned (fig. 54) ...... sansoni much thickened region cephalad of ductus bur­ Length of aedeagus about four times length of cor­ sae entrance; ostium bursae longer than wide nutus; costal margin of valve straight (fig. 44) (fig. 120) ...... _.. mappa ...... _...... pasiphaeia 12. Ostium bursae about as long as wide in ventral 11. Length of aedeagus about five times length of cornu­ aspect (fig. 131) ...... v-alba tus (fig. 53) ....•...... californicn Ostium bursae small, not as long as wide in ventral Length of aedeagus about six times length of cornu­ aspect (fig. 123) ...... •. . .. pasiphaeia tus (fig. 51) ...... metallica 13. Caudal end of ductus bursae with lateral appendage Length of aedeagus about seven times length of (fig. 134) ...... _. _...... 14 cornutus (fig. 57b) ...... pseudognmma Caudal end of ductus bursae without lateral appen­ dage (fig. 132) ...... speciosa 14. Main portion of corpus bursae disk shaped; elongated apical region appressed to side of corpus bursae KEY TO THE SPECIES BASED 0:-.' FEMALE (fig. 134) ...... metallicn GE:-.'ITALIA Corpus bursae not shaped as above ...... 15 15. Du. tus bursae smoothly strigate for most of its length (fig. 116) ...... • pswdogantntn 1. Entrance of ductus bursae on left side ...... 2 Ductus bursae not smoothly strigate for most of its Entrance of ductus bursae on right side ...... 4 length, strigae rugosissimus (fig. 125) 2. Apex of corpus bursae strigate and sclerotized (fig...... • . • ...... • . . . • • birnacll/nta 112) ...... _...... Jlrecationis Apex of corpus bursae clear or merely granulose, or if weakly strigate, not sclerotized (fig. 115) . 3 A utographa precationis (Guenee) 3. Corpus bursae constricted in middle (fig. 113) ...•...._ ...... • ...... • . califonrica Corpus bursae not constricted in middle (fig. 115) ...... •...•...... biloba Figure 182 4. Saclike protuberance or much thickened area ce­ Plusia precationis Guenee, 1852, p. 344. phalad of ductus bursae entrance into corpus Autographa precationis: Dyar, 1902, p. 8l. bursae (fig. 114) ...... •...... 5 No saclike protuberance or thickened area on corpus Phytometm precationis: Hampson, 1913, pp. bursae ...... •...... 13 524,525. 5. Ductus bursae with a sclerotized, strigated appen­ Autogmpha preccLtionis: McDunnough, 1916, p. dage near base of ostium bursae (fig. 114) .. 6 No appendage on ductus bursae near base of ostium 400. bursae .. _...... 8 Male genitalia (fig. 49).-Valve nal'rOW, elon­ 6. Saclike protuberance on corpus bursae with tubular gate, of equal width for most of its length, apex structures internally (fig. 121) ...... 7 narrower than median width and rounded; am­ Saclike protuberance on corpus bursae without tubu­ pulla typical, but arising basad of center of lar structures internally (fig. 114) ..... ampla valve about one-third of the total length of 7. Caudal one-half of corpus bursae, including apex, strigate (fig. 121) ...•...... /abrosa valve from the base; aedeagus basally with PLUSIINAE OF AMERICA NORTH OF MEXICO 29 many minute spines on vesica; other structures Phytometm bimaculata: Hampson, 1913, pp. typical of the genus. 530, 531. Female genitalia (fig. 112).-Ostium bursae Autographa bimaculata: McDunnough, 1916, p. about as wide as it is long; ductus bursae 400. membranous for caudal one-third, then pig­ Male genitalia (fig. 43).-Valve with apex mented and sclerotized to entrance into corpus somewhat narrowed to a blunt point and up­ bursae, which is near the meson, dorsad but turned; otherwise typical of genus. Refer to slightly toward the left, where it becomes heav­ descriptions of the genus and key to species. ily strigate, corpus bursae long and narrow, Female genitalia (fig. 125).-Ostium bursae curved dorsad toward cephalic one-half; apex of about as wide as long, dorsoventrally flattened, corpus bursae complex, with a sclerotized and and weakly sclerotized; ductus bursae initially strigate lobe projecting dorsally on left side and membranous, rugosissimus strigae, with caudal a membranous bulge on the right side, with the projection to the right, and sinuous, making a ductus seminalis arising on that narrow, mem­ dorsal loop like an upside-down U before enter­ branous, caudal portion produced below and ing the corpus bursae on the right side; corpus slightly to the right of the ductus bursae. bursae without protuberance or thickened area, Lanoae.-Integument with at least a subdor­ strigate, membranous throughout, expanded sal band of microspines, which includes pina­ cephalad, somewhat laterally compressed and cula of setae SD-2 of abdominal segments; ribs bent both ventrad and laterad toward the of mandible terminating in processes, not ex­ right; ductus siminalis arises from the apex, tending to cutting margin; crochets range from which is narrowed and rounded. 21 to 23 in number; rabuloid of hypopharynx Larvae.-Integument without microspines; with 16 ridges; otherwise typical for genus. mandible with ribs 2 and 3 terminating in Type.-Syntypes not seen by us, British Mu­ processes,- not extending to cutting margin; seum (Natural History), National Museum of Natural History. head without black, lateral stripe; raduloid of hypopharynx with 15 ridges, otherwise typical Type localitY.-North America. for the genus. Host plant.-Althea rusea, Aureolaria pedicu­ Type.-[?] British Museum (Natural History) laria, Carduus sp., Helianthus sp., Lactuca sp., (bimaculata). Syntypes, Barnes Collection in the Pastinaca sativa, Phaseolus sp., Plantago sp., National Museum of Natural History and BI"it­ Taml.·acum ofticinale, Crumb (1956); Arctiu'I'n ish Museum (Natural History) Cu-brevis). lappa, Brassica olemcea, Chenopodium album, Type locality.-[?] (bimaculata). New York (u­ Cirsium sp., Ipomoea pur-purea, Tietz (1972); brevis). Verbena sp. Host plant.-TarcLxacum ojjicinnie, Tietz Range (map 5).-Nova Scotia to Ontario; (197;2). south to Georgia; west to Wisconsin, and Kan­ sas. Ran.ge (map 18).-Nova Scotia to British Col­ umbia; south to New Jersey in the east and to Comments.-Laboratory cultures were initi­ New Mexico in the west. ated from females captured in Auburn, Ala.; Comments.-Parental females of laboratory Lincoln, Me.; East Lansing, Mich.; and Sha­ wano, Wis. cultures were collected at black lights in Strat­ ford, Coos County, N. H., and Mare du Sault Camp, Laurentides Park, Quebec. Autographa bimaculata (Stephens) A utographa sansoni Dod Figure 183 Figure 184 Plusia bimaculatn Stephens, 1832, p. 102. Autogmpha sansoni Dod, 1910, p. 349. Plusia u-brevis Guenee, 1852, p. 341.-Walker Phytornetm sansoni: Hampson, 1913, pp. 546, 1857, p. 905. 547. Plusia bimacu[atn: Grote, 1874b, p. 72. Autographa sctnsoni: McDunnough, 1916, p. 400. Male genitalia (fig. 54).-Typical, refer to de­ Autogmpha bimaculnta: Dyar, 1902, p. 81. scription of the genus and the key to species. 30 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE Female genitalia (fig. 117).-Ostium bursae with contrasting light pinacula and light spira­ wide, only lightly sclerotized; ductus bursae cular stripe; legs black; crochets range from 20 typical and without appendage near base of to 24 in number; raduloid of hypopharynx with ostium bursae, entrance to corpus bursae on 17 ridges. the right side; corpus bursae elongate, bent at Type.-Female, American Museum of Natu­ middle, with apex and fundus coming close ral History. together to the right; a large, darkly pig­ Type locality.-Winnipeg, Manitoba. mented, protuberance dorsad on corpus bursae Hosi plant.-Unknown. near entrance of ductus bursae. Range (map 5).-Nova Scotia to Alberta; Type.-Male, National Museum of Natural south to Pennsylvania and west to Illinois. History. Comments.-The specimens studied agreed Type locality.-Banff, Alberta. with the type. The larvae examined were from Host plant.-Unknown. the National Museum of Natural History. Range (map 15).-Alaska, British Columbia, Alberta, Montana, Wyoming, Oregon. Comments.-Forbes (1954) mentioned an iso­ A utographa biloba (Stephens) lated record from Quebec. The specimens stud­ ied agreed with the type examined in the N a­ Figure 186 tional Museum of Natural History. Plusia biloba Stephens, 1832, p. 104. Autographa biloba: Dyar, 1902, p. 81. Autographa rubida Ottolengui Phytometra biloba: Hampson, 1913, pp. 512, 513. Autographa biloba: McDunnough, 1916, p. 400. Male genitalia (fig. 45).-Ampulla on valve Figure 185 short, not reaching costal margin; aedeagus Autogmpha rubida Ottolengui, 1902, p. 67. without a cornutus; otherwise typical of the Phytometm ntbida: Hampson, 1913, pp. 491, genus. 492. Female genitalia (fig. 115).-Ostium bursae Autogmpha rubida: McDunnough, 1916, p. 400. small, weakly strigate and pigmented, membra­ Male genitalia (fig. 56).-Valve elongate, nar­ nous basally as is first one-third of ductus rower than is typical of the genus and curves bursae; ductus bursae short, straight, strigate, gradually dorsad to the apex; aedeagus with granulose, and weakly sclerotized, entering the patch of small cornuti near apex on vesica; corpus bursae at the apex slightly to the left otherwise typical of the genus, refer to generic and definitely caudad of the origin of the ductus description. seminalis; corpus bursae narrow, tapering Female genitalia (fig. 122).-Ostium bursae slightly to the fundus and somewhat hook typical; ductus bursae membranous and unpig­ shaped from a left lateral aspect; fundus mem­ mented except for that portion at entrance to branous but strigate. corpus bursae, which is on the right side just Larvae.-Integument with relatively long, cephalad of the apex; the main body of the conspicuous, dark microspines; head with black corpus bursae is disk shaped, with the apical lateral stripe including all ocelli in those speci­ region narrowed abruptly and curved to the mens seen; ribs of mandible extend to the right; dorsad to the right and cephalad of the margin with no processes; raduloid on hypo­ ductus bursae entrance is a circular, thickened, pharynx with 13 ridges; crochets range from 20 darkly pigmented, rugose area. to 22 in number; otherwise typical for genus. Larvae.-Integument with microspines; ab­ Type.-British Museum (Natural History). dominal segment 1 with SV-2 absent; two V-·l Type locality.,-Unknown. setae on abdominal segment 4 as far apart from Host plant.-Brassica oleracea, Medico .'Iativa, each other as each V-I seta is from SV-2; head Nicotiana tabacc1tm, Crumb (1956); Trifolium black except for two median light stripes; rib 2 incamatum, Eichlin and Cunningham (1969); of mandible with a sharp, raised ridge; general Centaurea solHtitialis, Cirsium horridulu,m, He­ larval ground is green overlayed with brown lianthus sp., Lactuca .'Sativa, Tietz (1972); Del­ PLUSIINAE OF AIHERICA NORTH OF MEXICO 31 phinium sp., Geranium sp., Gladiolus sp., Hedra type. The second female syntype is from Cali­ sp., Helxine soleirdii, Hordeum Vltlga're, Mimu­ fornia. lus cardinalis, Musa paradisiaca, Phacelia sp., Type locality.-{?] (ca,lifornica). Colorado (rus­ Phaseolus mtlgaris, Salvia leucophylla, Stachys sea). ajugoides, Verbena sp. Host plant.-Ceanothus sp., Chenopodium al­ Range (map 18).-Nova Scotia to Manitoba; bum, Cit'rullus Vltlgaris, CUC1tmis melD, C. sati­ United States (except Northwest); Neotropics. VltS, CUC1trbita maxima, Dahlia variabilis, Dau­ Cornrnents.-Parental females of laboratory cus carota, Doronicum sp., Helianthus sp., flex cultures were collected in light trap at Auburn, opaca, Kalmia sp., Lactu.ca ca.nadensis, Linum Ala. sulcatum, Malus pumila, Mal'va sp., M. rotundi­ folia, Medicago sativa, Melilotus alba, Mentha Autographa cali/ornica (Speyer) sp., M. spicata, Pisum satiVltm, Plantago sp., Rhododend1"On sp., Ribes grossularia, R. sati­ Figure 187 Vltm, Rosa sp., Rubus idaeus, Rumex sp., Sam­ bucus canadensis, Trifolium sp., Vicia sativa, Plusia gamma abo californica Speyer, 1875, p. 164. Viguiera deltoidea, Zea mays, Tietz (1972); Al­ lium cepa, Amelanchier florida, Amsinckia Plusia ou var. californica: Smith, 1893, p. 252. dougla,siana, Apium graveolens, Atriplex sp., Autogmpha californica var. 1"USsea (Edwards), Beta Vltigaris, Brassica oleracea varieties; B. 1886, p. 70; Ottolengui, 1902, p. 65. [= var. of rapa, Calendula officina lis, Ca1thamus tincto­ ou] '!'"ius, Castilleja sp., Citrus sinensis, Et·echtites Plusia californica: Hampson, 1908, p. 105. arguta, Fragaria sp., Gossypium sp., Hordeum Phytomet'ra californica: Hampson, 1913, pp. 540, 'llUlgare, Lactuca sativa, Lup1:nus sp., Lycopersi­ 541. con esculentum, Nemophila menzeisii, Phaseo­ Autographa californica: McDunnough, 1916, p. lus sp., Podocarpus sp., Raphanus sativus, 400. Rheum sp., Senecio cinera1'ia, S. jacobaea, So­ Male genitalia (fig. 53).-Valve tapering from lanum tubemsum, Spinacea oleracea" Triticum aestiVltm, Vitis sp. the broad median section to a narrower, rounded apex; otherwise typical of the genus, Range (map 6).-Alberta and British Colum­ refer to the description of the genus and key to bia; Kansas, Nebraska; Western United States; species. Mexico. I"emale genitalia (fig. 113).-Ostium bursae Cornments.-This species is commonly termed typical of the genus; ductus bursae short, mem­ the "alfala looper." Specimens of larvae were branous initially, then strigate, sclerotized and borrowed from the laboratory culture at the bending to the left before entering the corpus University of California, Riverside. bursae on the left side; corpus bursae straight, constricted around the middle, where it is stri­ Autographa pseudogamma (Grote) gate, produced into a thickened hump to the left, caudad, and tapering strongly to a narrow, Figure 188 blunt apex from which arises the ductus semin­ aliso Plusia pseudogamma Grote, 1875c, p. 203. Larvae.-Integument with minute graules; Autographa pseudogamma: Dyar, 1902, p. 81. ribs of mandible extend to cutting margin; head Phytometra pseudogamma: Hampson, 1913, pp. mayor may not have. lateral: black stripe; 539,540. crochets range from 18 to 20 in/number; radu­ Autographa pseudogamma: McDunnough, 1916, loid of hypopharynx with 12 ridges; otherwise p.400. typical for the genus. Male genitalia (fig. 57).-Typical, refer to de­ Type.-{?] (californica). A female syntype of scription of genus and key to species. russea from Colorado in the American Museum Female genitalia (fig. 116).-Ostium bursae of Natural History has been selected and la­ typical; ductus bursae typical with appendage beled, and is presently designated as the lecto­ near base of ostium bursae present, entrance to 32 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE corpus bursae on right side; corpus bursae elon­ may be a function of the overall smaller size of gate, somewhat pointed cephalad, and with no the moth; valve with straight costal border; thickened area or protuberance cephalad of otherwise typical of the genus, refer to generic ductus bursae entrance. description. Type.-British Museum (Natural History). Female genitalia (fig. 123).-Ostium bursae Type locality.-Cape Breton, Nova Scotia. short, about one-half as long as wide from a Host plant.-Unknown. ventral aspect; ductus bursae typical with no Range {map 15).-Nova Scotia to British Col­ caudal appendage, entering the corpus bursae umbia, north to Alaska; south to Maine, Michi­ on the right side well beyond apex; corpus gan, South Dakota, Montana, Wyoming, Ari­ bursae broadly rounded cephalad, tapering zona; and California. gently to a rounded apex; a semilunate, pig­ mented protuberance mesad on corpus bursae A utographa v-alba Ottoiengui and cephalad of entrance of ductus bursae. Type.-A male syntype of pasiphaeia in the American Museum of Natural History has been Figure 189 selected, labeled and is presently designated as Autographa v-ulba Ottolengui, 1902, p. 73. the lectotype: "Plusia pasiphaeia GrJNo. 11301, Syngrapha v-ulba: Hampson, 1913, pp. 429, 430. Collection Hy. Edwardsl1521California." Autographa v-ulba: McDunnough, 1938, p. 113. Host plant.-Stachys ajugo'icies, S. rigida. Male genitalia (fig. 52).-Valve slightly up­ Range (map 17).-California and Oregon. turned apically, cornutus of aedeagus short; otherwise typical of genus. Female genitalia (fig. 13l).-Ostium bursae A utographa flagellum (Waiker) longer than wide; ductus bursae typical, and with no caudal projection, entering corpus bur­ Figure 191 sae on right side; corpus bursae saclike, ovoid Plu.siaflagellum Walker, 1857, p. 909. and broadly rounded at the apex, with a large, Plusia monodon Grote, 1875c, pp. 202, 203.­ semilunate, thickened and pigmented protuber­ Smith, 1893, p. 25l. ance on the corpus bursae just cephalad and Autographa flagellum: Dyar, 1902, p. 81. slightly to the right of the ductus bursae en­ Autographa insolita (Smith), 1895, p. 340, pI. 51, trance. fig. l'7.-0ttolengui, 1902. p. 67. Type.-Female, American Museum of Natu­ Phytometra flagellum: Hampson, 1913, pp. 541, ral History. Type !ocality.-Yellowstone Park, Wyo. 542. Host plant.-Unknown. Autogmpha flagellum: McDunnough, 1916, p. Range (map 5).-Alberta and British C;()lum­ 400. bia; Northwestern United States. Male genitalia (fig. 47).-Valve slightly nar­ Comments.-The specimens studied agreed rowed and upturned apically; basal cornutus of with the type examined in the American Mu­ aedeagus may be curved or straight, but pro­ portionate length of cornutus to length of ae­ seum of Natural History. deagus remains about the same (see key to A utographa pasiphaeia (Grote) species); otherwise typical of genus. Female genitalia (fig. 119).-Ostium bursae typical; ductus bursae membranous initially, l<"'igure 190 then strigate, expanding abruptly to twice its Plusia pasiphaeia Grote, 1873a, p. 146. width for remainder of cephalic one-half, enter­ Autographa pasiphaeiu: Dyar, 1902, p. 81. ing corpus bursae on the right side; proturber­ Phytometra pasiphaeia: Hampson, 1913, p. 546. ance on corpus bursae cephalad of ductus bur­ Autographu pasiphueia: McDunnough, 1916, p. sae entrance large, at its base about one-fourth 400. length of corpus bursae. Male genitalia (fig. 44).-Overall size of geni­ Type.-Syntypes, British Museum (Natural talia smaller than typical for genus, but this History) Syngrapha ignea and an undescribed species have plates of several small cornuti, and a few species have no Syngrapha Hubner armament at all on the vesica. The female genitalia are described under the Generotype.-Noctua deve?yens Hubner [1818]. species descriptions that follow. Designated by Grote, 1895b, p. 62. The similarities of the larvae of Syngrapha species to the species of Autograph{t are the Syngrapha Hubner, 1821, p. 250. fusion of the pinacula of SV-1 and SV-2 on Autographa: Hampson (nee Hubner), 1913, p. abdominal segments 2-4; no vestige of prolegs 404, (pa?tim). on abdominal segments 3 and 4; the presence of Caloplusia: Kostrowicki, 1961, p. 381, (padim). a raduloid in some species; and the absence of Syngrapha: Ichinose, 1962, p. 28. SV-2 on the meso- and metathorax in some Anagrapha McDunnough, 1944, p. 202, (new species. The larvae of Syngmpha species differ synonomy). by having seta SV-2 on abdominal segment 1. Kostrowicki (1961) uses S. intenugationis (L.) One group of species has seta SV-2 on the as the generotype of Syng?-a7)ha, because he meso- and metathorax, which is considered by considered S. devergens to belong to Caloplusia Crumb to be a unique situation for noctuid Smith based on the presence of spines on all larvae. The raduloid is missing in several spe­ tibiae of the latter species. He also places S. cies. On those species with a raduloid, where alticola (Walker) in Caloplusia. However, S. alti­ larvae were examined, the number of ridges cola lacks spines on all but the hind tibiae. ranges from 10 to 12. 36 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE

