Volume 63 Number 1 29 Apr 2009 ISSN 0024-0966 Journal of the Lepidopterists' Society Published quarterly by The Lepidopterists' Society THE LEPIDOPTERISTS’ SOCIETY Executive Council John H. Acorn, President Lazaro Roque-Albelo, Vice President William E. Conner, Immediate Past President Michael E. Toliver, Secretary Jon Pelham, Vice President Kelly M. Richers, Treasurer Carmen Pozo, Vice President Members at large: Stephanie Shank Michelle DaCosta Charles Harp John H. Masters Todd Stout Michael G. Pogue Editorial, Board John W. B rovvn (Chair) Michael E. Toliver (Member, at large) Brian Scholtens (Journal) Lawrence F. Gall (Memoirs) Dale Clark (Neivs) John A. Snyder ( Website ) Honorary Life Members of the Society Lincoln P. Brower (1990), Frederick H. Rindge (1997), Ronald W. 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Journal of The Lepidopterists’ CSMIT HSOA^ ® 22 2017 ^gARiES^ Society Volume 63 2009 Number 1 Journal of the Lepidopterists’ Society 63(1), 2009, 1-10 A NEW SPECIES OF DRASTERIA HUBNER (NOCTUIDAE: CATOCALINAE: MELIPOTINI) FROM ARIZONA, WITH COMMENTS ON THE GENUS Jan Metlevski Department of Entomology, 123 West Waters Hall, Kansas State University, Manhattan, KS 66506-4004, U.S.A; email: jmetlevs@hsu.edu AND Gregory Zolnerowich Department of Entomology, Kansas State University, Manhattan, KS 66506-4004, U.S.A; email: gregz@ksu.edn ABSTRACT. Drasteria walshi n. sp. (Noctuidae: Catocalinae) from southeastern Arizona is described and illustrated. 'The new species be¬ longs to the D. fumosa-D. divergens species group. Drasteria walshi is distinguished from related species of Drasteria by wing color and pat¬ tern, and structure of male and female genitalia. Taxonomic problems in Drasteria are discussed. Additional key words: taxonomy, Drasteria, new species. Arizona, Nearctic The genus Drasteria Hubner, 1818, with more than 60 described species, is the largest genus in the tribe Melipotini (sensu Fibiger & Lafontaine 2005). It is exclusively Holarctic in distribution, with 27 species known from the Nearctic region and 35 species known from the Palearctic region. No species are known to occur in the tropics or Southern hemisphere. The only comprehensive study of the Palearctic species of the genus was published by John (1910) almost a hundred years ago. Since then, a dozen new species and subspecies of Drasteria have been described from the Palearctic region. The Palearctic species of Drasteria are in need of revision because the status of many taxa remains unresolved, and some widely distributed species may actually be complexes of closely related species, as is the case for Drasteria rada (Boisduval, 1848) (Metlevski et al ., unpublished data). Richards (1939) revised the Nearctic species of Drasteria, and few changes have occurred in the taxonomy of Nearctic species since then. Some taxonomic problems pointed out by Richards, such as the relationship between D. tejonica (Behr, 1870) andD. howlandii (Grote, 1864), or the status of D. nichollae (Ilampson, 1926), have not been resolved yet. Richards' 1939 revision downgraded to subspecies (race by Richards) a number of taxa originally described as separate species by other workers. Poole’s 1989 catalog of the world Noctuidae (Poole 1989) did not recognize any taxa at the subspecies level and synonymized all the names applicable to the taxa treated as subspecies by Richards. As a result, all Drasteria names synonymized by Poole have been excluded from the lists of Nearctic fauna, such as Nomina Insecta Nearctica (Poole & Gentili 1996). Nevertheless, the status of many taxa treated by Richards as subspecies of other Drasteria species remains unclear. Drasteria hastingsii (H. Edwards, 1878) is now recognized as a valid species, and Wagner et al. (in press) provide evidence that Drasteria graphica atlantica Barnes & McDunnough, 1918 represents a full species. It is likely that others, such as D. sabulosa abrupta (Barnes & McDunnough, 1918) or D. adumbrata alleni (Grote, 1877), are also valid species. In addition, our current investigations suggest that some North American species of Drasteria are actually complexes of closely related species. For example, populations of D. inepta (Edwards, 1881) from Colorado and northern New Mexico are not nonspecific with populations from southeastern Arizona and southwestern New Mexico. Finally, discoveries of new 2 Journal of the Lepidopterists’ Society Fig. 1. Medial surface of Drasteria male valve (based on a left valve of D. pallescens (Grote & Robinson, 1866)) (c - costa; co - conus; cp -costal process; dlcp - dorsal lobe of costal process; hrp - harpe; pulv - pulvinus; sel - sacculus; scle - saccular extension; vlvl - valvula). Fig. 2. Female genitalia of Drasteria (based on D. sinuosa (Staudinger, 1884)) (aa - apophyses anteriores; an - antrum; ap - apophyses pos- teriores; be - bursa copulatrix; col - colliculum; db - ductus bursae; la - lamella antevaginalis; Ip - lamella postvagin'alis; pa - papilla analis; pb - pseudobursa). Volume 63, Number 1 3 species of Drasterici in North America are still occurring. Drasteria convergens Mustelin, 2006, was described recently from California, and below we describe a new species from southeastern Arizona. The terminology used here for structures of the male valva follows the scheme shown in Fig. 1. We use the term “harpe” in a broad sense since the problem of homologies of valval structures, to which this term is applicable in various groups of Noctuidae, remains unresolved. For this reason we follow with some modifications the broad definition of the term “harpe” given by Kuznetzov & Stekolnikov (2001), and define the harpe as various and not necessarily homologous sclerotized structures on the inner surface of valva, that are not derived from the costa or sacculus. Here we use the term “harpe” for the long and slender process (processus basalis valvae in John 1910) arising from a sclerotised, angled ridge on the inner surface of proximal part of the valva. We also use the term “conus” for the small process located just below the haq:>e, and the term “pulvinus” for the shelflike process arising laterad from the inner side of the basal part of the saccular extension. The last two terms were introduced by John (1910) for Drasteria but have not been used since. Terminology for male genitalia other than valva follows Goater et al. (2003). Species of Drasteria have aedeagi with a veiy complex vesica, consisting of a number of diverticula of different sizes. The complexity of the vesica of species of Drasteria was shown only recently (Goater et al. 2003; Mustelin 2006); previous taxonomic works on the genus have not made mention of the vesica. Our current investigations show the number and relative position of diverticula are good characters for grouping members of the genus into species groups. Thus, we include a detailed description of the vesica in the species description. Terminology for female genitalia follows Kiihne (2005) and is shown in Fig. 2. Terminology for wing patterns is shown in Fig. 3. Fig. 3. Wing pattern of Drasteria. dorsal surface (based on D. christophi (Alpheraky, 1895)) (am - antemedial line; as - apical spot; ash - api¬ cal shadow; b - basal line; ba - basal area: cl - discal spot; in - medial shadow; ma - medial area; pin - postmedial line; pina - postmedial area; pmb - postmedial band; r - reniform spot; st - subterminal line; sta - subterminal area; t - terminal line; ts - terminal spot; tsh - terminal shadow). 4 Journal of the Lepidopterists’ Society AZ Graham Co. 9,000 ft. Pinaleno Mis, Cunimingham Campdgr, UV + MV lights 20 May 2002. B. Walsh leg. wa&ly WeTLftrsjci PARATYPE Onion Saddle Elv. 7600, ft. Chirachua Mts AZ June 14. 2007 Leg. Ray B. Nagle KSU-MEPAR Collection <? Gen. Prep. No J. Metlevski ,W4 . 2006 Gen. Prep. No. 1068 J. Metlevski 2007 Figs. 4-9. Drasteria toalshi. 4-6, holotype. 4, dorsal surface. 5, ventral surface. 6, pin labels. 7-9, female paratvpe, Chiricahua Mts. 7, dorsal surface. 8, ventral surface. 9, pin labels. Volume 63, Number 1 5 Figs. 10-13. D. divergens. 10-11, male, White Pine Co., Nevada. 10. dorsal surface. 11, ventral surface. 12-13, f. socia, female, Santa Bar¬ bara; California. 12, dorsal surface. 13, ventral surface. Abbreviations for institutions and collections used in the text are as follows: KSU-MEPAR = Kansas State University Museum of Entomological Prairie Arthropod Research; BW = Bruce Walsh collection (Tucson, AZ); RN = Ray B. Nagle (Tucson, AZ) collection. Drasteria walshi Metlevski, new species (Figs. 4-9, 18-20) Diagnosis. Drasteria walshi is most similar to D. divergens (Figs. 10-13, 21, 23), especially to the form with orange hind wings, D. divergens f. socia (Behr, 1870) (Figs. 12, 13). The following features can be used to differentiate D. walshi from D. divergens: in D. walshi the underside of the wings is pinkish, while they are yellow to pale yellow in D. divergens ; in D. walshi the terminal spots on the underside of the wings are obscured or completely absent, while in D. divergens there are strong black terminal spots on the underside of the wings; in D. walshi the discal spot is a thin dark comma-like line not connected to the postmedial band, compared to a strong black discal spot connected to the postmedial band (at least vein CuA2 is black from the postmedial band to the discal spot, but usually the connection is much stronger) in D. divergens. Among other species of Drasteria occurring in southeastern Arizona only one, D. tejonica (Behr, 1870), has forewings with similar pattern and colored hind wings. Drasteria tejonica however is noticeably smaller in size and has much stronger black pattern on the hind wings (Figs. 14-17). Males of D. tejonica (Figs. 14-15) 6 Journal of the Lepidopterists’ Society Figs. 14-17. Drasteria tejonica, Pima Co., Arizona. 14-15, male. 14, dorsal surface. 15, ventral surface. 16-17, female. 16, dorsal surface. 17, ventral surface. can be easily separated from D. walshi by the coloration of the hind wings, which are white basally in D. tejonica. In males, the costal processes of the valvae of D. walshi are robust, elongate, sinuous and pointed at the tip (Fig. 18). This readily distinguishes D. walshi from 1). divergens (Fig. 23), as well as from any other species in the genus. Female genitalia of the new species (Fig. 20) could be confused only with those of D. divergens (Fig. 21) and D. fumosa (Fig. 22). The antevaginal plate in D. walshi is more than 3 times longer than the colliculum, while just slightly longer in D. divergens and about 2 times longer in D. fumosa. The antevaginal plate of D. walshi is semi-rectangular in shape, compared to triangular with a blunt tip in D. fumosa. The shape of the antevaginal plate of D. walshi is similar to that of D. divergens : both species have a rectangular anterior portion and a slightly narrower posterior portion. However, in D. walshi the broader anterior portion is about half the length of the narrower posterior portion, while in D. divergens the length of both parts is subequal. Description. Head: Light brownish gray; labial palpus light grey, with some dark brownish gray scales on the external side of first and second segments; antenna filiform in both sexes. Thorax: Covered with long flattened and slightly spatulate hairs dorsally, with long simple hairs ventrally; patagia light brownish gray, with a dark brown longitudinal stripe; remainder of thorax little darker and with light pinkish tint; tegulae slightly rimmed with dark brown hairs on external margin; legs same color as thorax except for tarsi, which are darker brownish gray with a ring of light gray scales at distal end of each segment. Forewing: Length 19-23 mm; males and females similar; ground color light brownish grav with light pinkish tint; basal line double, black, enclosed area light brownish gray; basal area brownish gray in anterior half, dark gray in posterior; antemedial line double, internal Volume 63, Number 1 and external lines black, enclosed area filled with dark brown, internal line making a strong long tooth toward base of wing between veins A and Cu and two small teeth at veins A and Sc, entire antemedial line almost straight from costa to vein A, abruptly curving inward after vein A; medial area very light brownish gray, somewhat darker in outer part and at costa; medial fine brown, not strongly contrasting, double, the internal line thin, external one wider; both lines strongly expanded and becoming dark brown or almost black at costa; reniform spot dark brown, black rimmed, with strong whitish outline on outer margin and thin whitish outline followed by thin black line on inner margin; postmedial line double, area enclosed between internal and external lines dark brown except at costa where pinkish, internal line strong and black through its entire length, external one weaker, marked with dark brown or black scales, almost disappearing in the middle; postmedial line turning abruptly toward the apex at vein Rj and almost parallel to costal margin until vein R„, then bending again and curving outward, includes strong tooth pointed outward at vein and another smaller tooth at vein M3, curving inward in between; making an abrupt turn at vein CuAp continuing along this vein toward base of reniform spot then turning toward posterior margin of wing, curving slightly outward between base of reniform and vein A, turning toward anal angle of wing at vein A; area between reniform spot and postmedial line brownish grey to brown, filled with brown and light brownish grey scales, veins M3 and CuA, within this area dark brown with strong whitish outlines joined with similar outline on inner margin of reniform spot; postmedial area dark gray, pinkish along postmedial line at costa; subterminal line slightly incurved between costa and vein Rp then slightly waving and subparallel to inner margin of wing, at vein A, turning toward anal angle of wing, dark brown at veins, otherwise black in anterior half of wing, mostly brown, with some black scales between veins in posterior hatf; 4 strong black arrow marks arising from subterminal line between veins in anterior half of wing, and pointing toward base of wing; subterminal line with a thin pale brownish gray outline along outer side, immediately followed by dark brown line; subterminal area light brownish gray, somewhat darker in place where terminal shadow is located; apical shadow obvious, as a dark brown path, connected to dark brown outer outline of subterminal line; terminal line fine, wavy, obvious, dark brown to almost black; fringe pinkish at base, otherwise grayish brown, checkered. Underside pale pinkish to pinkish; reniform spot dark brown to black; dark brown transversal shadow extending from the middle of costa through inner edge of reniform toward the anal angle, ending well before anal angle between veins CuA., and A; subterminal line greasy pinkish with dark edges at costa, otherwise dark brown to black, convex, wide at costa, rapidly narrowing toward posterior edge of wing becoming a thin line at its posterior end, meeting transversal shadow at vein CuA, and not extending after this point, outer edge of subterminal line wavy, inner edge evenly convex; terminal and apical shadows brownish, obscured and sometimes barely visible; terminal line thin, dark brown, wavy, vanishing at anal angle; fringe pinkish at base, otherwise grayish brown, lighter toward anal angle. Hind wing: Ground color orange pink, a few dark brown scales at anal margin and occasionally a few' brownish scales near base of ufing and on Cu stem; discal spot a narrow dark grayish brown line along cross-vein, not connected to postmedial band; postmedial band dark grayish brown to black, narrow through most of its length but strongly widened between vein R, and costa, subparallel to outer margin between anal angle and vein CuA„ approaching vein CuA, curving abruptly mesad, and at vein CuA, angling toward apex, then turning toward costa between veins M, and Mp outer margin of postmedial band turning sharply outward at vein ftp terminal and apical spots narrow patches, dark grayish brown to black, with obscure inner edge, almost disappearing in some specimens; terminal line obvious through most of its length, vanishing near anal angle, dark grayish brown to black, thin, wavy; basal portion of fringe pink, outer portion pink orange, greasy orange brown at terminal spot, with some blackish scales between basal and outer layers at terminal and apical spots. Underside pale pinkish to pinkish, some brown or brownish scales along costal margin; discal spot similar to that on upper side but weaker; postmedial band intermittent, marked by few' brown scales at costal margin, remaining remnants present only between veins M, and M„ and between vein CuA, and anal angle; terminal spot a dark brown to blackish shadow, weak, in some specimens almost disappearing; apical spot absent; terminal line weaker than on upper side, otherwise similar; fringe same as upper side. Abdomen: Ochreous gray, with some scattered dark brown scales. Male genital capsule (Fig. 18): Uncus strong, curved, pointed at tip, dorsally with a low' longitudinal crest and dense hairlike setae in third distal quarter; scaphium long, narrow, articulated at base of uncus; valvae obovate, slightly asymmetrical, left valve slightly smaller, narrower in the middle and with more sinuous dorsal margin; valvula rounded, acute at ventro-distal angle, with a small tuft of setae on inner surface at ventro-distal end; area occupied by coremata small, subbasaly on outer side of valva, extending along sclerotized base of valva from ventral edge of base of costa to ventral side of distal half of sacculus; sacculus about 4 times as long as wide, its dorsal edge slightly concave; saccular extension strong, slightly curved, ending just before tip of valvula in a small free prong-like projection turned inward toward genital capsule; pulvinus strong, broad, broader on left valva, tip of pulvinus with robust elongate setae; harpe curved, slender, smoothly narrowing to tip, apex flattened and rounded; conus present on both valvae, left one tapered, about 2 times as long as wide, right one minute, short; tip of conus bearing few microscopic setae; costa moderately sclerotized, with costal process present on both valvae; costal process prominent, broad based, strongly elongated, sinuous, tapered, pointed at tip; costal process on left valva shorter, directed along longitudinal axis of valva; costal process on right valva longer, directed slightly dorsad, reaching dorsal margin of valvula; saccus short, V-shaped: juxta symmetrical, about 2 times as long as wide, with deep V-shaped incision on dorsal margin and two short, curved, and divergent ventroanterior arms. Aedeagus (Fig. 19): strong, straight, tubular, membranous in distal half dorsally but with a small sclerotized area at base of vesica, otherwise well sclerotized, its membranous portion finely spiculose; coecum short, slightly flattened dorsoventrally at tip; ventral plate of carina strongly sclerotized, long, narrow, slightly expanded at top, basal half of right margin w'ith a strong longitudinal fold; dorsal plate of carina absent; inflated vesica short, turned dorsad, with a complex system of diverticula (insertion on Fig. 19): diverticula arising on ventral side right