PROC. ENTOMOL. SOC. WASH. 110(2). 2008. pp. 504 515

SITOCHROA PALEALIS: A PALEARCTIC PYRAUSTINE (: : CRANIBIDAE) NEWLY INTRODUCED TO NORTH AMERICA

STEVEN PASSOA, GEORGE BALOGH, AND M. ALMA SOLIS (SP) U.S. Department of Agriculture, and Plant Health Inspection Service, Plant Protection and Quarantine, The Ohio State University, Museum of , 1315 Kinnear Road, Columbus, OH 43212, U.S.A. (e-mail: steven.c. [email protected] ); (GB) 6275 Liteolier Street, Portage, MI 49024, U.S.A. (e-mail: [email protected]); (MAS) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, do National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, MRC 158, Washington, DC 20013-7012, U.S.A. (e-mail: [email protected])

Abstract. ---Sitochroa pa/ca/is (Denis and Schiffermüller) is recorded from North America for the first time. Adults were collected in Illinois, Indiana, Michigan and Wisconsin. The capture of larvae and adults in four states over a six-year period is strong evidence that S. pa/ca/is is established in North America. Characters separating S. pa/ca/is from other Sitochroa in North America are given. The potential impact of S. pa/ca/is to prairies, agriculture, and biocontrol of noxious weeds is discussed. Ke y Words. , introduction, immatures, North America, Apiaceae

Palearctic frequently are dis- published in faunal studies. For example, covered in North America. A very Spodoptera /ittora/i,r (Bdv.) was listed in conservative list published by Leverton a study of owlet moths of Ohio (Rings et (2001) includes approximately 85 al. 1992) and an appendix of "Quaran- of Lepidoptera introduced from Great tine Interceptions" was published by Britain to North America. In some cases, Kimball (1965). As these authors cor- it is not clear if Old World species have rectly noted, species listed in such reports come from Europe or Asia (Leverton should not be automatically included as 2001: 253). The exact number of species additions to the North American fauna. included in any list of introduced Lepi- Regulatory entomologists consider an doptera depends upon how the word organism to be introduced only if there "introduced" is defined (Frank and is evidence of an established population. McCoy 1990), and if species occurring The best criteria to measure establish- in certain agricultural situations such as ment would reflect the populations weed biological control programs, heat- ability to breed and survive for multiple ed greenhouses, ports-of-entry, and generations. For example, successful stored grain warehouses are excluded. overwintering of a population over Intercepted organisms or eradicated in- several seasons, presence of several life festations of exotic species sometimes are stages, presence of the organism over a