KEY TO THE SPECIES BASED ON MALE GENITALIA Cornutus either uncurved or short and stubby (rose­ thorn) (fig. 79b) ...... 21 18. Ampulla with minute apical spine (fig. 77a) or repre­ 1. Costa of valve with preapical spine ...... 2 sented by a broad ridge (fig. 71a) ...... 19 Costa without preapical spine ...... 4 Ampulla extending almost to costal margin (fig. 64a) 2. Ampulla long, thin, and straight, extending beyond ...... abstrusa costal margin; one cornutus about two-thirds as 19. Ampulla merely a broad ridge truncate apically (fig. longs as aedeagus (fig. 58) ...... epigaea 71a) ...... cryptica Ampulla broadly triangula:-; cornutus not as above Ampulla short and tuberc1elike pointed apically 20 ...... 3 20. Base of sacculus greatly produced mesad and bluntly 3. Valve abruptly narrowed to a blunt point from triangular (fig. 78a) ...... alias preapical spine to apex (fig. 62a); aedeagus with Base of sacculus not produced mesad to form a blunt a patch of several small cornuti (fig. 62b) triangle (fig. 77a) ...... rectangula ...... (unnamed species) 21. Cornutus short and stubby (rose-thorn) (fig. 79b) 22 Apex of valve broadly rounded beyond preapical Cornutus long and straight ...... 23 spine (fig. 66a); aedeagus with one large apical 22. Base of costa with a sharp spine (fig. 79a) cornutus (fig. 66b) ...... octoscripta ...... interrogationis 4. Costa produced apically into sharp spine (fig. 72a) 5 Base of costa without spine (fig. 80a) ... u-aureum Costa without apical spine; apex merely sharply 23. Ampulla terminated by a bifid spine (fig. 74a) angled or rounded ...... 8 ...... surena 5. Aedeagus with a small basal and a curved median Ampulla not terminated by a bifid spine ...... 24 cornutus (fig. 69b) ...... viridisigma 24. Ampulla with a truncate apex and small terminal Aedeagus with only one cornutus ...... 6 spine ...... borea 6. Aedeagus with basal cornutus; ampulla with Ampulla tapering to a sharp pvint ...... 25 knobbed apex directed ventrad (fig. 87) 25. Valve subtruncate at apex (fig. 83a) ..... diasema ...... falcifera Valve tapering apically to bluntly rounded point (fig. Aedeagus with apical cornutus (fig. 60b); ampulla 81a) ...... lui a extending dorsad at least to costal margin (fig. 26. Juxta with strong apical spine; ampulla with broad 60a) ...... 7 spine base and terminating in a sharp spine (fig. 7. Cornutus more than one-third as long as aedeagus 75a) ...... alticola (fig. 72b) ...... selecta Juxta with a strong apical point but not a distinct Cornutus smaller, about one-fifth as long as aedea­ sclerotized apical spine; ampulla tapering to a gus (fig. 60b) ...... altera point but without distinct terminal spine (fig. 8. Aedeagus with cornuti ...... 9 67a) ...... parilis Aedeagus without cornuti ...... 26 9. Aedeagus with basal and apical cornuti ...... 10 (S. srwwi not seen. S. borea was not seen but Aedeagus with only one cornutus ...... 16 was placed in the above key based on genitalic 10. Many small cornuti projecting from basal and apical descriptions provided by MeDunnough (1944).) sc1erites (fig. 63b) ...... ignea Aedeagus with only two cornuti ...... 11 11. Basal cornutus large (fig. 73b) ...... 12 Basal cornutus minute (fig. 65b) ...... 15 KEY TO THE SPECIES BASED ON FEMALE 12. Ampulla short (fig. 73a) ...... sackeni GENITALIA Ampulla elongate (fig. 70a) ...... 13 13. Ampulla long and outcurved and on right valve more strongly outcurved, subparallel with costal mar­ 1. Entrance of ductus bursae on right side of corpus gin; ampulla on left valve more erect (fig. 70a) bursae...... 2 ...... orophila Entrance of ductus bursae on left side of corpus Ampulla not outcurved and approximately symmetr­ bursae ...... 10 ical ...... 14 2. Apex of corpus bursae heavily sclerotized or strigate 14. Ampulla tapering gradually to a point; apex of juxta or darkened ...... 3 produced to a strong, sharp point (fig. 76) Apex of corpus bursae not as described above .. 6 ...... angulidens 3. Base of ostium bursae with lateral appendage on Ampulla with a broad base, narrowed abruptly to a right side ...... ,...... 4 point at terminal one-third; apex of juxta only No appendage on ostium bursae ...... 5 slightly pointed (fig. 61a) ...... celsa 4. Projection off cephalic end of ostium bursae directed 15. Juxta with a spine (fig. 18) ...... microgamma caudad (fig. 124) ...... celsa Juxta without a spine (fig. 65a) ...... montana Projection off ceph&.lic end of ostium bursae directed 18. Aedeagus with basal cornutus (fig. 68b) ottolenguii laterad (fig. 126) ...... angulidens Aedeagus with apical cornutus ...... 17 5. Ventral surface of apex pigmented (fig. 19) . ignea 17. Cornutus strongly curved and with broad base (fig. Pigmentation of apex confined to area around base 78b) ...... 18 of ductus seminalis (fig. 129) ...... montana PLUSIINAE OF AMERICA NORTH OF MEXICO 37

6. Base of ostium bursae with short lateral process (fig. its own width in ventral aspect ...... 19 133) ...... 7 Caudal extension of ventral plate of ostium bursae Base of ostium bursae without process ...... 9 only slightly produced beyond ostium to a length 7. Projection off ostium bursae almost or entirely mem­ less than its own width in ventral aspect .. 21 branous (fig. 133) ...... rectangula 19. Cephalad extension of ostium bursae beyond origin Projection off ostium bursae sc1erotized ...... 8 of ductus bursae projected on right side, one-half 8. Width of ductus bursae at entrance to corpus bursae as wide as ostium bursae (fig. 141) . octoscripta greater than one-half of length of ductus bursae Cephalad extension of ostium bursae beyond origin (fig. 147) ...... cryptica of ductus bursae not confined to right side and Width of ductus bursae at entrance to corpus bursae larger than in fig. 141 ...... 20 one-half or less but greater than one-fifth of 20. Cephalad extension of ostium bursae mostly mem­ length of ductus bursae (fig. 130) ...... alias branous (fig. 142) ...... selecta Width of ductus bursae at entrance to corpus bursae Cephalad extension of ostium bursae mostly sc1ero­ about one-fifth of length of ductus bursae (fig. tized (fig. 144) ...... altera 145) ...... abstrusa 21. Apex of bursae a well-defined region since it 9. Ostium with lateral pouchlike structures (fig. 128) is comtricted at its a.rigin from the corpus bur­ ...... alticola sae proper (fig. 146) ...... ottolenguii Ostium with laterocaudad extension of a long, heav­ Apex of corpus bursae not distinctly separated by ily sc1erotized ostium bursae; juncture of ductus constriction from corpus bursae proper .... 22 bursae with ostium bursae a swollen, darkened 22. Corpus bursae sharply curved forming an acute an­ structure, the ductus bursae and ostium bursae gle (fig. 148) ...... u-aureum forming a right angle (fig. 127) ...... surena Corpus bursae if curved then angle obtuse (fig. 150) 10. Apex of corpus bursae heavily sclerotized, strigate, ...... interrogationis or darkened (fig. 135) ...... 11 23. Ductus bursae enters corpus bursae ventrad toward Apex of corpus bursae membranous or merely gran­ left (fig. 152) ...... parilis ulose (fig. 152) ...... 17 Ductus bursae enters corpus bursae laterad ... 24 11. Apex of corpus bursae strigate or with deep lateral 24. Base of ostium bursae produced to right as a short fold...... 12 membranous sac ...... 25 Apex of corpus bursae not strigate or infolded. 13 No membranous sac on right side at base of ostium 12. Apex of corpus bursae darkened and deeply infolded bursae...... diasema on right side (fig. 137) ...... epigaea 25. Ostium bursae produced caudad as two sublateral, Apex of corpus bursae not as above; ductus semin­ winglike extensions ...... borea alis arises from cephalic end of apical region (fig. Ostium bursae not produced caudad as winglike 135) ...... falcifera extensions (fig. 153) ...... lula 13. Ductus bursae entirely membranous; apex of corpus bursae shagreened and dark, curving to left (Syngrapha borea and diasema are placed in dorsad of ductus bursae (fig. 143) . viridisigma this key on the basis of genitalic descriptions Ductus mostly or entirely sc1erotized and strigate; provided by McDunnough, 1944.) apex of corpus bursae may be darkened but not curving to left dorsad of ductus bursae .... 14 Syngrapha ottolenguii (Dyar), new combination 14. Ductus bursae with a membranous swelling on right Figure 198 side (fig. 139) ...... 15 Ductus bursae without a membranous swelling on Autographa. arctica Ottolengui, 1902, p. 66. right side ...... 16 [preoccupied] 15. Apex of corpus bursae with brown, sc1erotized plate Autographa ottolengui Dyar, 1903, p. 200. [new extending down right side of ductus bursae l fig. 139) ...... orophila name pro arctica] Apex of corpus bursae heavily shagreened and dark Autographa vaccinii (Edwards), 1886, p. 170.­ (fig. 138) ...... microgamma McDunnough, 1921, p. 85. 16. Ductus bursae about as wide as base of ostium Male genitalia (fig. 68).-.Male genitalia of the bursae with membrallous bulge on left side near ostium bursae (fig. 136) ...... sackeni same general form as species of Autographa: Ductus bursae about twice as wide as base of ostium Valve broad toward apex; ampulla long and bursae, without membranous bulge, and curves slender, extending beyond 'costal margin; cla­ to the left (fig. 140) ...... snowi vus long, filiform; vincul urn longer tht.n in Syn­ 17. Ostium bursae, wide and long, extending cephalad grapha, tapered to a pointed apex, with rela­ beyond apex of corpus bursae ...... 18 Ostium bursae shorter, not extending as far as apex tively long, ventral projections near apex; of corpus bursae...... 23 aedeagus with strong, basal cornutus. 18. Caudal extension of ventral plate of ostium bursae Female genitalia (fig. 146).-Ostium bursae produced beyond ostium to a length greater than wide, heavily sclerotized, long, extending ce­ 38 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE phalad well beyond apex of corpus but-sae, and subequal in length to ostium bursae and sha­ the venter extending caudad beyond ostium; greened on apex. ductus bursae membranous, about as long as Type.-Female, American Museum of Natu­ greatest width of ostium bursae, entering cor­ ral History. pus bursae on right side near the middle; cor­ Type locality.-Lake Nipigon, Ontario. pus bursae in area around ductus bursae en­ Host plant.--Unknown. trance strigate, somewhat sclerotized; main Range (map I8).-Nova Scotia to Ontario; portion of corpus bursae membranous, no south to New York and Michigan. longer than ostium bursae, with apex a distinct Comments.-Adult specimens studied here membranous sac; ductus seminalis arises from agree with the type examined in the American apex of corpus bursae. Museum of Natural History. Type.-National Museum of Natural History. Type locality.-Alter Islands, Alaska. Host plant.-Unknown. Syngrapha octoscripta (Grote) Range (map Il).-Alaska and Yukon Terri­ tory. Comments.-This species has spines on the Figure 200 hind tibiae, which is typical of species of Syn­ Plusia octoscripta Grote, 1874a, p. 72. grapha, and the female genitalia are also like Autographa octoscripta: Dyar, 1902, p. 81. Syngrapha and not Autographa. Larval speci­ Syngrapha octo scripta: Hampson, 1913, pp. 425, mens are needed for further clarification. 426. Autographt~, pallida Ottolengui, 1902, p. 71; va'r­ iana Ottolengui, 1902, pp. 70, 71; zeta Ottolen­ Syngrapha altera (Ottolengui) gui, 1902, p. 70; epsilon Ottolengui, in Dyar, 1900, p. 495--Barnes and McDunnough, 1917, p.83. Figure 199 Autographa magnifica Ottolengui, 1919, p. 124- Autographa altera Ottolengui, 1902, p. 69. McDunnough, 1938, p. 113. Syngrapha altera: Hampson, 1913, p. 429. Syngrapha octoscripta: McDunnough, 1944, p. Autographa altera: Barnes and McDunnough, 198. 1917, p. 83. Male genitalia (fig. 66).-Valve with preapical Syngrapha variana (Ottolengui), 1902, p. 70- spine projecting from costal margin; apex of McDunnough, 1944, p. 197. valve round; ampulla triangular with a broad Male genitalia (fig. 60).-Valve somewhat nar­ base; aedeagus with apical cornutus; apex of rowed in caudal third, produced costoapically aedeagus externally shagreened. into a short spine; ampulla arises from a broad Female genitalia (fig. 141}.-Ostium bursae base and extends just beyond costal margin, long, flat, heavily sclerotized, extending ce­ slightly curved; clavus sharply bent, apex pro­ phalad beyond origin of the ductus bursae on jecting toward apex of valve; aedeagus with the right side and about half the width of the apical cornutus, about one-fifth as long as ae­ ductus bursae, and extending caudad as a ven­ deagus. tral plate well beyond the ostium about twice as Female genitalia (fig. 144).-Ostium bursae far as its own width from a ventral aspect; very wide, extending cephalad beyond the apex ductus bursae about as wide as ostium bursae, of corpus bursae to a blunt point beyond origin short, S-shaped from left lateral aspect, sclero­ of ductus bursae, and extending caudad well tized, and heavily strigate, entering corpus bur­ beyond ostium as a heavily sclerotized, strigate sae on the left side; corpus bursae membranous ventral plate longer than wide caudad of os­ and shagreened throughout. tium; ductus bursae short, strigate, originating 'fype.-{?] Syntype, British Museum (Natural from left side of ostium bursae, projected ven­ History) (octoscripta). Female, American Mu­ trad and bending cephalad to enter corpus seum of Natural History (pallida). Female, bursae on the left side; corpus bursae small, American Museum of Natural History (vaT­ PLUSIINAE OF AMERICA NORTH OF MEXICO 39 iana). Female, American Museum of Natural ostium bursae, directed ventrad from the os­ History (zeta). Male, National Museum of Natu­ tium bursae, becoming strigate and sclerotized ral History (epsilon). Female, Canadian N a­ after it becomes directed to the right, expand­ tional Collection, Ottawa, Ontario (magnifica). ing to a width of at least one-third the length of Type locality.-[?] (octosc1ipta). Salmonier, the corpus bursae, which it enters on the left Newfoundland (pallida). St. John, New Bruns­ side well cephalad of the apex; the midsection of wick (val"iana). Calgary, Alberta (zeta). Kodiak, the corpus bursae is constricted with the fun­ Alaska (epsilon). Ucluelet, Vancouver [British dus membranous and minutely granulose; the Columbia] (magnifica). corpus bursae is strigate on the left side from Host plant.-Vaccinium sp. the fundus to the apex of the corpus bursae; Range (map 12).-Newfoundland to North­ the right side of the apex is heavily shagreened west territory, Alaska south to New York, imparting a brownish color to it and deeply Michigan, Wisconsin, Idaho, British Columbia. infolded with the ductus seminalis arising Comments.-McDunnough (1944) spoke of the within the fold at the caudal end. form magnifica, a female type and a male Larvae.-Integument with microspines like specimen from Ucluelet, British Columbia. minute setae; crochets range from 23 to 30 in Though lighter in color, these two specimens number; seta SV-2 absent on meso- and meta­ proved to be conspecific with octosc1-ipta, based thorax; interval between the pinacula of setae on comparisons of the genitalia of each sex of V-Ion abdominal segment 4 more than one­ both forms. A male, labeled as magnifica in the half the interval between V-I and SV-2; radu­ American Museum of Natural History, from loid on hypopharynx with 12 ridges; otherwise the type locality of magnifica and superficially same as generic description. resembling it, is quite unique and not a subspe­ Type.-British Museum (Natural History). cies of octosc'1-ipta. Refer to the illustration of Type locality.-New York. the genitalia (fig. 64, 65). We have deferred Host plant.-Amelanchie1· alnifolia, Larix sp., naming this species until additional specimens Spiraea sp., McGuffin (1954); Pinus eontoria, can be examined. Populu,.s tremuloides, Vaecinium sp., Ross and Evans (1957); Alnus sp., Betula pendwa, Junipe­ rus sp., Pinus banksiana, Salix sp., Prentice Syngrapha epigaea (Grote) (1962); Vaeeinium augustifolium, Wood (1951); Gale palu,.stris, KalmiCL augustifolia. Figure 201 Range (map 19).-Newfoundland to British Plusia epigaea Grote, 1874c, p. 208. Columbia; south to New York, Wyoming, and Autogmpha epigaea: Dyar, 1902, p. 81. Colorado. Syngmpha epigaea: Hampson, 1913, p. 421. Comments.-Larvae of this species were col­ Autographa epigaea; Barnes and lVIcDunnou.gh, lected in commercial blueberry fields by work­ 1917, p. 83. ers in the Canada Department of Agriculture, Syngrapha epigaea; McDunnough, 1944, p. 200. New Brunswick. Larvae were reared to adults Male genitalia (fig. 58).-Valve wide; ampulla for identification and association with larval thin, straight, extending just beyond costal specimens. The specimens were obtained from G. W. Wood. margin; costa produced as a spine before apex; cornutus large, thick, one-half as long as aedea.. gus. Syngrapha selecta (Walker) .Female genitalia (fig. 137).-Ostium bursae long, about two-thirds length of corpus bursae, Figure 202 extending well past the apex of the corpus bursae, heavily sclerotized, expanded caudally Plusiu selectCL Walker, 1875, p. 912. and produced into two lateral extensions that Plusia viridisigmCL: Smith (nee Grote), 1893, p. taper to a point directed dorsad, extending well 255. beyond ostium; the initial portion of the ductus Autographa seleeta: Dyar, 1902, p. 81. bursae is membranous and narrower than the SyngraphCL seleeia; Hampson, 1913, pp. 421, 422. 40 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE A.utogmpha selecta: Dod, 1915, p. 130. dull luteous, with the outer one-third fuscous. Syngmpha selecta: McDunnough, 1944, p. 20l. In selecta, the hind wings tend toward being ochreous, contrasting more with the fuscous Male genitalia (fig. 72).-Valve slender, some­ border. Females of selecta can be identified by what tapered to apex from which arises a rela­ examining the tip of the abdomen. By brushing tively long spine, compared to the apical spine a few scales off, the ventral plate of the ostium of vi1'idis'igma; ampulla arises from a wide base, bursae shows as a dark, sclerotized plate from curves basad and extends just beyond costal under the ovipositor. There is no ventral plate margin; clavus sharply bent with apex directed on viridisigma females. The males of selecta are toward valve apex; aedeagus with a large, thick also recognizable without removing the geni­ apical corn utus, about one-third as long as talia. The costoapical spines of the valves are aedeagus. long and curved, frequently crossing each Female genitalia (fig. 142).-Ostium bursae other, as seen from the caudal aspect. Parental heavily sclerotized, sides infolded dorsad, ex­ female of the laboratory culture was collected tending cephalad far beyond apex of corpus in Schoolcraft County, Manistique, Mich., at bursae, the cephalic extension twice as wide black light. beyond origin of ductus bursae and a large, Refer to comments given for viridisigma. thick, mostly membranous sac dorsad of corpus bursae; ostium protected ventrally by a heavily sclerotized, scooplike plate which arises from Syngrapha viridisigma (Grote), ostium bursae near origin of ductus bursae; new combination ductus bursae very short, about as wide as long, mostly sclerotized and strigate, arising from membranous sac of ostium bursae, enter­ Figure 203 ing corpus bursae on the left side; corpus bur­ Plusia vir-idisig'l'na Grote, 1874a, p. 73. sae and membranous sac, somewhat narrowed Plusia vir-idisignata Grote, 1875c, p. 205. at apex, which is lightly shagreened. Male genitalia (fig. 69).-Valve produced at Larvae.-Similar to viridisigma with the fol­ apex into a distinct spine on costal margin; lowing exceptions: Crochets range from 21 to 24 ampulla short, curved slightly caudad, with in number; interval between pinacula of setae rounded apex; aedeagus with a large basal V-1 on abdominal segment 4 one-half the inter­ cornutus and a slightly smaller median cornu­ val between setae V-1 and SV-2; mesothorax tus. with pinaculum of D-2 equidistant from D-l Female genitalia (fig. 14:~).--Ostium bursae and SD-2; and raduloid on hypopharynx has 11 large, about as long as wide, sclerotized and ridges. creased ventrad, rounded to base; ostium oval, Type.-Female, British Museum (Natural unprotected; ductus bursae entirely membra­ History). nous except perhaps near base of ostium bur­ Type locality..-St. Martins Fa.ll, Albany sae, narrow caudad but widening to about River, Hudson Bay District, Ontario. three times this width as it enters corpus bur­ Host plant.-Unknown; some host records for sae on the left side; apex of corpus bursae vir-idisigma recorded in literature erroneously shagreened and darkly pigmented throughout, for selecta probably apply to selecta as here curving to the left dorsad of ductus bursae; corrected. ductus seminalis arises from apex far to the left Range (map 16).-Newfoundland, Ontario, near entrance of ductus bursae. and Alberta; Massachusetts, Michigan, and Larvae.-Integument with middorsal, longitu­ Wisconsin. dinal band formed of dark microspines; crochets Comments.-This species is superficially like range from 25 to 28 in number; seta SV-2 viridisigrna. The color pattern of the wings may absent on meso- and metathorax; interval be­ serve to select possible suspects from a mixed tween the pinacula of setae V-Ion abdominal series of both species. In viridisigma, the more segment 4 more than one-half the interval be­ common species of the two, the hind wings are tween V-I and SV-2; mesothorax with pinacu­ PLUSIINAE OF AMERICA NORTH OF MEXICO 41 lum of seta D-2 closer to D-1 than to SD-2; Male genitalia (fig. 70).-Valves with ampul­ raduloid of hypopharynx absent. lae asymmetrical; left valve with ampulla only 1.'ype.-A female syntype ofV-iTidisigma in the slightly curved caudad, pointed apex extending British Museum (Natural History) has been beyond costal margin; right valve with ampulla selected and labeled and is presently desig­ strongly curved, pointed apex directed almost nated as the lecto-type: "Type, Plusia v-iridi­ ventrocaudad; aedeagus with both basal and sigma Grote/Grote colI. 82--54JType (a white disc apical cornuti. with a red border)fViridisignata Grote 1875 is Female genitalia (fig. 139).-Ostium bursae unnecessary replacement name for mridisigma heavily strigate, extended caudolaterally as two Grote, 1874, det. 1. W. B. Nye, 1970." winglike processes somewhat protecting the os­ Type locality.-Quebec. tium; ductus bursae mostly sclerotized and stri­ Host plant.-Abies lasioca1pa" Picea glauca, P. gate except for a large membranous swelling on engelmannii, P. sitchensis, Pinus contoTia, the right side; ductus bursae wide as it enters Pseudotsuga menziesii, Tsuga heterophylla, corpus bursae on left side; apex of corpus bur­ Ross and Evans (1857); La1-iX lancina, Thuja sae with a brown pigmented, sclerotized plate plicata, Tsuga meriensiana, Prentice (1962); Pi­ extending cephalad along right side of corpus cea mariana, Tietz (1972). bursae. Range (map 11).-Canada and Alaska; south Larvae.-Larval exuviae were examined and to New York and Wisconsin; Washingt­ based on discernible characters, this species is southeast to New Mexico. similar to v'iridisigma. Refer to description of Comments.-McDunnough (1944) described the latter species. this species but erroneously called it selecta. Type.-Syntypes not seen by us, British Mu­ SyngTapha viridisigma is by far the more com­ seum (Natural History). mon of the two species, yet all specimens seen Type locality.-Brobokton Creek, Alberta. were labeled selecta. A female specimen was Host plant.-Vaccinium sp., Crumb (1956). sent to 1. W. B. Nye at the British Museum Range (map 12).-Alberta to Alaska; south to (Natural History) to compare with the types of Oregon and Wyoming. selecta and its junior synonym viridisigma. A subsequent communication confirmed that Grote's female syntype of viridisigma agreed with the descriptions and illustrations pre­ Syngrapha snowi (Edwards) sented for selecta by McDunnough, and the female syntype should be designated as lecto­ Figure 205 type. A specimen of the less common species Plusia snowi Edwards, 1884, pp. 44, 45. sent for comparison with the type, agreed with A.utographa snowi: Dyar, 1902, p. 81. Walker's holotype of seiecta, a female. Syngmpha snowi: Hampson, 1913, pp. 418, 419. Parental females of the laboratory cultures Autogmpha snowi: Ottolengui, 1919, p. 27. were collected at black lights in Mare du Sault Camp, Laurentides Park, route 54, Quebec. Re­ Syngrapha snowi: McDunnough, 1944, p. 188. fer to comments given for selecta. Male genitalia.-Not examined. Female genitalia (fig. 140).-Ostium bursae bell shaped, short and broad, expanded caudad; Syngrapha orophila (Hampson) ductus bursae bends to the left, wide, lightly sclerotized, and strigate mostly on the right l'-'igu re 204 side; the broadly rounded apex of corpus bursae PlUo''lia oTophila Hampson, 1908, p. 105. lightly pigmented. Type.-Male, American Museum of Natural Autogmpha orophila: Dod, 1913, pp. 236--238. History. Syngrapha orophila: Hampson, 1913, p. 416. Type locality.-Hot Springs, Las Vegas, N. Autographa orophila: Barnes and Mc- Mex. Dunnough, 1917, p. 83. Host plant.-Unknown. Syngrapha orophila: McDunnough, 1944, p. 184. Range (map 16}.-New Mexico. 42 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE Syngrapha sackeni (Grote) ductus bursae, and corpus bursae similar to sackeni, with the following exceptions: A thick­ Figure 206 ened membranous pouch is on right side of Plusia sackeni Grote, 1877, pp. 135, 136. ostium bursae, and the apex of the corpus Autographa sackeni: Dyar, 1902, p. 81. bursae is not darkened. Syngrapha sackeni: Hampson, 1913, pp. 417, Type.-British Museum (Natural History) 418. (lula). American Museum of Natural History Autographa sackeni: Barnes and McDunnough, (diversigna). 1917, p. 83. Type locality.-Canada (lula). Nordegg and Laggan, Alberta, (diversigna) Syngrapha sackeni: McDunnough, 1944, p. 187. Male genitalia (fig. 73}.-Valve wide, apex Host plant.-Unknown. Range (map 12).-Alberta, British Columbia, broadly rounded; clavus short and broad; am­ Alaska. pulla reduced, short, broad at apex, aedeagus with basal and apical cornuti, approximately equal in length; juxta produced to a sharp Syngrapha borea (Aurivillus) point, but not a distinct, sclerotized apical spine. Plusia diasema var. bore a Aurivillus, 1890, p. 4. Female genitalia (fig. 136}.-Ostium bursae Syngrapha borea: McDunnough, 1944, p. 187. short with no appendages or extensions pro­ This species was not examined, so the follow­ tecting ostium; ductus bursae bends to the left, ing male and female genitalic descriptions are its initial one-third membranous and continued reproduced from McDunnough (1944): on left side, remainder of ductus bursae stri­ "MALE GENITALIA. Juxta and aedeagus much as gate and sclemtized, where it enters corpus in lula. The harpe (= ampulla) is of moderate width bursae on left side, becoming about as wide as throughout with truncate apex from the underside of widest part of ostium bursae; corpus bursae which a minute spine projects; this truncate apex at once distinguishes borea from diasema and lula in uniformly shagreened except at apex, which is which the harpes taper to sharp points. partially sclerotized and darkened. FEMALE GENITALIA. Very similar to those of lula Type.-British Museum (Natural History) but with definite suggestions of weakly chitinized Type locality.-Idaho Springs, Colo. wings extending caudad from ostium margin. Ductus Host plant.-Unknown. funnel (= ostium bursae) weakly chitinous and stri­ gate, except for a strip on left side which shows Range (map 19).-Alberta south to Colorado. stronger chitinization; the base of the funnel is membranous and projects as a short sac below and to Syngrapha lula Strand the right of the continuation of the ductus; this continuation as well as the bursa are much as in lula." Figure 207 Type.-[?] Syngrapha snowi form lula Strand, 1916, p. 47 Type locality.-Greenland. Autographa l1tla: Barnes and Benjamin, 1923, p. Host plant.-Unknown. 212. Range (map 14).-Greenland, Yukon, and Autographa diversigna Barnes and Benjamin, Northwest Territory. 1923, p. 212. Syngrapha lula: McDunnough, 1944, p. 186. Syngrapha diasema (Boisduval) Male genitalia (fig. 81).-Valve tapers some­ Figure 208 what to a narrowly rounded apex; ampulla as in diasema; aedeagus spiculate apically, with Plusia diasema Boisduval, 1829, p. 160. apical cornutus perhaps not as thick as in Autographa diasema: Dyar, 1902, p. 81. diasema but with a broad base. Syngrapha diasema: Hampson, 1913, pp. 416, Female genitalia (fig. 153).-Ostium bursae, 417. PLUSIINAE OF AMERICA NORTH OF MEXICO 43