of carinal plate (2 large and 4-5 very small): 1. diverticulus next to the tip of carinal plate, broad, about twice as long as wade, distally rounded, spiculose on right side, directed caudad, tip slightly turned left; 2„group of 4-5 aligned small spiculose diverticula, next right to diverticulus 1, followed by diverticulus 3; 3. the biggest diverticulus, two times longer than next longest diverticulus, slightly narrowing to the tip, finely spiculose, directed right laterodorsad, perpendicular to longitudinal axis of aedeagus; diverticula arising on ventral side left of carinal plate (1 bigger and 1 veiy small): 4. diverticulus next to tip of carinal plate, short, bifurcated at tip, spiculose, directed left laterad. perpendicular to longitudinal axis of aedeagus, one of tips turned cephalad; 5. very small, lobelike, with no spicula, between diverticulus 4 and tip of aedeagus; diverticula arising at left dorsolateral angle of vesica (2 bigger and 1 veiy small): 6. globular, expanded distally, as long as wide, finely spiculose, directed dorsocaudad, with a narrow fingerlike projection arising left ventrolaterally and directed ventrad; 7. second longest, tapering, finely spiculose, curved, directed cephalad, tip curving right dorsocephalad; 8. very small, short, narrowing at tip, with no spicula, between diverticulus 7 and base of vesica; top of vesica exposed dorsad, bearing 1 small diverticulus and 1 small lobe: 9. minute diverticulus, with no spicula, arising on right side close to diverticulus 2: 10. small lobe with no spicula, close to the base of diverticulus 6; posterior side of vesica : 11. bearing a small lobe with no spicula between diverticulus 1 and top of vesica. Female genitalia (Fig. 20): Ovipositor telescopic; papillae analis with moderately sclerotized longitudinal band dorsally, weakly sclerotized in other parts, narrow, about 5 times as long as wide, strongly narrowing in distal third, tips rounded, membranous; surface of papillae with short scattered setae, which are denser on tips, some long setae at the base and tips of papillae analis; apophyses long and slender, posterior ones about 1.7 times longer than anterior; lamella postvaginalis (postvaginal plate) a V-shaped sclerotized band, its ends not connected to eighth segments sclerotization; antrum wide, with almost parallel lateral edges, about 2 times as long as wide, dorsoventrally flattened, with large sclerotized antevaginal plate ventrally, otherwise membranous; antevaginal plate occupying almost whole ventral side of antrum, semi-rectangular in shape, slightly narrowing distally, with slightly convex posterior edge; ductus bursae short, slightly longer than wide, about 2 times narrower and 2.5-3 times shorter than the antrum; colliculum as long as wide, its left edge slightly longer than the right one, edges of colliculum not overlapping; bursa copulatrix asymmetrically ovate, membranous, with some Fig. 18-22. 18. Male genital capsule of D. walshi (posterior), holotype. 19. Aedeagus of D. walshi with vesica everted (ventral view), holo- type. Inset: vpc - ventral plate of Carina; 1-7 - numbers of diverticula according to the description. 20, Female genitalia oi D. walshi (ventral view), paratype, genital prep, by Jan Metlevski No. 945. 21, Female genitalia of D. divergens (ventral view), Tuolumne Co., California, genital prep. b\ Jan Metlevski No. 774. 22, Female genitalia of D. fumosa(ventr;i\ view), Pima Co., Arizona, genital prep, by Jan Metlevski No. 963. Volume 63, Number 1 9 Fig. 23. Male genital capsule of D. divergent (posterior), genital prep, by Jan Metlevski No. 794. longitudinal wrinkles on left side anterad of insertion of ductus bursae, otherwise smooth; insertion of ductus bursae posteriorly, slightly left; internal wall of bursa copulatrix in posterior half of bursa with very minute spines, most visible at wrinkled area, becoming smaller and barely visible laterad, and disappearing anteriorad; pseudobursa arising posteriorly, right of insertion of ductus bursae, coiling dorsad and to right, making 0.5 coil; ductus seminalis arising from the tip of pseudobursa. Holotype. d, “AZ, Graham Co. Pinaleno Mts. Cunningham Campgrd. 9000 ft. UY + MV lights. 20 May 2002. B. Walsh leg.”, “Genitalia Prep, by J. Metlevski No 766”, deposited in KSU-MEPAR (Figs. 4-6, 18, 19). Paratypes. (3 d and 3 9): 1 d, same label data as holotvpe, in BW; 1 9, “AZ, Graham Co. Pinaleno Mts. Cunningham Campgrd. 9000 ft. Ponderosa pine habitat. 14 June 2005. UV + MV lights. B. Walsh leg.”, “Genitalia Prep, by J. Metlevski No 945”, in BW (Fig. 20); 1 d, “AZ, Graham Co. Pinaleno Mts. Cunningham Campgrd. 9000 ft. Ponderosa pine habitat. 5 June 2007. B. Walsh leg.", in BW; 1 9, “AZ, Cochise Co., Chiricahua Mts. Onion Saddle, 7700 ft, Ponderosa Pine/Oak habitat, 14 June 2007. B. Walsh leg.”, “Genitalia Prep, bv | Metlevski No 1084”, in BW; 1 9, “AZ, Cochise Co., Chiricahua Mts. Onion Saddle, 7600 ft, June 14 2007. Leg. Ray B. Nagle”, “Genitalia Prep, by J. Metlevski No 1068”, in RN (Fig. 7-9); 1 d, same label data as above, “Genitalia Prep, by |. Metlevski No 1069”, in RN. Variation. Specimens from the Chiricahua Mountains have the hind wings a slightly brighter ground color and with a stronger dark pattern compared to specimens from the Pinaleno Mts. Biology. Unknown. Specimens have been collected in the second half of May and first half of June. Distribution. Pinaleno Mountains (Mt. Graham) and Chiricahua Mountains in southeastern Arizona. Etymology. Drasteria walshi is named in honor ol Bruce Walsh, Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, in recognition of his invaluable efforts in studying the lejridopteran fauna of southeastern Arizona. Discussion. Female genitalia of Drasteria walshi have a greatly enlarged antevaginal plate, which occupies most of the ventral side of the antrum and is longer than the colliculum. This condition of the antevaginal plate links D. walshi with D. fumosa (Strecker, 1898) and D. divergens (Behr, 1870), which are the only other Nearctic species in the genus sharing this character state. In the male genitalia, the aedeagus also provides characters linking D. walshi with the two species mentioned above in that the general structure of the vesica, such as the relative position and number of 10 Journal of the Leridopterists’ Society diverticula, of D. walshi is most similar to that of D. divergens and D. fumosa. All I mown specimens of Drasteria walshi have been collected in the Pinaleno and Chiricahua Mountains at altitudes between 7500-9000 feet (approximately 2300-2750 m) in ponderosa pine or mixed ponderosa pine/oak forest habitats. Both the Pinaleno Mts. and Chiricahua Mts. are isolated high mountain massifs surrounded by lowland plains grasslands and deserts. AC KN OWLE DG E ME NTS We thank P. A. Opler and B. C. Kondratieff (Colorado State University), J. H. Boone (Field Museum, Chicago), E. H. Met- zler (Alamogordo, New Mexico), and R. B. Nagle (Tucson, Ari¬ zona) for loans of specimens used in this study. We are grateful to T. Mustelin (San Diego, California), D. L. Wagner (University of Connecticut) and two anonymous reviewers for reviewing the manuscript. This article is Contribution No. 0S-395-J from the Kansas Agricultural Experiment Station (KAES) and was sup¬ ported in part by KAES Hatch Project No. 583 (Insect System¬ atic^ ). Literature Cited Fibiger, M. & [. D. Lafontaine 2005. A review of the higher classifi¬ cation of the Noctuoidea (Lepidoptera) with special reference to the Holarctic fauna. Esperiana 11: 7-92. Goater, B., L. Ronkay & M. Fibiger. 2003. Catocalinae and Plusi- inae. Noctuidae Europaeae, Volume 10, Sor0, 452 pp. John, O. 1910. Eine revision der gattung Leucanitis Gn. Horae Soci- etatis Entomologicae Rossicae 39: 585-640. Kuhne, L. 2005. Revision und Phylogenie der Gattungsgruppe Crvp- sotidia Rothschild, 1901. Tachosa Walker, 1869, Hypotacha Hampson, 1913, Audea Walker, [1858] 1857 und Ulotrichopus Wallengren, 1860 (Lepidoptera, Noctuidae, Catocalinae). Esperi¬ ana Memoir 2: 1-220. Kuznetzov, V. I.. & A. A. Stekolnikov. 2001. New approaches to the system of Lepidoptera of world fauna (On the base of the func¬ tional moi-phologv of abdomen). St. Petersburg, Nauka. 462 pp. Lafontaine, J. D. & M. Fibiger. 2006. Revised higher classification of the Noctuoidea (Lepidoptera). Canadian Entomologist, 138 (5): 610-635. Mustelin, T. 2006. Taxonomy of southern California Erebidae and Noctuidae (Lepidoptera) with descriptions of twenty-one new species. Zootaxa 1278: 1—47. Poole, R. W. 1989. Noctuidae 1-3. In Heppner. |. B. (ed.), Lepi- dopterorum Catologus (New Series) fasc. 118. Leiden, New York, Kopenhavn, Koln. 1314 pp. Poole, R. W. & P. Gentili (eds.). 1996. Nomina Insecta Nearctica. Volume 3. Diptera, Lepidoptera, Siphonaptera. Entomological Information Sendees, Rockville, Maryland, 1143 pp. Richards, A. G. 1939. A revision of the North American species of the Phoberia-Melipotis-Drasferia group of moths (Lepidoptera, Phalaenidae). Entomologica Americana 19: 1-100 Received for publication 1 July 2008; revised and accepted 9 December 2008 Volume 63, Number 1 11 Journal of the Lepidopterists’ Society 63(1), 2009, 11-20 BIOLOGY AND DISTRIBUTION OF A RECENTLY REDISCOVERED ENDEMIC HAWAIIAN LEAFROLLER MOTH, OMIODES CONTINUATALIS (CRAMBIDAE) Cynthia King University of Hawaii, Dept, of Plant and Environmental Protection Sciences, 3050 Made Way, Gilmore Hall 310, I lonolulu, III 96822 e-mail: cbaking@hawaii.edu Daniel Rubinoff University of Hawaii, Dept, of Plant and Environmental Protection Sciences, 3050 Maile Way, Gilmore I Pall 310, Honolulu. HI 96822 e-mail: rubinoff@hawaii.edu and William Haines University of Hawaii, Dept, of Plant and Environmental Protection Sciences, 3050 Maile Way, Gilmore Hall 310, Honolulu, III 96822 e-mail: whaines@hawaii.edu ABSTRACT. The endemic Hawaiian leafroller moth Omiodes continuatalis was documented by early entomologists to be one of the most common species encountered during initial collecting trips in the Hawaiian Islands at the end ol the 19th century. The species was declared ex¬ tinct in 1982, and while subsequent surveys have resulted in their “rediscovery”, it is evident that O. continuatalis populations have experienced declines and localized extinctions throughout their historical range. The objective of our research was to document the life history, biology, gen¬ eral morphology, and behavior of this little studied species. To complete the investigation, we observed both wild caught and laboratory reared O. continuatalis. We also conducted light trap surveys to document the current distribution of O. continuatalis across the Hawaiian Islands. We have integrated our findings from this study with all available historical data on O. continuatalis, to create a document which can be referenced for future research relating to the biology or conservation of this species. Additional Key words: pyraloid, light trap, fossorial, development, extinction The genus Omiodes Guenee, 1854 (Lepidoptera: Crambidae) occurs in tropical and subtropical regions of Oceania, Southeast Asia and the Americas, and contains over 100 species, including 23 species endemic to the Hawaiian Islands. The subject of this research, Omiodes continuatalis (Wallengren, 1860), is a generalist grass- feeding species, originally documented from all of the high Hawaiian Islands, with the exception of Kahoolawe (Zimmerman 1958). The Reverend Thomas Blackburn found O. continuatalis to be one of the most common moths encountered during collecting trips in Hawaii prior to 1880 (Zimmerman 1958). Swezey reported the species from “drier, lower” habitats, but does not provide specific collection localities beyond a single reference to a population in Olinda, Maui (Swezey 1907; Zimmerman 1958). Despite their early documented widespread abundance, 100 years later the species was declared extinct by the U.S. Fish and Wildlife service, and other agencies (Gagne & Howarth 1982; IUCN 2007). Omiodes continuatalis and 13 other Omiodes species were listed as “extinct or possibly extinct” based on reviews of museum collections which indicated that specimens had not been collected for several decades. Non-target impacts of the biological control agents introduced for the coconut and sugar cane leafroller pest species were blamed for the extinctions (Gagne & Howarth 1982). However, data from surveys of more recent museum collections completed in 2003 revealed extant O. continuatalis populations (Haines et al. 2004). Specimens were identified from light trap samples collected at five mesie forest studv sites on both the leeward and windward J sides of Hawaii Island (Haines et al. 2004). In addition, field collections completed in 2003 confirmed O. continuatalis to be abundant at several localities on East Maui (Haines unpublished data). In the past, O. continuatalis larvae have been collected from native pili grass Heteropogon contortus (L.), as well as introduced species such as kikuyu grass Pennisetum clandestinum (Hochst. ex Chiov.) and Hilo grass Paspalum conjugation Berg. (Zimmerman 1958). Like other species in the group, O. continuatalis larvae are leaf-rollers, and use silk to bind plant foliage together to create a refuge in which they feed and take shelter. Early observations by Swezey describe how O. continuatalis larvae typically feed near the base of their 12 Journal of the Lepidopterists’ Society host plants, within the grass matrix or in leaf litter at the soil surface (Zimmerman 1958). This behavior is uncharacteristic of other Hawaiian Omiodes larvae, which generally feed higher on host plant foliage, fastening or rolling the tips of leaves together (Swezey 1907). Omiodes continuatalis larvae were also noted to be physically distinct from larvae of other Omiodes species, in that they attain a greater body width and often exhibit pinkish-green coloration in contrast to the monochromatic green coloration displayed by larvae of other Omiodes species (Zimmerman 1958). Unlike the sugarcane leafroller O. accepta (Butler, 1877), O. continuatalis is not considered a pest and consequently few data have been collected to document its life history. Omiodes continuatalis appear to share similar habitat requirements, feeding habits and physical characteristics with O. accepta (Zimmerman 1958), and initially seemed likely to have comparable biology and development. The objective of this chapter is to describe the complete life history of O. continuatalis , providing new information on its biology, ecology, general moqdiology and life history, and to integrate recent findings with information gathered by early naturalists, to create a document that can serve as a reference for future research on the biology and conservation of O. continuatalis. Materials and Methods We collected data for this laboratory and field study from February 2005 to March 2008. Insect rearing. We established laboratory colonies of O. continuatalis using wild female moths captured from UV light traps set on Kailiili Road in Kokomo, Maui (484m) and at the University of Hawaii Kula Agricultural Research Station (975m), Mani. Moths were collected alive in plastic specimen vials, and held at ambient temperature until being transferred into rearing cages. Moths were easily identified using the key to Omiodes species in Zimmerman (1958), as O. continuatalis is a veiy distinctive species. Rearing cages were composed of hand-made 61cmx46cmx31cm PVC- framed sleeve cages covered with fine insect mesh sleeves. Moths were fed a 30% honey/70% water solution, and were provided potted sugarcane, pili grass and/or Hilo grass, on which to oviposit. We obtained young sugarcane starts from Hawaii Commercial and Sugar (IICS) and Hawaii Agricultural Research Center (I4ARC) fields, while pili grass and Hilo grass were propagated from seed. These plants provided food and served as host plants for colony larvae. The substrate in each host plant pot consisted of a 50/50 blend of Sunshine Mix 5® potting soil and vermiculite. All moth colonies and host plants were housed in the University of Hawaii at Manoa, Gilmore Hall greenhouses, and University of Hawaii, Kula Agriculture Research Station greenhouses. In Manoa, temperatures ranged seasonally from a mean low of 18.7°C in the winter to a mean high of 24.3°C during the summer (Manoa Lyon Arboretum weather station), and plants were watered three times each day. In Kula, temperatures ranged from a mean low of 11.1°C in the winter 2006, to a mean high of 24.4°C during the summer 2006 (UI4 Kula weather station), and plants were watered twice daily. We collected additional wild O. continuatalis moths and integrated them into laboratory colonies periodically to replenish adult stock and maintain genetic diversity. Biology and morphology. To elicit oviposition, we isolated female moths in vials lined with wax-paper and temporarily denied them access to a food source. These stressful conditions commonly stimulate female moths to oviposit. Eggs deposited in each egg mass were then counted under a Leica dissecting microscope to determine the total number present, and the minimum and maximum number of eggs per mass. In order to quantify mean egg size, we selected 50 eggs randomly from 50 O. continuatalis egg masses, and measured them at the widest point using an ocular micrometer in a Wild Heerbrugg microscope. We obtained data on O. continuatalis larval development and moiphology by observing 100 individual larvae develop from egg to pupal stage. Larvae were stored in individual 2-oz Solo® cups (ventilated with 3-6 holes in the lid). Cups were cleaned anti larvae provided with fresh sugarcane foliage eveiy 2-3 days. We measured the head capsule width of each larva daily to document larval growth; head capsule widths were measured at their widest point, as above for eggs. We also recorded changes in larval moiphology following each larval molt. Only head capsule widths of larvae which survived from egg to pupal stage (n = 52) were used to assess instar duration and size ranges. We completed a one-way analysis of variance to document variation in head capsule width by instar. Behavior. Data relating to O. continuatalis larva! behavior were collected based on observations of individual larvae contained in laboratory colonies, or from colony larvae deployed on potted host plants at selected Maui field sites. We collected data opportunistically, observing behaviors in the field during larval deployment and retrieval, and in the laboratory during daily colony maintenance and larval feedings. Adult O. continuatalis were isolated and observed individually to assess adult lifespan under laboratory colony conditions. Distribution. Field sites were selected to be surveyed based on accessibility and the presence of O. Volume 63, Number 1 13 continuatalis host plants. We collected Omiodes continuatalis adults using two types of UV light traps: sheet light traps and bucket light traps. Sheet light traps consisted of an 18” UV light bulb placed in front of a large, white bed sheet suspended vertically. Bucket light traps were constructed with an 18” UV light bulb attached to three 10.2cmx25.4cm clear plastic flight intercept vanes, and secured on top of an 8 liter bucket with bungee cords. Each bucket trap was baited with a killing agent such as cyanide or a Hot Shot No-Pest Strip®. Individual