VOLUME 110, NUMBER 2 505 wide area, or the sudden colonization of METHODS AND MATERIALS a new host are all indications that a Sitochroa pa/ca/is specimens examined population is established in a region during this study are deposited in the (Pender 1983). following collections: BSC = Brian G. In this paper we report Sitochroa Scholtens collection, Charleston, South pa/ca/is (Denis and Schilfermüller) Carolina; GBC = George J. Balogh (: Pyraustinae) from North collection, Portage, Michigan; LFC = America for the first time. In 2004, Leslie A. Ferge collection, Middleton, specimens of S. pa/ca/is were brought to Wisconsin; NMNH = National Museum The Ohio Lepidopterists Society identi- of Natural History, Washington, D.C.; fication meeting called "ID day." Based SPC = Steven Passoa collection, Colum- on the male genitalia, they were tenta- bus, Ohio; and SMTD = Staatliches tively identified as S,tochroa pa/eat/s by Museum für Tierkunde, Dresden, Ger- SP. This was confirmed by comparing many. larval and adult specimens of S. pa/ca/is Genitalia were dissected using the from Illinois with specimens from Eu- procedure given by Clarke (1941). Pho- rope and Asia in the National Museum tographs of the larvae were taken by SP of Natural History, Smithsonian Institu- using a Minolta X-700 camera with an tion, Washington. DC, by MAS. 80PX ringflash. Adults were photo- Sitochroa pa/ca/is occurs from eastern graphed by Todd Gilligan with a Canon Russia west to Great Britain and Portu- D60 camera and Aristo DA- 10 light box. gal, and south to Greece, Italy (Karsholt Genitalia were photographed using a and Razowski 1996) and North Africa Nikon DXM 1200 camera on a Nikon (Balachowsky 1972). In Asia, Park Optophot 2 microscope (by SP) or a pa/ca/is from Korea, (1979) records S. Nikon Coolpix 4500 on a Leica MZ 12.5 Japan, and China. The feeds on Apiaceae (= Umbelliferae). Recorded stereoscope (by GB). host genera are Daucus L., Foenicu/um MATERIAL EXAMINED L., Laserpitium L., Mill., All records represent a single adult Peucedanum L., Seseli L., and Si/aiim Mill. (Hasenfuss 1960, Gaedike 1980). capture unless otherwise noted. Recent checklists (Fletcher and Nye ILLINOIS: Cook Co., Air Station 1984, Maes 1994, Karsholt and Razowski Prairie, 1 Aug 2002, R. Panzer (SMTD); 1996) placed pa/ca/is in Sitochroa, but Chicago Ridge Prairie N. P., 22 Aug previous authors have used an assortment 2003, G. Derkovitz (USNM); 18 Aug of other genera, e.g., Ph/yctaenodes 2003, R. Panzer (SPC); Orland Tract wet (Hampson 1899). Pvrausta (Hasenfuss remnant, 8 Aug 2003, R. Panzer (SPC); 1960, Gaedike 1980), and Paintbrush Prairie, 21 Jun 2004, R. (Balachowsky 1972, Bollmann 1955). Panzer (GBC); 29 Jul 2004, R. Panzer The following synonymy is currently in (SMTD); 12 Sep 2004, R. Panzer (SPC); The Global Lepidoptera Name Index Santa Fe Prairie, 6 Jul 2005, G. Derko- website (Beccaloni et al. 2005): Pha/aena vitz (SPC); 15 Jul 2005, R. Panzer, 2 Jlaveo/ata Rott., Pvra/is se/ena/is Hbn., specimens (SPC). DuPage Co.. W. Chi- Botys anaxisa/is WIk., and Spilodes a/gir- cago Prairie, 7 Aug 2004, R. Panzer a/is Allard. A subspecies with poorly (GBC); Il Jul 2005, R. Panzer (SPC). developed dark scaling along the wing Livingston Co., 1.5 mi W Kankakee veins, Loxostege (Ph/vctaenodes) pa/ca/is county line on Hwy 17, roadside adja- var. extrema/is Caradja, was described cent to agricultural fields, larvae collect- from China (Caradja 1916). ed from heads, 27 Aug 506 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

2006, G.J. Balogh (SPC. NMNH). Will aureola/is and S. chortalis have forewings Co., Midewin Deg Upland, 26 Jul 2003, with the fully developed pattern of the R. Panzer (GBC). ; the ground color is dark orange INDIANA: Lake Co., Dupont Dune yellow in the former, whitish buff in the and Swale N. p., 18 Jul 2004, R. Panzer latter. The forewings of S. dasconalis are (S PC). cream to yellow brown with orange MICHIGAN: Allegan Co., T3N scales at the base of the costa (Fig. 1). R14W S23, Heath Twp, larvae collected Sitochroa pa/ca/is has pale yellow fore- from Daueus carota heads along roadside wings (described by Goater (1986) as 3 Sep 2006, adults emerged 13 Jun 2007 "pale sulphur-yellow") with a faint and 18 June 2007, G.J. Balogh, 2 greenish tint and no orange scales on specimens (GBC). Charlevoix County, the costa! vein (Fig. 3). There are no Beaver Island, Sand Bay, Lake Michigan species of Pyraustinac north of Mexico shoreline. 22-26 July, 67 specimens, BL with a faint greenish tint on the fore- traps, G.J. Balogh (GBC). Cheboygan wings like S. pa/eat/s. Other differences Co., Wildwood Road S of Indian River, are that the underside of S. pa/ca/is tends jack pine plain, nectaring on IVIonarda to be more strongly marked than S. sp. during day along with Pirausta spp., dasconahs (Figs. 2, 4), and S. palea/i,r 27 Jul 2006, B.G. Scholtens, 2 specimens often has a patch of dark scales at the (BSC); Hogback Road, at Carp Creek, N end of the forewing cell (Fig. 3). of Burt Lake, mature woods, 31 Jul Some variation was noted in the wing 2006, B.G. Scholtens (BGS). Eaton Co.. markings of S. pa/eat/s. A limited num- 7 mi N. of Charlotte, T3N, R5W, S32, ber of Old World specimens, mostly mature beech/maple forest, at UV light, from China and Japan (at the Staatliche 26 July 2006, J.H. Wilterding (GBC); at Naturhistorische Samml ungen, Museum UV light, 28 Jul 2006, J.H. Wilterding, 2 für Tierkunde, Dresden Germany, (M. specimens (GBC). Nuss, pers. comm) and The National WISCONSIN: Door County, Baileys Museum of Natural History, Smithso- Harbor Twp [township], 25 August 2004, nian Institution, Washington, DC (ex- J. Stiefel; 15 July 2005, J. Stiefel; 29 July amined by MAS), also Inoue et al. 1982, 2006, J. Stiefel (all GBC). Iowa Co, T8N, plate 42, fig. 16) have a forewing pattern R1E, Sec 10, 29 July 2007, flushed from that varies strongly from conspicuously Queen Annes Lace, L.A. Ferge (LFC). dark-colored veins (on both the dorsal and ventral surfaces) to being dorsally Identification of Sitoc/iroa pa/ca/is unmarked and uniformly yellowish. The Adult.—Munroe (1976) stated that apex of the hindwing sometimes has light Sitochroa is easily recognizable by the to dark apical markings. This variation is wing pattern that includes "weakly not the result of scales being lost in worn dentate or crenulate bands parallel to individuals. We are unable to determine the termen" which are "often connected the source of the North American along the veins." He noted "the pattern introduction based on wing markings. is recognizable even in S. dasconalis Further study, using molecular methods (Grote), where it is weakly developed." and documenting geographical variation Compared to the three native U.S. in Old World populations, would be species (S. aureolalis (Huist), S. chortalis helpful in this regard. (Grote) and S. dasconalis), the color Because S. pa/ca/is and S. dasconalis pattern of S. pa/ca/is is most similar to have reduced wing patterns, they are that of S. dascona/is because only the easily confused if the forewings are postmedial line is recognizable. Both S. damaged. This often occurs in bucket 507 VOLUME 110, NUMBER 2