Autographa diasema: Barnes and Mc­ Plusia reetangula: Walker (nee Kirby), 1857, p. Dunnough, 1917, p. 83. 908. Autographa diversigna Ottolengui, 1919, p. Autographa falcifera: Dyar, 1902, p. 81 121.-McDunnough, 1921, p. 85. Autographa simplex (Guenee), 1852, p. 47.- Ot­ Syngrapha diasema: McDunnough, 1944, pp. tolengui, 1902, p. 74. 185-186. Syngrapha falcifera: Hampson, 1913, pp. 419, Male genitalia (fig. 83).-Valve broad at apex 420. and slightly rounded; ampulla extending to cos­ Autogra.pha simplieima Ottolengui, 1902, pp. 74­ tal margin, rathel abruptly narrowed in distal 75.-Dod, 1913, p. 239. one-fourth to a sharp pointed apex; aedeagus with a thick apical cornutus. Anagrapha faleifem: McDunnough, 1944, p. 202. The female of this species was not examined, Male genitalia (fig. 87).-Valve broad, widest so the following female genitalic description at truncate apex; ampulla downcurved, with was reproduced from McDunnough (1944): knobbed apex directed ventrad; aedeagus with a sharply pointed basal cornutus. "FEMALE GENITALIA. Ostium considerably Female genitalia (fig. 135).-Ostium bursae broader than in oTophila; the two sul:>lateral wings narrow and short, ostium unprotected; ductus project caudad but their apices are broadly rounded bursae relatively thick, sclerotized and strigate and not pointed. The ductus-funnel (= ostium bur­ sae) is strongly chitinized and strigate, gradually throughout, entering corpus bursae on the left; tapering, and noticeably longer than in oTophila. main body of corpus bursae membranous and The continuation of the ductus bends to the left and saclike, but the apicaltegion sclerotized, darkly is short and broad, less heavily strigate and chitin­ pigmented, and spiculate, with the ductus sem­ ized than in orophila, the strigae continued for only inalis arising from the c~phalic end of the apical a very short distance along the left side of the bursa, section. below the entrance. The bursa consists of the usual large, membranous sac; the rounded apex projects Larvae.-Very similar to mridisigma with the caudad along the right side of the ductus for a following exceptions: Integument with micro­ greater distance than in orophila and shows none of spines present but inconspicuous; head without the chitinization found in this species, being merely spots; crochets range from 21 to 24 in number; finely granulate; the membrane of the remainder of the bursa is smooth." interval between pinacula of setae V-Ion ab­ domin~1 segment 4 one-half the interval be­ Type.-A male syntype of diasema in the tween V-I and SV-2. National Museum of·Natural History has been Type.-(?] British Museum (Natural History) selected and labeled and is presently desig­ (faleifera). Syntypes, Barnes Collection, Na­ nated as the lectotype: "Ex Musaeo, Dris"Bois­ tional Museum of Natural History, and British duvalJDiasema(l'ype, Plusia diasema Bdv., alc Museum (Natural History) (simplex). Female, Hofer/Fig. par J. Culot, Noct. et Geom. d' Eu­ American Museum of Natural History (simpli­ rope, PI. 72 Fig. 6/0berthur ColIectionJO geni­ eima). talia on slide, Aug. 21, 1936, J.F.G.C. -558." Type locality.-Nova Scotia (faldfera). New Type locality.-Saponie, Lapland. York, Canada (simplex). Oregon (not Washing­ Host plant.-Unknown in North America; in ton) (simplicima). Europe feeds on Betula spp. and Populus spp., Host plant.-Apium graveolens, Beta vulgaris, Kostrowicki (1961). Brassica oleracea, Lactuca sativa, Zea mays, Range (map 13).-Labrador to Alberta, and Crumb (1956); Althaea rosea, Arctium lappa, north to Alaska; Scandinavia. Cirsium sp., Daucu.s earota, Plantago sp., Tar­ axae1~m offieinale, Vaeeinium macroearpan, Tietz (1972); Viburnum sp. Syngrapha falcifera (Kirby) Range (map 3).-NovaScotia to British Col­ umbia; United States. Figure 209 Comments.-This species is generally called Plusia faleifera Kirby, 1837, p. 308. the celery looper. Parental females of labora­ 44 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE tory cultures were collected in a light trap at Comments.-Parental females for laboratory Auburn, Ala., and others were collected at light cultures were collected during daylight hours in Pittsburgh, Pa. while they were feeding on flowers of a small species of Solidago at 4,000 feet elevation on Mt. Washington, N. H. Syngrapha u-aureum (Guenee)

Syngrapha interrogationis (Linnaeus) Figure 210 Plusia 1~-aureu,m Guenee, 1852, p. 354. Figure 211 Autographa u-aureum: Dyar, 1902, p. 81. Noctua interrogationis Linnaeus, 1758, p. 513. Syngrapha u-aureum: Hampson, 1913, pp. 424, Noctuainterrogationis: Fabricius, 1781, p. 228. 425. Autographa interrogationis: Hubner, 1816, p. Autographa vaccinii (Edwards), 1886, p. 170.­ 251. Barnes and McDunnough, 1917, p. 83. Pl1~ia interrogationis: Treitschke, 1823, p. 190. Autographa u-aureum: McDunnough, 1938, p. 113. [= interrogationis] Syngrapha interrogationis: Hampson, 1913, p. 430,431. Syngrapha u-a1~re'Um: McDunnough, 1944, p. 193. Autographa intermgationis var. he?'schelensis Male genitalia (fig. 80).-Similar to interroga­ Benjamin, 1933, pp. 61, 62. tionis, but on all specimens examined, with no Autographa u-aureum (Guenee), 1852, p. 354; sharp projection basad on costal margin; with­ vaccinii (Edwards), 1886, p. 170.-Mc­ out projection on basal third of ampulla; cos­ Dunnough, 1938, p. 113. toapical edge of valve not as sharply angled as Syngraphainterrogationis: McDunnough, 1944, in interrogationis; and aedeagus much less spi­ p.195. culate apically. Ml:l,le genitalia (fig. 79).-Valve with costal Female genitalia (fig. 148).-Structures the margin basad produced as a small, sharp spine; same as in interrogationis exctpt for the follow­ ampulla broad from base, tapering to a pointed ing: The elongate apical region of the corpus apex and curved toward base, \\;th a small bursae is recUl-ved to the left forming an angle point on edge of basal one-third of ampulla on of 90° or an acute angle with the corpus bursae, the European specimens examined; costoapical and in some specimens directed somewhat ce­ edge of valve sharply angled, and may even be phalad. slightly produced as a spine in some specimens; Larvae.-Crochets range from 27 to 30 in aedeagus heavily spiculate apically, with a number; raduloid on hypopharynx with 10 small, stubby apical cornutus on an irregularly ridges; otherwise similar to viridisigma. shaped base. Type.-A female syntype of u-aureumin the Female genitalia (fig. 150).-Ostium bursae National Museum of Natural History has been long and wide, heavily scierotized, extending selected and labeled and is presently desig­ cephalad beyond the origin of the ductus bur­ nated as the lectotype: "Ex Musaeo. Dris Bois­ sae as a membranous projection, and extending duvaVDelecarlielu. aureum/Type, Plusia u-au­ caudad beyond ostium ventrally as a plate reum' Bdv.', alc Hofer/Oberthur Collectionl often bilobed, the length of the extension be­ genitalia on slide, 3 Nov. 1942, C.H. #3." Syn­ yond ostium less than its own width; ductus types, American Museum of Natural History bursae strigate and short, about one-third the (vaccinii). length of the ostium bursae, entering the cor­ Type locality.-Dele carlie, Laponie (= Lap­ pus bursae on the left; corpus bursae may be land) (u-aureum). Mt. Washington, N. H. (vacci­ bent to the right apically, but always at an nii). obtuse angle; apex minutely granulose. Host plant.-Vaccinium sp., Forbes (1954). Type.-British Museum (Natural History) (in­ Range (map ll).-Newfoundland, Manitoba to terrogationis). Male, National Museum of Natu­ Alaska; south to New York and Colorado. ral History Ch,e?'schelensis), PLUSIINAE OF AMERICA NORTH OF MEXICO 45 Type locality.-[?] (intermgationis). Herschel Syngmpha alias: McDunnough, 1944, p. 189. Island, Yukon Territory (herschelensis). Male genitalia (fig. 78).-Tegumen about one­ Host plant.-Unknown. half length of valve; uncus broad, shorter than Range (map 13).-Labrador to Alberta and tegumen; valve narrow near middle, widest in Alaska. apical one-fifth, sacculus near base of valve produced into a triangUlar-shaped protuber­ ance, which IS broad at base and round at apex; Syngrapha su,rena (Grote) clavus typical for genus, ampulla reduced to short stub with apex variable, sometimes blunt, Figure 212 or pointed, or blunt with a minute spine; aedea­ Pl'USia surena Grote, 1882, p. 585, gus with a single apical cornutus on vesica, Autogmpha S'!.t1·ena: Dyar, 1902, p. 81. which is thick at base, sharply curved and Syngrapha surena: Hampson, 1913, p. 430. tapered to a point, and aedeagus basad with Autographa SU1'ena: Barnes and McDunnough, vesica granulate. 1917, p. 83. Female genitalia (fig. 130).-Ostium bursae in Syngmpha 8urena: McDunnough, 1944, p, 200. two distinct parts; the caudal portion short and Male genitalia (fig. 74).-Valve broadly trun­ wide, slightly wider than long, sclerotized and cate at apex, sharply angled on costoapical strigate, and the second portion slightly longer edge; ampulla with broad base, tapering some­ and narrower than caudal portion, rounded ?t what to apex, which is a bifid spine; aedeagus base, and produced on right side as a short with a large, thick apical cornutus; vesica when rounded appendage; ductus bursae mostly everted forming a complete loop near the mid­ membranous initially, becoming more sclero­ dle. tized and strigate, entering corpus bursae on Female genitalia (fig. 127).-Ostium bursae right side where it is wider than one-fifth, but wide, as long as ductus bursae, scooplike, ex­ no more than one-half of the total length of the tended caudad just beyond ostium, bilobed on ductus bursae; corpus bursae membranous, sac­ the end; ductus bursae arises from base of like shagreened at apex from which ductus ostium bursae as a thick heavily pigmented, seminalis arises. swollen area becoming strigate and sclerotized Larvae.-Crochets range from 25 to 27 in to entrance into corpus bursae on the right number; microspines of integument present; side; corpus bursae membranous with apex interval between the pinacula of setae V-l on curved to left. abdominal segment 4 more than one-half the Type.-Not known to us. interval between V-I and SV-2; otherwise simi­ Type locality.-Orono, Me. lar to cei.r.;a, which also has raduloid absent. Host plant.-Unknown, Type.-Male, American Museum of Natural Range (map 16).-Newfoundland to Ontario; History (alias). Female, American Museum of Maine. Natural History (intemlia). Comments.-This species is rare in collections. Type locality.-Jefferson, N. H. (alias). Nor­ No more than a dozen specimens were seen by deg-6'. Alberta (interalia). us. Host plant.-Pice(L en[Jelmanni, P. glaHca, Ross and Evans (1957); Abies wmubilis, A. buis(t­ Syngrapha alias (Ottolengui) mea, A. lasiocal'pa, Lab·i.r laricina, Picea »tW'­ ianet, P. l'ubens, Pinus banksiana, P. monticola, Figure 213 P.'leuciotsuga menzie81:i, Thuja plicota, Tsuga heterophylla, Prentice (Z 9(j2). Alltogmpha alias Ottolengui, 1902, p. 69. Range (map U).-Newfoundland to Alaska~ Syngmph(t alias: Hampson, 1913, p. 424. south to New York, Wisconsin, and California; Autographa alias: Barnes and McDunnough, Rocky Mountains south to Arizona. 1917, p. 83. Comments.-The specimens studied here Autogmplw interalia Ottolengui, 1919, p. 122.­ agree with the type examined in the American McDunnough, 1938, p. 113. Museum of Natural History. The type of inter­ 46 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE alia, a female, was examined and determined to Secondaries dull luteous, basal half or less be alias. Parental female of laboratory culture heavily shaded, fuscous, with a somewhat con­ was collected in Schoolcraft County, near Man­ trastingly lighter, narrow band between; fringe istique, Mich. Other larvae were provided by A. white with some fuscus bands. H. Rose, Canada Department of Forestry, Sault Length of primaries from wing base along Sainte Marie, Ontario. costa to apex: 14 ± 1.0 mm (10 specimens). Male genitalia (fig. 64).-Tegumen about one­ SYTt,grapha abstrusa n. sp. half length of valve; uncus shorter than tegu­ men, thick, produced as a sharp spine at apex; Figure 214 juxta not produced at apex into a distinct, This species, like cryptica, is related to alias, sclerotized spine; valve tapering somewhat at which is the predominant species in those col­ tip to a rounded apex, with clavus broad at lections examined to date. The existence of base, short, narrowing to blunt, setose apex; abstrusa as a distinct species and not as part of sacculus at base smoothly rounded, not pro­ the inherent variability of alias was discovered duced into a broad, somewhat triangular protu­ by examination of the male genitalia. berance as with alias; ampulla arising from a Adult description.-Primaries basically gray, broad base, tapering to a sharp point, and not darkest in median area, dark gray mixed with extending to costal margin; aedeagus with a dark brown, dark area extending outward to single apical cornutus on vesica, which is thick postmedial (pm.) line, in some specimens at base, sharply curved, and tapered to a point, slightly obscuring line, and basad to antemedial aedeagus basad with vesica granulose. (am.) line, becoming slightly lighter gray with Female genitalia (fig. 145).-Ostium bursae in black specks to inner margin; above median two distinct parts, the caudal portion about as dark area, becoming slightly lighter gray, light­ wide as long, sclerotized and strigate, and the est around orbicular and on distal side of pm. second portion narrower at origin, sclerotized, line; basal third between am. line and basal line widening to base where it is rounded, and (b.!'), gray with small amounts of silver or gold produced on the right side as a round projection scales. Fringe white, alternating with dark (may be appressed to ostium bursae in some gray bars; terminal line (t.!.), forming white preparations); ductus bursae about one-half crescents, the points of which join at bases of width of ostium bursae, sclerotized and strigate bars on fringe, except at apex of wing where it throughout, only slightly wider at entrance to fades out; subterminal Cst.) line black almost to corpus bursae, about one-fifth of its total anal angle, thinly bordered on outer side with length; corpus bursae membranous, saclike, white, becoming mostly white only near anal shagreened at apex from which ductus semin­