moths landing on sheet light traps were collected alive, however insects collected from bucket traps were always dead. We set at least one of the two trap types at each field site to survey for (). continuatalis adults. When an initial survey yielded numerous O. continuatalis specimens, we revisited sites to replenish and diversify laboratory colonies. Visual surveys of and sweep netting for adult O. continuatalis were also completed at field sites to complement data collected from light trapping. To quantify total survey effort, the number of nights and the number of traps that were used, were multiplied to obtain the total number of “trap nights” completed at each survey location. Results and Discussion Biology. Omiodes continuatalis are highly attracted to UV light, and both male and female moths were regularly collected from light traps. In addition, O. continuatalis appeared to be more attracted to UV light traps than other Omiodes species. Once drawn to a light, O. continuatalis adults frequently remained around the light for long periods of time, in contrast to the sedge-feeding species O. anastrepta Meyriek, 1899, which are not as readily attracted to light traps, even when adults are observed flying in close proximity to a trap. Over the course of one year, adult O. continuatalis were collected, on multiple occasions, from light traps at Kokomo, Makawao, and Kula, on Maui (Table 1). Data from these surveys indicate that O. continuatalis is multivoltine, with more than three generations per year. Considerable climatic seasonal variation exists at mid- high elevation localities, so if O. continuatalis populations are mutivoltine at these three mid-high elevation field sites, it is likely that populations are also multivoltine and not seasonal at low elevation sites, where seasonal changes in temperature and rainfall may be even less restrictive to O. continuatalis development. Based on mitochondrial genetic data, O. continuatalis populations do not exhibit much structure, either within or among islands (Haines, unpublished data), suggesting that moths are strong dispersers. Because of this, we do not suspect that populations on islands other than Maui exhibit very different morphology or life history characteristics from those observed in our colonies. Laboratory colonies of O. continuatalis were maintained continuously from July 2005-December 2005, and from March 2006-March 2008. In laboratory colonies, the moths were most successfully reared on sugarcane host plants, as compared with pili grass and hilo grass, which the moths were previously documented feeding on in the field. At one point during rearing of the colony, some larvae escaped from cages and were discovered feeding on Canavalia pubescens , Hook. & Arn. (Fabaceae) an endemic Hawaiian legume which was being grown in the same greenhouse as food for another caterpillar species (O. monogona Meyriek, 1888). The O. continuatalis larvae were caged with C. pubescens, and surprisingly, multiple generations were reared on the plant. There are no previous records of this grass-feeding species surviving on a dicot host, yet it survived through four generations on C. pubescens with no obvious moi-phologieal or reproductive disadvantages. The success of the moth on this host plant over multiple generations indicates that its host range in the wild might be broader than previously expected, although opposition preference may be more important than nutritional suitability as a determinant of host range. While the utilization of this host plant was unexpected given the documented life history of the species, it may be explained by recent molecular analysis of the genus. Phylogenetic analysis based on mitochondrial DNA shows that O. continuatalis is the sister species, and the closest relative, of O. monogona, the sole Hawaiian Omiodes species which feeds on fabaceous host plants (Haines unpublished data). Other tropical Omiodes species are documented to feed on Fabaceae, thus this dietaiy shift may reflect a reversion to an ancestral host plant retained in the evolutionary ‘memory’ of some Hawaiian Omiodes. Evolutionary memory is the idea that larvae may retain an ability to feed on ancestral hostplants on which they are no longer found in nature (Janz & Nylin 1998). This discovery has important implications not only for the conservation of the species, but also for the evolution of host plant shifts in the genus. Adult O. continuatalis (Fig. 1) lived a maximum of 18 days under laboratory conditions. In cages, O. continuatalis females were observed ovipositing on both the upper and lower surfaces of host plant vegetation, and females did not avoid ovipositing on material such as wood, plastic, wax paper and screen mesh. Egg masses ranged in size from a single egg, to a maximum of 73 eggs, with a mean of 7.25 (± 0.81 S.E.) per individual egg mass (n = 115). A limited survey of O. continuatalis females (n = 33) indicated that gravid 14 Journal of the Lepidopterists’ Society Fig. 1. Adult Omiodes continuatalis moth. females can lay more than 400 eggs over their lifetime, and up to 122 eggs in a 24 hour period. Egg masses are composed of even rows or clusters of Battened eggs, slightly overlapping one another (Fig. 2). Eggs have a mean width of 1.22mm (± 0.025) (n = 50), are light yellow in color, and exhibit fine surface reticulations and an iridescent sheen. Omiodes continuatalis eggs took four days to hatch at 32°c, and eight days to hatch 22°c. Morphology. Swezey broadly described the life history and physiology of O. continuatalis (summarized by Zimmerman 1958). Zimmerman (1978) later provided limited descriptions and illustrations of larval moiphology, including a larval key to most Hawaiian Omiodes spp. However, no detailed research has been reported that fully characterizes the life history and larval stages of O. continuatalis. Based on head capsule width measurements from 52 individuals, we observed O. continuatalis larvae transitioning through seven to nine developmental instars (Fig. 2). In a previous study, O. continuatalis larvae were observed transitioning through as many as 10 instars prior to pupation (King & Rubinoff 2008). Analyses of variance of head capsule widths by instar indicates that mean head capsule widths for instars 1 through 9 are significantly different from one another (Figure 2; df = 8; F = 1823.68; P = <0.001). Due to considerable overlap in head capsule widths in each instar (Fig. 3), it is difficult to distinguish discrete larval instars at any stage of development based on measurements alone. Upon eclosion, larval head capsules are reddish- brown in color and have a mean width of 0.35mm (range: 0.32-0.38). The reddish-brown coloration persists throughout the entire larval stage. Unlike the larvae of other Hawaiian Omiodes species which maintain bright green body coloration (Swezey 1907), O. continuatalis larva! coloration is comparatively less vivid. The abdomen is a gradient of off-white to light olive, and once larvae begin feeding they acquire the approximate pigment of the vegetation they are digesting. Larvae may molt to the second instar (mean: 0.45mm, range: 0.40-0.50) and the third instar (0.64mm, 0.50-0. 74mm) after three to five days. In the third instar, larvae develop a single black head spot on each of the head capsule lobes, as well as two black spots on the prothoracic shield. O. continuatalis larvae continue to molt every three to five days (4th: 0.92mm, 0.66-1. 10mm; 5th: 1.28mm, 0.90-1. 42mm; 6th: 1.78mm, 1.3-1. 96mm), growing in size and developing more intricate patterns of melanization on the prothoracic shield, as well as tubercles on the 2nd and 3rd thoracic segments and the 7th-10th abdominal segments. During the penultimate instar (6th; 7th: 2.32mm, 1.8-2. 52mm; 8th: 2.84mm, 2. 34-3. 56mm), O. continuatalis larvae acquire a faint pink tint, while the Fig. 2. Omiodes continuatalis larvae exhibit changing pat¬ terns of melanization on the prothoracic shield during nine de¬ velopmental instars. A. Cluster of eggs. B. First instar. C. Sec¬ ond instar. D. Third instar. E. Fourth instar. F. Fifth instar. G. Sixth instar. H. Seventh instar. I. Eighth instar. J. Ninth instar. Volume 63, Number 1 15 head, thoracic and abdominal spots are reduced. Molts to the final instar (7th; 8th; or 9th: 3.19mm, 2. 86-3. 50mm) also occur in three to five days. At this stage, head spots disappear completely, and the thoracic and abdominal segments become veiy pink. Immediately prior to pupation, O. continuatalis larvae become pale and begin to shrink in size to form pre¬ pupae. It is common for lepidopterous larvae to change color prior to pupation, when the procuticle increases in area in the cuticle column above the epidermal cells (Chapman 1998). Pre-pupae develop into pupae over 1-2 days, and remain as pupa for 13-17 days. In total, immature O. continuatalis development was 36-57 davs. J Behavior. Early instar larvae feed on the surface and inner tissue of host plant foliage, often near the midrib of the leaves. Omiodes continuatalis larvae cannot be characterized as gregarious, like the larvae of O. blackburni (Butler, 1877) which share the same shelters throughout most of their development, however early instar O. continuatalis larvae often feed in close proximity to one another. Aggregations of larvae on host plants especially in the first and second instars are a natural result of larvae emerging from eggs laid in clusters. General observations from this research suggest that first instar O. continuatalis larvae are more likely to survive when feeding together. First instar larvae that were separated, and made to feed independently, frequently failed to feed successfully and progress to the subsequent instar. The reasons for this increased mortality are not clear, but may involve the cumulative benefit of aggregated larvae feeding cooperatively during early instars. Research indicates that cooperative feeding behavior has many advantages for lepidopterous larvae, including increasing feeding efficiency on physically tough host plant material (Clark & Faeth 1997). It is possible that O. continuatalis larvae exhibit this type of cooperative feeding due in part to the fact that they have evolved on grasses, which are tougher than many dicot host plants and contain higher concentrations of silica (Falin 1982). When O. continuatalis larvae reach the third instar they continue to consume the surface and inner tissue of host plant leaves, but fed independently from other larvae more frequently than in the early stages. Third instar larvae began to fasten silk threads across the foliage and/or roll-up foliage to create a protected shelter in which to retreat. During the fourth instar and/or when larvae attain a head capsule width of 1mm, they begin to feed on the entire leaf structure. Some larvae were observed to feed predominantly inside their shelters, while others migrated back and forth between their shelter and feeding areas elsewhere on the host plant. Fate instar larvae primarily feed outside the shelters they construct. Feeding patterns were easily discerned based on areas of feeding damage on field deployed host plants, and host plant material provided to isolated O. continuatalis larvae. In colony, O. continuatalis larvae occupied all available space on the host plants provided, from the sheaths at the veiy base of the sugarcane, to the top of each blade. In contrast, when placed at lower densities on host plants in the field, larvae generally stayed on the green sugarcane foliage on the mid to upper portion of the plant. When foliage was available at the base of the plant, larvae often constructed shelters there, using both fresh and dead vegetation as materials. This behavior is consistent with early observations of larval feeding habits made by Swezey (Zimmerman 1958). Additional observations by King & Rubinoff (2008) indicate that some O. continuatalis larvae display a more unusual habit, burrowing beneath the soil surface at the base of their host plants for shelter. In some cases, larvae were recovered up to 14cm below the soil surface. This fossorial behavior was observed in early, middle and late instar larvae, and is not pupation behavior. Another behavior we observed during laboratory colony rearing was larval cannibalism. O. continuatalis larvae, specifically late instars, were observed feeding on other live O. continuatalis larvae. This behavior was more common on occasions when larval density was high and host plant resources were decreasing, but still available. Most frequently, late instar larvae were observed feeding on middle instar larvae. In one case a ninth instar larva was found feeding on another ninth instar larva. In all cases, the larvae being consumed did not appear to fight or tiy to avoid the predation, but remained stationary during the attack. The cannibalistic Instar Fig. 3. Boxplot of headcapsule widths of Omiodes continu¬ atalis larvae by instar. Medians and inter-quartile ranges are rep¬ resented for each instar, and box widths are proportional to sam¬ ple size. 16 Journal of the Lepidopterists’ Society Table 1. Locations surveyed for O. continuatalis on the main Hawaiian Islands. Elevation Trap O. continuatalifi Island Location (m) Method Date nights present Surveyorfs) Kauai Pu'u Ka Pele Forest Reserve 855 light trap 8/6/08 1 No W Haines Koke'e State Park, Camp Sloggett 1060 light trap 5/16/05, 10/6/07 3 No W Haines, C. King Mohihi Road Koke'e State Park 1110 light trap 6/2/04 2 No W. Haines Koke'e State Park, Koke'e Museum 1130 light trap 8/7/07 2 No W Haines Na Pali Kona FR, Pihea trail 1140 light trap 8/7/07 i No W Haines Koke'e State Park, Nualolo Trail 1170 light trap 8/8/07 i No W I Lines, M. Medics Na Pali Kona FR, Alakai Swamp Trail 1180 light trap 8/7/07 3 No King, Haines, Rubinoff Na Pali Kona FR, Pihea trail 1200 light trap 5/18/05, 7/23/06, 8/18/06 4 No Eiben, Haines, Rubinoff Koke'e State Park, Near Awaawapuhi Trail 1245 light trap 6/3/04 1 No W Haines Na Pali Kona FR, Kalalau Lookout 1255 light trap 8/17/06 2 No |. Eiben, D. Rubinoff Oahu Aina Haina, Niuiki Circle 2 light trap 3/24/05 i No C. King Mokuleia Beach 2 light trap 5/1/05 i No W Haines, C. King Sandy Beach 3 light trap 3/6/05. 6/20/05 2 No W Haines, C. King Ewa, open grass lot 15 light trap 5/21/05 i No C. King Diamond Head Crater, wetland 80 light hap 4/3/05, 2/15/08 4 No W Haines Palolo Valley 80 light hap 2/25/08, 2/26/08. 2/27/08, 2/28/08 4 No W Haines HARC Maunawili 160 light trap 5/26/05, 6/22/05 3 No C. King Lyon Arboretum, sugarcane patch 228 light hap 1/27/07 2 No C. King Lanipo Trail 240 light hap 5/29/05 i No W Haines, C. King Dole pineapple field 256 light trap 3/12/05 i No Haines, King, Vorsino I Iawaiiloa Ridge Trail 425 light hap 6/14/06 i No J. Eiben, W Haines Wiliwilinui Trail, access 440 light trap 6/6/06 i No W Haines Wa'ahila Ridge 475 light trap 6/8/06 i No W Haines Kahanahaiki Valiev 605 light trap 2/24/07 i No C. King Mt. Tantalus 613 light hap 6/3/05, 6/22/05 2 No W Haines, C. King Wiliwilinui Trail, Summit 731 light hap 6/24/06 i No W Haines Ko'olau Moutains, Konaliuanui Trail 762 light trap 6/1/06 i No W Haines Palikea Trail 777 light trap 5/15/06 i No W Haines Mokuleia FR, ;dong Mt, Kaala Road 1100 light trap 6/15/07 2 No W Haines Waianae-Kai FR, near Ka'ala NAR boundary 1190 light hap 6/15/07 i No W Haines Ka'ala NAR, hog 1200 light hap 6/15/07 i No W Haines Ka'ala NAR. DOFAWshed 1210 light trap 6/15/07 i No W Haines Maui Kanaha Beach Park 3 light trap 3/20/05 i No C. King Maliko Gulch sugarcane field 76 light trap 6/1/05, 7/13/06 2 No C. King West HCS sugarcane field 137 light trap 6/6/06 1 No C. King Iao Vdley 183 light trap 6/5/06 1 No C. King Waiehu Valley Trail 183 light hap 7/6/06 1 No C. King Haliimaile sugarcane fiekl 274 light hap 6/1/05 1 Yes C. King University of Hawaii Agricultural Park 305 light trap 6/1/06 1 No C. King Waihe'e Ridge Trail 305 \ismd survey 6/8/05 0 Yes W Haines 2955B Kaili'ili Rd, Kokomo 484 light trap 3/19/05“ 15 Yes W Haines, C. King Pu'u Kukui Watershed Kaluanui Valley 485 light hap 45/06 2 No W Haines Waihe'e Ridge Trail 670 light hap 6/8/05 i No W Haines “dates =3/19/05, 5/25/05, 6/1/05, 7/19/05, 9/3/05, 9/23/05, 2/8/06. 3/7/06, 4/18/06, 7/8/06, 3/9/07, 4/19/07, 5/18/07, 8/17/07, 2/6/08 Volume 63, Number 1 Table 1. Continued Island Location Elevation (ni) Method Date Trap nights O. continuatalis present Surveyor(s) Maui Makawao Forest Reserve, Site 1 750 light trap 3/23/05 1 No W. Haines Makawao Forest Reserve, banana patch 762 light trap 6/19/06,7/14/06, 7/15/06 8 Yes W. Haines Makawao Forest Reserve, Site 5 825 light trap .3/23/05 1 Yes W. Haines Kipahulu Valley Delta Camp, HALE 860 light trap 1/15/04,9/14/04 4 No W. I Iaines, D. Rubinoff Makawao Forest Reserve 914 light trap 3/19/05,5/25/05, 6/1/05 4 Yes W. Haines, C. King Makawao Forest Reserve, Site 2 915 light trap 3/23/05 1 Yes W. Haines Makawao Forest Reserve, Site 6 930 light trap 3/23/05 1 Yes W. Haines UH Kula Field Station 975 light trap '3/9/07° 5 Yes C. King Makawao Forest Reserve, Site 4 1050 light trap 3/23/05 i No W Haines Waipoli Road, Kula 1065 light trap 6/20/04 2 Yes W Haines Haleakala Ranch 1280 visual survey 623/06 0 Yes C. King Makawao Forest Reserve 1280 light trap 63/06, S/17/07 2 Yes W. Haines, C. King Makawao Forest Reserve, Site 3 1330 light trap 3/23/05 i Yes \ Y. Haines Kula FR 2073 light trap 64/06 i No C. King Molokai Moomomi Preserve 10 light trap 7/5/05 i No W. Haines Molokai Forest Reserve, TNC Barracks 915 light trap 7/7/05, 1230/05 3 No W. Haines Kamakou Preserve, Kamakou Flats 1105 light trap 5/1 604 2 No W. Haines Kamakou Preserve, Puu Kolekole 1130 light trap .5/19/04,7/7/05 2 No W. Haines Kamakou Preserv e, Pepeopae Bog 1240 light trap 5/19/04, 7/605 2 No W. Haines, D. Rubinoff Lanai Lanai Game Management Area .365 ligjit trap 7/3/05 i No W. Haines Kanepuu Preserve 520 light trap 7/2/05 i No W. Haines Kanepuu Preserve .544 light trap 3/608 2 Yes D. Rubinoff, R Schmitz Munroe Trail 1000 light trap 7/3/05 i No W. Haines Kahoolawe Base Camp, south of Kanapou Bay 5 light trap 1/14/06 i Yes S. Meyers Big Island Pahala sugarcane field 292 light trap 610/05 i No C. King Honokaa sugarcane field 303 light trap 69/05 2 Yes C. King Glenwood, Omega Rd. 700 light trap 2605 2 No W Haines Big Island Glenwood Near Kahaualea NAR 700 light trap 2605, 7/30/06 6 No W. Haines Ka'u, Kaiholena 770 light trap 7/31/06 2 No W Haines, J. Giffin Ola'aFR 822 light trap 25/05 i No W. Haines Ola'a FR 975 light trap 25/05 i No W. Haines Ka'u, Kaiholena 975 light trap 7/31/06 i No W. Haines, J. Giffin Kahuku Ranch, HAVO 1005 light trap 5/18/06 3 No W. Haines, D. Rubinoff Volcano Village 1128 light trap 6605, 11/2605 2 No C. King Ola'a Tract, HAVO 1150 light trap .5/17/06 4 No W Haines Escape Rd/Thurston. HAVO 1158 light trap 4/1605,629/05, 5/18/06 3 No W. Haines, C. King Kilauea Field Station, HAVO 1158 light trap 25/05 1 No W. Haines Kipuka Puaulu, HAVO 1200 ligjit trap 10/15/04 1 Yes W. Haines Ola'a Tract, Pu'u Unit 1300 light trap 5/17/06 1 No W. Haines, D. Rubinoff Waildi 1432 light trap 6/11/05 1 No C. King Mauna Loa Strip Roacl HAVO 1500 light trap 10/14/04 1 Yes W Haines Kona Forest Unit. Hakalau Natl Wildlife Refuge 1615 light trap 4/1205 2 No C. King Keamoku flow; H AM ) 1707 light trap 4/15/05,62605 3 No C. King ° dates = 3/9/07, 4/7/07, 4/19/07, 5/10/07, 5/18/07 18 Journal of the Lepidopterists’ Society Fig. 4. Map of current distribution of Omiodes contimiatalis in the Hawaiian Islands, based on recent field survey data. larvae began their attacks at the tenth abdominal segment, working forward and consuming the entire contents of the abdomen. The only structures not consumed by the cannibalistic larvae were the head capsule and the cuticle. Thus, the end product of such an encounter resembled a shed head capsule and cuticle following a developmental molt. Perhaps the behavior has never been documented in Omiodes larvae because Hawaiian Omiodes species have not been reared in colony prior to this investigation. Cannibalism, prompted by competition for limited resources such as host plant resources and over-wintering sites, has been documented in non- Hawaiian crambid species (Baskauf 2003), so it is possible that O. contimiatalis larvae may exhibit the behavior in the natural environment under high density conditions. Distribution. We completed more than 166 trap nights of surveys at 85 field sites for populations of O. contimiatalis across the Hawaiian Islands (Table 1). While previous collecting records show O. contimiatalis to be present in mostly diy or mesic habitats (Zimmerman 1958; Haines et al. 2004), our surveys show that O. contimiatalis populations are present in diy, mesic and also wet habitat types (Table 1) (Fig. 4). On Hawaii Island, O. contimiatalis was collected from three mesic habitats: Kipuka Puaulu (1195m) and Manna Loa Strip Road (1492m) on Mauna Loa in Hawaii Volcanoes National Park, and a remnant sugarcane field near the town of Honokaa (303m). All of these sites receive a mean annual rainfall (MAR) of 1500-2000mm. On Maui, O. contimiatalis were collected from seven distinct field sites: Waihee Ridge trail (305m). Haliimaile/Haleakala Hwy sugarcane field (274m), Kokomo (484m), UH Kula field station (975m), Makawao Forest Reserve (914m, 1280m), and Haleakala Ranch (1280m). With the exception of Waihee Ridge, which is located on West Maui, all of the field sites are located on East Maui. The two Makawao Forest Reserve sites and the Haleakala Ranch site are wet habitats (2QQ0-3000mm MAR). Waihee is a mesic habitat (1500mm MAR), as is Kokomo (1500-2000mm MAR), while Haliimaile/Haleakala Hwy sugarcane field and the UH Kula field station are dry habitats, due to their position in the rain shadow of East Maui (MAR Volume 63, Number 1 19 <750mm). O. continuatalis were also collected on the islands of Kahoolawe and Lanai. Both of these islands are exceedingly dry (630mm MAR) as they are also located in the rain shadows of Maui and Molokai. Based on survey data of O. continuatalis , it is clear that this species can persist in diverse habitat types ranging from diy to wet. O. continuatalis also occurs in habitats exhibiting varying levels of alteration and disturbance. The species is present in predominantly native forest habitat (e.g. Mauna Loa, Makawao Forest Reserve), as well as areas where native habitat has been converted to pastureland (e.g. Haleakala Ranch), and/or agricultural land (e.g. Honokaa and Haliimaile sugarcane fields). Their persistence appears possible so long as the habitat contains sufficient host plant resources, in the form of native or introduced grasses, to support larval development. Given their widespread distribution, their palpable ability to adapt to significant changes in habitat, and their ability to utilize invasive grasses as host plants, O. continuatalis would seem an unlikely candidate to face high risk of declines and/or extinction. Nevertheless, the current distribution of O. continuatalis on the islands of Maui, Kahoolawe, Lanai and Hawaii represents a significant range reduction for a species which was previously common, but is now extirpated from, Kauai, Oahu, and Molokai. It is possible that the reduction in distribution is not as great as these survey data might indicate. Even small variations in season, moon phase, air temperature, precipitation and wind speed can affect light trap catch on a given day. Therefore, it is possible that O. continuatalis may have gone undetected at sites where light trapping occurred on only one occasion. The Kailiili Road study site in Kokomo, Maui was surveyed frequently due to the consistent O. continuatalis trap catch. Even so, there were at least two occasions where O. continuatalis adults were not observed. Furthermore, during surveys on Hawaii Island, O. continuatalis was not collected from light traps at the Keanroku How on Mauna Loa, where the species has been previously recorded (Haines et al. 2004). Thus, it is possible, that O. continuatalis are present at additional field sites on Maui, Kahoolawe, and Hawaii Island. Accordingly, the species may remain undetected on other islands despite extensive negative data from these survey efforts. This hypothesis seems more plausible at sites where traps were only set-up for 1-2 trap nights, as opposed to field sites like Kokee State Park or the Alakai Swamp on Kauai where more than 18 trap nights have cumulatively been completed, and no O. continuatalis have yet been detected. Potential factors influencing the decline and restricted distribution of O. continuatalis were not quantified under the scope of this research. Given the persistence of the moth in severely degraded habitats, it seems unlikely that habitat destruction or habitat alteration alone could be significant limiting factors for the species. There are habitats on the islands of Kauai, Oahu and Molokai, both pristine and disturbed, which we presume contain suitable resources to support O. continuatalis , and yet the species has still not been “rediscovered” on these islands. If habitat destruction and alteration are not consistently affecting the persistence of this species across the Hawaiian Islands, then what other factors may be contributing to their declines? The intentional and accidental introduction of natural enemies into the Hawaiian Islands has been viewed suspiciously by entomologists for over a centuiy (Perkins 1897; Zimmerman 1958; Howarth 1983; Henneman & Memmott 2001), and some of these parasitoids and predators appear to suppress populations of other Omiodes species (Zimmerman 1958; Bess 1974). Additional research is currently underway to examine the mortality factors, specifically parasitism, which may be affecting O. continuatalis populations. However, the individual and compound effects of introduced predators, such as ants and spiders, are also likely mortality factors, and their effects on Omiodes species remain to be investigated. Acknowledgements We thank A. Arcinas (Hawaii Agricultural Research Center) and M. Nakahata (Hawaii Commercial & Sugar Company) for providing plant material, and D. Oka and G. Otani (University of Hawaii Kula Agricultural Research Station) for field preparation and greenhouse assistance. J. Patu provided essential laboratory assistance. Access permits were granted by S. Meidel (Haleakala Ranch), G. Shishido and B. Gagne (Department of Land and Natural Resources, Division of Forestry and Wildlife), and C. Haines and G. Haines (Pacific Produce). Special thanks to M. Wright, C. Hooks, J. Eiben, L. Leblanc and L. Kaufman (Uni¬ versity of Hawaifi, Department of Plant and Environmental Pro¬ tection Sciences) for valuable input on research methods, and also W. Haines (University of Hawai i, Department of Plant and Environmental Protection Sciences) for access to unpublished data. Anonymous reviewers provided valuable feedback on this publication. Literature Cited Baskauf, S. f. 2003. Factors influencing population dynamics of the southwestern corn borer (Lepidoptera: Crambidae): A reassess¬ ment. Environmental Entomology 32(5): 915-928. Bess, H. A. 1974. Hedylepta blackbumi, a perennial pest of coconut on wind-swept sites in Hawaii. Proceedings of the Hawaiian En¬ tomological Society 21: 343-353. Chapman, R. F. 1998. The insects, structure and function. Cam¬ bridge, Cambridge University Press. Clark, B. & S. Faeth. 1997. The consequences of larval aggregation in the butterfly Chlosyne lacinia. Ecological Entomology 22: 408-415. Fahn, A. 1982. Plant anatomy, 3rd ed. New York, Pergamon Press. Funasaki, G. Y.. P Y. Lai. L. M. Nakahara, J. W. Beardsley, & A. K. 20 Journal of the Lepidopterists’ Society Ota. 1988. A review of biological control introductions in Hawaii: 1890 to 1985. Proc. Hawaii. Entomol. Soc. 28: 105-160. Gag^e, W. C. & F. G. Howarth. 1982. Conservation status of en¬ demic Hawaiian Lepidoptera. Proc. 3rd Congr. eur. Lepid., Cam¬ bridge. Haines, W. P, J. Giffin, & D. Foote. 2004. Rediscovery of five species of Omiodes Guenee (Lepidoptera: Crambidae) on Hawaii Island. Bishop Museum Occasional Papers 79: 45^49. Henneman, M. L. & J. Memmott. 2001. Infiltration of a Hawaiian community by introduced biological control agents. Science 293: 1314-1316. Howarth, F. G. 1983. Classical biocontrol: Panacea or Pandora's box. Proc. Hawaii. Entomol. Soc. 24(2): 239-244. 1UCN. 2007. IUCN - The World Conservation Union. Janz, N. & S. Nylin. 1998. Butterflies and plants: A phylogenetic study " Evolution 52(2): 486-502. King, C. & D. Rubinoff. 2008. First record of fossorial behavior in Hawaiian leafroller moth larvae, Omiodes continuatalis (Lepi¬ doptera: Crambidae). Pacific Science 62(1): 147-150. Perkins, R. C. L. 1897. The introduction of beneficial insects into the Hawaiian Islands. Nature 55: 499-500. Swezey, O. II. 1907. The sugar cane leaf-roller ( Omiodes accepta): With an account of allied species and natural enemies. Honolulu, Experiment Station of the Hawaiian Sugar Planters Association: 1-59. Zimmerman, E. C. 1958. Insects of Hawaii, Lepidoptera: Pyraloidea. Honolulu, University of Hawaii Press. _ . 1978. Insects of Hawaii, Microlepidoptera I. Honolulu, Uni¬ versity of Hawaii Press. Received for publication 23 May 2008; revised and accepted 9 Dec 2008. Volume 63, Number 1 21 journal of the Lepidopterists’ Society 63(1), 2009, 21-26 A NEW SPECIES OF MORRI SONIA (NOCTUIDAE) FROM SOUTHEASTERN NORTH AMERICA J. Bolling Sullivan 200 Craven St., Beaufort, North Carolina 28516; e-mail: sullivanl4@earthlink.net AND James K. Adams Department of Biology, Dalton State College, Dalton, Georgia 30720; e-mail: jadams@daltonstate.edu ABSTRACT. We describe Morrisonia triangula. (Noctuidae: Hadeninae: Orthosini) from material collected from Virginia to Texas. Illus¬ trations of adults and their genitalia are provided. The generic placement is discussed. Additional key words: Achatia, Egira, Himella, Orthosia For at least 40 years collectors in the southeastern United States have been aware of an undescribed species of hadenine provisionally placed in the genus Morrisonia bv Jack Franclemont. Apparently, Franclemont never collected the species but was aware of it from material sent to him for identification by Charles Kimball prior to the latter’s publication of the Lepidoptera of Florida in 1965. Kimball (1965) mentions 5 specimens collected in Escambia Co., Florida, 4 in April of 1961 and one in May of 1962. Additional material, much of which is in the U. S. National Museum, was collected in the period from 1960-80 in Virginia, the Carolinas, Louisiana and Texas by several eastern collectors. Kimball (1965) indicated that Franclemont would describe the species but a manuscript was never completed. In this paper we describe the new species and discuss some of the current taxonomic problems posed by the “genus” Morrisonia. Materials and Methods Specimens were examined from the National Museum of Natural History, Washington, DC (USNM) and Furman University, Greenville, SC (FU). Data were provided by Steve Roble on material in the Virginia Department of Natural Resources (VDNR), Richmond, VA; by Steven Hall on specimens in the North Carolina Heritage Program (NCIIP), Raleigh, NC; and by Richard Brown on material in the Mississippi State Museum (MSM), Starkville, MS. Data were also provided from the personal collections of James K. Adams; Vernon Brou, Abita Springs, LA; Chuck Ely, Nacogdoches, TX; living Finkelstein, Atlanta, GA; Howard Grisham, Maysville, AL; Ed Knudson, Houston, TX; Hugo Kons, Gainesville, FL; Tom Neal, Gainesville, FL; Brian Scholtens, Charleston, SC; Jeff Slotten, Gainesville, FL; and J. Bolling Sullivan. Genitalia were prepared by digestion in 10% potassium hydroxide, dissected in water, and photographed with a Nikon Coolpix 995 camera attached to a Nikon SMZ800 microscope. Additional photographs were provided by Donald Lafontaine (see Lafontaine 1998). Recently collected specimens were sent to Paul Hebert at the University of Guelph for barcode COl analyses (Ratnasingham & Hebert, 2007). Morrisonia triangula sp. nov. Sullivan & Adams (Figs. 1-2) Description. Head: Antennae fasciculate, setae exceeding shaft diameter at base in male, shorter than shaft diameter in fe¬ male; scape straw with rusty scales intermixed; antennal dorsum scaled with alternating straw and rusty bands. Palps upturned, slightly longer than eye diameter, tan, dark brown and reddish scaled. Frons straw above tongue, short scaling but dorsal half of frons protruding with ventral part deep maroon, dorsal part straw, tan with reddish scales; a distinct dark line visible across upper part of frons. Area lateral to eye blackish scaled, tan scales below tongue. Thorax: Neck with a mixture of reddish and straw scales, largely erect. Collar a mixture of white, straw, tan and deep maroon scales forming a series of horizontal bands; scaled tuft behind collar. Tegulae reddish brown with some white scales; scaling underneath mostly white. Legs with reddish scal¬ ing dorsallv, brown with cream scales ventrallv; single pair of spurs on L2, two pairs on L3. Abdomen: two reddish dorsal scale tufts on first two segments, 2nd tuft small. Dorsum of abdomen tan with scattered reddish scales; underneath more reddish but similar. Forewing: Pattern without obvious cross lines, dominated by a black basal and anal dash and a black triangular wedge at mid costa; whitish scaling in costal area before wedge, distal to wedge are 5 whitish costal spots, the most distal the largest; cream elliptical mark between dashes, marginal line black, scal¬ loped; anal dash with smaller dash on costal side, whitish scaling on vein along anal dash; wing base above and below basal dash Journal of the Lepidopterists’ Society darkened; orbicular and reniform spots lightly outlined, filled with the surrounding light gray ground color of the forewing; or- bicular oblong, posterior to triangular black wedge; reniform kidney shaped, just distal to triangular wedge. Hindwing : Fuscus with marginal black band and tan marginal scales forming fringe. Underside: With distinct discal spots; perceptible PM (post- medial) line. Reddish on marginal half of forewing; black scaling on veins distal to PM line giving a striated appearance; basal half of forewing with blackish scaling. Hindwing with similar scaling, black scales densest on costal edges of both wings. Male genitalia (Fig. 6): Valvae symmetrical, extending beyond uncus when viewed laterally, flared toward apex; costal region sclerotized, saccular region membranous; digitus arising from costa and curving ventrally well beyond saccular edge; clasper C-shaped, deriving from well developed medial plate; flared api¬ cal portion of valva setose with corona; uncus gently tapering and arching ventrally, slender, subbasal setae on dorsal side; gnathos and socii absent; anal tube slightly sclerotized ventrally; tegumen arms broad, forming inverted V; penieillum well devel¬ oped and located on pointed flap at posterior end of tegumen; vinculum V-shaped, well developed; pleurite slender, curving up to dorsal side of tegumen; juxta flaring posteriorly, with deep V in middle, posterior edges rounded; aedeagus rounded at base, sclerotized and tapering to a gentle curv ing apex; vesica with 3 or more spirals, basal spiral with longitudinal row of short cor- nuti, largest cornuti in short row above long, longitudinal row, diverticula on first and second spirals, ribbon-like band of em¬ bedded cornuti beginning on second spiral, cornuti patch (6-10 long cornuti) ventral to second diverticulum, distal end of vesica membranous. Female genitalia (Fig. 9); Lightly sclerotized anal papillae, an¬ terior and posterior hypophvses equal in length; ostium a simple flared opening, ductus sclerotized to bursa; accessory bursa aris¬ ing ventrally, spiralled with sperm duct arising near tip: bursa a well defined sack with 4 evenly spaced longitudinal signa ex¬ tending 2/3 toward anterior end, walls of bursa striated. Types: Holoti/pe: Male (Fig. 1): U. S. A. North Carolina: Craven Co., Croatan National Forest, Road 169, 2 iv. 2000, J. B. Sullivan. Paratypes: (11 males, 8 females) - North Carolina: Col¬ lection data same as holotype (2 males); Craven Co., Croatan National Forest, Road 147, 1 1 iv. 2000 (2 males, 1 female); 23 iv. 2006 (2 males, 1 female); 12 v. 1996 (1 female); 6 vi. 2006 (1 fe¬ male); Croatan National Forest, Road 3046, 26 vi. 1998 (1 fe¬ male) J. B. Sullivan. Georgia: Lumpkin Co., Blood Mountain, .25 mi. S of Neel’s Gap, 3100', 27 v. 2006 (1 female). Louisiana: St. Tammany Parish, sec 24T6SR12E. 4.2 mi. NE of Abita Springs, 30 iii. 1998 (1 male); 10 iv. 1998 (1 male); 25 iii. 2002 (1 male); 28 iii. 2004 (1 female). Texas: Tyler Co., Big Thicket Na¬ tional Preserve, Turkey Creek Unit, 15—16 iv. 1994 (2 males); 8 iv. 1995 (1 female). The holotvpe will be deposited into the U. S. National Museum and paratypes into the Canadian National Collection, British Museum, Florida State Collection of Arthro¬ pods, Texas A&M University, and the Texas Lepidoptera Survey (with Charles Bordelon and Edward C. Knudson, Houston, TX). Etymology. The name triangula is an adjective describing the distinctive triangular mark along the forewing costa. We suggest Triangle-barred Mor¬ risonia as the common name. Diagnosis. Morrisonia triangula is abundantly distinct in maculation and genitalia from other species of Morrisonia and from other hadenine species encountered in the Eastern United States. In pattern it resembles less heavily marked forms of M. evicta (Grt.) but is larger, has the distinct costal triangle and except perhaps for a few mountain localities, is allopatric. If the two species do fly together, one would expect M. evicta to he on the wing earlier in the spring. M. triangula more closely resembles M. latex (Gn.) (Figs 3, 4) in size and ground color, and, to some extent, in the major dark markings of the forewing. The two species certainly occur in similar habitats in the mountains and possibly on the coastal plain. When the wing pattern is obscured, the genitalia of either sex can be used to distinguish the species. Males of M. triangula lack hair pencils at the base of the abdomen as do M. confusa (Hbn.) and M. latex. The digitus is almost basal in M. confusa hut crosses the membranous sacculus more distally in M. latex and triangula. In M. latex the membranous area is swollen distally, in triangula it tapers evenly. The everted vesica of triangula is unique in the genus because it has no large cornutus nor comb-like concentrations of smaller cornuti. Instead, it has a ribbon-like band of small cornuti . Distribution and Habitat. This species occurs from the southeastern corner of Virginia along the coastal plain to all of Florida and west to east Texas. There are veiy few piedmont records (which may reflect a lack of collecting in this area). In the mountains it occurs from Averv Co. in North Carolina to the foothills in NE Alabama. County records are as follows: VIRGINIA: City of Suffolk, City of Virginia Beach, Isle of Wright; NORTH CAROLINA: Bladen, Brunswick, Carteret, Craven, Cumberland, Hoke, Macon, Martin, Moore, Onslow, Pender, Union; SOUTH CAROLINA: Charleston, Greenville, Oconee; TENNESSEE: Cocke; GEORGIA: Fulton, Long, Lumpkin, Rabun, Towns, Union; FLORIDA: Alachua, Escambia, Liberty, Monroe, Okaloosa; ALABAMA: Bibb, Conecuh, Sumpter; MISSISSIPPI: Forrest, Franklin, George, Hancock, Harrison, jackson. Pearl River, Pike; ARKANSAS: Ashley, Union; LOUISIANA (Parishes): Iberville, St. Tammany, West Feliciana, Winn; TEXAS: Hardin, Jasper, Polk, Sabine, Tyler. Collections of this moth are not limited to a specific habitat. It occurs in swamps, savannahs, pinelands and mesic hardwood sites throughout the Coastal Plain. Mountain sites are equally undistinguished. Flight Season. In the Coastal Plain of North Carolina the extended flight period runs from late March into }une and perhaps early July. Most males fly in mid April and most females are in flight in May-June. Mountain records (above 4000’) are May through mid July. For other states with numerous records, the recorded flight periods are as follows: Mississippi: Volume 63, Number 1 23 Figs. 1-4. Adult Morrisonia triangula Sullivan and Adams. Male. Little Deep Creek Rd., Croatan National Forest, Craven Co, N. C. April 2, 2000, J. Bolling Sullivan 2. Morrisonia triangula Sullivan and Adams. Female. Croatan National Forest Rd. 147, Craven Co., N. C. April 1 1, 2000, }. Bolling Sullivan. 