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/oM units, Adult and lar\aI hahitiis ol Sil l iii. I S. /aoiu//Ls. imiL. dortl \ \\ . S. male, ventral view. 5. S. pa/ea/is, late male, ventral view. 3, S. palealis, male, dorsal vics. 4.S. pa/ca/is, instar larva on Daucus carola seed head, lateral view. 508 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

light traps commonly used in fauna] bursae in S. dascona/js is round to oval surveys. Thus, the male and female (Fig. 15) whereas that of S. pa/ecu/s is genitalia are figured and briefly com- elongate and bean-shaped (Fig. 14). This pared for use with doubtful specimens shape difference is best seen in water from trap samples. before the genitalia are slide mounted. The male genitalia of S. pa/ca//s are Larva.—Sitochroa pa/ca/is is one of slightly different from those of other the few crambid larvae associated with North American species. The incus in the flower heads of Apiaceae in North the native species is rounded compared to America (Allyson 1981, 1984 Robinson S. pa/ca/is where it is more acute (Figs. 6- et al. 2002). It is most likely to be 7). All members of the genus have on the confused with Depressaria pastinace//a valvae a large, arched, scierotized clasp- ( Duponchel)() the parsnip er" with two ventrally directed thornlike webworm, which sometimes is recorded processes, and a prominent strongly from Dauctus carotcu L. (Robinson et al. scierotized process from one side of the 2002: 530). Early to middle instars of S. aedeagus tip (Munroe 1976). The sinus pa/ca/is (Fig. 16) are similar to D. between the prongs of the clasper on the paslinacc//a because both species have valvae is rounded in S. dascona/is ( Fig. 7) uniformly dark body pinacula. At matu- and quadrate in S. palea/is (Fig. 6). Just rity several color differences are appar- basal to the clasper, both species have a ent. The most obvious is that S. pci/ca//s mesally directed process which is elongate has a pale area in the middle of the and fingerlike in S. dascona/is, but round- abdominal L and SV pinacula (Fig. 5) in ed or triangular in S. pa/ca/is. The D. pastinace//a all the body pinacula are posterior margin of the juxta is U-shaped uniformly dark (Stehr 1987: fig. 26.49). in S. pa/ca/is but widely V-shaped in S. The easiest structural character to sepa- dasconalis (Figs. 6-7). The distal ventral rate the two species is the number of sclerotized process of the aedeagus of S. setae in the prothoracic prespiracular dascona/is is rounded in cross section and pinaculum. Sitochroa pa/ca/is has a less than one-third the width of the bisetose prespiracular group while aedeagus. In S. D. pa/ca/is, the process is pastinacc//a has a trisetose prespiracular stouter, laterally flattened, and from one- group. This difference is valid for larvae third to one-half the aedeagus width when of both species from the 2nd through the viewed laterally (Figs. 8-9). Both species last instars (first instar larvae were not have deciduous cornuti, thus some vari- examined). For a larval description of D. ation in the number of cornuti is expected. pastinc,cc//a consult Passoa (1995). The Females of both S. dascona/is and S. larva of S. pa/cc//is was described by pa/ca/is have a long ductus bursae with Bollmann (1955) (translated to English the caudal end, adjacent to the ostium by Ericson 1956) and Hasenfuss (1960). bursae, widened and partially sclerotized Users of the Ericson (1956) translation (Figs. 10-13). In S. dascona/ir, the cau- should note that one character present in dal end of the ductus bursae is abruptly S. pa/ca/is was called "sklerotisiertes widened, being as wide or wider than Grubchen" by Bollmann (1955: fig. long, with strongly developed lateral 225) and "Grubchen" by Hasenfuss pockets. The caudal end of the ostium (1960: fig. 203). This was translated by bursae in S. pa/cc//is is longer than wide Ericson (1956) as "scierite of the pit." It and without large lateral pockets. Both is more correctly called a tonofibrillary species have a membranous appendix platelet which in S. pa/cc//is is located on bursae extending from the corpus bur- A3-6, posterodorsad of the spiracle, in a sae. When fully inflated, the appendix shallow depression. qqw-