angle, costal portion wavy, incised above M2, alis arises. then irregularly dentate, curving outward to Type.-Holotype, male, Lake Katherine, margin just above fringe of scales on anal Oneida County, Wis., july 4, 1938; genitalia slide angle; pm. line silvered throughout, bordered AU#051; deposited in Los Angeles County Mu­ with black, concave opposite cell and fold, in­ seum of Natural History, Los Angeles, Calif. curved to costa; am. line and b.1. silvered and Allotype, female, same as type, H. M. Bower; wavy. Renifonn elongate oval with black out­ genitalia slide AU#053; deposited in Los Ange­ line thin or faded out at top and bottom, some­ les County Museum of Natural History, Los what silvered on both sides; orbicular outlined Angeles, Calif. in silver, surrounded by light gray area, dark in Paratypes, two males, one female from Rain­ the center; stigma variable, irregular in shape, bow Falls Park near Rossport, Thunder Bay thickly outlined in silver, dark in the center, County, Ontario, M. C. Nielson, deposited in expanded on lower, inner side as a silver spot, Michigan State University, East Lansing; one with anothe?O silver spot on lower, outer side male from Jefferson's Notch, near Jefferson, completely separated jmm stigma in all speci­ N.H., T. D. Eichlin, deposited in National mens so jar exarnirled. Museum of Natural History, Washington, D.C. PLUSIINAE OF AMERICA NORTH OF MEXICO 47 Host plant.-Unknown; probably some host margin just above fringe of scales at anal an­ records for alias should apply to abst1'USa. gle; pm. line unsilvered, light bmwn, outline Range (map 2).-Ontario, Maine, New Hamp­ jUscous for costal half, becoming somewhat sil­ shire, Michigan, and Wisconsin. vered for inner one-half to margin, concave Comrnents.-The italicized phrase in the pre­ opposite cell and fold, incurved to costa; am. ceding adult description should enable one to line and b.!. silvered and wavy. Reniform nar­ select suspects of this species from a mixed row, approximating a parallelogram, outlined in series containing alias and c1yptica, but defini­ black with at most a few silver scales inter­ tive identification must be made by examina­ mixed; orbicular outlined in silver, dark brown" tion of genitalic characters. in the center; stigma variable, irregular in shape, thickly outlined in silver with dark brown in the center, extended on its lower two Syngrapha cryptica n. sp. corners in rounded, silver spots, variable in size, the outer one of which may be disjoined as Figure 215 a separate spot on one or both wings. Secondar­ This species, like abstrusa, is related to alias. ies dull luteolls, basal one-half heavily shaded, The existence of c1Yptica as a distinct species, fuscous, with somewhat contrastingly lighter, and not as part of the inherent varibility in the narrow band between; fringe white with fus­ maculation of alias, was discovered by exami­ cous bands. nation of the male genitalia. The following dif­ Length of primaries from wing base along ferences in wing patterns were subsequently costa to apex: 16.5 ± 1.5 mm (12 specimens). noted. Male genitalia (fig. 7l).-Tegumen less than Adult descIription.-Primaries basically one-half length of valve; uncus shorter than brown, darkest in median area, dark area ex­ tegumen, thick, produced as a sharp, curved tending outward, slightly across pm. line, par­ spine at apex; juxta without sclerotized apical tially obscuring line, and basad to am. line, spine; valve widest in apical one-fifth, narrowed becoming gray-brown, flecked with black specks toward middle, with clavus broad at base, short, to inner margin; above medial dark area, becom­ narrowing to blunt, setose apex, sacculus at ing light brown, slightly lighter mi.-xed with gray base not produced into a broad, somewhat tri­ in some specimens just distad of pm. line; basal angular protuberance as with cLlias, and am­ one-third between am. line and b.l. may be pulla represented by a broad, short ridge with silver-gray or gold tinged with green or yellow; a truncate apex and no apical spine; aedeagus somewhat 1·ectangular patch of slightly irides­ with a single a.pical cornutus on vesica, which is cent, silver-gray or a shade of gold, c01Tespond­ thick at base, sharply curved, and tapered to a ing -roughly to basal hue ofthat particular speci­ point, and aedeagus basad with vesica granu­ men, extending from vein Mz almost to anal lose. angle, and extending basad f1'om t.l. to just Female genitalia (fig. 147).-Ostium bursae in beyond st. line; dark scales on veins in the latter two distinct parts, the caudal portion short and area may show as thin, dark, stmight streaks, wide, slightly wider than long, sclerotized and particularly on M3 and Cu.! and on Mz and" CUI strigate, and the second portion nan'ower at on some specimens. origin, sclerotized, becoming as wide as caudal Fringe white, alternating with dark gray portion of ostium bursae, rounded cephalad and bars; t.!. forming white crescents, the points of produced on right side as a short, round appen­ which join at bases of bars on fringe, except at dage near base of cephalic portion of ostium apex of wing where it is dark gray and incurved bursae; ductus bursae at origin mostly mem­ to costal margin; st. line with costal one-half branous, about one-half the width of ostium either entirely black or with a thin border of bursae, gradually widening and becoming more silver distad, becoming mostly silver toward sclerotized and strigate to entrance into corpus inner margin as it passes through iridescent bursae on right side, at which point the ductus area, costal portion wavy, incised above Mz, bursae is wider than one-half of its total length; then irregularly dentate, curving outward to corpus bursae membranous, saclike, sha­ 48 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE greened at apex from which ductus seminalis bursae cephalad of corpus bursae apex, but in arises. alias it enters about one-third of total length of Type.-Holotype, male, Lake Katherine, corpus bursae cephalad of corpus bursae apex, Oneida County, Wis., July 13, 1946, H. M. perhaps being subject to more variation than Bower; genitalia slide AU#059; dep'[}sited in has been observed in this study. Los Angeles County Museum of Natural His­ Larvae.-Refer to description of alias to tory, Los Angeles, Calif. which rectangula is similar. Allotype, female, same as above, H. M. Type.--[?] British Museum (Natural History) Bower; genitalia slide AU#062; deposited in ('rectangula). Syntypes, American Museum of Los Angeles County Museum of Natural His­ Natural History (nargenta). British Museum tory, Los Angeles, Calif. (Natural History) (mortuorum). Paratypes, two males, one female, Roscom­ Type locality.-Canada (rectangula). Kaslo mon County, Mich., V. Warczunski, deposited in and Vancouver, British Columbia (nargenta). Michigan State University, East Lansing; one New York (mortuorum). male, Wausau, Wis. and two males, two females Host plant.-Pseudotsuga menziesii, Crumb from type locality, H. M. Bower, deposited in (1956); Abies lasiocarpa, Picea engelmanni, Pi~ Los Angeles County Museum, Los Angeles, nus monticola, Tsuga heterophylla, Ross and Calif. Evans (1957); Abies amabilis, A. grandis, Larix Host plant.-Unknown. laricina, Picea rubens, P. stichensis, Tsuga mer­ Range (map l).-Wisconsin, Michigan, and tensiana, Prentice (1962); Abies balsamea, Picea New Hampshire. glauca, P. ma'riana, Tietz (1972). Comments.-The italicized passages in the Range (map 19).-Nova Scotia to British Col­ preceeding description indicate diagnostically umbia; south to Pennsylvania and west to significant characters differing from the outer Washington. species of the alias complex, alias and abstr>vtsa. Conunents.-Ova were obtained from females collected at black lights in Lincoln, Me., Strat­ ford, N. H.; and Manistique, Mich., but the Syngrapha rectangula (Kirby) larvae on artificial diet fed very little and died as first or second instal" larvae. Larvae were provided by A. H. Rose, Canada Department of .Figure 216 Forestry, Sault Sainte Marie, Ontario. Plusia rectangula Kirby, 1837, p. 306. Autographa rectangula: Ottolengui, 1902, p. 68 and mortuorum (Guenee), 1852, p. 353. Syngrapha angulidens (Smith) Syngrapha rectangula: Hampson, 1913, p. 423. Autographa rectangula: Barnes and Mc­ Figure 217 Dunnough, 1917, p. 83. Plu.sia angulidens Smith, 1891, pp. 111, 112. Autographa rectangula race nargenta Ottolen­ gui, 1919, p. 122. Autographa angulidens: Dyar, 1902, p. 81. Syngrapha rectangula: McDunnough, 1944, p. Syngrapha angulidens: Hampson, 1913, pp. 427, 188. 428. Male genitalia (fig. 77}.-Similar to alias, ex­ Autographa angulidens: Barnes and Mc­ cept that rectangula does not have the sacculus Dunnough, 1917, p. 83. produ.ced mesad as a broad triangular projec­ Syngrapha angulidens var. excelsa (Ottolengui) tion. 1902, pp. 71, 72.-McDunnough, 1944, pp. 191, Female genitalia (fig. l33).-Same as for alias 192. with the following exceptions: The small projec­ Male genitalia (fig. 76).-Valve with ampulla tion to the right at the base of the ostium arising from broad base, tapering to a pointed bursae is membranous; in all specimens studied apex, the length and shape of ampulla appar­ the ductus bursae enters the corpus bursae ently variable; juxta produced to a strong, about one-fourth of the total length of corpus sharp point apically, aedeagus with an apical PLUSIINAE OF AMERICA NORTH OF MEXICO 49 cornutus about one-third as long as aedeagus, from 25 to 28 in number; seta SV-2 on abdomi­ and with a much smaller basal cornutus. nal segment 1; seta SV-2 on meso- and meta­ Female genitalia (fig. 126).-Similar to celsa thorax; mesothol'ax with pinaculum of D-2 but with membranous projection off base of equidistant from D-1 and SD-2; pinacula and ostium bursae shorter and produced laterad, area surrounding base of seta dark; raduloid of not directed caudad; overall size of genitalia hypopharynx absent. smaller than celsa. Type.-A male syntype of celsa in the Na­ Type.-Lectotype female, National Museum tional Museum of Natural History has been of Natural History (angulidens), designated by selected and labeled and is presently desig­ Todd (1972). Female, American Museum of Nat­ nated as the lectotype: "Plusia celsa Hy. Ed., ural History (e.tcelsa). Type/Collection Brklyn. MusJType No. 34106, Type locality.-Colorado (angulidens). Jeffer­ U.S.N.M./Col. B. Neumoegen/o genitalia on son, N. H. (e.l'celsa). slide, Aug. 19, 1936, J.F.G.C. #535." Female, Host plant.-Abies lasiocarpa, Ross and Ev­ American Museum of Natural History (sierrae). ans (1957). Type locality.-Mount Hood, Oreg. (celsa). Range (map 18).-Nova Scotia, south to New Lake Tahoe, Calif. (sierrae). Hampshire; Rocky Mountains in Canada, south Host plant.-Abies grandis, A. lasiocarpa, to Arizona. Crumb (1956); Picea glauca, Tsuga heterophylla, Ross and Evans (1957); Picea engelmanii, Pinus monticola, Prentice (1962). Syngrapha celsa (Edwards) Range (map 13).-Alaska; south to California and Utah. Figure 218 Comments.-The adult specimens studied Plu..sia celsa Edwards, 1881, p. 101. agreed with the type of "sierrae" examined in Autogmpha celsa: Dyar, 1902, p. 81. the collection of the American Museum of Nat­ Syngrapha celsa: Hampson, 1913, pp. 422, 423. ural History and celsa in the National Museum Autographa celsa race sie1'?'ae Ottolengui, 1919, of Natural History. Larvae of this species were p.123. collected by investigators in the field in the Canada Department of Agriculture, British Col­ Syngrapha celsa: McDunnough, 1944, p. 189. umbia. Larvae were reared to adults for identi­ Male genitalia (fig. 61).-Similar to anguli­ fication and association with larval specimens. dens, but with ampulla on valve shorter on all The specimens were obtained from D. A. Ross specimens examined, abruptly narrowed at and B. A. Sugden. apex, or stuhby with tiny, apical wart or spine; juxta oply slightly produced to an apical point at best; apex of aedeagus heavily spiculate, but only weakly spiculate and more granulose in Syngrqpha montana (Packard) angulidens. Figure 219 Female genitalia (fig. 124).-Ostium bursae about equal length to the ductus bursae, we1l Plusia montana Packard, 1874b, p. 313. sclerotized, tapering toward the base from Autographa montana: Barnes and Benjamin, which is produced a membranous, fingerlike 1923, p. 213. process to the right and directed caudad; the Syngrapha montana: Ferguson, 1955, pp. 23-27. ductus bursae sclel'otized and strigate, entering Male genitalia (fig. 65).-Similar to mic?'o­ the corpus bursae on right side about half way gamma, but with clavus tapering to a narrowly from apex; corpus bursae minutely granulose, rounded apex in all specimens examined; juxta becoming more concentrated on the extended, without sclerotized, apical spine; aedeagus only much darkened apex: ductus seminalis origi­ slightly spiculate apically, and with apical cor­ nates on dorsal side of the rounded apex. nutus longer and thinner than that of micro­ Larvae.-Integument thickly covered with gam?nCL. dark microspines, which form middorsal and Female genitalia (fig. 129).-Ostium bursae subdorsal longitudinal bands; crochets range small, about as long as wide; ductus bursae 50 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE straight, strigate, but mostly membranous, en­ Host plant.-Unknown in North America; in tering corpus bursae on the right side well Europe feeds on Salix myrtilloitks and S. ros­ beyond apex at about the middle of corpus marinifolia, Kostrowicki (1961). bursae; apex of corpus bursae granulose and Rang~ (map 12).-Nova Scotia west to British darkly pigmented around base of ductus semin­ Columbia and north to Yukon Territory; south aliso to Maine, New Hamphsire, Michigan, Wiscon­ Type.-Lost (Ferguson, 1955). sin, and Colorado. Type locality.-White Mountains, N. H. Comments.-Parental females of laboratory Host plant.-Unknown. cultures were collected while they flew during Range (map 19).-Nova Scotia to Saskatche­ daylight hours over sphagnum-heath bogs in wan; Maine, New Hampshire, and Wisconsin. Schoolcraft County, near Manistique, Mich. Al­ though adults were common in these situations, they were elusive and difficult to capture. Syngrapha mrcmgamma :(I;J;iib'ner)

Figure 220 Syngrapha alticola (Walker) Noctua microgamma Hiibner, 1823, p. 151, figs. 698,699. Figure 221 Plusia mwrogamma: Treitschke, 1823, p. 198. Plusia alticola Walker, 1857, p. 912. Syngrapha microgamma: Dod, 1910b, p. 118. Plusia alticola Walker, packard, 1974a, p. 554. Autographa microgamma: Strand, 1916, p. 47. [= hochenwarthi] Autographa montana (Packard), 1874b, p. 313.- Plusia ignea Grote, 1863, p. 274.-Grote, 1874a, Barnes and Benjamin, 1923, p. 213. p.31. Syngrapha microgamma: McDunnough, 1944, p. Syngrapha alticola (Walker), Ottolengui, 1902, p. 184. 63. [= tkvergens] Male genitalia (fig. 18).-Ampulla on valva Plusia alticola: Hampson, 1908, p. 106. broad basally, then narrow and tapered to a point in apical third; clavus short and broad; Caloplusia alticola: Hampson, 1913, p. 407. juxta with a distinct, sclerotized, sharp, apical Syngrapha alticola: Dod, 1913, p. 241. spine; aedeagus spiculate apically, with a Syngrapha alticola (Walker), McDunnough, strong apical cornutus and a small basal cornu­ 1938, p. 112. [= devergens] tus. Syngrapha alticola: McDunnough, 1944, p. 182. Female genitalia (fig. 138.).-Ostium bursae Male genitalia (fig. 75).-Valve with ampulla small about as wide as long, sclerotized but arising from a broad base, tapering to near incompletely on venter; posterior two-thirds of costal mal"gin, with a small, sharp, terminal ductus bursae sclerotized, swollen on the right spine projecting just over costal margin; juxta cephalad before becoming strigate as it enters with a distinct, sclerotized, sharp, apical spine; the corpus bursae on the left side; an additional aedeagus without cornuti. membranous bulge is over the swollen area on Female genitalia (fig. 128).-Ostium bursae the ductus bursae; the apex of the corpus bur­ subcylindrical, slightly longer than wide, sae shagreened and darkly pigmented. weakly sclerotized; ostium with a thick, circu­ Larvae.-Integument with microspines ex­ lar, membranous, pouchlike structure on each cept on the spiracular line; crochets range from side; d'lctus bursae mostly membranous and 24 to 26 in number; interval between the pina­ granular, except for some sclerotization near cula of setae V-Ion abdominal segment 4 one­ entrance to corpus bursae, bending somewhat half the interval between V-I and SV-2; other­ to the left in the middle, about one-half the wise similar to celsa, which also lacks a radu­ width and twice the length of ostium bursae, loid. widening slightly where it enters corpus bursae Type.--[?]. on the right side; corpus bursae somewhat J­ Type locality.--[?]. shaped from ventral aspect, with the cephalic PLUSIINAE OF AMERICA NORTH OF MEXICO 51 end tapering to a point, and the apex of uni­ bands including setae D-l; interval between form diameter, granulose, rounded caudad, the pinacula of setae V-Ion abdominal seg.. with the ductus seminalis arising in the center ment 4 about subequal to interval between V-I of apex. and SV-2; all setae black; head brown, some­ Type.-Male, British Museum (Natural His­ what mottled; otherwise similai" to celsa, which tory) (alticola). [?] (devergens). also has the raduloid absent. Type locality.-Rocky Mountains (alticola). [?] Type.--{?] British Museum (Natural History) (devergens). (parilis). Female, British Museum (Natural His­ Host plant.-Unknown. tory) (quadriplaga). Range (map 14).-Manitoba west to Alaska, Type locaiity.-{?] (parilis). Fraw Islands, Arc­ British Columbia; south to Montana, Wyoming, tic North America (quadriplaga). and Colorado, with isolated records in Sierra Host plant.-Unknovm. Nevadas of California. Range (map 11).-Arctic south to Labrador in the East, and Alberta and British Columbia in the West. Syngrapha panlis (Hubner) Comments.-The larvae were borrowed from the Canadian National Collection. These were Figure 222 the same specimens used by MacKay (1972) in her a-=:scription of the larval form of parilis. Noctua Parilis Hubner, [1808-1809], pI. 90, fig. Based on characters examined in this study, 422. the larvae of parilis, rectangula, alias, and celsa Autographa parilis: Hubner, 1816, p. 251. could not be separated. Plusia parilis: Boisduval, 1829, p. 251. Plusia qUadriplaga Walker, 1857, pp. 911, 912.­ Smith, 1893, p. 256. Syngrapha ignea (Grote) Syngrapha parilis: Dyar, 1902, p. 82. Autographa parilis: Hampson, 1913, pp. 404, Figure 223 405. Plu,cn:a ignea Grote, 1863, p. 274. Syngrapha parilis: McDunnough, 1916, p. 400. Plusia ignea: Grote, 1874a, p. 31. [= alticola] Male genitalia (fig. 67).-Similar to alticola, Syngrapha ignea: Ottolengui, 1902, p. 76. but apex of ampulla does not appear to have a Caloplusia, ignea: Hampson, 1913, pp. 409, 410. distinct spine, and the juxta is produced api­ cally into a strong point, but not a distinct Syngrapha ignea: McDunnough, 1916, p. 400. sclerotized spine. Syngrapha ignea: McDunnough, 1938, p. 113. [= Female genitalia (fig. 152).-Ostium bursae lwchenwarthi] short, about as wide as long; ostium unpro­ Caloplusia ignea: McDunnough, 1944, pp. 178, tected; initial one-half of ductus bursae mem­ 179, and val'. simulans McDunnough. branous and granulose, the extreme caudal Male genitalia (fig. 63).-Valve rounded at portion curved sharply to the right, then ab­ apex; ampulla thick, sharply pointed and ruptly widened, then curved back left and be­ strongly curved, apex projecting ventrocaudad; comes wider, strigate, and sclerotized as it en­ juxta with large, sclerotized, apical spine; ae­ ters the corpus bursae ventrad toward the left deagus with a small, sclerotized, basal, spinose side; the corpus bursae dorsoventrally flattened plate, and a larger, rounded apical plate, from and approximately L-shaped in ventral aspect, which project several small cornuti. rounded on the cephalic end, and tapered to a Female genitalia (fig. 19).-Ostium bursae blunt point at apex, from which the ductus about twice as long as wide, sc1erotized, and seminalis arises; corpus bursae uniformly, mi­ somewhat strigate; ostium unprotected; ductus n utely granulose. bursae mostly membra.nous and granulose ex­ Larvae.-Crochets range from 21 to 23 in cept just prior to entering corpus bursae where number; microspines of integument present, it becomes strigate and sclerotized as it enters except on spiracular line and small longitudinal the corpus bursae on the right side; corpus 52 TECHNICAL BULLETIN 1567, U.~. DEPARTMENT OF AGRICULTURE bursae lightly shagreened with the caudoven­ sae, dorsoventrally flattened, sclerotized, with tral edge of the apex heavily pigmented. the caudal end on the venter folded and project­ Type.-[?] (ignea). Male, Canadian National ing cephalad, and dorsad with a granulose plate Collection, Ottawa, Ontario (simulans). extending caudad well beyond ostium. The duc­ Type locality.-Pike's Peak (ignea). Rocky tus bursae enters the corpus bursae on the Bay, Quebec (simulans). apex, ventrad and to the left of the origin of the Host plant.-Unknown in North America; ductus seminalis. The corpus bursae is elongate hosts for European hochenwarthi (Hochen­ but bulbous on the cephalic end. It is membra­ warth) include Umbelliferae spp., Kostrowicki nous with a darkly pigmented patch on the (1961); Antennaria dioica, Carex filifolia, Castil­ dorsal surface near the middle. leja lemmonii, Gentiana newberryi, Mimulus The larvae of C. putnami and contexta have primuloides, Penstemon confertus, Salix sp., been examined and no stable diagnostic charac­ Vaccinium sp., Tietz (1972). ters could be found to separate them. They are Range (map 13).-Newfoundland to Alaska; the only group having uniordinal crochets. The south to California and Colorado. crochets range from 15 to 25 in number. The mandible is unusual, having the ventral edge incurved, making a deep pocket near this edge on the mesal surface, an apparent modification Chrysaspidia Hiibner for feeding on grasses. Although the male genitalia tend to show a close relationship to Autographa spp., the setal Generotype.-Noctua festucae Linnaeus, 1758. patterns of Chrysaspidia lar.vae are very simi­ Designated by Grote, 1895a, p. 417. lar to Syngrapha spp. Seta SV-2 is present on Chrysaspidia Hiibner, 1821, p. 252. the abdominal segment 1, SV-2 is absent on the Euchal.cia Dyar (nec Hiibner), 1902, p. 81. meso- and metathorax, and SV-l and SV-2 Phytometra Hampson (nec Haworth), 1913, p. have their pinacula partially fused on abdomi­ 452 (partim). nal segments 1-4. On the abdominal segment 4, Palaeoplusia Hampson, 1913, p. 581. SV-2 is twice as far from V-I as each of the V­ I's are to each other. On the mesothorax, D-2 is Chrysaspidia: McDunnough, 1944, p. 208. equidistant from D-1 and SD-2. Microspines This is a very homogeneous group for the are present on the integument. The raduloid of most part on the basis of larval characters and the hypopharynx is not present as is the caE)e male and female genitalia. The males of put­ for certain of the Syngrapha species. nami, venusta, and nichollae cannot be sepa­ rated by the genitalic characters we have ex­ amined. No differentiating structures of the female genitalia of putnami and venusta could KEY TO THE SPECIES BASED ON MALE GENITALIA be found. The species do show characteristic Costa produced apically into a di:;tinct spine (fig. 84a) color patterns on the forewings, which enables ...... •...... contexta one to identify most species (see photographs). Costa not produced apically into a spine (fig. 82a) ...... •...... putnami 1'he male genitalia are similar to the species venu8ta of Autographa, having long filiform clavi. The nichollae tegumen is less than one-half as long as the No structures could be found on the male genitalia which valve. The ampullae of the valves extend be­ would differentiate the latter three species. Refer to the yond the costal margin, are not spined apically, plates of the adults (fig. 225, 227, 226). and are generally wider than the ampullae of the speciei.; of Autographa. The vinculum is KEY TO THE SPECIES BASED ON FEMALE relatively long and bluntly pointed. The juxta is GENITALIA strongly produced at the apex. The aedeagus is thick, about as long as the valve, and with a 1. Dorsal, sclerotized, pigmented area of corpus bursae somewhat circular (fig. 151)...... nichollae short cornutus basad. Dorsal, sclerotized, pigmented area of corpus hursae The female genitalia has a wide ostium bur­ elongate (fig. 155) ...... 2 PLUSIINAE OF AMERICA NORTH OF MEXICO 53 KEY TO THE SPECIES BASED ON FEMALE GENITA LlA-Continued Phytometra putnami: Hampson, 1913, pp. 509, 510. 2. Pigmented area approximately one-third length of Autographa putnami: McDunnough, 1916, p. corpus bursae; corpus bursae straight (fig. 154) 400...... contexta Chrysaspidia putnami: McDunnough, 1944, pp. Pigmented area approximately one-half length of cor­ pus bursae; cephalic half of corpus bursae curved 208,209. ventrad (fig. 155) ...... putnami Male genitalia (fig. 82).-Typical for the ge­ venusta nus, refer to the generic description. [For differentiation of putnami and venusta refer to Female genitalia (fig. 149).-Very similar to plates of the adults (figs. 225, 227).] venusta having the elongate, narrow, pig­ mented area on corpus bursae about one-half as long as the corpus bursae. Otherwise, the Chrysaspidia contexta (Grote) female genitalia are typical for the genus. Refer to the generic description. Figure 224 Larvae.-Refer to the generic description. Type.-Male, British Museum (Natural His­ Plusia cantexta Grote, 1873b, p. 193. tory). E1l..cho.lcia contexta: Dyar, 1902, p. 81. Type locality.-Albany, N. Y. Phytometra cantexta: Hampson, 1913, pp. 510, Host plant.-Grasses, Tietz (1972); sedges and 511. Sparganium sp., Forbes (1954). Autographa contexta: McDunnough, 1916, p. Range (map 17).-Nova Scotia to Alberta; 400. Maine to Rocky Mountains, south to Virginia in Chrysaspidia cantexta: McDunnough, 1944, pp. the East, to Colorado in the West. 208,209. Comments.-Larvae of this species responded Male genitalia (fig. 84).-Typical for the genus poorly to the artificial diet. The parental female except that the valve is produced into an apical was t· ken in Schoolcraft County, Manistique, spinelike projection. Refer to the generic de­ Mich. scription. Female genitalia (fig~ 154).-Typical for the genus except that the pigmented patch on cor­ pus bursae is elongate and narrow and only Chrysaspidia nichollae (Hampson) about one-third as long as corpus bursae. ~efer to the generic description. Figure 226 Larvae.-Refer to the generic description. Type.-Male, British Museum (Natural His­ Phytometra nichollae Hampson, 1913, p. 510. tory). Autographa nichollae: McDunnough, 1916, p. Type locality.-Albany, N. Y. 400. Host plant.-Grasses, Tietz (1972). Chrysaspidia nichollae: McDunnough, 1944, pp. Range (map 16).-Quebec and Ontario; Maine, 208,209. New York west to the Dakotas. Male genitalia (fig. 85).-Typical for the ge­ Comments.-The larvae and associated pa­ nus; refer to the generic description. rental female were borrowed from Canadian Female genitalia (fig. 151).-Pigmented area National Collection, Ottawa, Ontario. on corpus bursae circular, otherwise typical for the genus; refer to the generic description. Type.-Female, British Museum (Natural Chrysaspidia putnami (Grote) History). Type locality.-Pass D. Hope, British Colum­ bia. Figure 225 Host plant.-Unknown. Plusia putnami Grote, 1873a, p. 146. Range (map 17).-Alaska to northern Califor­ Eitchalcia putnami: Dyar, 1902, p. 81. mao 54 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE Comments.-We are unable to distinguish be­ Plusia striatella Grote 1873b. p. 194.-Smith, tween nichollae and putnami on the basis of 1893, p. 248. color patterns or male genitalia, but can sepa­ Euchalcia venusta: Dyar, 1902, p. 81. rate them by comparing female genitalia. In a Paleoplusia venusta: Hampson, 1913, p. 582. recent publication (Eichlin, 1975), putnami was Ch1ysaspidia venusta: McDunnough, 1944, pp. said to occur in California. Subsequently, it was 208,209. learned that this statement is incorrect; only Male genitalia (fig. 86).-Typical for the ge­ nichollae occurs in the Pacific coastal regions. nus; refer to the generic description. Apparently, the ranges of these two species do Female genitalia (fig. 155).-See discussion of not overlap. The type specimen of the form female genitalia under putnami. "mendocinensis" Strand was not seen, but ~'J}e.-Male, British Museum (Natural His­ should, in fact, be considered a color form of tory) (venusta). Syntypes not seen by us, British nichollae and not putnami a5 with previous Museum (Natural History) (striatella). authors, based on its type locality in Mendocino Type loca.lity.-Orilla, Canada (venusta). County, Calif. Washington, D.C., New York, Canada (stria­ teila). Chrysaspidia venusta (Walker) Host plant.-Swamp sedges and grasses, Forbes (1954). Figure 227 Range (map 17).-Nova Scotia to Alberta; Plusia venusta Walker, 1865, p. 836. south to District of Columbia and the Dakotas.