3. Morrisonia latex (Guenee). Male. East side of Carp Ridge, 45°24.63"N, 76° 03.45'W, Ontario, 14 June 2003, j. Troubridge. 4. Morrisonia latex (Guenee). Female. Temiscouata Co., Quebec, 16 June 1993, Henry Hensel. March 27-June 2; Louisiana: March 12-June 15; Texas: April 8-May 22; South Carolina: May 2 1— [uly 25 (the latest recorded specimen, from the mountains [Oconee Co.]); Georgia: April 23-May 27; Florida: March 17-May 23; Alabama: June 3- June 12. There is no evidence for a second brood. Generic Placement. The genus Morrisonia is attributed to Grote (1874), with evicta as the type species, but no characters are given in his work to help delineate the genus. Hampson (1898-1913) included taxa from North America and New Zealand but species from New Zealand have since been removed (Dugdale 1988). Forbes (1954) characterized the genus as follows: “vestiture always mixed, the division of the tuft on upper part of face obvious; collar usually with suggestion of a central ridge, and concave on either side; double dorsal tufts of thorax distinct; several distinct tufts on abdomen.” Since Forbes’ treatment of the genus, M. distincta Hbn. has been moved to the genus Achat ia (Hodges et. al. 1983) and Mamestra latex (Gn.) has been moved into Morrisonia (Poole 1989). Other North American genera placed in the tribe Orthosiini by Lafontaine (2007) are Acerra, Stretchia , Perigonica, Orthos ia , Himella, Egira and Achatia. Members of the tribe are united by characters of the larval hypopharynx and spinneret (Fibiger & Lafontaine 2005), characters which we have not evaluated. Genitalic variation in this tribe and within several of the genera is extreme (Ronkay et. al. 2001). The genus Morrisonia is no exception. What characters then seem to place this new species in Morrisonia? Two external characters other than wing pattern support placement of the new species in Morrisonia. First, the vestiture is coarse, a mixture of scales and hairs, and in most of the other genera only hair scales are present. Second, there is a well- differentiated line across the Irons that occurs in the other Morrisonia species but is extremely faint when present (rarely) in other genera. Comparison of related species and genera ( Egira alternans (Wlk.), variabilis (Sm.), dolosa (Grt), crucialis (Harv.), mbrica (Harv.), cognata (Sm.); Stretchia inferior (Sm.); Perigonica angulata (Sm.); Himella fidelis Grt.; Orihosia praeses 24 Journal of the Lepidopterists’ Society Figs. 5-7. Male genitalia of Morrisonia. 5a,b. Morrisonia evicta (Grote). a. Valves. Dunrobin, Ontario (CNC 13538). b. Aedea- gus. Same data as male valves. 6a, b. Morrisonia triangula Sullivan and Adams, a. Valves. Wedge Plantation, 7miN McClellanville, South Carolina (CNC 13964). b. Aedeagus. same data as male valves. 7a,b. Morrisonia latex (Guenee). a. Valves. North Gower, Ontario (CNC 13539). b. Aedeagus. same data as male valves. 7a 1 mm Volume 63, Number 1 25 8 lmm Figs. 8-10. Female genitalia of Morrisonia spp. 8. Morrisonia evicta (Grote). Female. Aweme, Manitoba (CNC 13560). 9. Mor¬ risonia triangula Sullivan and Adams. Female. Key Largo, Florida (CNC 13542). 10. Morrisonia latex (Guenee). Female. Lac Mon- dor, Ste Flore, Quebec (CNC 13540). (Grt), rubescens (Wlk.), hibisci (Gn.), alurina (Sm.), pacifica (Han/.), revicta (Morr.), pulchella (Han/.), mys (Dyar), erythrolita (Grt.), garmani (Grt.) and segregata (Sm.)) and all 5 Morrisonia species using COl barcoding techniques (Hebert et. al. 2006) reveals that, whereas most species sort with their supposed congeneric members, there are often inclusions of one or two additional non congeneric species in each generic cluster. In the case of Morrisonia both evicta and triangula sort together but may be separated from the other Morrisonia species by Orthosia hibisci or rubescens. Examination of the male and female genitalia (Figs. 5, 6, 8, 9) shows that M. evicta and triangula are quite similar in all characters and the pair is most similar to M. macens (Hbn.). M. confusa and M. latex have genitalia which are quite divergent from the macens, evicta, triangula complex. Significant divergence is seen in the COl data from M. latex but not M. confusa. Diverse assemblages of species are characteristic of Orthosia, Egira, and perhaps other orthosiine genera with multiple species (Ronkay et al. 2001). Finding good generic characters is difficult at best. For M. mucens, evicta (Fig. 5) and triangula (Fig. 6) we could list the spiral vesica with groups of cornuti but no major spine, a short diverticulum near the apex of the vesica, and the valva, diaphanous along its ventral edge, indented from the ventral side past the midpoint and expanded distallv. The valve process derives from the costal plate; the medial plate often produces a small process. In females (Figs. 8, 9) the appendix bursa is coiled, the ductus bursa well sclerotized, and the ostium simple. These same characters are either missing or highly modified in M. confusa and latex (Fig. 10). 26 Journal of the Lepidopterists’ Society Morrisonia confusa and latex are polyphagous (Covell 2005; Rings et al. 1992). Most records for Morrisonia evicta are from Primus , although Kalmia and Cornus are listed also, perhaps erroneously (Wagner et al 2008). Jack Franclemont reared Morrisonia mucens from Water Oak ( Quercus nigra L.) as reported in Godfrey (1972). While most orthosiines require tender new foliage, both M. confusa and M. latex develop comparatively slowly on older, hardened summer foliage. M. confusa is exceptional among noctuids in taking as long as 6-7 months to mature (Wagner 2005). The habits of M. confusa and M. latex are strikingly different from each other and completely different from the other spring feeding orthosines. M. confusa begins the year feeding externally on new growth as do the other species, but in the last instar it fashions a shelter on old summer foliage, densely lines it with silk, and continues to feed for upwards of 5-6 months. Morrisonia latex also feeds throughout the summer but rests on bark by day (at least in the last instar), a habit completely unique to other orthosines (Wood & Butler 1989; Wagner et al 2008). Based on the sum of differences in larval morphology, behavior, and life history, M. evicta , confusa and latex could easily be placed in three separate genera. Thus, everything points to inclusion of triangula in the genus Morrisonia , considering that it seems to be the closest relative to evicta , the type species. In fact it is a better fit than some of the other species currently placed there. When the larval biology of triangula is known, hopefully, it will confirm this placement rather than add more heterogeneity to yet another genus in the Orthosiini. Acknowledgements YVe would like to acknowledge the following persons who contributed data, insight or both: Vernon Brou, Richard Brown, Chuck Ely, Irving Finkelstein, Howard Grisham, Steven Hall, Paul Hebert, Ed Knudson, Hugo Kons, Don Lafontaine. Tim McCabe, Tom Neal, Steve Roble, Dale Schweitzer, Brian Scholtens, Darryl Searcy, Jeff Slotten, John Snyder and Dave Wagner. Figures were prepared by Jocelyn Gill. Literature Cited Coyell, C. V., Jr. 2005. A field guide to the moths of eastern North America. 2nd Ed. Houghton Mifflin Col, Boston, Mass. Dugdale, J. S. 1988. Lepidoptera-annotated catalogue, and keys to family group taxa. Fauna of New Zealand 14: 1-262. Fibiger, M. & J. D. Lafontaine. 2005. A review of the higher classi¬ fication of the Noctuoidea (Lepidoptera) with special reference to the Holarctic fauna. Esperiana (Buchreihe zur Entomologie), Band 11, pp. 7 - 92. Forbes, W. T. M. 1954. Lepidoptera of New York and neighboring states. Noctuidae, Part III. Cornell University Agricultural Ex¬ perimental Station. 433 pp. Godfrey, G. L. 1972. A review and reclassification of larvae of the subfamily Hadeninae (Lepidoptera, Noctuidae) of America north of Mexico. Tech. Bull. No. 1450. U. S. Dept. Agriculture, Wash¬ ington, D. C. 265 pp. Grote, A. R. 1874. List of the Noctuidae of North America. Bull, of the Buffalo Society of Nat. Hist. 2: 1-77. Hampson, G. F. 1898-1913. Catalogue of the Lepidoptera Phalaenae in the British Museum. Volumes 1-13. Hebert, P. D. N., E. H. Penton, |. Al. Burns, D. H. Janzen & W. Hallwachs. 2006. Ten species in one: DNA barcoding reveals cryptic species in the neotropical skipper butterfly Astraptes fttl- gerator. Proc. Natl. Acad. Sci. 101: 14812-14817. Hodges, R. W„ T. Dominick, D. R. Davis, D. C. Ferguson, J. G. Franclemont, E. G. Munroe, J. A. Powell (eds). 1983. Check list of the Lepidoptera of America north of Mexico (including Greenland). E. W. Classey Ltd. and The Wedge Entomological Research Foundation, London. 284 pp. Kimball, C. P. 1965. The Lepidoptera of Florida: an annotated check¬ list. In Arthropods of Florida and neighboring land areas. Vol. 1. Fla. Dept. Agile., Gainesville. 363 pp. Lafontaine, J. D. 1998. Noctuoidea: Noctuidae (part), Noctuinae (part-Noctuini). Fascicle 27.3. In R. B. Dominick et al. (eds). The moths of America north of Mexico. The Wedge Entomological Research Foundation. Washington, D. C. 348 pp. _ . 2007. Updated electronic checklist of the moths of America. Poole, R. W. 1989. Lepidoptorum catalogus. Fascicle 118, Noctuidae, 3 volumes. Ratnasingham, S. & P D. Herbert. 2007. Bold: The barcode of life data system. Molecular Ecology Notes 7(3): 355-364. Rings, R. W, E. H. Metzler, F. J. Arnold. & D. H. Harris. 1992. The owlet moths of Ohio (Order Lepidoptera: (Family Noctu¬ idae). Ohio Biol. Survey Bull., New Series, Vol. 9, College of Bio¬ logical Sciences, Ohio State University, Columbus, Ohio. 219 pp. Ronkay, L.. J. L. Yela & M. Hreblay. 2001. Noctuidae Europaeae, Hadeninae II. Soro: Entomological Press. 452 pp. Wagner, D. L. 2005. Cateipillars of eastern North America: A guide to identification and natural history, Princeton University Press. 496 pp. _ , D. F. Schweitzer, J. B. Sullivan, & R. C. Reardon. 2008. Owlet caterpillars of Eastern North America (Lepidoptera: Noc¬ tuidae). Princeton Universitv Press (in preparation). Wood, P. S., & L. Butler. 1989. Biology and immature states of Po- lia latex (Guenee) (Noctuiidae). [. Lepid. Soc. 43: 299-304. Received for publication 31 January 2008; revised and accepted 20 October 2008 Volume 63, Number 1 27 Journal of the Lepidopterists’ Society 63(1), 2009, 27-30 TINACRUCIS NOROESTA, NEW SPECIES, NORTH AMERICA’S LARGEST TORTRICINE MOTH (TORTRICIDAE: ATTERIINI) [erry A. Powell Essig Museum of Entomology, University of California, Berkeley, 94720, e-mail: poweIlj@nature.berkeley.edu ABSTRACT. The Neotropical tribe Atteriini is briefly characterized, and Tinacrucis noroesta is newly described in order to make the name available for a forthcoming book on moths of western North America. This species, the northernmost of the tribe, occurs in the Sierra Madre Occidental, Mexico, and mountains of southern Arizona, where it was first collected in 1927. The larvae are believed to be polvphagous leaf rollers on broadleaf trees and shrubs but have not been discovered in the field. Additional Key words: Egg mass adornment scaling, sexual dimorphism. Chihuahua, Durango, Arizona, leaf-roller. The Atterini comprises one of the smallest and most well defined tribes of the Tortricinae. Its monophyly is convincingly supported by a suite of uniquely derived morphological and behavioral characters associated with oviposition (Powell 1986). The adults are among the largest tortricid moths in the Western Hemisphere, FW length ranging to 22 mm. Females are conspicuously larger than males, and there is pronounced sexual dimorphism in wing shape and color patterns in some genera (. Anacrusis , Tina, Tinacrucis ), whereas the sexes are similar in other genera (e.g., Atteria, Templemania). The eggs are laid in imbricate, overlapping patches, presumably a synapomorphy with Archipini and Sparganothini. Females possess obvious patches of corethrogyne scales covering the venter of A6 and A7; a thick pad of long, thin, dark scales on A6, which are spread over the egg mass and its periphery, and an enormous patch of upright, thicker scales on A7 that are curled at the tips, which are glued to the substrate in clumps, like a fence surrounding the egg mass. Most atteriines are nocturnal, but Atteria are diurnal and brightly colored, evidently specific Batesian mimics of Pseudatteria (Tortricinae: Polyorthini). Atteriine larvae are similar to sparganothines but are much larger leaf rollers. The distance between V setae on A9 is greater (1.5-2X) than on A8, the crochets are bi- or triordinal, and there is a well developed anal fork. I have reared several species of Templemnaia, Tina, and Tinacucis from eggs to maturity on synthetic diet, which suggests atteriines are polyphagous, and Dan Janzen and others have reared field collected larvae of Costa Rican Anacusis from several unrelated plant families, enhancing that assumption. The Atteriini includes about 40 described species assigned to eight genera, in addition to numerous undescribed taxa in Central and South America. A few species range into northern Mexico, one into Arizona. Tinacrucis Powell, 1986, is distinguished by having elaborate antennal setulae in males, the costa sinuate in both sexes (unmodified in male Anacrucis and Templemania), markedly different size and FW pattern between the sexes (though not as extreme as in Tina), greatly enlarged A8 tergum, forming a hood enclosing the male genitalia. Tinacrucis includes six named species and several undescribed in Mexico and Central America. I have reared broods of several of them from single female egg masses, enabling association of the markedly differing sexes. I have been aware of the species described here for 50 years, based on males taken by J.A. Kusehe in the Chiricahua Mountains, Arizona, in 1927, but until recent years, we lacked sufficient material to confirm conspecificity of the Arizona species with specimens collected in Durango, Mexico. This species is described now because we discuss it in our forthcoming book. Moths of Western North America (Powell & Opler in press), scheduled for publication by the University of California Press in May, 2009. A treatment of the Sparganothini and Atteriini, by J. W. Brown and Powell, is in preparation for Moths of America North of Mexico, an appropriate place for this description, but it will be published later. Both of those volumes will include color images of the new species. Tinacmcis noroesta Powell, new species (Figs. 1, 2, male, female from El Salto, Durango, Mexico) This is the largest tortricine moth in America north of Mexico. Males have a relatively narrow FW with sinuate costa, pale beige to yellowish reticulated with rust lines and irregularly mottled with rust-brown. Females are larger with broader wings, pale chocolate- to rust-brown with faint darker and white banding in costal area when fresh and with huge masses of modified scales ventrally on abdominal segments 6 and 7. 28 Journal of the Lepidopterists’ Society Fig. 1 . Tinacrucis noroesto Powell <$ — Mexico, Durango, YV of El Salto, VII. 5. 1964 (J. Martin) CNC. Fig. 2. T. noroesta Powell Mexico, Durango, YV of El Salto, VII. 1.1964 (J. Martin) CNC. Volume 63, Number 1 29 Fig. 3. Tinacrucis noroesta, male genitalia, ventral aspect, valvae spread; aedeagus, lateral aspect shown to right; a, aedeagus with vesica extended, revealing the deciduous comuti. Fig. 4. T. noroesta, female genitalia, ventral aspect; a, detail of translucent ventral plate of A9; b, detail of signum, dorsal aspect. 30 Journal of the Lepidopterists’ Society Description. MALE. Forewing. Length 14.7-16.3 mm (n = 25), narrower than female, length 2. 6-2. 8 x width, costa sinu¬ ate, concave beyond middle, apex produced and subtended by concave terminal margin; ground color pale beige to yellow, reticulate with rust-brown along veins and numerous cross lines between veins, confined by rust-brown clouding to irregular blotches, basally, in cell, and subterminal area (resembling skeletonized spots on a fallen leaf); costal area beyond middle whitish in fresh specimens. Hindwing. Broad, pale cream col¬ ored, tinged with rust apically; underside variably mottled by rust in costal and apical areas, which is visible from above. Ab¬ domen. Scaling pale beige to rust tinged caudally, venter paler, with segmental dark, medial spots. Genitalia . As in fig. 3 (drawn from paratype, El Salto, JAP slides 2893; n = 8); uncus only slightly flared apically, socii densely covered with elongate scales; joined tip of gnathos elongate, slightly flared; a pair of dentate projections on posterior margin of transtilla usually widely separated but adjacent in one specimen from Jalisco; basal half of valva with weakly projecting saccular ridge re¬ cessed from anterior margin; vesica with a compact group of 8 bladelike, deciduous cornuti situated in two rows; after dehis¬ cence, the attachment scars are visible at high magnification (100X) on the withdrawn vescia near the phallobase/aedeagus junction. FEMALE. Forewing. Length 17.0-20.0 mm (n = 6); broad, length 2.45-2.55 x width; costa weakly concave beyond middle; ground color milk chocolate- to rust-brown, showing faint transverse, whitish and dark bands on costal half beyond middle on fresh specimens; underside rust tinged; fringe narrowly rust. Hindwing. As in male. Abdomen. Scaling pale beige dorsally; venter paler, with large, medial, black spots on segments 3—5; A6 witli a thick patch of thin, dark brownish scales; A7 with a conspicuous patch of upright, pale to dark scales, each bent at the tip. Genitalia. As in fig 4, a, b (drawn from paratype, Sum- merhaven, JAP slide 9080; n = 3); papillae anales broad, darkly pigmented, with raised setiferous nipples; sterigma a broad bowl subtending A9, which is modified as a translucent plate covered with tiny spicules, more densely at margins (4a); signum a densely wrinkled patch at bursa surface, preceded by a small, blind pouch, projected inwardly as a broad, weakly wrinkled, hollow blade (4b). Holotype female: ARIZONA, Pima Co., Summerhaven, 7,800', Santa Catalina Mountains, emerged 10 December 1997, larva reared on synthetic diet from egg deposited by female col¬ lected 29 August 1997 (R. B. Nagle. JAP lot 97H9); in Essig Museum of Entomology, University of California, Berkeley. Paratypes (35 6, 7 9): MEXICO, CHIHUAHUA, Sierra de la Catarina, 7,900', 18 rd. mi. NE Buenaventura, 1 6 VIII. 21. 