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palealo, ventral view. 7. S. Figs. 6-9. Male genitalia of Sitec/,roa with aedeagus removed. 6. S. dasconalis, aedeagus, lateral view. Iasconalis. ventral view. 8, S. palea/is, aedeagus. lateral view. 9, S. many of the same characteristics, except Separation of S. palealis from other North American pyraustine larvae on that the body does not have a black Apiaceae across the continental United middorsal stripe, but rather a grayish or States is more problematic because of the reddish stripe in some specimens. Also, paucity of morphological knowledge on S. pa/ca/is does not have prominent the larvae associated with these plants. longitudinal stripes, and although the prothoracic shield is heavily pigmented, For example, Robinson et al. (2002) L. recorded Loxostege sticticalis (L.) from the pattern is not two lateral lines as in Daucus carota. Allyson (1984) stated that sticticalis (see Allyson 1984: fig. 147). the last instar larva of L. sticticalis has 2 Pupa.—Mosher (1916) did not include SV setae on Al, prominent longitudinal Sitochroa in her North American pupal stripes on the body, a prothoracic shield study, but Patoka and Turááni (2005) with heavy pigmentation, the SD! pina- described and illustrated the pupa of S. cula fully pigmented on Al and AS, and pa/ealis. They noted that pilifers are a black middorsal stripe from the pro- present, the maxillary palpi are round thorax to A9. Sitochroa pa/ca/is has and widely separated from the maxillae.

510 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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10 11

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12 13 •L I

14 15

Figs. 10-15. Female genitalia 01 .Siiuc Iii a. M. pa/ea/i.. ventral \ ivy . I I. S. /a.sc000/jv. 12. S. pa/calLs. ventral view. caudal end of ductus bursae, ventral view. 13, S. ilasvo,za/js, caudal end of ductus bursae, ventral view. 14, S. pa/ca//v, appendix bursae of the corpus bursae, fully inflated, ventral view. 15, S. dascona/Lv, appendix bursae of the corpus bursae, fully inflated, ventral view. 511 VOLUME 110, NUMBER 2

. I •• fi

, • 16 Daucus carota seed head, lateral view. lie 16 Sitaclirea pa/euilix. Early instar larva from