PHYLOGENY

Richards (1932), in his arrangement of the additional characters, also lack tibial spines noctuid subfamilies, based on skeletal morphol­ while subfamilies more advanced than plusiines ogy of the tympanum, placed the Plusiinae as a have tibial spines, indicates Abrostola is a pri­ derivative of the tropical subfamily Stictopteri­ mitive genus in the subfamily. Males lacked a nae. Forbes (1954) stated that stictopterines are clavus. Forbes states that the clavus of male very closely related to plusiines and perhaps genitalia is almost limited to the more primitive the two groups should be combined. Larvae of Agrotinae. This character must have evolved the Stictopterinae have many characters in independently in the Plusiinae. common with members of the plusiine genus The ancestral larval form had ventral prolegs Abrostola, on abdominal segments 3-6, a character com­ Forbes (1954), utilizing both adults and lar­ mon to the large majority of noctuids, and vae, and Crumb (1956), working with noctuid therefore considered to be primitive. Crochets larvae, agreed that the Catocalinae were were biordinal, which is typical of all but one closely related to the Plusiinae and exhibited small genus, Ch?ysaspidia, in the extant Plusi­ characters more advanced than the plusiines. inae. This biordinal condition is rare in the We are postulating (figs. 156 and 157) that the other noctuid subfamilies, but occurs in the ancestor of the Plusiinae had the following more primitive subfamily, Cuculliinae, and ap­ characters: The adult had the alula of the pears to represent another independently tympanum with two small hoods. Richards evolved specialization in the Plusiinae. Seta (1932) says that the Plusiinae alone possesses SV-2 was absent on the meso- and metathorax. two hoods on the tympanum, and in the genus The presence of SV-2 is found only in plusiines Abrostola, they are smaller than in the other in the Noctidae, and is confined to a group of species of Plusiinae, indicating that Abrostola is plusiines considered to be advanced, based on the most primitive group in the subfamily. other characters of the larvae and adults; the Adults were without spines on the tibiae. The absence of SV-2 on the meso- and metathorax fact that more primitive subfamilies, based on is therefore considered to be a primitive condi­ PLUSIINAE OF AMERICA NORTH OF MEXICO 55 tion in the plusiines. Seta SV-2 was absent on talia particularly places it closest to the more abdominal segment 1; pinaculum of seta D-2 advanced and specialized Mouralia tinctoides closer to D-1 than SD-2 on mesothorax; pinacu­ line. Mouralia tinctoides is the most specialized la of setae SV-1 and SV-2 separated on ab­ species in the Abrostolini based on several mod­ dominal segment 2. The setal relationships are ifications of the genitalia. typical of the subfamilies more primitive than A common ancestor from the tinctoides line the plusiines. Raduloid on the hypopharynx gave rise to the Argyrogrammini and Autogra­ was present, and found in other extant noctuids phini lines. The mandibles of the larvae became in only four species of Lithocodiinae (Acontiinae modified with ribs two and three terminating in of Forbes), considered to be a more primitive processes before reaching the cutting margin, subfamily and in Phyprosopus callitrichoides and the prolegs were reduced on abdominal Grote (filament looper), a species whose place­ segments 3 and 4. A split occurred in this line ment is uncertain, Forbes putting it in Plusi­ with the Argyrogrammini being the more pri­ inae, Crumb in Herminiinae. mitive, having the prolegs of the larvae reduced The plusiines are divided into three tribes, to peglike structures lacking crochets, but to­ based chiefly on the condition of the prolegs of tally absent in the more advanced Autogra­ the larvae. In Abrostolini, all the prolegs are phini line. present though slightly reduced on abdominal The Argyrogrammini contains three genera. segment 3, which already indicates the tend­ The ancestral larval form of this line had the ency toward reduction of the prolegs and which first two pairs of prolegs vestigial, and terminal is progressively more pronounced in the other processes on the ribs of the mandible; the adult two tribes. The Argyrogrammini have the pro­ males had long, lateral tufts of scales caudad on legs reduced to peglike v('stiges on abdominal the abdomen, and a long, filiform clavus on the segments 3 and 4. However, according to Ichi­ valve. In the genus Argyrogramma, the larvae nose (1962), a species belonging to this tribe, have retained the mandibular processes, and Podioplusia peponis (Fab.), has minute prolegs the males retained the lateral, abdominal, setal on abdominal segments 3 and 4 as a first-instar tufts, indicating that this is the most primitive larva, but not on subsequent instars. This is a genus in the tribe. The females have the unu­ further indication that the reduction and. even­ sual condition of the ductus seminalis arising tual loss of prolegs was an evolutionary trend from the fundus of the corpus bursae, a condi­ in the Plusiinae. The Autographini have no tion found in only one other species, Autoplusia vestige of prolegs on abdominal segments 3 and illustrata. Three species are in Argyrogramma. 4. Trichoplusia, with two species, lost the mandi­ The prototype species gave rise to the Abros­ bular processes of the larvae, and has a highly tolini line. This most primitive line retained all specialized ductus bursae in females, which is of the ventral prolegs on the larvae. The Abros­ longer than the total length of the abdomen. In tolini consists of the genera Abrostola and t.he monotypic genus Pseudoplusia, the larvae Mouralia, the larvae of which are similar to retained the primitive mandibular character, each other. The most sensitive indices to the but the female corpus bursae has a greatly relationships among the five species in the tribe elongated apex, almost twice the length of the seem to be the characters of the female geni­ main body of the corpus bursae, and the males talia (see description of female genitalia for lost the lateral tufts of long scales on the each species in the Abrostolini group). The abdomen. Both characters indicate that this urentis line was determined to be the most genus is the most advanced of the Argyrogram­ primitive, based on the fact that this line re­ mini. tained more of the ancestral characters than In the Autographini, the prolegs were com­ the other four species. Some specialization of pletely lost on abdominal segments 3 and 4. The both the female and male genitalia indicates first major branching of the Autographini line, that parvala and microvalis are an offshoot of occuned with the migration of seta SV-2 to­ the urentis line. Abro.c;tola ovalis shows further ward SV-1, to the point where their pinacula modifications of the genitalia, the female geni­ are partially fused on abdominal segment 2. In 56 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE the more primitive line, where the pinacula of therefore, interpreted the shorter clavus as a the two setae are separated, setae SV-l, SV-2, specialization in this line. The tendency in this and V.··l are grouped together on abdominal line was toward an overall reduction in the size segment 4. This condition is considered primi­ of the male genitalia. Species of Pseudeva and tive, because the arrangement of the setae is Adeva have the clavus reduced to a mere the same as it is in the Argyrogrammini, where raised, setose ridge, which indicates the species the three setae are positioned around the ves­ in these two genera are more highly specialized tigial prolegs on abdominal segments 3 and 4. than the species of Polych7ysia. An examina­ The degree to which the two V-I setae have tion of the larvae of Pseudeva and Adeva might migrated away from the SV-group and toward help to clarify the relationships among- the each other on the venter is apparently a factor three genera. of how long prolegs have been lost in the The Ch7ysanympha and EosphoTopte7Yx evolutionary history of a particular line. This is lines are placed near the three preceding gen­ based upon the additional fact that in larvae of era because of the general similarities of the other subfamilies, where the prolegs have been male genitalia and upon data from the tym­ lost, this same tendency of the V-l setae to panal studies of Richards (1932). This arrange­ migrate mesad has been observed. Also, on ment is tentative pending studies of the larvae. those segments where prolegs are assumed to The next major modification leading to the have existed on lepidopteran progenitors, these Auiogmpha-Syngmpha line is indicated by the two setae are close together. On one species evolution of seta SV-2 on abdominal segment 1 seen, the two V-I setae migrated mesad, occur­ of the larvae. The Alltogmpha line retained the ring close together on abdominal segment 1 and mandibular processes on the larvae, whereas becoming progressively farther apart on seg­ these processes were lost in the SyngTapha­ ments 2-4. Mainly on this basis, Alltoplusia is Chr1.jsaspidia line. considered most primitive of this line, followed The Autogmpha line, which contains 16 spe­ by Rachiplllsia, then Plusia and Allagmpha. cies, is a homogeneous group with similar larval The placement of Autoplusia and Rachiplusia forms as well as similar genitalia in the males is complicated by the spination found on the and females. The Autographa line is divided tibiae, a character of the most advanced plu­ into two major branches based on the presence siines. In Autoplusia, spines occur only on the or absence of seta SV-2 on abdominal segment hind tibiae, this line giving rise to Rachiplus1:a 1. The more primitive branch, which lacks SV-2, in which all tibiae are spined. The other charac­ is subdivided into three groups. ters of adults and larvae, which are the same as The most primitive of these three species mentioned for this evolutionary line, tend to groups is so designated, b~cause the larvae justify its placement here, assuming that tibial have retained the mandibular processes on spining arose independently in the Autoplu..sia­ both ribs 2 and 3. The larvae of the second Rachiplusia line. branch lost the process on rib 3, and larvae The genus Polych?Ysia is an offshoot of the comprising the third branch lost both processes, main line in which the larvae have the pinacula indicating that the biloba group is the most of SV-l and SV-2 setae partially fused on ab­ highly specialized of these three lines. dominal segment 2. The setal arrangement sub­ The most advanced major branch of the Au­ ventrad on abdominal segment 4 is primitive as togmpha line, which has evolved seta SV-2 on with the group just discussed, and the raduloid abdominal segment 1, gave rise to the Syngra­ on the larval hypopharynx has been lost. pha-Cll1ysaspidia lineage. The latter is charac­ The Pseudeva and Adeva lines probably arose terized by the loss of the mandibular processes from the Polych?Ysia line since all three have in the larvae. male genitalia which in general are structur­ The line comprising the genus Syngmpha ally quite similar. The clavi of Polychrysia spe­ includes all species with tibial spines, at least cies are shorter than in species of more primi­ on the hind tibiae. Syngrapha ottolenguii is tive genera and are shorter than in the species considered to be the most primitive member of of the more advanced genus, AutognLpha. We, the genus because the male genitalia are more PLUSIINAE OF AMERICA N01:

tropical fauna. We hope, however, that this species will be useful as a frame·,rrork for such proposal for the systematic relationships of the needed future work.

CHECKLIST OF NORTH AMERICAN PLUSIINAE

ABROSTOLINI, new tribe AUTOGRAPHA Hubner ABROSTOLA Ochsenheimer precationis (Guenee}-Temperate North America, urentis Guenee-General in temperate North Amer­ east of Rocky Mountains ica bimaculata (Stephens}-Nearctic Boreal parvula Barnes and McDunnough-Arizona and Cali­ sansoni Dod-Western Nearctic Boreal fornia (low deserts) rubida Ottolengui-Nearctic Boreal micro valis Ottolengui-Texas and Arizona biloba (Stephens}-Temperate and subtropical North ovaHs Guenee-New England and Mid-Atlantic America States californica (Speyer}-Western North America MOURALIA Walker pseudogamma (Grote}-Nearctic Boreal tinctoides (Guenee}-Neotropics and Subtropics v-alba Ottolengui-Western Nearctic Boreal ARGYROGRAMMINf, new tribe pasiphaeia (Grote}-Oregon and California ARGYROGRAMMA Hubner flagellum (Walker}-Nearctic Boreal verruca (Fabricius)-Neotropics, Subtropics, and At­ metallica (Grote}-Western Nearctic Boreal lantic coastal plain speciosa Ottolengui-Vancouver, British Columbia basigera (Walker}-Atlantic and gulf coastal plains, labrosa (Grote}-Central coast of California sUbtropical Mexico corusca (Strecker}-Western Nearctic Boreal TRICHOPLUSIA MeDunnough mappa (Grote and Robinson}-Nearctic Boreal ni (Hubner}-General in Western Hemisphere south ampla (Walker}-Nearctic Boreal of Arctic zones SYNGRAPHA Hubner oxygramma (Geyer}-Subtropical and eastern mild ottolenguii (Dyar}-Arctic North America Temperate zones altera (Ottolengui}-Eastern Nearctic Boreal octoscripta (Grote}-Nearctic Boreal PSEUDOPLUSIA McDunnough epigaea (Grote}-Nearctic Boreal includens (Walkerl-Tropics, Subtropics, and eastern selecta (Walker}-Nearctic Boreal mild Temperate zones viridisigma (Grote}-Nearctk Boreal AUTOGRAPHINI, new tribe orophila (Hampson}-Western Nearctic Boreal AUTOPLUSIA McDunnough snowi (Edwards}-Boreal New Mexico olivacea (Skinner}-Southern coastal California sackeni (Grote}-Boreal in Rocky Mountains illustrata (Guenee}-Neotropics, subtropical Florida lui a Strand-Western Nearctic Boreal egena (Gueneel-Neotropics, Subtropics borea (Aurivillus}-Artic North America RACHIPLCSIA Hampson diasema (Boisduval}-Holarctic ou (Guenee)-Subtropical and temperate North falcifera (Kirby}-Temperate North America America u-aureum (Guenee}-Nearctic Boreal, Scandinavia PLUSIA Ochsenheimer interrogationis (Linneaus}-Holarctic balluca (Geyer}-Nearctic Boreal surena (Grote)--Eastern Nearctic Boreal ALLAGRAPHA Franclemont alias (Ottolengui}-Nearctic Boreal aeroides (Grote}-Nearctic Boreal abstrusa n. sp.-Eastern Nearctic Boreal aerea (Hubner}-Nearctic, east of Rocky Mountains cryptica n. sp.-Eastern Nearctic Boreal POL YCHRYSIA Hubner rectangula (Kirby}-Nearctic Boreal moneta (Fabricius}-Holarctic angulidens (Smith}-Nearctic Boreal morigera (Edwards}-Western Nearctic Boreal celsa (Edwards}-Nearctic Boreal ADEVA McDunnough montana (Packard}-Eastern Nearctic Boreal albavitta (Ottolengui}-Mountains and deserts of microgamma (Hubner}-Nearctic Boreal Western United States alticola (Walker}-Western Nearctic Boreal PSEUDEVA Hampson parilis (Hubner}-Holarctic palligera (Grote)-Mountains of Western United ignea (Grote}-Nearctic Boreal States CHRYSASPIDIA Hubner purpurigera (Walker}-Nearctic Boreal contexta (Grote}-Eastern Nearctic Boreal CHRYSANYMPHA Grote putnami (Grote}-Nearctic Boreal formo!>a (Grote}-Nearctic Boreal nichollae (Hampson}-Coastal northwestern United EOSPHOROPTERYX Dyar States, British Columbia thyatyroides (Guenee}-Nearctic Boreal venusta (Walker}-Mostly eastern Nearctic Boreal PLUSIINAE OF AMERICA NORTH OF MEXICO 59 LITERATURE CITED

AURIVILUUS, C. CUNNINGHAM, H. B. 1890. GRONLANDS INSEKTFAUNA, 1. LEPIDOP­ 1969. See EICHLIN, T. D., and H. B. CUN­ TERA, HYMENOPTERA. Bihang Till K. NINGHAM. Svenska Vetenskaps-Akademiens Handlin­ DOD, F. H. W. gar, Band 15, Afd. IV., No.1: pp. 1-34, 3 1910a. A NEW AUTOGRAPHA FROM THE ALBERTA plates. ROCKIES. The Canadian Entomologist BARNES, WM., and J. McDUNNOUGH. 42(11): 349. 1916. SOME NEW RACES AND SPECIES OF NORTH AMERICAN LEPIDOPTERA. The Canadian 1910b. NOTES OF CA PTURES, LEPIDOP- Entomologist 48: 225--226. TERA. Fortieth Annual Report of the En­ tomological Society of Ontario, pp. 114­ 1917. CHECK LIST OF THE LEPIDOPTERA OF BO­ 121. REAL AMERICA. Herald Press, Decatur, Ill., pp. i-viii, 1-392. 1913. FURTHER NOTES ON ALBERTA LEPIDOP­ BARNES, W., and F. H. BENJAMIN. TERA. The Canadian Entomolo~ist 45: 1923. NOTES ON ABERRATIONAL NAMES OMITTED 236-244. FROM THE BARNES AND MCDUNNOUGH CHECK-LIST (LEPID.). The Canadian Ento­ 1915. FURTHER NOTES ON ALBERTA LEPIDOP­ mologist 55: 217-213. TERA. The Canadian Entomologist 47: BENJAMIN, F. H. 122-134. 1923. See BARNES, W., and F. H. BENJAMIN. DUFAY, C. 1970. INSECTES LEPIDOPTERES NOCTUIDAE PLU­ 1932. NEW SYNONYMY (LEPID. PHALAENIDAE SIINAE. Faune De Madagascar 31: 1-198. AND LASIOCAMPIDAE). Proceedings of the DUPONCHEL, P. A. J. Entomological Society of Washington 34: 1829. HISTORIE NATURELLE DES LEPIDOPTERES, 124-127. ETC. Paris, 7: 71. DYAR, H. G. 1933. NOTES ON PHALAENIDAE FROM ARCTIC 1902. A GENERIC SUBDIVISION OF THE GENUS AMERICA, SUPPLEMENTED WITH MATERIAL PLUSIA. Journal of the New York Entomo­ FROM HIGH ALTITUDES. The Pan-Pacific logical Society 10: 79-82. Entomologist 9(2): 5~2. BERGER, R. S. 1903. A LIST OF NORTH AMERICAN LEPIDOPTERA 1968. SPECIF1CITY OF THE CABBAGE LOOPER SEX AND KEY TO THE LITERATURE OF THIS OR­ A'ITRACTANT. Joul'T)al of Economic Ento­ DER OF INSECTS. United States National mology 61(2): 45~54. Museum Bulletin, No. 52: i-xix, 1-723. BIRCH, M. EDWARDS, H. 1970. PERSUASIVE SCENTS IN MOTH SEX 1881. DESCRIPTIONS OF SOME NEW SPECIES OF LIFE. Journal of American Museum of NORTH AMERICAN MOTHS. Papilio 1: 100­ Natural History 79: 34-39; 72. 101. BOISDUVAL, J. A. 1829. EUROPAEORUM LEPIDOPTERORUM INDEX 1882. DESCRIPTIONS OF NEW SPECIES OF NORTH METHODIC US. 1-103. Paris. pp. AMERICAN HETEROCERA. Papilio 2: 123­ BROWN, A. W., and W. C. McGUFFIN. 130. 1942. NEW DESCRIPTIONS OF LARVAE OF FOREST INSECTS III. The Canadian Entomologist 1884. SOME NEW FORMS OF NORTH AMERICAN 75: 52--56. MOTHS. Papilio 4: 43-48. CRAMER, P. 1782. PAPILONS EXOTIQUES DES TROIS PARTIES 1886. APPARENTLY NEW FORMS OF NORTH DU MONAE, L' ASIE, ET L' AMERIQUE. Part AMERICAN HETEROCERA. Entomologica 4, 1-252, plates 289-400 (text 33-253, and Americana 2: 165--171. plates after 253 evidently by Stoll), S. J. EICHLIN, T. D. Baalde, Amsterdam. 1975. GUIDE TO THE ADULT AND LARVAL PLUSI­ CRUMB, S. E. INAE OF CALIFORNIA (LEPIDOPTERA: NOC­ 1956. THE LARVAE OF THE PHALAENI­ TUIDAE). Occasional Papers in Entomol­ DAE. United States Department of Agri­ ogy, California, State Department of Food culture, Technical Bulletin 1135: 1-356. and Agriculture, No, 21: 1-73. 60 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE

--and H. B. CUNNINGHAM. 1969. CHARACTERS FOR IDENTIFICATION OF SOME 1865. DESCRIPTIONS OF NORTH AMERICAN LEPI­ COMMON PLUSINAE CA1'ERPILLARS OF THE DOPTERA, NO.6. Proeeedings of the Ento­ SOUTHEASTERN UNITED STATES. Annals mological Society of Philadelphia 4: 315­ of the Entomological Society of America 330. 62(3): 507-510. EVANS, D. 1873a. DESCRIPTIONS OF NOCTUIDAE PRINCIPALLY 1957. See ROSS, D. A., and D. EVANS. FROM CALIFORNIA. Bulletin of the Buf­ FABRICIUS, J. C. falo Society of Natural Science 1: 129-155. 1781. SPECIES INSECTORUM 2: 1-517. E. Hohn, Hamburg. 1873b. ON EIGHT SPECIES OF NOCTUIDAE. Bulletin of the Buffalo Society of Natural Science 1: 1787. MANTISSA INSECTORUM 2: 1..,'382. Hafniae. 190-194.