1976 (J. & K. Donahue, LACM). DURANGO. 10 mi. YV of El Salto, 9,000', 1 6 VI. 13.1964, 5 d, 19 VII.5/29.1964 (J. Martin, CNC), 1 6 VII. 30. 1964 (Chemsak & Powell, EME), 1 6 VIII. 8. 1964 (Powell, EME). JALISCO, 18 km SW Sayula, 1 6 X.18.1992 (J. McCarty, EME). USA; ARIZONA, Apache Co., Greer, White Mts., 3 6 VHI.6/8.1962 (E. & I. Munroe, CNC), 2 6 VIII. 5. 1986 (R. Leuschner, RL), 6 Alpine, 8,200', VIII. 3. 1986 (R. Leuschner, RL). Cochise Co., Barfoot Ridge, 8,500'. Chiricahua Mts., 2 6, VIII.6.1927 (J. Kusche, CAS); Barfoot Park, 1 9 IX.27.1987 (N. McFarland, EME); Rustler Park, 3 6 VII.30.1987, 1 6 VII.24.1989, 1 i VIII. 7.1997 (R. & J. Robertson, EME), 1 d VIII.6.1991 (Robertson & Powell, EME); Pinery Canyon, 6,500', Chiricahua Mts., 1 d VIII.6.1991 (Robertson, EME); Pinery Canyon 6800’ 1 9 VIII. 1.2005 (K. Richers, KR); Miller Canyon, 6,200’, Huachuca Mts., 1 6 VIII. 7. 1974 (Powell, EME). Graham Co., Soldier Creek Camp, 9,350', Graham Mts., 1 9 VIII. 16.1974 (R. & J. Wielgus, USNM); Mt. Graham, Cunningham Campgr., 3 d VII. 20.2007 (C. Ferris, CF, EME). Pima Co., same data as holotype, 5 d, 2 9 VII.24 to VIII.29, 1991 to 1999 (EME, RN), 1 9 VIII. 1996, eggs infertile (EME), 1 d VII.21.1998, (D. Fer¬ guson, USNM)."" Diagnosis. The geographically nearest congener is Tinacmcis apertana (Walker), with which T. noroesta Hies in the mountains of Durango, Mexico. I have reared T. apertana in Nuevo Leon from eggs, confirming association of the sexes. The two are similar, but the FW of apertana males has more extensive network pattern on a paler, cream yellowish ground, defining a broad, brown costal triangle; females of apertana have cream-tan FW, more or less uniformly strigulate with thin, brownish, transverse lines, with a brownish bar from mid-costa curving towards the tornus, and a poorly defined subterminal patch, contrasted with the more uniform rust tan of noroesta. T. sebasta (Walsingham), which was described from Guatemala and occurs in Mexico, has the reticulate pattern restricted to the basal half of the male FW, anti the female FW pattern is similar to that of apertana but darker tan. The uncus of T. apertana has a wide, T- shaped apex, and that of sebasta is broadly spatulate subapieally, tapering to the tip, whereas it is narrow, only weakly flared in noroesta. T. apertana and sebasta have narrower valvae than does noroesta , with a more distinct saccular ridge. The female signum is short and broad in T. apertana , elongate in sebasta , more slender than in noroesta. In addition we have two male specimens from Mexico, Sinaloa, 8 road miles W of El Palmito, 6,400', X. 12.1975 (Chemsak & Powell, EME), which are similar but differ in several respects. They are slightly larger than the other T. noroesta specimens, and their colors are richer, more contrasting yellow and brown. The male genitalia have a slender uncus, not flared. The dentate projections of the transtilla are closely adjacent, and there are slight differences in the valva. These specimens are provisionally considered to represent another population of T. noroesta. Acknowledgements Carolyn Mullinex made the drawing of the male genitalia. Sev¬ eral collectors made special efforts in response to my pleas to col¬ lect this remarkable species in Arizona and Mexico; the late John Chemsak, Ron Leuschner, Ray Nagle, the late Ron Robertson, and Ron Wielgus. Paratypes are deposited in the California Academy of Sciences (CAS); Canadian National Collection, Ottawa (CNC); Essig Museum of Entomology, UC Berkeley (EME); Los Angeles County Museum of Natural History (LACM); Ron Leuschner, Manhattan Beach, California (RL); Ray Nagle. Tucson, Arizona (RN); K. Richers, Bakersfield, California (KR); and U.S. National Museum of Natural History, Washington, D.C. (USNM). Literature Cited PowelI, J. A. 1986. Synopsis of the classification of Neotropical Tor- tricinae, with descriptions of new genera and species (Lepi- doptera: Tortricidae). Pan Pacific Entomologist 62:372-398. Received for publication 27 Aug 2008: revised and accepted 17 Nov 2008. Volume 63, Number 1 31 Journal of the Lepidopterists' Society 63(1), 2009, 31-36 MELITARA WALKER (PYRALIDAE) IN WESTERN CANADA: THE DOCUMENTATION OF M. SUBUMBRELLA (DYAR) IN THE PRAIRIE PROVINCES DEMONSTRATES THE VALUE OF REGIONAL COLLECTIONS AND SPECIES LISTS. Thomas J. Simonsen Department of Biological Sciences, University of Alberta, Edmonton, AB, Canada; email: thomas.simonsen@ualberta.ca Jason J. Dombroskie Department of Biological Sciences, University of Alberta, Edmonton, AB, Canada. AND Gregory R. Poiil Natural Resources Canada, Northern Forestry Centre, Edmonton. AB, Canada. ABSTRACT. The moth genus Melitara comprises 8 recognized species of medium sized to large micromoths that all feed on Opuntia cacti as larvae. One species, M. dentata is widespread in western North America, including the southern parts of the Canadian provinces British Co¬ lumbia, Alberta and Saskatchewan. We report the presence of a second species, M. subumbrella in southern Alberta and Saskatchewan, a species hitherto widely believed to extend no farther north than southern Idaho and Wyoming. Closer examinations of Melitara collections in major Canadian museums and public collections revealed several additional M. subumbrella specimens from Alberta, including specimens from the 1940s. Two specimens were actually reported by K. Bowman (1951) under a previous taxonomic arrangement, but were' subsequently over¬ looked by later workers on North American Phycitinae. Combined with subtle differences in mitochondrial DNA, this leads us to conclude that M. subumbrella has been present in Canada for a long time. This provides an excellent example of the scientific value of continuous insect col¬ lecting, well-curated regional collections, and regional faunal lists. Additional key words: Disjunctive distribution, mitochondrial DNA, cactus-feeding The pyralid genus Melitara Walker, a prominent group of cactus-feeding phycitines, is found throughout much of the USA, southwestern Canada, and northern Mexico. One species, M. prodenialis Walker, is widespread throughout the east, and another species, M. dentata (Grote), is widespread in the west. The remaining 6 species are till restricted to Mexico and the southwestern USA with only M. subumbrella (Dyar) extending north to Wyoming and southern Idaho, and west to southern California (Heinrich 1956; Mann 1969; Neunzig 1997). Bowman (1951) reported M. dentata and Olycella [ =Melitara ] nephelepasa from Alberta, but his records were ignored by later workers. The latter is referable to M. subumbrella, which was considered to be a synonym of M. nephelepasa at the time (McDunnough 1939). A full list of currently recognized species of Melitara and their reported range is given in Table 1. Neunzig (1997) included the genus Olycella Dyar in Melitara, with the three species O. subumbrella, O.junctionella (Hulst) and O. nephelepasa (Dyar) (Heinrich 1939, 1956). This synonymy has later been shown to be systematically justified (Simonsen 2008). It is not currently clear whether Melitara and Olycella (both sensu Heinrich 1956) are reciprocally monophyletic and thus deserve subgenus status. All species in the genus feed strictly on prickly pear cactus in the genus Opuntia (Mann 1969) and are thus restricted to habitats with Opuntia. Identifying Melitara subumbrella. Both M. subumbrella and M. dentata are large micromoths with wingspans exceeding 50 mm. With their stout bodies and rectangular wings they have a superficial similarity to medium sized Noctuoidea. The resting moth with its large porrect palps, and the wings coiled cigar-like around the body is fairly easy to identify as a phycitine (Fig. 1). Fresh specimens can fairly easily be identified to species without dissection (at least in areas where only M. subumbrella and M. dentata occur). Both species have white hind wings and gray forewings (sometimes with a yellowish cast — especially in older collection specimens). Melitara dentata has a fairly elaborate zigzag pattern across the forewing, whereas M. subumbrella has uniformly gray forewings almost devoid of any pattern (Fig. 2a, b). Old, faded, and especially denuded specimens can be much harder to identify. In this case dissection is often necessary. Females are easy to identify based on genital ic dissections as M. subumbrella. The species formerly 32 Journal of the Lepidopterists’ Society Fig. I . Live Melitara dentata displaying typical Phycitinae posture with the wings rolled around the abdomen and the an¬ tennae pressed down along the back. CAN: AB: Buffalo: 50.848°N -110.696°W: 16 viii 2006. Photo: JJD. grouped in Olycella are the only members of Melitara with a signum on the corpus bursae (Heinrich 1939, 1956; Neunzig 1997). The males are harder to identify, but the shape of the valves differs slightly between the two species. In M. dentata the valve has a fairly pointed apex with curved dorsal and ventral margins. The costa extends almost to the apex of the valve (Fig. 3a). In M. subumbrella the valve has a much broader and blunter apex, and the costa does not extend as far (Fig. 3b). Finally, the flight period is often a veiy good way to tell the species apart. All M. subumbrella from Canada were collected between May 15 ami July 2, whereas all M. dentata were collected between July 27 and October 25, so there is only a small risk of an overlap in flight time between the two species. This pattern where M. subumbrella is a late spring to early summer flyer, and M. dentata is a late summer to fall flyer seems to be the same in the western USA, though some M. dentata are reported from early Julv (Neunzig 1997). Herein we report on an examination of M. subumbrella specimens, to determine if Canadian populations are distinct from those farther south. To do so, we examined morphological and genetic characters. Materials and Methods Morphology. To search for morphological differences, two male and one female M. subumbrella from Grasslands National Park SK, one male and one female M. subumbrella from central USA, and several male and female M. dentata were examined. Abdomens were dissected and macerated in 10% KOH, and stained in Chlorazol Black in a 70% ethanol solution by TJS, both for use in this study and others (e.g. Simonsen 2008). Terminology follows Heinrich (1956), Klots (1970) and Neunzig (1997). Molecular analysis. To test for genetic differences between the Canadian and USA populations, we sequenced 740 bp from the cytochrome oxidase subunit I (COI) gene from three specimens from Grasslands National Park and one from Colorado. DNA was extracted from legs with the QIAgen’s DNEasy extraction kit (QIAgen Sciences, Maryland, U.S.A). PCR reactions were performed using the Jerry - Pat primer pair (Simon et al. 1994), and the PCR cycling profile was: 95.0°C for 2 minutes followed by 35 cycles of denaturizing (95.0°C for 30 sec.), annealing (45.0°C for 30 sec.) and extension (72.0°C for 2 minutes). PCR products were purified using the QIAgen’s PCR Purification Kit, and the PCR primers were also used for direct sequencing. Sequencing was done with an ABI Prism 377 DNA sequencer using Big Dye®. Consensus sequences from the two sequencing directions were constructed using Sequencher 4.1 and aligned by eye. The sequences are deposited on GenBank under accession numbers FJ422995-FJ423000. Phylogenetic analysis. An exhaustive maximum parsimony (MP) analysis was carried out in PAUP0 4.10b (Swofford 2002). The haplotype distributions were analyzed in MacClade 4.0 (Maddison & Maddison 2000). The tree was rooted using a M. dentata specimen from Alberta as outgroup. Fig. 2. Melitara dentata (a) and M. subumbrella (b) display¬ ing subtle, yet clear differences in forewing pattern (arrows point to the zigzag pattern on the forewing of M. dentata). Volume 63, Number 1 33 Table 1. Currently recognized species of Melitara and their distributions following Heinrich (1956), Mann (1969), and Neunzig (1997). Species Distribution Melitara prodenialis Walker Southeastern U.S.A. from Texas and Florida to New York Interior western North America from north western Mexico, Arizona and western Texas to Melitara dentata (Grote) southwestern Canada. Melitara texana Neunzig Texas Melitara doddalis Dvar Southern Arizona, southern New Mexico and western Texas. Melitara apigramella Blanchard & Knudson Southwestern Texas Melitara junctionalla Hulst Southern Texas and eastern Mexico. Melitara nephelepasa (Dyar) Central Mexico Southwestern USA from southeastern California and western Texas to southern Idaho, Melitara subumbrella (Dyar) Wyoming and Nebraska. Results Morphology. When dissected, males from Grasslands National Park were found to be indistinguishable from males from Colorado, USA. The dissected female from Grasslands National Park, showed a very subtle difference. The signum on the corpus bursae was less developed than the corresponding structure in specimens from Colorado. Molecular analysis. The MP analysis resulted in a single tree 62 steps long (Fig. 4). The four specimens from Grasslands National Park all shared the same haplotvpe; whereas the specimen from Colorado differed in three base pairs, corresponding to a difference of 0.4%. M. clentata differed in 59 base pairs compared to all four M. subumbrella specimens, corresponding to a difference of 8.0%. Discussion Melitara in Canada. In 2004 GRP collected five specimens of M. subumbrella in Grasslands National Park, Saskatchewan. In his annotated list of Lepidoptera in Alberta, Bowman actually reported two specimens of M. subumbrella from the Medicine Hat region (Bowman 1951), though he listed them as Ohjcella nephelepasa (following McDunnough 1939). Melitara subumbrella is technically a new record for Canada. However, a simple update of Bowman's (1951) checklist, following the synonymy reported in Munroe (1983) would yield the report of this species in Alberta. Both the specimens and Bowman’s paper have been overlooked by later workers on the genus in North America (e.g. Heinrich 1956; Mann 1969; Neunzig 1997). One of Bowman's specimens was located in the University of Alberta Strickland Museum. Four other specimens were misidentified as M. dentate in the same collection, including another specimen collected by Bowman in 1940 near Medicine Hat. The latter is obviously Bowmans second specimen which later must have been misplaced in a tray with M. dentate. The three remaining specimens were collected in 2005 at two different localities in southeastern Alberta. This inspired us to examine other public collections that we felt could hold either overlooked or misidentified specimens of M. subumbrella from western Canada. Examinations of the materials in the collections of the Canadian Forest Service (Edmonton, AB), Royal Alberta Museum, (Edmonton, AB), Olds College (Olds, AB), Royal Saskatchewan Museum (Regina, SK) and the Canadian National Collection, (Ottawa, ON) revealed one M. subumbrella from Lethbridge, AB collected by C. E. Lilly in 1949, and one from Onefour, AB collected by GRP in 1995. The same examinations also showed that M. dentate occurs from south-central Fig. 3. Differences in male genitalia between Melitara den- tata (a) and M. subumbrella (b). Note the more pointed apex of the valve and that the costa extends to the apex in M. dentata. 34 Journal of the Lepidopterists’ Society Saskatchewan to the southern interior of British Columbia (Table 2), a more widespread Canadian distribution than reported recently by Neunzig (1997), who only listed the species from southern Alberta. The distributions of both species based on the collection data are shown in Fig. 5. These records underscore the importance of regional collectors, and their published lists and collections, in determining species ranges. Local collectors typically have extensive knowledge of productive collecting localities, and often amass extensive collections of excellent quality (Ferris 1986; Hendra 2005, 2007). These collections and publications are a neglected resource that can yield many exciting finds to the diligent taxonomist. Two prickly-pear species, Opuntia fmgilis (Nutt.) Haw. and O. polyacantha Haw., occur (sometimes abundantly) throughout the southern parts of Canada’s Prairie Provinces (personal painful observations). Though Fragile Prickly-pear Cactus (Opuntia fmgilis), a possible host plant for M. clentata (Mann 1969; Neunzig 1997), occurs scattered in remnants of the Peace River grasslands in northwestern Alberta (Royer & Dickinson 2007), there are no records of M. clentata from that region. It is, however, quite possible that M. clentata Table 2. Records of Melitara in Canada based on the collections listed in the text. Collection acronyms: CNC: Canadian National Collection of Insects, Arachnids and Nematodes; NFRC: Northern Forestry Centre Research Collection; OC: Olds College: RSM: Royal Saskatchewan Museum; UASM: University of Alberta, Strickland Museum. Species Locality Coll, period Year No. Collection M. clentata Grasslands National Park, SK 24/7-12/9 2002 6 RSM M. clentata Buffalo Pond Provincial Park, SK 7/8-25/8 1974, 1984-85 3 RSM M. clentata Wood Mountain Park, SK 4/9 2002 2 RSM M. clentata Fort Qu'Appelle, SK 12/8-31/8 1976, 1991 2 RSM M. clentata St. Victor, SK 14/8 2002 i RSM M. clentata Matador, SK 7/8 2002 i RSM M. clentata Eastend, SK reared 1974 i RSM M. clentata Saskatchewan Landing Provincial Park, SK 10/8 1981 i RSM M. clentata Scout Lake, SK 27/8 1968 4 CNC M. clentata Swift Current, SK 8/8 1936 1 CNC M. clentata Tolinan Bridge, AB 3/8-25/10 2000-2, 2006 13 UASM, NFRC, OC M. clentata Taber, AB 14-15/8 1999, 2005 2 OC M. clentata Medicine Hat, AB 17/8 1943 1 UASM M. clentata Edgerton, AB 24-25/8 2002 1 UASM M. clentata Steveville Bridge, AB 13/8 1999 2 UASM. NFRC M. clentata Buffalo, AB 16/8 2006 i UASM M. clentata Champion, AB 3/8 1961 12 CNC M. clentata Coaldale, AB 4/8 1961 8 CNC M. clentata Manyberries, AB 10/8 1951 1 CNC M. clentata Lethbridge, AB 26/7-24/8 1921-22, 1949, 1986 6 CNC M. clentata Willow Creek Canyon, AB 29/7 1961 4 CNC M. clentata Kamloops, BC 5/8-9/8 1937, 1956 4 CNC M. clentata Nicola, BC 29/8-6/9 1922-23 4 CNC M. clentata Oliver, BC 1/8-15/9 1923, 1953 113 CNC M. subumbrella Grasslands National Park, SK 2/6 2004 5 NFRC M. subumbrella Onefour, AB 19/6 1995 1 NFRC M. subumbrella Lost River Valley w. of Onefour, AB 20-21/5 2005 1 UASM M. subumbrella Medicine Hat, AB 31/5 1940 2 UASM M. subumbrella