the appendages extend past forewing entian Great Lakes suggests that ports-of- apices, the metathoracic legs are hidden, entry are a likely possibility. Adults could the prothoracic femora are exposed, the have been trapped accidentally in ships or maxiliac extend to the caudal margins of planes destined for North America from the wings, and the hooked spines of the Europe or other parts of the Old World. poorly developed cremaster are arranged Crambid moths are sometimes collected in two widely spaced groups. "Punctura- from planes at U.S. ports-of-entry, al- tion" (see Patoëka and Turëáni 2005: though infestation rates of European 273, couplet 41) is "absent" in S. planes are low (Dobbs and Brodel 2004). Another possible pathway is that larvae, pa/ca/is. This refers to the texture of the cuticle of the middle abdominal seg- or perhaps eggs, may have been intro- duced with herbs in the Apiaceae. Several ments, which in S. pa/ca/is is shagreened and lacks punctures. Sitochroa pa/ca/is is genera in this family often are imported into the U.S. as fresh foliage (J. Brusch, similar to Loxostegc from Europe and North America (Heppner 1975, Patoèka pers. comm.) including two known hosts and Turèáni 2005: plate 154, 154a) and (E.iaucus. Focnicu/uin) of S. pa/ca/is. Using criteria listed by Pender (1983), to S. dasconalis from Illinois (specimen in SP coil.) in that the spiracular furrows there is little doubt that S. pa/ca/is is of A2 and A3 are fused to form a shiny established in the United States. The scierotized, slightly raised, curved ridge. capture of adults over a six-year period must be over- Characters to separate the pupa of S. indicates that S. pci/ca/is wintering successfully from year to year. palealis from other North American Pyraustinae are not available, but the As would be expected in a breeding form of the spiracles on A2 and A3 will population. larvae were present. The likely be part of the diagnosis of this colonization of Daucus is apparently a taxon. In Europe. Loxostege differs from recent one because previous surveys of Apiaceae in central Illinois by members Sitochroa in having pointed maxillary palpi. Dl not on the dorsum of the of the Berenbaum lab (University of cremaster, and usually exposed metatho- Illinois, Urbana) since the 1980s failed racic legs (Patoëka and Turöãni 2005). to discover any crambid larvae on this host (T. Harrison, pers. comm.). Lastly, DiscussioN as expected in a spreading established pa/ca/is It is unclear how S. palealis entered the population, the range of S. United States. The distribution near major already spans four states (Illinois, In- airports and shipping lanes of the Lar- diana, Michigan. and Wisconsin). 512 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Although the complete life cycle of S. decades. Some are serious pests, such as pa/ealis in the United States is unknown, nub/la/is (l-Ibn.) (Pyraustinae) most aspects of the biology (based on first officially reported from North collection data in the Material Examined America in 1917 (Glover et al. 1991). section) agree well with the European Other introductions, for example, literature. Moths were collected from Sc/er- ocona acute/la (Evers.) (Wagner et al. June through September in the U.S. 2003), are not expected to be agricultur which is similar to the midsummer (June ally significant. The potential impact of to August) flight period in Denmark S. pa/ca/is in North America is more (Palm 1986). The American population difficult to characterize. colonizes multiple sites (woods, prairies, Balachowsky (1972) claimed that car- and weedy fields) as does S. pa/ca/is in rot plants grown to seed, presumably for Denmark where the favored habitats are seed selection and genetic improvement open woods, disturbed fields, and other were completely destroyed b y S. pa/cali,s sunny places (Palm 1986). Monarda L. in Europe. Carrots grown for human (Lamiaceae) is a nectar source for S. consumption were not affected. An palealis in Michigan, and adults were argument could be made that £ pa/ca/is captured in light traps throughout its is a potential minor pest of related root range in the United States. This parallels crops (e.g., parsnip) and some seeds used the capture of S. pa/ea/is at lights for spices (e.g., Herac/eum). (Goater 1986) or flowers (Palm 1986) in S. Fortunately, pa/ca/is is easily controlled with Europe. GB collected larvae of S. insecticides (Balachowsky 1972) which palea/is in heads of Daucus from Michi- should limit any pest outbreaks. gan and Illinois during August where Apiaceae is well represented in eastern they fed in a silken pouch sometimes tallgrass praries (Metzler et al. 2005). covered by plant fragments. Pupation Some of these genera could be at risk if and adult emergence occurred in spring their seed heads are attacked by high of the following year. A similar larval populations of S. and pupal phenology was noted by pa/ca/is. Sitocliroa dascona/is is considered a "conservation Goater (1986) and Bollmann (1955) Europe. in priority" in Illinois (Illinois Department of Natural Resources 2005). Confusion We do not have enough information of adult S. pa/ca/is with S. da.sconalis to