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GARDNER, J. C. M. 1875c. ON NORTH AMERICAN SPECIES OF 1947. ON THE LARVAE OF THE NOCTUIDAE (LEPI­ PLUSIA. The Canadian Entomologist 7: DOPTERA) III. Transactions of the Royal 202-205. Entomological Society of London 98(4): 59­ 60. 1877. A NEW PLUSIA ALLIED TO HOCHEN­ GMELIN, J. F., ed. WARTHI. The Canadian Entomologist 9: 1789. SYSTEMA NATURAE ... 1: 2556. Laurentii 135-136. Salvii. GEi"ER, C. ] 881. NEW NOCTUIDS, WITH A LIST OF THE SPE­ 1832. ZUTRAGE ZUR SAMMLUNG EXOTISCHER CIES OF ONCOCNEMIS. Papilio 1: 38-35. SCHMETTERLINGE, VIERTER HUNDERT, AUGSBURG. Pp. 1-48. 1882. NEW MOTHS, PRINCIPALLY COLLECTED BY GROTE, A. R. MR. ROLA:-ID THAXTER IN MAINE, WITH 1863. ADDITIONS TO THE CATALOGUE OF U.S. NOTES ON NOXIOUS SPECIES AND REMARKS LEPIDOPTERA, NO.5. Proceedings of the ON CLASSIFICATION. Bulletin of the United Entomological Society of Philadelphia 2: Stat.es Geological and Geographical Survey 273-276. of the Territories 6: 579 ....591.

1864. DESCRIPTIONS OF NORTH AMERICAN LEPI­ 1895a. THE HYPENOID MOTHS AND ALLIED DOPTERA, NO.3. Proceedings of the Ento­ GROUPS. Proceedings of the American mological Society of Philadelphia 3: 73-96. Philosophical Society 34: 416-436. PLUSIINAE OF AMERICA NORTH OF MEXICO 61

KIMBALL, C. P. 1895b. LIST OF NORTH AMERICAN EUPTEROTIDAE, 1965. ARTHROPODS OF FLORIDA AND NEIGHBOR­ ETC. Abhandlugen des Naturwissen­ ING LAND AREAS. Lepidoptera of Florida, schaftlichen Vereins zu Bremen 14: 61. an Annotated Check List. Pp. i-v, 1-363, iIIus. Gainesville, Fla. GROTE, A. R., and C. T. ROBINSON. KOSTROWIGK, A. S. 1868. DESCRIPTIONS OF AMERICAN LEPIDOPTERA 1961. STUDIES OF THE PALEARCTIC SPECIES OF NO. 4. ~ransactions of the American En­ THE SUBFAMILY PLUSIINAE (LEPIDOPTERA, tomological Society 2: 179-206. PHALAENIDAE). Acta Zoologica cracovien­ GUENEE, A. sia 6: 367-472. 1852. SPECiES GENERAL DES LEPIDOPTERES. LATREILLE, P. A. Noctuelites 2: 1-444. Libraire Ency­ 1825. FAMILLES NATURELLES DU REGNE ANI­ c10pedique de Roret, Paris. MAL ... Paris, Bailliere. HABECK, D. H. LINNAEUS, C. 1975. See HEPPNER, J. B., and D. H. HABECK. 1758. SYSTEMA NATUREA '" 10th edition, 1-S24. HALE, R. L. Laurentii Salvii, Holmiae. 1965. See SHOREY, H. H., and R. L. HALE. LINTNER, J. A. HAMPSON, G. F. 1885. PLUSIA DYAUS GROTE. Second Report on 1908. NOTES ON NOCTUIDAE. The Canadian En­ the Injurious and Other Insects of the tomologist 40: 102-107. State of New York. pp. 94-97. MACKAY, M. R. 1913. Catalogue of the Lepidoptera Phalaenae in 1922. THE LARVAE OF CANADIAN ARCTIC NOC­ the British Museum 13: i-xiv, 1-609. TUIDAE (LEPIDOPTERA). The Canadian HEPPNER, J. B., and D. H. HABECK. Entomologist 104: 859-S72. 1976. INSECTS ASSOCIATED WITH POLYGONUM McDUNNOUGH, J. (POLYGONACEAE) IN NORTH CENTRAL 1916. See BARNES, W., and J. McDuNNOUGH. FLORIDA. 1. INTRODUCTION AND LEPIDOP­ TERA. The Florida Entomologist 59(3): 1916. ON THE TYPES OF CERTAIN NOCTUA GEN­ 231-239. ERA OCCURRING IN NORTH AMERICA HINTON, H. E. (LEPID.). Entomological News 27: 393-400. 1946. ON THE HOMOLOGY AND NOMENCLATURE OF THE STATE OF LEPIDOPTEROUS LARVAE 1921. NOTES ON A COLLECTION OF LABRADOR WITH SOME NOTES ON THE PHYLOGENY OF LEPIDOPTERA. The Canadian Entomolo­ THE LEPIDOPTERA. Transactions of the gist 53: 81-S7. Royal Entomological Society of London 97: 1-37. HUBNER,J. 1938. CHECKLIST OF THE LEPIDOPTERA OF CAN­ [1800]-{1803]; [1808)-[1809]; [1823]. SAMMLUNG ADA AND THE UNITED STATES OF AMERICA. EUROPAISCHER SCHMETTERLINGE. (One PART I. MACROLEPIDOPTERA. Memoirs of volume of text [1814H1823] and eight vol­ the Southern California Academy of Sci­ umes of plates, 1796-{1838]), Augsburg. ences. pp. 1-274.

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WALKER, F. WOOD, G. W. 1857. LIST OF SPECIMENS OF LEPIDOPTEROUS 1951. AN ANNOTATED LIST OF LEPIDOPTEROUS INSECTS IN THE COLLECTION OF THE BRIT­ LARVAE FROM COMMERCIAL BLUEBERRY ISH MUSEUM 12: 765-982. FIELDS, CHARLOTTE CO., N.B. The Cana­ dian Entomologist 83: 241-244. 1858. LIST OF SPECIMENS OF LEPIDOPTEROUS INSECTS IN THE COLLECTION OF THE BRIT­ ISH MUSEUM 15: 1521-1888. --and W. NEILSON. 1960. NOTES ON LIFE HISTORY AND PARASITES 1865. LIST OF SPECIMENS OF LEPIDOPTEROUS OF SYNGRAPHA EPIGAEA (GRT.) (LEPI­ INSECTS IN THE COLLECTION OF THE BRIT­ DOPTERA: NOCTUIDAE). The Canadian ISH MUSEUM 33: 707-1120. Entomologist 92: 759-761. WALTER, E. V. 1928. SOME NEW MOTHS FROM ARIZONA. Proceed­ ZIMSEN, E. ings of the Entomological Society of Wash­ 1964. THE TYPE MATERIAL OF I. C. FABRICIUS. pp. ington 30(8): 137-144. 14i56. Munksgaard, Copenhagen. 64 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE

MAP 1

T Autop/usia il/ustrata o Argyrogramma basigera • Autographa amp/a o Syngrapha cryptica

MAP 2

• AllagrupilCl CH'rtl(l T Autograpilll iabrosa o Autogruph(l I1l1lppa o Syllgruphfl Ilbs/m.la PL USIINAE OF AMERICA NORTH OF MEXICO 65

MAP 3

• Adeva albavitla ... Rachiplusia OIL o Syngrapha falcifera o Autoplusia egena

MAP 4

• ;!rgyrogramma verucca • Pseudop/usia indudcrrs o Chrysarrympha fonnosa o Eosophoroptcryx thyatyroid,'-, 66 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE

MAP 5

• Autographa v-alba • Autographa precationis o Autographa ntbida o AuCographa speciosa

MAP 6

• Autograp/w califomica ~ Trichoplusia IIi PLUSIINAE OF AMERICA NORTH OF MEXICO 67

MAP 7

• tlutopiu.

MAP 8

• 1~'('tul('t'll pllLLiW'nt • P{{'Ud,Jl'(l flu rpu rtg" (rI \ o .. Ibrmlo/u p"rru/II 68 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE

MAP 9

• Abruslolll micromli.~ • Abrostalll oralis

MAP 10

• • A/mlstoill ur('fI ti.~ \ • .\IolLraiia tiflcloir/I'S \) PLUSIINAE OF AMERICA NORTH OF MEXICO 69

MAP 11

• SYIIW"p/W oilOLl'lIgllii ... Syllf!,rtlp/w [xlriLis o SYllWllp/W u·aUrt'ut1l ° Synf!,mp/w eiridisL:t.{lrlU •.-IIl/OWllphll /IIe/Ill/iell

MAP 12

• S_nlWllplw tllLIl o SyllWllp/1Il nUt,~r()J!(lnlllUI ... S.l'"Wllf)/w oetosrriptll O·'l)-f1Wllp/W orophilll • Trich O[lllt_lill ox,.,...·!!r({llllll11 70 TECHNICAL BULLETIN 1567. U.fl DEPARTMENT OF AGRICULTURE

MAP 13

• SYrlgrnplw ["elsa " Syrlgrup/ta diast'fTw o Syll!!ruplw l{!fl(fU <> SyTlgrap/ta illterroWttioflis

MAP 14

• PolyciJrysia mOlleta " Sy/lgrapiJa alias o SyngrupllU altieo/a <> Syllgrap/ta bon·a PLUSIINAE OF AMERICA NORTH OF MEXICO 71

MAP 15

• luloJ{fllp/1IJ

('om.~rll T "III/.ogmp/w jlll,!!{*llulTI o "III/o,!{rtlp/Itl Po;()Uc/Ol-!.(IIUrtlfl o IlIloWIl/)//(! .'ij(JflSO/U

MAP 16

• ,'i\'II,!{fIl(l/W "tunlllli T SYII/!fIIl'illI "'/""/11 o Chn'''''I',dw ( olll('rio 72 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE

MAP 17

: "'-,..... - \ ' ...... j ~-, . I : ~

• Chrys(lspidia "ie/lOlllle • ChryslIspidia plJlflami o Chrysa.~pidill t'('llusllJ o Alllol(mpha pasip/wcia

MAP 18

• tllllogmp/w biloba • Alllogmpha bim(lcu/alll o SYIIgrap/w angulidefls o SYflgrap/w u/tpm PL USIINAE OF AMERICA NORTH OF MEXICO 73

MAP 19

.. '~!--~

/ . ~

• "yllgmp/w ()fJiJ!u('(J • SYIIgr

r I:)

/ Ul Cl CIl

-0C l_ Ola. OJ Ul i!) \ >

FIGURES 1-7.-Larval chaetotaxy, right mandibles (mesal view): 1, Trichoplusw, ni; 2, P8eudoplu8ia includens; 3, Chrysuspidiu contexta; 4, Abro.~tola urent'isj 5, Mouraliu tinctoides; 6, Polychrysiu morigeru;7, Trichoplusia ni. PLUSIINAE OF AMERICA NORTH OF MEXICO 75

15t abd. 1St abd.

3 rd thor. 3rd thor.

SV-3

SV-l

<3 SV-2

2 abd TV "d r.. (J .....­

SV-l

SV-2

SV-l .' SV-2

10 • SV-3

/'~ , r- SV-l SV-2 .,/ 13

FIGURES 8-13.-Larval chaetotaxy, hypopharyngeal complex (lateral view): 8, SV-2 absent on abdominal segment 1 (lateral view); 9, SV-2 present on abdominal segment 1 (lateral view); 10, pinacula of SV-1 and SV-2 fused on abdominal segment 2 (lateral view); 11, pinacula of SV-l and SV-2 separated on abdominal segment 2 (lateral vi~w); 12, hypopharyngeal complex of Trichoplusia oxygramma; 13, SV setae equidistant apart on abdominal segment 2 (ventral view). 76 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE

UNCUS ANAL COSTAL TUBE MARGIN I VALVE i I 17 AMPULLA -

JUXTA / " SACCULUS DUCTUS SPINE' EJACULATO RI US VINCULUM ~, , I ~./~.:l AEDEAGUS 18 " ." ...i

BASAL CORNUTUS / APICAL CORNUTUS

FIGURES 14-18.-Larval chaetotaxy (ventral view), spermatophore, male genitalia: 14, SV-1 and SV-l! closer to each other than to SV-J on abdominal segment 2; 15-16, relative distances between SV_l and V-Ion abdominal segment 4; 17, SYllgraplll1. altieola; 18, S. microgamma (ventral view), aedeagus (lateral view). PLUSIlNAE OF AMERICA NORTH OF MEXICO 77

DUCTUS SEMINALlS...... /'

APEX...... ' OSTIUM BURSAE -- i

FUNDUS

" OSTIUM

---." \ " " ' "- DUCTUS BURSAE

19

- CORPUS BURSAE

FIGURES 19-22.-<.remtalla lventral vIew): l~, Sy7I{JTupha 19mu (female); 20, Abro8tola parvula (male); 21a,b, A. urentis (male); 22, A. microvalis (male). 78 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE

25

FIGURES 23-25.-Male genitalia: 23, Mouralia tinctoide.~ (ventral view); 24, Argyrogramma basigera (lateral view, left valve removed); 25, A. verruca (lateral view, left valve removed). PLUSIINAE OF AMERICA NORTH OF MEXICO 79

FIGURES 26-31.-Male genitalia (lateral view, lett valve removed): 26, Rachiplusia OU; 27, Pseudeva p'urpurigera; 28, Pseudoplusia includens; 29, Plusia balluca; 30, Trichoplusia oxygramma; 31, Pseudeva palligera. 80 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE

..

FIGURES 32-38.-Male genitalia (32-35, 38, lateral view, left valve removed; 36-37, ventral view): 32, Adeva albavitta; 33, Allagrapha aeroides; 34, Trichoplusia ni; 35, Allagrapha aerea; 36a,b, Polychrysia moneta; 37a,b, P. morigera; 38, Autoplusia egena. PLUSIINAE OF AMERICA NORTH OF MEXICO 81

43

FIGURES ii!l-44.-Male genitalia (lateral view, left valve removed); 39, AutoplU8ta olwacea; 40, Chrysanympha!ormoaa; 41, Eosphoropteryx thyatyroides; 42, Autographa mappa; 43, A. bimaculata; 44, A. pasiphaeia. 82 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE

45

FIGURES .i5--49.-Male genitalia (lateral view, left valve removed): 45, Autographa biloba; 46, A. BpecioBa; 47, A. f/agf

FIGURES 50-54.-Male genitalia (50, ventral view; 51-54, lateral view, left valve removed): 5Oa,h, Autographa labrosa; 51, A. metallica; 52, A. v-alba,· 53, A. californica; 54, A. sansoni. 84 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE

.~ '.~6

FIGURES 55-59.-Maie genitalia (55-56, lateral view, left valve removed; 57-59, ventral view): 55, Autographa ampla; 56, A. rubida; 57a,b, A. pBeudogamma; 58a,b, Syngrapha epigaea; 59, Autoplusia illuBtrata. PLUSIINAE OF AMERICA NORTH OF MEXICO 85

l"IGURES 60--64.-Male genitalia (ventral view, aedeagus lateral view): 60a,b, Syngrapha altera; 61a,b, S. celsa; fi2a,b, S. sp.; 63a,b, S. ignea; 64a,b, S. abstrusa. 86 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE

FIGURES 65-69.-Male genitalia (ventral view, aedeagus lateral view): 65a,b, Syngrapha 77Wntana; 66a,b, S. octoBcripta; 67a,b, S. parili8; 68a,b, S. ottolenguii; 69a,b, S. viridiBigma. PLUSIINAE OF AMERICA NORTH OF MEXICO 87

70

FIGURES 70-73.-Male genitalia (ventral view, aedeagus lateral view): 70a,b, Sy7I{/rapha orophila; 71a,b, S. c1yptica; 72a,b, S. B6lecta; 73a,b, A. Backeni. 88 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE

FIGURES 74-78.-Male genitalia (74,75,77,78, ventral view, aedeagus lateral view; 76, lateral view, left valve removed); 74a,b, Syngrapha 8Urenaj 76, S. angulidensj 75a, b, S. alticola; 77a,b, S. rectangula,· 78a,b, S. alias. PLUSIINAE OF AMERICA NORTH OF MEXICO 89

FIGURES 79--83.-Male genitalia (ventral view, aedeagus lateral view): 79a,b, Sy'I1{Jrapha interrogationis; 80a,b, S. u­ aureum; 81a,b, S. lula; 82a,b, Chrysaspidia putnami; 83a,b, S. diasema. 90 TECHNICAL BULLETIN 1567. U.S. DEPARTMENT OF AGRICULTURE

FIGURES 84-88.-Genitalia (84, ventral view, aedeagus lateral view; 85-87, lateral view): 84a,b, Chrysaspidia contexta (male); 85, C. nichollae (male, left valve removed); 86, C. venU8ta (male, left val1Je removed); 87, Syngrapha falcifera (male, left valve removed); 88, Abrostola ovalis (female). PLUSIINAE OF AMERICA NORTH OF MEXICO 91

FIGURES 89-96.-Female genitalia: 89, Abrostola partmla (dorsal view); 90, Mouralia ti7lCtoides (ventral view); 91, Argyrogramma basigera (lateral view); 92, Abrostola microvalis (dorsal view); 93, A, urentis (dorsal view); 94, Plusia balluca (lateral view); 95, Autoplusia illustrata (lateral view); 96, Argyrogramma verruca (lateral view). 92 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE

FIGURES 97-104.-Female genitalia (97, ventral view, 98-104, lateral view): 97, Hachlplusla ou; 98, Allagrapha aerea; 99, P8eudoplusia includens; 100, P8eudeva palligera; 101, P. purpur.gera; 102, TricJwplusia ni; 103, E08pJwropteryx thyatyroide8; 104, Allagrapha aeroide8.