Oldman River Hwy. 36, s ofVauxhall, AB 18/5 2005 2 UASM M. subumbrella Lethbridge, AB 15/5 1949 i CNC Volume 63, Number 1 35 does occur in the Peace River region, as suitable habitats in the region are fairly poorly collected, especially during M. dentata’ s main flight period (G. G. Anweiler pers. com.). The status of Melitora subumbrella in Canada. The three oldest specimens of M. subumbrella from southern Alberta show that the species was present in the province 60-70 years ago. The presence of recently collected individuals from different localities in southeastern Alberta and Saskatchewan clearly demonstrates that M. subumbrella occurs at multiple localities in western Canada. It is also not unthinkable that the species still is overlooked and is more widespread, not only in southwestern Canada, but also in Montana, though an examination of the Melitara specimens in the entomological collection of Montana State University in Bozman did not result in additional specimens. Are Canadian M. subumbrella different from M. subumbrella in the U.S.A.? Ideally, many more specimens from both Canada and the U.S.A. should have been included to strengthen the results of the molecular analysis. Unfortunately such material was not available. Nonetheless, the difference reported here is well within variations that have been found in other tarn subumbrella in Canada and west central USA. The closed circle marks the clade comprising the four Canadian specimens. The six digit code after the species name is the specimen ID. The numbers indicate the number of unique base pair changes. Fig. 5. Distribution map for Melitara in western Canada based on the records listed in Table 2. An open circle marks a M. subumbrella locality, and a closed triangle marks a M. dentata lo¬ cality. insect populations considered to be conspecific (e.g. Simonsen et al. 2008; Cognato 2006; Laffin et al. 2005a, b; Scheffer & Grissel 2003). The slight difference in female genitalia may well represent individual variation. It is not known whether the species is present in Montana, and the variation thus could be continuous. Despite the apparent geographical isolation, these differences do not in our opinion justify separate taxonomic status (subspecies or otherwise) for the Canadian M. subumbrella populations. The species should be sought after in Montana to determine whether the distribution is continuous or the Canadian populations are disjunct. Acknowledgements We wish to thank Paul Opler, of the Cillette Museum, Col¬ orado State University for loans of specimens from the U.S.A. and for permission to extract DNA from these. We thank the fol¬ lowing people and institutions for help and assistance: |ean- F ranyois Landry, Canadian National Collection, Ray G. Poulin, Roval Saskatchewan Museum, Tyler Cobb, Royal Alberta Mu¬ seum, Ken Fiy, Olds College and Michael A. Ivie, Montana State University, Bozman. Felix A. II. Sperling, Gary G. An¬ weiler, Danny Shpeley, Marie Djernaes and Lisa Lumley, all of the University of Alberta, and Brenda Laishley, Canadian Forest Sendee, Edmonton. AB, are thanked for help, advice and dis¬ cussions. Financial support was provided by The Carlsberg Foundation, Copenhagen, Denmark (post doc salary forT. J. Si¬ monsen). NSERC (a Discovery Grant to Felix A. H. Sperling paid for lab work and various expenses), and Natural Resources Canada (support for faunistics work by G. R. Pohl). Literature Cited Bowman, K. 1951. An annotated list of the Lepidoptera of Alberta. Can. J. Zool. 29: 121-165. Cognato. A. I 2006. Standard percentage DNA sequence difference for insects does not predict species boundaries. (. Econ. Entomol. 99: 1037-1045. Ferris, C. D. 1986. Presidential address, 1986: unexplored hori¬ zons — the role of the amateur lepidopterist. J. Lepid. Soc. 40: 247-254. 36 Journal of the Lepidopterists’ Society I Ieinrich, C. 1939. The cactus-feeding Phycitinae: a contribution to¬ wards a revision of the American pyralidoid moths of the family Phycitidae. Proc. U. S. Nat. Mus. 86: 331^13. _ . 1956. American moths of the subfamily Phycitinae. U. S. Nat. Mus. Bull. 207: i-viii, 1-581. Hendra, L. 2005. Kenneth Bowman, Alberta Lepidopterist. Blue Jay 64:168-174. _ . 2007. Kenneth Bowman, Alberta Lepidopterist, part 2. Alberta Lepidopterists' Guild website: http://www.biology.ua]berta. ca/old_site/uasm/aIg/downloads/KBowman_article_Part2_Sept20 07.pdf Klots, A. B. 1970. Lepidoptera. Pp. 115-130. In S.L. Tuxen (ed.). Taxonomists glossary of genitalia in insects (second edition). Munksgaard, Copenhagen, Denmark. Laffin, R. D., L. M. Dosdall, & F. A. II. Sperling. 2005a. Popula¬ tion structure of the cabbage seedpod weevil, Ceutorhynchus ob- strictus (Marsham) (Coleoptera Curculionidae): Origins of North American introductions. Environ. Entomol. 34: 504-510. _ , _ . & _ . 2005b. Population structure and phvlogenetic relationships of Ceutorhynchus neglectus (Coleoptera: Cur¬ culionidae). Can. Entomol. 137: 672-684. Maddison, W. P. & D. R. Maddison. 2000. MacClade: version 4.0 PPC. Sinauer, Sunderland. Mann, J. 1969. Cactus-feeding insects and mites. U. S. Nat. Mus. Bull. 256: v-x, 1-158. McDunnough, [. II. 1939. Check list of the Lepidoptera of Canada and the United States of America. Mem. South. Cal. Acad, of Sci. 1: Part II. Munroe, E. 1983. Pyralidae (except Crambinae). pp. 67-85. In: Hodges, R. W., Dominick, T., Davis, D. R., Ferguson, D. C., Franclemont, J. G. Munroe, E. G., & Powell, J.A. 1983. Check list of the Lepidoptera of America North of Mexico. E. W. Classey Ltd. and the Wedge Entomological Research Founda¬ tion, London, UK. 284 pp. Neunzig, II. H. 1997. Pvraloidea, Pyralidae (part). Fascicle 15.4 In Dominick, R. B., et al. (eds). The moths of America north of Mex¬ ico. The Wedge Entomological Research Foundation. Washing¬ ton, D.C. 157pp. Royer, F. & R. Dickinson. 2007. Plants of Alberta. Lone Pine Pub¬ lishing. Edmonton, AB, Canada. 527pp. Scheffer, S. }. & E. E. Grissell. 2003. Tracing the geographic ori¬ gin of Megastigmus transvaalensis (Hymenoptera: Torymidae): an African wasp feeding on a South American plant in North America. Molec. Ecol.12: 405-414. Simon, C., F. Frati, A. Beckenbach, B. Crespi, H. Liu. & P. Flook. 1994. Evolution, weighting, and phylogenetic utility of mito¬ chondrial gene sequences and a compilation of conserved poly¬ merase chain reaction primers. Ann. Entomol. Soc. Amer. 87: 651-716. SlMONSEN, T. J. 2008. Phylogeny of the cactus-feeding phycitines and their relatives (Lepidoptera: Pyralidae) based on adult morphol¬ ogy: evaluation of adult character-systems in phvcitine systemat- ics and evidence for a single origin of Cactaceae-feeding larvae. Insect Syst. Evol. 39: 303-325. _ . R. L. Brown, & F. A. H. Sperling. 2008. Tracing an invasion: phylogeography of the cactus moth, Cactoblastis cactonun (Berg) (Lepidoptera: Pyralidae) in the U.S.A. based on mitochondrial DNA. Ann. Entomol. Soc. Amer. 101: 899-905. Swofford, D. L. 2002. PAUP°: Phvlogenetic Analysis Using Parsi¬ mony (“and Other Methods) (Version 4.0bl0). Sinauer Associ¬ ates, Sunderland. Received for publication 5 June 2008; revised and accepted 27 October 2008. Volume 63, Number 1 37 Journal of the Lepidopterists’ Society 63(1), 2009,37-47 NOT ALL BILATERAL GYNANDROMORPH BUTTERFLIES ARE INTERSPECIFIC HYBRIDS: NEW PAPILIO SPECIMENS FROM FIELD POPULATIONS Mark Scriber Department of Entomology, Michigan State University, E, Lansing, MI 48824; research associate McGuire Center for Lepidoptera and Diversity, University of Florida, Gainesville, FL 3261 1-2710; email: scriber@pilot.msu.edu Rodrigo Mercader Department of Entomology, Michigan State University, E. Lansing, MI 48824 Howard Romack 28 Irish Lane, Cambridge, NY 12816, AND Mark Deering Lepidoptera Curator, Sophia Sachs Butterfly House, 15193 Olive Blvd., Chesterfield. MO 63017 ABSTRACT. Bilateral gynandromorphs (half male, half female) have previously been reported in interspecific hybrids. However, we report 6 new nearly perfect bilateral gynandromorphs, three (one Papilio canadensis , and two P. glaucus) of which were collected from areas not near hybrid zones. New interspecific gynandromorphs also are described for apparent P. nitulus x P. multicaudatus hybrids. Bv using hand-paired in¬ terspecific hybrids we have generated both diapausing and direct developing (non-diapausing) gynandromoiphs. While X (or Z)-linked regula¬ tion of diapause in these Papilio may be similar throughout the male and female halves of the individual, the dark melanic color [Y (or W)- linked] is distinctly different. Wing color regulation is differentially implemented on dorsal and ventral surfaces as seen in sexual mosaics described here. The developmental irregularities resulting in bilateral gynandromorphs and sexual mosaics are not solely due to interspecific ge¬ netic incompatibilities. However, genetic factors seem to play a major role in their expression. The existence of dark melanic female patchy pat¬ terns supports the suggestion that control of the W (=Y)-linked b* melanic gene is both cell-specific and sex-specific in its ability to suppress the BAS enzyme activity that normally shunts dopamine to the yellow papiliochrome pathway. Interspecific hybrids involving dark mothers (with the Y-linked b* gene) mated to P multicaudatus , P. nitulus , and P. canadensis males (all species lacking the dark morph) have also produced hy¬ brid daughters with “intermediate” suppression of melanism giving a dusty colored appearance (differing from color mosaics). This uniform “dusty” coloration often has been referred to as “cinnamon,” and may reflect incomplete genetic penetrance. Additional key words: melanic wing color, sex chromosomes, Papilio glaucus, P. canadensis, P. nitulus, P. multicaudatus, P. appalachiensis Lepidoptera that have developed abnormally to produce mosaic color patterns and a mixture of other traits, often representing intersexes (part female and part male) have frequently been described (Clarke & Ford 1980; Ayala & Kiger 1984; McCafferty & Bloodgood 1986). If such developmental errors occur early in development (perhaps from fertilization of a binucleate oocyte; Blanchard & Descimon 1988) the result might be a perfect (or near-perfect) bilateral gynandromoiph with one half being male and one half female (Clarke & Clarke, 1983). A large number of gynandromoiphs have been reported in Lepidoptera from at least 10 different species of Papilionidae, eight species of Lycaenidae, four Hesperiidae, ten Pieridae, four Nymphalidae, and several families of moths including Saturniidae, Pyralidae, Geometridae, and Lymantriidae (reviewed in Scriber & Evans 1988b; see also Sibatani 1980, 1983a, b; Shuey & Peacock 1985). Cockayne (1935) provides numerous additional records, and the Milwaukee Public Museum’s fames R. Neidhoefer Collection has several hundred mosaics and at least 150 bilateral gynandromorphs. David and Marc Perlman in Illinois have also compiled a large collection of mosaics and gynandromorphs. A number of suggestions have been advanced to potentially explain these developmental abnormalities including: extreme temperatures (Standfuss 1900; Drescher & Rothenbuhler 1963; Riotte 1978), ultraviolet light (Nekrutenko 1965), viral infections (Gardiner 1972; Blau 1978; Sevastopulo 1973), translocations of parts of the sex-chromosomes and/or 38 Journal of the Lepidopterists’ Society autosomes (Blanchard & Descimon 1988; Seri her & Evans 1987; Marec et al. 2001), and background uranium radiation-induced somatic mutations and the Chernobyl nuclear power plant disaster (Dantchenko et al. 1995). Interspecific hybrid crosses have also been suspected as a potential cause of mosaic/gynandromorphs (Whicher 1915; Cockayne 1935; Clarke & Sheppard 1953; Clarke et al. 1977; Platt 1983; Blanchard & Descimon 1988; Scriber & Evans 1988a, b). In Papilio such gynandromorphs have been found to be most often the result of interspecific (or intersubspecific) crosses (Scriber & Evans 1988b). Asymmetrical color mosaics may also result from such interspecific crosses, but are more often seen from the central parts of a species geographic range, or from intraspecific pairings, which suggests that mosaic gynandromorphs could be the result of more complex and basically different developmental processes than bilateral gynandromorphs (Clarke & Clarke 1983; Blanchard & Descimon 1988). The evidence we had collected from 1979-1986 with the Papilionidae in eastern North America suggested that perhaps only interspecific hybridization would generate bilateral gynandromorphs, and at that time we had never found a “natural” (i.e. field-collected) bilateral gynandromoiph (Scriber & Evans 1988b). However, we have subsequently collected interesting mosaics and near-perfect bilateral gynandromoiphs from field populations, some of which were not clearly near a hybrid zone where introgression from different species was obvious. We report on some of these interesting aberrations here. Tiger Swallowtail Gynandromorphs and Color/Sexual Mosaics. For an earlier study (Scriber & Evans 1988b), we reared nearly 30,000 butterflies from more than 4,000 families of Papilio glaucus and P. canadensis (previously considered subspecies P. glaucus canadensis ; Hagen et al. 1991). Of these individuals we observed 6 of 34 color mosaic adults, all of which possessed basic dark (mimetic) wing patterns of females interrupted by yellow patches or blotches on one to three of the individual wings. These yellow patches represent either the non-mimetic female, or male type pattern. Such color mosaics may represent intersexes or gynandromorphs with part male and part female asymmetric traits. They may be either gynandromoiphs or sexual mosaics (Blanchard & Descimon 1988; Cockayne 1935; Ford 1955; Sevastopulo 1973). In Papilio glaucus and P. alexiares Hopffer, both polymoiphic yellow and black forms are found (Scriber et al. 1996). Only these two Papilio species have the polymorphic dark and yellow morph females, clearly showing the color contrast in gyndromoiphism/ mosaicism. One such P. glaucus female (with one yellow half and one mostly dark half) has recently been figured and described by Bradshaw (2000). In addition to these 34 color mosaics, we also observed three perfect bilateral gynandromoiphs (including one that is half dark female and half yellow male) and two “nearly” perfect bilateral gynandromoiphs basically half dark and half yellow (Scriber & Evans 1988a, b). These perfect bilateral gynandromoiphs were all offspring from interspecific hybrid crosses. Furthermore, while the dark color of the female half was expressed in the pairing of a dark Ohio female with a P. alexiares Mexican male (Fig. 1), the female half of the second gynandromoiph [a dark female P. glaucus from South Carolina (with the b+ gene) mated with a P. canadensis male from Marinette Co., Wisconsin] had the dark potential suppressed (Fig. 2; Scriber et al. 1996). The third gynandromoipih (a yellow P glaucus from Pennsylvania mated with a P. canadensis from Juneau Co., Wisconsin) had the female half yellow because it likely lacked the Y-linked dark gene from the mother and also probably received an X- linked suppressor gene from the P canadensis father (Scriber et al. 1996). Such gynandromoiphs are not only fascinating in appearance, but also with regard to physiology. Both of these P. glaucus x P canadensis bilateral gynandromoiphs emerged in the second year and did not develop directly as did many of their “normal¬ looking” brothers. This reflects an X-linked obligate diapause that is hemizygous in the female hybrids (and the female half of these gynandromoiphs; XY or ZW), but heterozygous in its sibling male hybrids (XX or WW), which allows direct development and adult male eclosion without diapause. Apparently the two halves of both gynandromoiphs were cued to eclose as adults in the next season rather than directly (at the 16:8 L:D photophase, 23°C rearing conditions; Scriber & Evans 1988b). Of the 9,000 or more field collected specimens during this study (1979-1987; Scriber & Evans 1988a, b) we never collected an obvious bilateral or nearly bilateral gynandromoiph. The closest was one reared from a field-collected female from Jasper Co. Texas (Fig. 4). However, we collected many color mosaics (dark and yellow patterns being the most obvious). In fact, 33 of the 39 color mosaics were field-captured individuals (6 were lab-reared from hand-paired hybrids). While most of these field-collected color mosaics were from Georgia, Ohio, Illinois, and Texas, one family from a female captured in Dane Co., Wisconsin produced 17 of the color mosaics. It is possible that some introgression from P canadensis Volume 63, Number 1 39 Figs. 1-3. 1. A bilateral gynandromorph (from brood #4210) from a lab cross of a dark morph P. glaucus female (Adams Co., Ohio) mated to a wild collected male P. alexiares garcia from Nuevo Leon, Mexico, (a) dorsal, (b) ventral. 2. A bilateral gynandromorph (from brood #628) from a lab cross of a dark morph P. glaucus female (Pickens Co., South Carolina) mated to wild collected male ( P. canadensis from Marinette Co., Wisconsin (collected 1-5 July 1983 by Don Caine), (a) dorsal, (b) ventral. 3. A gynandromorph, or sexual mosaic, which appears to be more than 60% male (from brood #2830) from a dark morph P. glaucus female (reared from eggs obtained from a dark morph female wild collected in Clay Co., West Virginia) bred to a wild collected P. rutulus male (reared from eggs obtained from a wild female collected in Sacramento Co., California), (a) dorsal, (b) ventral. could have occurred in this geographic area since it is very near the hybrid zone (Luebke et al. 1988). In fact, the famous 19th century Strecker collection (Strecker 1878; Walsten 1977; Ehle 1981; Shapiro 1981a, b; Clarke & Clarke 1983) of P. glaucus color (or sexual) mosaics (which had been on loan from the Chicago Field Museum to the Allyn Museum) may reflect such introgression in those mosaics near the Papilio glaucus/ P. canadensis hybrid zone in the Midwest and Pennsylvania (Scriber et al. 2003). Here we review our hypothesis in light of new information. We reared these hybrid larvae only through to pupation on black cherry (Primus serotina) leaves, since this host is used by both P. canadensis and P. glaucus (and by all four other species in the P. glaucus group; Scriber 1996). If we had attempted to rear these gynandromorphs on tulip tree, Liriodendron tulipfera (this P. glaucus favorite is toxic to P. canadensis; Scriber 1982) or on quaking aspen, Populus trenudoides (this P. canadensis favorite is toxic to P. glaucus, Lindroth et al. 1988), these insects might have might have survived to become adults, as normal hybrids, since detoxification enzymes are inherited autosomally, and are used inside the gut, they can be expressed throughout the insect regardless of whether they are produced by one or both halves (male or female) of the gynandromorphs (Scriber 1986; Scriber et al. 1989). Results We collected a female Papilio canadensis from Cheboygan Co., Michigan during the period 9-12 June 1997. Larvae were reared on black cherry. One of the resulting adults was a nearly perfect bilateral gynandromoiph with blue hindwing color and female genitalia on the right, and with male genitalia and coloration on the left (Fig. 5). Another near-perfect bilateral gynandromorph was actually collected in the field on 15 June 1996 in southwestern Vermont near the New York border (II. R.). This specimen has basically female blue 40 Journal of the Lepidopterists’ Society Figs. 6-8. 