characterize the population dynamics could affect monitoring programs for the of S. palea/is in the U.S. with certainty, latter species. Larval and adult collections suggest one Because some introduced Apiaceae are generation per year with the prepupa toxic to humans (Gleason 1952), S. pa/eu/is being the overwintering stage. Popula- may be somewhat beneficial as a biocontrol tion numbers vary widely in Denmark, in agent of these exotic weeds. A similar some years this species is common, in example is the introduced moth other years it is absent (Palm 1986). Agonop- terix aistroelneriana (Depressariinae) which There is one generation per year in Great has sometimes destroyed stands of the Britain (Goater 1986), but Balachowsky introduced weed (onium macu/alum (Cas- (1972) suspected that S. pa/ca/is may tells and Berenbaum 2006). have two generations in some Mediter- In summary, S. pa/ea/is could be a ranean countries. Multiple generations minor pest and a source of confusion per year might be possible if S. pa/ealis with a native species of special concern. spreads into the southern United States. However, it may also be beneficial as a Introduced species of Crambidae have weed biological control agent. Future been known in North America for many careful survey will be needed before the VOLuME I tO. NUMBER 2 true effects, if any, of this introduction for his fabulous moth photographs and are known. preparation of the plates. Jonathan Brusch (U.S. Department of Agriculture, Besides S. palealis, several other cram- bid species merit attention because of Animal and Plant Health Inspection their potential impact on North Ameri- Service, Plant Protection and Quarantine. can agriculture. Two examples are Mar- Boston, Massachusetts) provided impor- tant data from interception records at uca iitrata (F.) and Diaphania inc//ca (Saunders), both likely established in U.S. ports. Useful comments on this southern Florida (Heppner 2003). An manuscript, and the reared S. dasconalis invasive European species, Duponchelia used in this study, are yet two more examples of the many significant contri- Jiealis (Zeller), was found on cut flowers in three Ontario greenhouses butions I. Harrison (University of Illi- (Canadian Food Inspection Agency nois, Urbana, lillinois) has made to 2005) and a California greenhouse (Ep- Illinois Lepidoptera. J. Fraudenstein and stein 2005), in additon to being inter- J. Wenzel (The Ohio State University, cepted at ports-of-entry in England Columbus, Ohio) translated Danish and (Clark 2000). At U.S. ports, D. tbvcalis French literature, respectively, into En- is most frequently associated with cut glish for us. Stuart McKamey (Systematic flowers, herbs, and peppers originating Entomology Laboratory, Plant Sciences from several European and Mediterra- Institute, Agricultural Research Service, nean countries (USDA/APHIS/PPQ Pest U.S. Department of Agriculture, Wa- ID database). The only rearing record at shington, D.C.) improved the manuscript a U.S. port is a malformed adult with his editorial suggestions. We thank emerged from peppers intercepted from N. Johnson (The Ohio State University, the Netherlands (S. Passoa, unpublished Columbus, Ohio) for use of the Lepidop- data). If not detected early and eradicat- tera collection during "ID day." This project was easier thanks to the adminis- ed, there is a danger D. Josealis will become the next Pyraustinae accidentally trative support provided by J. Cavey and established in North America. the equipment provided by J. Aronoff (U.S. Department of Agriculture. Animal ACKNOWLEDGMENTS and Plant Health Inspection Service, We thank B. G. Scholtens (College of Plant Protection and Quarantine. River- Charleston, Charleston, South Carolina), dale, Maryland). J. H. Wilterding (Olivet College. Olivet, LITERATURE CITED Michigan), G. Derkovitz (Frankfort, Illi- Allyson, S. 1981. Last instar larvae of Pyraustini nois), R. Panzer (Oak Forest, Illinois), K. of America north of Mexico (Lepidoptera: Gnaedinger (Highland Park, Illinois), J. ). Canadian Entomologist 113(6): Stiefel (Baileys Harbor, Wisconsin), C. 463-518. Rudy (Calumet County, Wisconsin) and 1984. Description of last-instar larvae of 22 species of North American (Lepi- L. A. Ferge (Middleton, Wisconsin) for doptera: Pyralidae: Pyraustinae) with a key to sharing their specimens and data with us. species. Canadian Entomologist 116(10): Matthias Nuss (Staatliche Naturhisto- 1301-1334. rische Sammlungen, Museum für Tier- Balachowsky, A. S. 1972. Entomologie appliquée a kunde, Dresden, Germany) provided use- lagriculture. Tome II. Lépidoptères. Masson ful comments on the manuscript, assisted et Cie. Paris, France. 575 pp. Beccaloni. G. W., M. J. Scoble, G. S. Robinson, with the adult identification, and helped and B. Pitkin. eds. 2005. The Global Lepidop- characterize wing pattern variation. Todd tera Names Index (Lepindex). World Wide Gilligan (University of Colorado, Love- Web electronic publication. http://www.nhm.ac. land, Colorado) deserves special praise uklentomologyflepifldex [accessed October 2007]. 514 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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