PLUSIINAE OF AMERICA NORTH OF MEXICO 93

108 109

FIGURES 105-111.-Female genitalia: 105, Trichoplu8ia oxygramma (lateral view); 106, Adeva albavitta (lateral view); 107, Autoplu8ia olivacea (ventral view); 108, Polychry8ia morigera (lateral \,j;r-w); 109, Autoplu.8ia egena (ventral view); 110, Polychry8ia moneta (lateral view); 111, Chry8anymphajormo8a (lateral view). 94 TECHNICAL BULLETIN 1567. U.S. DEPARTMENT OF AGRICULTURE

FIGURES 112-118.-Female genitalia (112-116, 118, lateral view; 117, dorsal view); 112, Autographa precationisj 113, A. californicaj 114, A. amplaj 115, A. biloba; 116, A. p8eudogammaj 117, A. 8ansoni; 118, A. corusca. PLUSIINAE OF AMERICA NORTH OF MEXICO 95

FIGURES 119-124.-Female genitalia: 119, Alttogral)hu jZuO"llu1Il (lateral view); 120, A. mUl'prL (lateral view); 121, A. labroBa (lateral view); 122, A. rubida (dorsal view); 123, A. pu.~iphaeia (lateral view); 124, Syngrapha ceL~a (dorsal view). 96 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE

FIGURES 125-131.-Female genitalia: 125, Autographa bimaculata (lateral view); 126, Syngrapha a1!gldiden.~ (dorsal view); 127, S. Burena (lateral view); 128, S. alticola (dorsal view); 129, Syngrapha montana (ventral view); 130, S. alias (ventral view); 131, Autographa v-alba (ventral view). PLUSIINAE OF AMERICA NORTH OF MEXICO 97

FIGURES 132-138.-Female genitalia: 132, Autographa speciosa (ventral view); 133, Syngrapha orectangula (ventral view); 134, Autographa metallica (lateral view); 135, S-yngrapha falciJera (ventral view); 136, S. sackeni (ventral view); 137, S. epigaea (dorsal view); 138, S. microgamma (dorsal view). 98 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE

FIGURES 139-146.-Female genitalia: 139, Syngrapha orophila (ventral view); 140, S. 81WWi (ventral view); 141, S. octo8eripta (lateral view); 142, S. Beleeta (lateral view); 143, S. 1JiridiBigma (dorsal view); 144, S. altera (lateral view); 145, S. ab8truBa (ventral view); 146, S. ottoler!{Juii (ventral view). PLUSIINAE OF AMERICA NORTH OF MEXICO 99

FIGURES 147-155.-Female genitalia: 147, Syngrapha cryptica, (ventral view); 148, S. u-aureum (ventral view); 149, ChryBaBpidia putnami (lateral view); 150, Syngrapha interrogationis (ventral view); 151, ChryBaspidiu nichallae (dorsal view); 152, S. pariliB (lateral view); 153, S. lula (ventral view); 154, C. contexta (lateral view); 155, C. venusta (lateral view). 100 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE ,. ., = ~g ?;; ." <0"0 l: :: .. tJl 0 E; en :;'B!.~ 5 g OJ g ~ ~""=V1 o ~ lii a~ ~ e.8 ~ gg r§ g ~ t: ... :a tt~ ~~a::a ~'E ~~§t9 rJ0'>- Cl 2~8 ~~ ~~ 6 ri ~ ~~ ~~ ~ ;;;'r;~1JI r. 2..;' ~i g~"'m [~M U~~.. c .. tt:n i'n Co gfj 0" g g:~ - 3- il§ ~ 0 ~~i Hi ~ ~ " ~ OJ 'a ~~»- ~ s;: ~ "1:- ~ " ;; "51 s;: :;: "~ uS! "~ t tlfi " n ~ " "" !i' ~ " ~g

Alr::otwHIr:r ...... 'pinaCUla DfSV~1 and ~ _---pro-leg::: ::"ost on fused on abd. se~~ 2 -!!..:.. aega. j ar..u ...

pro.:.egs re'J.u;;ed nr. .l4.:',~. sego. 3 ar.d :"j 1.ar.-ii::le r..a:t:.!iC'..1 .r,i::h riLs ? and J ":,en-iml:ting m processes

tlblae un!>plIlcd

proleg'1 on abd. !Jcgs_ J ;lnd "; ~t'O.:"ct$ blOrdln.1.l, SV~~ abs~nt on m~50· ilnd r.u!tathora.(.. S\·<~ absent on ftrs:t nbd. seg.;. tiltiulol.u prescnt 156

FIGURES 156.-Dendrogram (part I): Phylogeny of the subfamily Plusiinae less species of Autographa, Syngrapha, and Chrysaspidia.

; ~~E~:~ ~ ~'! "" '~j

:n .: t"."l~f,tI .m f; ro;;r ,lhd "(>1; AUTOGRAP~INI (in part) p'O::l~-::::C': m n;'U1IU:( 157

FIGURES 157.-Dendrogram (part II): Phylogeny of the species of Autographa, Syngrapha, and Chrysaspidia. PLUSIINAE OF AMERICA NORTH OF MEXICO 101

PN"-<;682

FIGURE 158.-Abrostola urentis. PN-5685 FIGURE 161.-Abrostola ovali8.

PN"-<;683 FIGURE 159.-Abrostola parvula.

PN-fi686 FIGURE 162.-Mouralia tinctoideR.

PN-5684 'FIGURE 160.-Abrostola microvalis.

PN-0687

FIGURE 163.-Argyrogramma. verrUca. 102 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE

PN-li691 FIGURE 167.-Pseudoplusia includen8.

PN-li688 FIGURE 164.-Aryy·roy·ramma basiyera.

PN-li692 FIGURE 168.-Autoplusia olivacea,

J .",

PN-5689 FIGURE 165.-T·richoplusia ni.

PN-6693 FIGl'RE 169.-Autoplusia illustrato..

FIGl'RE 166.-Trichoplu8ia oxyymmma. PLUSIINAE OF AMERICA NORTH OF MEXICO 103 -

FIGURE 170.-Autoplusia egen,a. PN-5697 FIGURE 173.-Allagrapha aeroide8.

PN";;695 FIGURE 171.-Rachiplusia ou.

F1GURE 174.-Allagrapha aerea.

PN-5696 FIGURE 172.-Plusia balluca.

PN";;699 FIGURE 175.-Po[ychry8in. moneta. 104 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICUL'l'URE

PN-5703 PN-5700 FIGURE 176.-Polych·rysia m07-igera. FIGURE 179.-Pseudeva purpurigera.

PN-5704 FIGURE 180.-Chrysanympha formosa. P1'.· .5701 FIGURE 177.-Adeva alba1,itta.

PN-5706 FIGURE 181.-Eosphoropt.e·ryx: thyatyroides.

FIGURE 178.-Pseudeva palligera. PLUSIINAE OF AMERICA NORTH OF MEXICO 105

PN-5709 FIGURE 185.-Autographa rubida. PN-5706 FIGURE 182.-Autographa precationis.

PN-57l0 FIGURE 186.-Autographa biloba.

PN-5707 FIGURE 183.-AutograJlha bimaculata.

PN-r,711 FIGURE 187.-Autographa cal~rornica.

PN-5708 FIGURE 184.-Autographa Ranson'i. 106 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE

PN-5712 FIGURE 188.-Autographa pseudogamma. PN-5715 FIGURE 191.-Autographa flagellum.

PN-5713 FIGURE 189.-Autogrupha v-alba. PN-5716 FIGURE 192.-Autographa metallica.

PN-5714 PN-5717 FIGURE 190.-Autographa pasiphaea. FIGURE 193.-Autographa speciosa. PLUSIINAE OF AMERICA NORTH OF MEXICO 107

PN-5721 PN-ii718 FIGURE 197.-Autographa ampla. FIGURE 194.-Autographa /abrosa.

PN-5722 FIGURE 198.-Syngrapha ottolenguii.

PN-5719 FIGURE 195.-Autographa corusca.

FIGURE 199.-Syngrapha altera variana. PN-5720 FIGURE 196.-Autographa mappa. 108 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE

PN-5724 PN-5727 FIGURE 200.-Syngrapha oct08cripta. FIGGRE 203.-Synnrapha viridiBinma. ..

PN-5728 FIGt;R::: 204.-Syngrapha orophila. PN-5725 FIGURE 201.-Syngrapha epigaea.

,

PN-5729 FIGURE 205.-Syngrapha 8now1.

PI ,72" FIGURE 202.-Syngrapha selecta. PLUSIINAE OF AMERICA NORTH OF MEXICO 109

PN-5730 FJGl'RE 206.-Syngrapha sackeni.

FIGURE 209.-Syngrapha falcifem.

FIGt'RE 207.-SyngraphCL lulCL.

PN -5734 FIGURE 210.-SyngraphlL u-auremn.

FIGl'RE 208.~"yngTCLphCL diCL.~ema.

Pt'· 57~o FIGl'RE 211.--..')yng rapha interroga.tioni.q lio n:tLli.q. 110 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE

PN-5739 FIGt:RE 215.-Syngrapha cryptica. PN-5736 FIGURE 212.-Syngrapha SlLrena.

PN-5740 FIGt~RE 216.-..")yngrapha 'rectanguia.

PN-573~ FJUt:RE 213.-SYllgrapha alias.

PN-574! FIGl'RE 217.-..'>yngrapha angulidens.

FHH"RE 214.-SYIl{/rapha ablltrllsa. PLUSIINAE OF AMERICA NORTH OF MEXICO 111

PN-S742 FIGURE' 218.--Syngrapha celsa. PN-S'i45 FIGURE 221.--Syngrapha dticola.

PN-5743 FIGURE 219.--Syngrapha montana.

P!\ -5746 FIGURE 222.--SY1!f} ra1)ha po.rili,q.

PN-S744 FIGURE 220.--Syngrapha microgamma.

FIGt'HE 223.-SYllr!1·uph« iYIl!'a. 112 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE

PN-5748 FIGURE 224.-Chrusaspidia contexta.

Pl':-5750 ,FIGURE 226.-Chrysaspidia nichoUae .

.. PN-574£ FIGl:RE 225.-Chrysaspidia putnam'i.

PN-5751 FIGURE 227.-Chrysaspidia venu.~ta. PLUSIINAE OF AMERICA NORTH OF MEXICO 113

INDEX TO GENERA, SPECIES, AND LOWER CATEGORIES OF PLUSIINAE

[Valid names are set in roman type and synonyms are set in italics] ABROSTOLA Ochsenheimer, 1, 3, 4, 7, 9, 11, 26, 54, 55 monoav:' (Grote), 32, 33 mariana Walter, 10 au var. californ'ica (Speyer), 31 microvalis Ottolengui, 9, 10, 55 pasiphaeia (Grote), 28, 32 ovalis Guenee, 8, 9, 10, 11, 55 precationis (Guenee), 8, 28 parvula Barnes and McDunnough, 9, 10, 55 pseudogamma (Grote), 28, 31 urentis Guenee, 8, 9, 11, 55 rubida Ottolengui, 8, 28, 30 abstrusa n. sp., 3, 6, 37, 46, 47, 48, 57 rUBBea (Edwards), 20 ADEVA McDunnough, 7, 17, 23, 24, 25, 56, 57 sansoni Dod, 28, 29 albavitta (Ottolengui), 24, 25 scapularis (Edwards), 33 Jmtsonii (Smith), 24, 25 speciosa Ottolengui, 27, 28, 33 aerea (Hubner), 9, 21, 22 'u-brevis (Guenee), 29 aeroides (Grote), 9, 22, 23 v-alba Ottolengui, 27, ,28, 32, 33 albavitta (Ottolengui), 24, 25 AUTOPLUSlA McDunnough, 2, 7,17,18,19,56 alias (Ottolengui), 8, 36, 37, 45, 46, 47, 48, 51, 57 egena (Guenee), 8, 18,19 ALLAGRAPHA Franclemont, 17, 21, 22, 56 illustrata (Guenee), 18,19,55 aerea (Hubner), 9, 21, 22 olivacea (Skinner), 8, 18 aeroides (Grote), 9, 22, 23 balluca (Geyer), 9,21 altera (Ottolengui), 36, 37, 38, 57 basigera (Walker), 8, 12, 13, 14 alterna (Strecker), 34, 35 biloba (Stephens), 9, 27, 28, 30, 56 alticola (Walker), 35, 36, 37, 50, 51 bimaculata (Stephens), 8, 28, 29, 34 ampla (Walker), 8, 27, 28, 34, 35 borea (Aurivillus), 36, 37, 42, 57 angulidens (Smith), 36, 37,48, 49, 57 brasBicae (Riley), 14, 15 a'rctica (Ottolengui), 37 californica (Speyer), 9, 28, 31 ARGYROGRAMMA Hubner, 2, 7, 12, 19, 55 cal-ifonLica var. 'ruBsea (Edwards), 31 basigera (Walker), 8, 12, 13, 14 callitrichoides Grote laticlavia (Morrison), 13, 14 celsa (Edwards), 8, 36, 37,49, 50, 51, 57 omega Hubner, 13 celsa race sierrae (Ottolengui), 8, 49 omicron (Hubner), 13 CHRYSANYMPHA Grote, 7, 17, 26, 56, 57 00 (Cramer), 13 formosa (Grote), 26 verruca (Fabricius), 8, 12, 13,14 CHRYSASPIDIA Hubner, 1,4,7,17,52,54,56,57 AUTOGRAPHA Hubner, 5, 7, 12, 17,20,26,27,35,37,38, contexta (Grote), 8, 52, 53 52,56,57 nichollae (Hampson), 52, 53, 54 alterna (Strecker), 34, 35 putnami (Grote), 8, 52, 53, 54 ampla (Walker), 8, 27, 28, 34, 35 st·riatella (Grote), 54 biloba (Stephens), 9, 27, 28, 30, 56 venusta (Walker), 52, 53, 54 bimaculata (Stephens), 8, 28, 29, 34 contexta (Grote), 8, 52, 53 californica (Speyer), 9, 28, .31 corusca (Strecker), .27, 28, 33, 34 califont'ica var. rUBsea, 31 cryptica n. sp., 36, 37, 46, 47, 48, 57 corusca (Strecker), '2:1, 28, 33, 34 culta, (Lintner), 16, 17 flagellum (Walker), 28, 32, 33 deve'rgens (Hubner), 35, 50, 51 gamma (Linneaus), 27 diasema (Boisduva\), 36, 37,42,43,57 gamma abo califomiea (Speyer), 31 diasema, var. OOrea (Aurivillus), 42 fnBolittJ. (Smith), 32, 33 diversignu (Ottolengui), 42, 43 kas/oenBes (Strand), 33 dyalls (Grote), 16, 17 labrosa (Grote), 27, 28, 33 echinocystidis (Strecker), 14, 15 Lenzi! French, 33 egena (Guenee), 8, 18, 19 mappa (Grote and Robinson), 8, 28, 34 EOSPHOROPTERYX Dyar, 7, 17,26,56,57 metallica (Grote), 28, 33 thyatyroides (Guenee), 26 114 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE epigaea (Grote), 8, 36, 37, 39, 57 'PLUSIA Ochsenheimer, i, 2, l'l, 21, 22, 56 epsilon (Ottolengui), 38, 39 balluca (Geyer), 9, 21 excelsa (Ottolengui), 48, 49 POLYCHRYSlA Hubner, 7, 17, 23, 25, 56 falcifera (Kirby), 8, 35, 36, 37, 43, 57 moneta (Fabricius), 23, 24 flagellum (Walker), 28, 32, 33 morigera (Edwards), 8, 23, 24 formosa (Grote), 26 trabea (Smith), 23, 24 frateUa (Grote), 20 precationis (Guenee), 8, 28 gamma (Linnaeus),27 PSEUDEVA Hampson, 7, 17,24,25,56,57 gamma abo californica (Speyer), 31 pnlligera (Grote), 25, .26 hamifera (Walker), 16, 17 purpurigera (Walker), 25, 26 herschelensis (Benjamin), 44, 45 rubigera Hampson, 25 hochenwarthi (Hochenwarth), 50, 51, 52 pseudogllmma (Grote), 28,31 hutsonii (Smith), 24, 25 PSEUDOPLUSIA McDunnough, 2, 7, 12, 16, 55 ignea (Grote), 35, 36, 37, 50, 51, 52, 57 culta (Lintner), 16, 17 ignea var. simulans (McDunnough), 51, 52 dyaus (Grote), 16, 17 illustrate (Guenee), 18, 19, 55 hamifera (Walker), 16,17 includens (Walker), 3, 8, 12, 16,17 includens (Walker), 3, 8, 12, 16, 17 indigna (Walker), 15, 16 00 (Cramer), 16, 17 " insolita (Smith), 32, 33 rogationis (Guenee), 16,17 interalia (Ottolengui), 45 purpurigera (Walker), 25, 26 interrogationis (Linnaeus), 35, 36, 37,441., 45, 57 putnami (Grote), 8, 52, 53, 54 interrogationis var. herschelensis (Benjamin), 44, 45 quadriplaga (Walker), 51 kasloenses (Strand), 33 RACHIPLUSIA Hampson, 2, 7, 17, 20,56 labrosa (Grote), 27, 28, 33 fratella (Grote), 20 laticlavia (Morrison), 13, 14 ou (Guenee), 9, 20 lenzii French, 33 pedalis (Grote), 20 lula Strand, 36, 37,42,57 rectangula (Kirby), 8, 36, 37, 43, 48, 51, 57 magnifica (Ottolengui), 38, 39 recta7lgula race nargenta (Ottolengui), 48 mappa (Grote and Robinson), 8, 28, 34 rogationis (Guenee), 16,17 mariana Walker, 10 rubida Ottolengui, 8, 28, 30 metallica (Grote), 28, 33 rubigera Hampson, 25 microgamma (Hubner), 3, 8, 36, 37, 49, 50 TU8sea (Edwards), 20 microvalis Ottolengui, 9, 10, 55 sackeni (Grote), 36, 37, 42, 57 monodon (Grote), 32, 33 sansoni Dod, 28, .29 moneta (Fabricius), 23, 24 scapularis (Edwards), 33 montana (Packard), 36, 37, 49, 50 selecta (Walker), 8, 39, 40, 41, 57 morigera (Ed ...vards), 8, 23, 24 sierrae (Ottolengui), 8, 49 mortuorum (Guenee), 48 simplex (Guenee), 43 MOURALlA Walker, 4, 7, 11, 55 simplicima (Ottolengui), 43 tinctoides (Guenee), 8, II, 55 simulans (McDunnough), 51, 52 nargenta (Ottolengui), 48 snowi (Edwards), 36, 37, 41, 57 ni (Hubner), 3, 8, 12, 14, 15 snawi form lula Strand, 42 nichollae (Hampson), 52,53, 54 speciosa Ottolengui, 27, 28, 33 octoscripta (Grote), 36, 37, 38, 39, 57 striatelia (Grote), 54 olivacea (Skinner), 8, 18 surena (Grote), 36, 37, 45, 57 omega (Hubner), 13 SYNGRAPHA Hubner, 1, 2, 5, 7, 17, 35, 38, 52, 56, 57 00 (Cramer), 13, 16, 17 abstrusa n. sp., 36, 37, 46, 47, 48, 57 orophila (Hampson), 8, 36, 37, 41, 57 alias (Ottolengui), 8, 36, 37, 45, 46, 47, 48, 51, 57 ottolenguii (Dya.r), 35, 36, 37, 56 altera (Ottolengui), 36, 37, 38, 57 ou (Guenee), 9, 20 alticola (Walker), 35, 36, 37, 50, 51 ou var. calijornica (Speyer), 20, 31 angulidens (Smith), 36, 37, 48, 49, 57 ovalis Guenee, 8, 10, 11, 55 arctica (Ottolengui), 37 oxygramma (Geyer), 8, 14, 15,16 borea (Aurivillus), 36, 37, 42, 57 pallida (Ottolengui), 38, 39 celsa (Edwards), 8, 36, 37, 49, 50, 51, 57 palligera (Grote), 25, 26 celsa race sierrae (Ottolengui), 8, 49 parilis (Hubner), 8, 36, 37, 51, 57 cryptica n. sp., 36, 37, 46, 47, 48, 57 parvula Barnes and McDunnough, 9, 10, 55 devergens (Hubner), 35, 50, 51 pasiphaeia (Grote), 28, 32 diasema (Boisduval), 36, 37, 42, 43, 57 pedalis (Grote), 20 diasema var. borea (Aurivillus), 42 PHYPROSOPUS Grote, 7 diversigna (Ottolengui), 42, 43 callitrichoides Grote, 55 epigaea (Grote), 8, 36, 37, 39, 57 PLUSIINAE OF AMERICA NORTH OF MEXICO 115

epsilon (Ottolengui), 38, 39 s'im1llans (McDunnough), 51,. 52 excelsa (Ottolengui), 48, 4l} snowi (Edwards), 36, 37, 41, 57 falcifera (Kirby), 8, 35, 36, 37, 43, 57 8nowi from htla Strand, 41, 42 herschelensis (Benjamin), 44, 45 surena (Grote), 36, 37, 45, 57 hochenwarthi (Hochenwarth), 50, 51, 52 u-aurem (Guenee), 3,8, 36,37,44,57 ignea (Grote), 35, 36, 37, 50, 51, 52, 57 vaccinii (Edwards), 37, 44 ignea var. s'imulans (McDunnough), 51, 52 va'riana (Ottolengui), 38, 39 'intemlia (Ottolengui), 45 viridisigma (Grote), 8, 36, 37, 39, 40, 41, 43, 44, 57 interrogationis (Linneaus), 35, 36, 37, 44,45, 57 ;zeta (Ottolengui), 38, 39 ·interrogationis var. herschelensis (Benjamin), 44, 45 "hyatyroides (Guenee), .26 lula Strand, 36, 37, 42, 57 tinctoides (Guenee), 8, 11 magnifica (Ottolengui), 38, 39 trabea (Smith), 23, 24 microgamma (Hubner), 3, 8, 36, 37, 49, 50 montana (Packard), 36, 37, 49, 50 TRICHOPLUSIA McDunnough, 2, 7, 12, 14,55 mo·rtuorl.Lm (Guenee), 48 bra8sicae (Riley), 14, 15 nargenta (Ottolengui), 48 ech'inocystidis (Strecker), 14, 15 octoscripta (Grote), 36, 37, 38, 39, 57 indigna (Walker), 15, 16 orophila (Hampson), 8, 36, 37, 4}, 57 ni (Hubner), 3, 8, 12, 14, 15 ottolenguii (Dyar), 35, 36,37,56 oxygramma (Geyer), 8, 14, IS, 16 pall·ida (Ottolengui), 38, 39 u-aureum (Guenee), 3, 8, 36, 37,44, 57 parilis (Hubner), 8, 36, 37, 51, 57 u-brevis (Guenee), 29 quad:riplaga (Walker), 51 urentis Guenee, 8, 9, II, 55 rectangula (Kirby), 36, 37, 43, 48, 51, 57 v-alba Ottolengui, 27, 28, 32, 33 ·rectangula race nargenta. (Ottolengui), 8, 48 vaccini'i (Edwards), 37, 44 sackeni (Grote), 36, 37, 42, 57 va'riana (Ottolengui), 38, 39 selecta (Walker), 8, 39, 40, 41, 57 venusta (Walker), 52, 53, 54 sierrae (Ottolengui), 8, 49 verruca (Fabricius), 8, 12, J3, 14 simplex (Guenee), 43 viridisigma (Grote), 8, 36, 37, 39, 40, 41, 43, 44, 57 simplicima (Ottolengui), 43 ;zeta (Ottolengui), 38, 39 116 TECHNICAL BULLETIN 1567, U.S. nEPARTMENT OF AGRICULTURE

INDEX TO HOST PLANTS The species for which host plants r.:.ave been Amsinckia douglasiana recorded are listed under the scientific name of Autographa californica, 31 Antirrhinum sp. the food plant. A few species are listed under Triehoplusiani,15 the common name of the plant, if the common Apium graveolens name could not be defmitely associated with Autographa californica, 31 one plant species. For ease of reference, both Autograph egena, 19 the common and scientific names of the plants Syngrapha faleifera, 43 Triehoplusia ni, 15 are listed. apple (see Malus pumila) Abies amabilis Arctium lappa Syngrapha alias, 45 Autographa preeationis, 29 Syngrapha -reetangula, 4D Syngrapha falcifera, 43 Abies balsamea Argemone platyceras Syngrapha alias, 45 Triehoplusia ni, 15 Syngrapha reetangula, 48 arrowleaf (see Sagittaris sp.) Abies gradis Asclepias sp. Syngrapha eelsa, 49 Triehoplusia ni, 15 SyngraphareeLangula, 48 Asparagus officinalis Abies lasioea'rpa Triehoplusia ni, 15 Syngrapha alias, 45 asparagus (see Aspa'ragus offidnalis) Syngrapha angulidens, 49 aspen, quaking (see Populus tremuloides) Syngrapha celsa, 49 aster (see Aster sp., Aster umbellatus) ., Syngrapha reetangula, 48 Aster sp. Syngrapha viridisigma, 41 Allagrapha aeroides, 22 Acacia greggii Triehoplus'ia oxygramma, 16 I' Triehoplusia ni, 15 Aster umbellatus Agapanthus sp. Allagrapha aerea, 23 Autoplusia egena, 19 Atriplex sp. Ageratum eonyzoides Autographa ealiforn'iea, 31 Triehoplusia ni, 15 Atriplex canescens alder (see A/niLS sp.) Autographa ealifomiea, 31 \ alder, red (see Alnus oregona) Au'reolaria pedieularia alfalfa (see Medieago sativa) Autographa preeafion'is, 29 Allium eepa avocado (see Persea americana) Autographa ealiforniea, 31 baby bl ue eyes (see Nemophila menziesi·i) Alnus sp. babys' tears (see Helxine solei'rollii) Autographa ampla, 35 barley (see H01ue1Lm 1nLlgare) Autographa eOT1LSea, 34 basil (see Typtis sp.) Syngrapha epigaea, 39 beans, unspecified (see also Phaseo/us sp.) Alnus oregona Autog'rapha preeation'is, 29 A1dographa ampla, 35 Syngrapha falcifera, 43 Althaea rosea beans, green (see Phaseolus mLlgaris) . Autographapreeationis, 29 beans, lima (see Phaseolus limensis) Autop/usia egena, 19 beans, sieva (see Phaseolus lunatus) Syngrapha faleifera, 43 beans, snap (see Phaseolus V1Llgaris) Triehoplusia ni, 15 beets (see Beta t'Ulgaris) Amela1Lekier florida Be'rgenia sp. Autographa ampla, 35 Triehoplusia ni, 15 Autographa eaviforniea, 31 Beta t'Ulga'ris Amelanehier alnifolia Autographa eal-iforniea, 31 Syngrapha epigaea, 39 Syngrapha falcifera, 43 • PLUSIINAE OF AMERICA NORTH OF MEXICO 117

Trichoplusia ni, 15 Autogl'ap}w; .jJ7·ecat-ionis, .29 Betula pendula Trichoplusia n'l, 15 Autographa ampia, 35 Chenopodium ambrosioides Synyrapha ep'igaea, 39 Raclviplusia ou, 20 birch, European white (see Betula pe?ldula) cherry, "...ild (see Prun'lls sp.) blueberry (see Vaccinium sp.) chinchweed (see PeeNs papposa) blueberry, commercial (see VacC'inium, angltstijolimn) Chrysant,/tenmm sp. Brassica campest'ris Autopiusia egena, 19 Trichoplusia 7ti, 15 PSl'udoplusiainci1tdens, 17 Brass'ica kina T,.ichoplusia ni, 15 T'richophtSia nt, 15 Chrysanthemum frutescens Brassica napltS A,utoplusia e{lena, 19 T'rickoplusia 'Ii, 15 Cineraria sp. Brassiea n'igra Trichopillsia nt, 15 Triehoplusia nt, 15 Ci'rsium sp. Brassica oleracea Aulogmp/ta precationis, 29 Autographa biloba, 30 Syn{lraplta falcife'm, 43 Autographa califonliea, 31 Cinlium horriduLum Autographa p'reeationis, 29 A uto{lrapha biloba, 30 A1Ltoplusia egena, 19 Citr·ullu.s vuigcwis Pseudoplusia includens, 17 Aufographa caU/ornica, 31 Syngrapha faleife'ra, 43 T1"ic}wpl!/"~ia ni, 15 Triehoplusia 'ni, 15 Citrus g inensis Brassiea Tapa Autographa ca,Lifomica, 31 Autogmpha californiea, 31 Trichopltl,qia ni, 15 Triehoplusia nt, 15 Cilr1/"~ sp. broccoH (see Braslliea oleraeea) Trichopl!l,~ia ni, 15 brussels sprouts (see Brass'iea oleracea.) clover (see Trofoliwn sp.) buffaloberry (see Shephe'rdia cancldensi.~) clover, crimson (see Trifolium inca'matum) burdock, great (see Arctiu:m lappa,) Coleus sp. bur-reed (see Spa'ryan'ium sp.) PSt'udoplu.qia, irlclu(iens, 17 cabbage (see Bras8ica oleracea) Trichoplusiarli, 15 Calendula officinalis eoleus (see Coleu$ sp.) Autographa californica, 31 colza (se€' Bl"tLssi("(l napus) Trichoplusia ni, 15 comfrey (see Sympltytwrl sp.) Carduus sp. Commelina sp. Autogmpha precationis, 29 Pselldoplullicl inclildens, 17 Carex sp. corn (see Zea ma,y,~) Chrysaspi(/:ia 'putnami, 53 cornflower (see ('entlLU'rea solstitialis) carnation (see Dianthus caryophylltt:~) cosmos (see ('O~IIW8 spJ carrot (see DaltCus carota) Cosmos sp. Cartha:1rtltS t indo'rias RfJ.chiplll~ia ou, 20 Autographa californica, 31 cosmos (see Co.mlO8 sp.) cascara (see Rhammts sp.) cotton (see GOIlSl/piuiI hirsutum) Castilleja sp. cowpea (see Viga1l unguiculata) Autographa californica, 31 cranberry (see Vacclniwn ntCLcrocarpon) catclaw (see Acacia greggii) C1"('l1i8 Sp. cauliflower (see Brassica o/era,cea) Trichol1lll.~ia )Ii. 15 Ceanoth'us sp. Cm/alaria mllllldiJolia Autographa caUforrdea, 31 T)·iclwpillsio. ni. 15 cedar (see Juniperus sp.) Cr%ll rflJIi/o/WI cedar, western red (see Thuja plica/a) PIII'Ua()l.{lIsio includens, 17 celery (see Apium {rraJleo/e1•.· \ cucumber (SE'E' ('1«'1I111i8 satil'lts) Centcmrea solslilia.lis Cllrllmi.~ /tIl'/" Autograp/ta bilobct, 30 AU/oglVphcl (·ali/rlmica., 31 Chaenactis slel'ioides Trichof!/u.qia ni, 15 T'richoplusia ni, 15 Cucumi,q sa,/intH Chenomeles jllPonica flu(ograpJw calijorn-ica, 31 Trichoplusia 'Ili, 15 Trichopl!(.~ia ni, 15 Chenopodium album ('urcltl"bita maximo Autographtl califomica, 31 AII/lIgrapJ/(I. cal((ol"llic(L, 31 118 T.ECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE

Trichoplusia ni, 15 Gladiolus sp. Curcurbita pepo Autographa biloba, 30 TrichopltLsia ni, 15 Glycine max current (see Ribes sativum) Aflagrapha aerea, 23 Dahlia variabilis Autoplusia egena, 19 Autographa californica, 31 Pseudopl1UJia includens, 17 DauclLs carota goldenrod (see Solidago sp.) Autographa califo1nica, 31 gooseberry (see Ribes grossularia) Autoplusia agena, 19 Gossypium sp. Syngrapha falcifem, 43 A utographa .californica, 31 Trichoplusia ni, 15 Trichoplusia ni, 15 day-flower (see Commelina sp.) Gossypium hirsutum Delphinium sp. Argyrogram7na verruca, 13 Autographa biloba, 30 Pseudoplusia includens, 17 A utoplusia egena, 19 grape (see Vitis sp.) Polychrysia moneta, 24 grasses, various unspecified (see also Poaceae) Polychrysia morigera, 24 Chry8aspidia contexta, 53 Dianthus caryophyUus Chrysaspidia putnami, 53 Trichoplusia ni, 15 groundsel (see Senecio sp.) dock, curly (see Rumex crispus) hawksbeard (see Crepis sp.) Doronicum sp. hedge nettle, bugle (see Stachys ajugoides) Autographa californica, 31 hedge nettle, rigid (see Stachys rigida) dusty miller (see Senecio cineraria) Hedra sp. Erechtites arguta Autographabiloba, 30 Autographa californica, 31 Helianthus sp. Erigeron canadensis Autographa biloba, 30 Trichoplusia oxygramma, 16 Autographa californica, 31 Erodium sp. A utographa jlagellum, 33 Trichoplusia n

Juniperus sp. marigold, big (see Taaetes erecta) Syngrapha epigaea, 39 marigold, pot (see Calendula officinalis) Justicia americana Medicago sativa Argyrogramma verruca, 13 Autographa biloba, 30 kale (see Brassica oleracea) Autographa cali/ornica, 31 Kalmia angustifolia Pseudoplusia includens, 17 Syngrapha epigaea, 39 Trichoplusia ni, 15 Kalmia sp. Meliolotus alba Autographa californica, 31 Autographa californica, 31 Lactuca .sp. melon (see Cucumis melo) Autographacalifornica, 31 Mentha sp. Autollrapha precationis, 29 Autographa cali/ornica, 31 Lactuca canadensis Autoplusia egena, 19 Trichoplusia ni, 15 Argyrogramma verruca, 13 Lactuca sativa Rachiplusia ou, 20 Autographa biloba, 30 Trichoplusia ni, 15 Autographa californica, 31 Mentha arvensis Pseudoplusia includens, 17 Allagrapha aero ides, 22 Syngrapha falcifera, 43 Mentha spicata Trichoplusia ni, 15 Autographa californica, 31 lamb-kill (see .Kalmia angustifolia) A1doplusiaegena, 19 lamb's quarters (see Chenopodium album) mexican tea (see Chenopodium ambrosioides) Lantana sp. mignonette, common (see Reseda odorata) Autoplusia olivacea, 18 milkweed (see Asclepias sp.) Pseudoplusia includens, 17 Mimulus cardinalis Larix sp. Autographa biloba, 30 Syngrapha epigaea, 39 Autoplusia oliwcea, 18 Syngrapha viridisigma, 41 mint (see Mentha sp.) Larix laricina mint, wild (see Mentha arvensis) Syngrapha alias, 45 monkey flower, scarlet (see Mimulus cardinalis) Syngrapha rectangula, 48 morning glory (see Ipomoea purpurea) Syngrapha viridisigma, 41 mustard, black (see Brassica nigra) larkspur (3ee Delphinium sp.) mustard, field (see Brassica campestris) Lathyrus odoratus mustard, white (see Brassica hirta) Trichoplusia ni, 15 nasturti urn (see Tropaeolum sp.) lettuce (see Lactuca sativa) Nemophila menziesii lettuce, wild (see Lactuca sp.) Autographa cali/ornica, 31 Liatris sp. nettle, stinging (see Urtica sp., Urtica diotica) Autographa flagellum, 33 nettle, Lyall (see Urtica lyallii) lilac, wild (see Ceanothus sp.) Nicotianc. glauca Linum sulcatum Trichoplusia ni, 15 Autographa californica, 31 Nicotiana tabaccum lupine (see Lupinus sp.) Autographa biloba, 30 Lupinus sp. Argyrollramma verruca, 13 Autographa cali/ornica, 31 Pseudoplusia includens, 17 Trichoplusia ni, 15 Rachiplusia ou, 20 Lycopersicon esculentum Trichoplusia ni, 15 Autographa californica, 31 Trichoplusia oxygramma, 16 Pseudoplusia inclwiens, 17 Nicotiana trigo1wphylla Trichoplusia ni, 15 Trichoplusia ni, 15 maize (see Zea mays) okra (see Hibiscus esculentus) mallow (see Malva sp.) onion (see A.ilium cepa) Malus pumila orange, sweet (see Citrus sinensis) Autogra'Pha californica, 31 painted cup (see Castilleja sp.) Malva sp. Papaver sp. Autographa cali/ornica, 31 .Trichoplusia ni, 15 Autoplusia egena, 19 parsnip (see Pastinaca sativa) Trichoplusia ni, 15 Pastinaca sativa Malva rotundi/olia Autographa precationis, 29 Autographa californica, 31 Trichoplusia ni, 15 marguerite (see Chrysanthemum jrutescens) pea, black-eyed (see Vigna unguiculata) 120 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE peas (see Pisum eativum) Pinus contorta pea, sweet (see Lathyrus oooratuB) Syngrapha alias, 45 Pectis pappoBa SY11lJrapha t'iridiBigma, 41 Trichoplusia ni, 15 Pinus nwnticola Pelargonium sp. Syngrapha alias, 45 Autographa biloba, 30 Syngrapha celBa, 49 Argyrogramma verruca, 13 Syngrapha rectangula, 48 PBeudoplusia includenB, 17 PiBum Bativum Persea americana Autographa californica, 31 PBeudoplusia includens, 17 Plantago sp. Petunia sp. Autographa biloba, 30 Trichoplusia ni, 15 Autographa californica, 31 petuna (see Petunia sp.) Autographa precationis, 29 Phacelia sp. Syngrapha falcifera, 43 Autographa biloba, 30 TrichopluBia ni, 15 Phaseolus sp. plantain (see Plantago sp.) Autograp1u:L californica, 31 plum, India (p1"'.tnus sp. 1) Autoplusia egena, 19 Poaceae sp. PBeudoplusia includens, 17 ChryBaspidia putnami, 53 Syngrapha falcifera, 43 Podocarpus .sp. Trichoplusia ni, 15 Autographa califarnica, 31 Phaseolus limensiB poinsettia (see Euplwrbia pulchemma) Autoplusia egena, 19 poplar, balsam (see Populus balsamifera) Autoplusia olivacea, 18 poppy (see Papaver sp.) Trichoplusia ni, 15 poppy, California (see EschBClwlzia californica) PhaBeolus lunatus Populus balsamife·m Tr'ichoplusia ni, 15 Autographa ampla, 35 PhaBeoluB vulgaris Populus tremuloideB Autographa biloba, 30 Autographa ampla,35 Autoplusia egena, 19 Plusia balluca, 21 Autoplusia olivacea, 18 Snygrapha epigaea, 39 Trichoplusia ni, 15 potato (see Solanum tuberoBum) Philodendron sp. Prunus sp. Trichoplusia ni, 16 Autographa ampla, 35 Picea engelmannii PBeudotsuga menzieBii Syngrapha alias, 45 Syngrapha alia8, 45 Syngrapha celsa, 49 Syngrapha rectangula, 48 Syngrapha rectangula, 48 Syngrapha viridisigma, 41 Syngrapha viridiBigma, 41 puncture weed (see Tribulus sp.) Picea glauce purple queens (see Tradescantia jluminensiB) Syngrapha alias, 45 quince, dwarf Japanese (see ChaenomeleB japonica) Syngrapha celsa, 49 radish (see Raphanus BativuB) Syngrapha rectangula, 48 ragwort, tansy (see Senecio jacobaea) Syngrapha viridiBigma, 41 Raphanus Bativus Picea mariana Autographa califarnica, 31 Syngrapha aliaB, 45 Triclwplusia ni, 15 SY71grapha rectangula,48 raspberry (see Rubus idaeus) Syngrapha viridisigma, 41 ReBeda odorata Picea rubens Triclwplusia ni, 15 Syngrapha alias, 45 Rhamu8 sp. Syngrapha rectangula, 48 Autographa mappa, 34 Picea BitchensiB Rheum sp. Syngrapha rectangula, 48 Autographa californica, 31 .r Syngrapha viridiBigma, 41 rhubarb (see Rheum sp.) pine, jack (see Pinus bankBiana) Rhododendron sp. pine, lodgepole (see Pinus contorta) Autographa californica, 31 pine, western white (see PinuB monticola) Ribe8 gros8ularia Pinus banksiana Autographa californica, 31 Syngrapha alias, 45 Ribe8 8ativum Syngraphaepigaea, 39 Autographa californica, 31 PLUSIINAE OF AMERICA NORTH OF MEXICO 121

Rosa sp. spiderwort (see Tradescantia fluminensis) Autographa californica, 31 Spinacea oleracea roses (see Rosa sp.) Autographa californica, 31 Rubus idaeUlJ Trichoplusia ni, 15 Autographa californica, 31 spinach (see Spinacea oleracea) rue, meadow (see Thalictrum sp.) spiraea (see Spiraea sp., Spiraea salicifolia) Rumex sp. Spiraea sp. Autographa californica, 31 Syngrapha epigaea, 39 Trichoplusia ni, 15 Spiraea salicifolia Rumex crispus AUagrapha aero ides, 22 Argyrogramma verruca, 13 spruce (see Picea sp.) safflower (see Carthamus tinctorius) spruce, black (see Piceamariana) sage (see Salvia sp.) spruce, Engelmann (see Picea engelmannii) Sagittaria sp. spruce, red (see Picea rubens) Argyrogramma verruca, 13 spruce, sitka (see Picea sitchensis) Salix sp. spruce, white (see Picea glauca) Autographa ampla, 35 squash (see Curcurbita maxima) Syngrapha epigaea, 39 squash (see Curcurbita pepo) saltbush, four-wing (see Atriplex canescens) Stachys ajugoides Salvia sp. Autographa biloba, 30 Trichoplusia ni, 15 Autographa pasiphaeia, 32 Salvia leucophyUa Stachys rigida Autographa biloba, 30 Autographa pasiphaeia, 32 Autographa califarnica, 31 storksbill (see Pelargonium sp.) Sambucus canadensis strawberry (see Fragaria sp.) Autographa californica, 31 sugar beet (see Beta vulgaris) saskatoon-berry (see Amelanchier alnifalia) sunflower (see Helianthus sp.) Scrophularia californica sweet gale (see Gale palustris) Autographa labrosa, 33 Symphytum sp. sedge (see Carex sp.) Autoplusia egena, 19 Senecio sp. Tagetes erecta Autoplusia egena, 19 A utoplusia egena, 19 Autoplusia olivacea, 18 tamarack (see Larix laricina) Senecio cineraria Taraxacum officinale Autographa californica, 31 Autographa bimaculata, 29 Senecio cruentis Autographa precationis, 29 Trichoplusia ni, 15 Syngrapha falcifera, 43 Senecio jacobea Trichoplusia ni, 15 Autographa californica, 31 Thalictrum sp. Autoplusia olivacea, 18 EOBphoropteryx thyatyroides, 27 Senecio mikanioides Pseudeva purpu1"igera, 26 Trichoplusia ni, 15 thistle (see Carduus sp., Cirsium sp., Cirsium .horridu­ service berry (see Amelanchier florida) lum) Shepherdia canadensis thoroughwort (see Euphatorium sp.) Autographa ampla, 35 Thuja plicata Smilax rotundifolia Syngrapha al·ias, 45 Trichoplusie ni, 15 Syngrapha viridisigma, 41 snapdragon (see Antirrhinum sp.) tobacco, tree (see Nicotiana glauca) Solanum tuberosum tobacco (see Nicotiana tabaccum) Autographa californica, 31 tomato (see Lycopersicon esculentum) Trichoplusia ni, 15 Tradescantia fluminensis Solidago sp. Argyrogramma verruca, 13 Pseudoplusia includens, 17 Mouralia tinctoides, 11 Trichoplusia oxygramma, 16 Pseudoplusia includens, 17 soybean (see Glycine max) Tribulus sp. Sparganium sp. Trichoplusia ni, 15 Chrysaspidia putnami, 53 Trichosporum sp. S'p«thiphyllum sp. Argyrogramma verruca, 13 Argyrogramma verruca, 13 Trifolium sp. spearmint (see Mentha spicata) Autographa californica, 31 122 TECHNICAL BULLETIN 1567, U.S. DEPARTMENT OF AGRICULTURE

Rachiplusia ou, 20 Vaccinium macrocarpon Trichoplusia ni, 15 Syngrapha faleifera, 43 Trifolium incarnatum verbena (see Verbena np.) Autographa biloba, 30 Verbena sp. Triticum aestivum Autographa biloba, 30 Autographa californica, 31 Autographa precationis, 29 Rachiplusia au, 20 Autoplusia egena, 19 Tropaeolum sp. vetch (see V·icia sativa) Rachiplusia ou, 20 Viburnum sp. Tropaeolum majus Syngrapha fakifera, 43 Trichoplusia ni, 15 Viburnum cassinoides Tsuga heterophylla Autographa ampla, 35 Syngrapha alias, 45 Vicia sativa SY7IfJrapha celsa, 49 Autographa californica, 31 Syngrapha rectangula, 48 Vigna sinensis Syngrapha viridisigma, 41 Trichoplusia ni, 15 Tsuga mertensiana Vigna unguiculata Syngrapha rectangula, 48 Argyrogramma 'Verruca, 13 Syngrapha viridisigma, 41 Vigulera deltoidea turnip (see Brassica rapa) Autographa califarnica, 31 Unica sp. Viti sp. Abro6tola ova lis , 11 Autographa californica, 31 Allagrapha aBrea, 23 wandering jew (see Tradescantia fluminensis and Ze­ Unica diotica brena pendula) Abrostola urentis, 10 watermelon (see Citrullus vulgaris) Urlica lyaUii wheat (see Triticum aestivum) Abrostola urentis, 10 willow (see Salix sp.) Autog-rapha mappa, 34 willow, water (see Justicia americana) Syngrapha falJifera, 43 withe-rod (see Viburnum cassino ides) Vaccinium sp. wormseed (see Chenopodium ambrosioides) Ghrysanympha formosa, 26 Zeamays Syngrapha epigaea, 39 Autographa californica, 31 Syngrapha octo scripta, 39 Syngrapha faleifera, 43 Syngrapha orophila, 41 Trichoplusia ni, 15 Syngrapha u-aureum, 44 Zebrina pendula Vaccinium angustifolium Mouralia tinctoides, 11 Syngrapha epigaea, 39 Pseudoplusia includens, 17

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