6. A bilateral gynandromorph of P. canadensis collected (H.R.) in Bennington Co., Vermont on 15 June 1996 (a) dorsal, (b) ventral. 7. A bilateral gynandromorph collected (H.R.) on 3 June 1989 in Lincoln Co., Wyoming. Possibly P. rutulus female (left half) and P. multicaudatus male (right half). 8. A hybrid (lab-paired, lab-reared) of a P. rutulus female x P. multicaudatus male (brood #7806). 5 (enlarged) Figs. 4-5. 4. A nearly perfect gynandromorph or sexual mosaic (from brood #1091) from a wild collected dark morph ( P. glaucus female from Jasper Co., Texas, (a) dorsal, (b) ventral. 5. A bilateral gynandromorph lab-reared from a family that was derived from a female P. canadensis collected in Charlevoix Co., Michigan, 9-12 June 1997 (a) dorsal, (b) close-up (In collection of Mark Deering). Volume 63, Number 1 41 coloration on the right hindwing and slightly heavier (wider) black bands (as in females) on the right forewings (Fig. 6; this specimen appears to possess 2 male claspers and no ovipositor). The third nearly perfect bilateral gynandromorph is one collected from Lincoln Co. (Wyoming) on 3 June 1989. This individual appears to be half female P. rutulus and half male P. multicaudatus (Fig. 7). Both species lack the dark morph females and probably the Y- linked b+ gene (Seriber et al. 1990, 1996). The right hindwing has the extra “tail " and the right forewing appears somewhat similar to P. multicaudatus in banding. The left female half looks rutulus- like. For comparison, we also present a female from an interspecific lab pairing of P. rutulus female x P. multicaudatus male (Fig. 8, brood #7806). In this specimen, the two species traits appear somewhat intermediate throughout and did not segregate into the two halves (one female and one male, as in the interspecific gynandromorph; Fig. 7). A field-captured, nearly-perfect bilateral gynandromorph between the nymphalids Limenitis arthemis arthemis (Drury) and L. a. astyanax (F.) was captured in the same area of the southwestern Vermont/New York border as the P. canadensis gynandromoiph (not figured) and may be part of an active hybrid zone. These Papilio and Limenitis species (sub-species) complexes have significant introgression and exhibit centers within their hybrid zones in New England, as they do across the Midwest in Michigan and Wisconsin (Platt & Brower 1968; Seriber 1996). We have also lab-reared a dark nearly perfect bilateral gynandromorph from a dark, east Texas P. glaucus mother (brood #8858, Janice Bossart). A specimen photo sent to us by Loretta Lutman of Ashboro, North Carolina (Randolf Co.) also appears to be a wild, bilateral gynandromoiph of P. glaucus, with the female half dark (Fig 9). Perfect bilateral gynandromorphs, with half expressing male traits and the other female traits, are extremely rare. Color/sexual mosaics with asymmetrical blotches or patterns of color (or other mixed gender traits) are somewhat more common than bilateral gynandromorphs among these swallowtails, but they are still quite rare. Earlier studies with nearly 30,000 lab- reared North American Papilio resulted in 30-40 color mosaics (approximately 0.001 % of reared adults was an obvious color and/or sexual mosaic; Seriber & Evans 1988b). It is easiest to find color (or sexual) mosaics in P. glaucus when the female is of the dark morph, but the hindwing blue color of females is also helpful in spotting them (Fig. 10). One thing that is clearly characteristic of color mosaics is that the dorsal and ventral patterns are determined independently; a few examples are illustrated in Figs. 11-16. Another mosaic from Illinois shows asymmetry (Fig. 17). It is also possible to have bilateral differentiation as in size or color for the same sex and single species individual larvae, pupae, and adults (see Seriber & Hagen 1990). Discussion We describe six new and nearly perfect bilateral gynandromoiphs from field collections in northern Michigan, southwestern Vermont, Montana and Texas. Four of these are likely to represent interspecific or intersubspecific hybrids (Figs. 5-7, and the Limenitis from Bennington Co. Vermont, collected by HR). While perfect and near-perfect bilateral gynandromorphs appear to be often generated by interspecific hybridization, we report two gynandromoiph specimens from Texas and North Carolina (Fig. 9) and East Texas (]. Bossart and JMS, not figured here) that were not near the P. canadensis/ glaucus hybrid zone. The somewhat unique ability of individual butterflies with one half male and one half female to develop as larvae, pupae, and live adults capable of flight (e.g. Seriber et al. 1995; Conners 2000) is fascinating, but still poorly studied and understood. The dramatic differences in wing color with partial dark morph females and yellow-patterned males enables us to easily detect individual bilateral gynandromorphs or asymmetrical sexual mosaics, but the yellow morph female gynandromorphs of tiger swallowtail butterflies are no less fascinating. Here we have presented one example of interspecific bilateral gynandromoiphs that were bred from Papilio glaucus female and P. canadensis male parents (Fig. 2; see also Seriber et al. 1995). This interspecific pairing united genes in the resulting gynandromorph that have segregated into the two gender-differentiated halves with the heterogametic female [XY (=ZW)] possessing the Y-linked black (b+) gene of dark morph females plus the X-linked suppressor (s+; Seriber et al. 1996), resulting in the yellow (suppressed) tiger pattern for the female half (Fig. 2). This contrasts with the interspecific (P. glaucus x P. alexiares) gynandromoiph with the dark melanic (b+) gene and no suppressor (s', an enabler), where the female half is black (Fig 1). The biochemical pathways (temporally and spatially regulated by enzymes, including dopa carboxylase) are apparently coordinated by multiple enzyme activities of the melanin (black) and papilioehrome (yellow) specific pathways. These enzyme regulations must be conducted simultaneously in the same gynandromorphs (Koch et al. 2000a), all of which is coordinated by the single major Y-linked factor (Seriber et al. 1996) on the female 42 Journal of the Lepidopterists’ Society Figs. 9-11. 9. A natural bilateral gynandromorph of P. glaucus from Randolph Co., North Carolina (photo from Loretta Lutman). 10. A sexual mosaic (gynandromoiph) with mostly male characters (except the blue coloration in the left hindwing) brood 2025, from a wild-collected yellow female, Clark Co., Georgia. 11. A color/sexual mosaic (lab-reared from a female collected in Ohio, 1989 (a) dorsal, (b) ventral. Figs. 12-13. 12. A gynandromoiph or sexual mosaic, which appears to be more than 60% female (from brood #3622) from a pure P. glaucus lineage (in which the mother was the daughter of a dark morph female collected in Hocking Co., Ohio and mated to a male from Adams Co., Ohio), (a) dorsal, (b) ventral. 13. A gynandromorph or sexual mosaic, which appears to be more than 75% female but with male claspers (from brood #3935) from a wild collected dark morph P. glaucus female (Scioto Co., Ohio), (a) dorsal, (b) ventral. Volume 63, Number 1 43 side of the individual; but these genes are either not expressed, or not present, on the male half oi the individual. The activity of other sex-linked traits such as obligate diapause regulators would explain the direct development (non-diapause) of both halves of the P. glaucus/P. alexiares gynandromorph and the one year delay (diapause) of the P. glaucus/P. canadensis gynandromorph (with both the hemizygous female and the heterozygous male having the X-linked obligate diapause gene, od+). Other sex-linked traits such as opposition preferences could not be analyzed, but in the future researchers could examine the sex-linked diagnostic allozymes and mitochondrial DNA variation with regard to distribution in the two halves of such gynandromorphs. In melanic females, black melanin replaces the background yellow' papilioehrome (Koch et al. 1998). The key enzyme involved is BAS (n -(5- alanyl- dopamine-synthase), which shunts (suppresses) dopamine from the melanin pathway into the production of the yellow papilioehrome pigment (probably in response to dosage of the Z (=Y)-linked b+ gene products; Scriber et al. 1996). The mixed mosaics we figure here appear to reflect the fact that melanism is both sex-specific and pattern-specific (cell autonomous), and therefore melanic specimens of P. glaucus are not just null mutants as in some ebony Drosophila strains (Koch et al. 2000b). The suppressor/enabler gene (s'/s+) that permits dark female expression of the b+ melanin gene has been identified on the Z (=X) chromosome (Scriber et al 1996), and may involve tyrosine hydroxylase, or linked factors that are involved in production of dopa (common to both melanin and papilioehrome pathways) (Koch et al. 2000a, b). The late incorporation of tyrosine into melanin may also be involved in the incomplete penetrance (where yellow scales are brownish) suggesting that sex-linked control genes and their sex- linked modifiers exert their effects by differential interpretation of the patterning signal (e.g. the delay in development of melanic scales is female-specific) (Ffrench-Constant & Koch 2003). The specific melanin (b+) modifier genes (s+/s') on the Z-chromosome, remain to be identified, but we have isolated the tyrosine hydroxylase gene from the Z- chromosome off! glaucus (Putnam et al. 2007). The Z- chromosome also carries other genes of major significance in this species, including oviposition preferences (Scriber 1993; Mercader & Scriber 2007; Mercader et al. 2009), diapause regulation (Rockey et al 1987a, b), post-diapause emergence delays (Scriber et al. 2008), and allozymes (lactate dehydrogenase (Ldh), and phosphoglueonate dehydrogenase (Pgd)), which appear to be species-diagnostic for P. glaucus, P. canadensis, and P. appalachiensis (Hagen et al. 1991; Scriber & Ording 2005; Scriber et al. 2008). The genetic incompatibilities of some recombinant hybrid genotypes appear to involve the Haldane effect in the heterogametic female pupae (Hagen & Scriber 1995). The delayed emergence of recombinant hybrid swarms (P. appalachiensis: Pavulaan & Wright 2002; Scriber & Ording 2005) may be controlled by the Z-linked obligate diapause gene (od+; Scriber et al. 2008) or a closely linked post-diapause developmental rate controlling gene on this Z-chromosome, such as identified for the delayed mid-July flight of European corn borer (Dopman et al. 2005). The tpi allozyme locus appears to correspond closely with this factor in the corn borer, and we have determined that it is also Z- linked in P. glaucus/P. canadensis (Hagen & Scriber 1989; Scriber 1994). Females of P. glaucus that are intermediate in their expression of melanism (with different degrees of “dusted” or “cinnamon” appearance; Fig 18) can be a result of rearing and/or pupal eelosion at higher temperatures (Ritland 1986). Such intermediate melanism may reflect variable penetrance, suggesting that the black melanin is superimposed over the yellow papilioehrome (Koch et al. 1998), which may appear brown in color (Ffrench-Constant & Koch 2003). However, we have also seen such intermediate melanism in hybrid daughters of dark females mated to P. multicaudatus (Fig 9), to P. rutulus (Scriber et al. 1995), and to P. canadensis (from Dane Co. WI, not figured here). Most P. canadensis males have X-linked suppressors (s+) that completely prevent the expression of melanism in hybrid daughters of dark mothers (Scriber et al. 1996). Here we report the partial expression of melanism (“cinnamon" with striations) in daughters of a primary F-l hybrid pairing between a dark P. glaucus mother mated with a male P. canadensis from central Michigan (in Isabella Co. on the cooler side of the historical hybrid zone; see below). These are not color mosaics as described earlier. These “cinnamon” intermediates of this Michigan P. canadensis father and dark P. glaucus mother, reared and emerged at 22° C, were also not due to high temperatures (28-30° C) as in Fig. 18 (see Ritland 1986). This result suggests that the wild Isabella Co. male parent of this pairing (#20011) had at least one Z-linked enabler (s ), which permitted expression of the Y-linked dark gene (b+). The introgression of the X-linked enabler has also been documented in daughters of another Isabella Co. P. canadensis male mated to a dark P. glaucus mother (#20015; Scriber et al. in prep.) and 44 Journal of the Lepidopterists’ Society Figs. 14-15. 14. A gyriandromorph or sexual mosaic which appears to be more than 50% female (from brood #4196) from a lab cross of a dark morph P. glaucus female (Champaign Co., Illinois) mated to a hybrid male (mother of which was the daughter of a dark morph P. glaucus female collected in Georgia and whose father was a P. canadensis collected in Lincoln Co., Wisconsin, (a) dorsal, (b) ventral. 15. A female color mosaic (from brood #3770) from a dark morph P. glaucus female (the daughter of a wild dark morph P. glaucus female, #3104, collected in Athens Co., Georgia) mated to a wild P. canadensis male (collected in Tompkins Co., New York), (a) dorsal, (b) ventral. Figs. 16-18. 16. A field-reared P. glaucus from Lancaster Co. PA 2004 (pupa from Win. Houtz). 17. A specimen from David and Marc Perlman (photo provided, pers. comm, to JMS). 18. An intermediate “dusty” or “cinnamon” type reared from Dane Co., Wisconsin at high temperatures (28-30° C). Volume 63, Number 1 45 also in daughters of several males of the hybrid swarm late flight at the New York/Vermont border where no dark females (with the W-linked b+ gene for melanism; Seriber et al 1996) have ever been collected (the nearest collection of a dark female is roughly 150 miles south; Seriber et al. in prep). The Z-linked enabler has also been detected in males captured from the hybrid swarm of late flight individuals in P. appalachiensis populations from West Virginia, but here, on the warmer side of the hybrid zone (Seriber et al. 2009b), the W-linked gene for melanism (b+) does also occur as evidenced by recombinant hybrid-like dark females (Pavulaan & Wright 2004; Seriber & Ording 2005; Seriber et al. 2009a, b). Individual species diagnostic traits moving northward across the historical hybrid zone include: 1) autosomal tulip tree detoxification (Seriber 1986, 2002; Seriber et al. 1999), 2) several wing color traits (Luebke et al. 1988), 3) some X-linked traits such as the allozyme allele Pgd-100 (Hagen & Seriber 1989; Hagen et al. 1991; Seriber et al. 2008), and 4) opposition preferences (Seriber et al. 1991; Mercader & Seriber 2007; Mercader et al. 2009). However, other X-linked species diagnostic traits (including Ldh-100 allozyme alleles and bivoltine potential (ock); Rockey et al. 1987) as well as the Y-linked dark gene (b+) and certain mitochondrial DNA alleles have not moved northward across the historical hybrid zone (Stump et al. 2003; Seriber et al. 2008). The divergent selection pressures on different parts of the X-chromosome (and differential movement of these parts) in the genomes of both P. glaucus and P. canadensis may be historically old (Putnam et al. 2007). However, recent origins of interspecific introgression and recombination combined with strong divergent selection on different regions of the X-chromosome may have been catalyzed by recent climate warming, possibly leading to the Mountain Swallowtail (potential hybrid species), P. appalachiensis (Pavulaan & Wright 2002; Seriber & Ording 2005). We will continue to investigate the linkage relationships of the dark morph enabler gene (s-) with other parts of the X-chromosome and how it may be involved in the easily detected sexual color mosaics. Acknowledgements This research was supported in part by the College of Natural Science, and the Michigan Agricultural Experiment Station (Project MICL #01644), NSF grants (DEB 9220122, DEB 9981608, DEB 0716683), and the Ray and Bernice Hutson En¬ dowment of the Department of Entomology. 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Received for publication 26 February 2008; revised and accepted 9 December 2008. 48 General Notes Journal of the Lepidopterists’ Society Journal of the Lepidopterists’ Society 63(1), 2009, 48-51 PITHING AND MINING BY A PUNKIE: THE UNUSUAL FEEDING STRATEGIES OF NOLA CILICOIDES (GROTE, 1873) (NOCTUIDAE: NOLINAE) Additional key words: Lysirnachia ciliata, leafmining, plant defense, hypertrophied tactile setae, leaf shelter Herbivorous insects have evolved innumerable strategies for coping with plant defenses. Among these are efforts made by insects to incapacitate a plants ability to transport defensive substances through its vascular or lactifer systems to sites of feeding injury. One means of doing such is to trench, vein cut, or otherwise breach the delivery systems of the host so that milky latexes, resins, and gums cannot be transported to the feeding site (Dussourd & Eisner 1987; Dussourd 1993). Another strategy is to girdle the host, either as an adult (e.g., ovipositing females) or as a larva (Forcella 1982; Dussourd 1993; Hanks 1999; Albanese et al. 2007). A third option is mass assault: bark beetles are a classic example. Adults attack host trees in such numbers that individuals under siege cannot individually “pitch out" all the invading beetles (Wood 1982). Here we report an example of a fourth option available to herbivores for thwarting a host’s defensive system: the hollowing (pithing) of non-woody stem tissues. In early July 2007 we found young (approximately 3rd instar) larvae of Nola cilicoides (Grote) (Noctuidae: Nolinae) pithing upper stems of fringed loosestrife (. Lysirnachia ciliata L.) (Primulaceae). Larvae tunneled into the side of new shoots, removed pith and associated vascular tissues, backed out, then crawled further up the compromised stem to feed on flowers and young leaves. Nola cilicoides (Fig. 1) occurs from southern Canada to northern Florida and west to Arkansas (Forbes I960; Handheld 1999; Ferge & Balogh 2000; Coveil 2005). In the Northeast the moth is local and uncommon in swamps, floodplains, wet meadows, damp woods, and other mesic to wet community types. Dyar (1899) did not provide life history information for N. cilicoides in his study of the early stages of Nearctic nolines. McCabe, in Rings et al. (1992), reported that the larva of N. cilicoides feeds on new leaves and flowers of fringed loosestrife ( Lysirnachia ciliata). Collection dates for adults in Connecticut and New York normally range from mid June into July, with the occasional specimen taken as early as late May or as late as August and September — we are unsure of the number of generations. In Georgia there are least three full broods (James Adams pers. comm.). Study Site and Methods. Observations were made on a colony of N. cilicoides located in Lebanon, Connecticut along a powerline cut south of Burnham Road and north of Route 87 (N41° 39’ 15.5” W72° 14’ 07.6”). Rearing was done in plastic bags and 40 dram plastic vials. Laivae were reared at 21-23° C with photoperiods set to match those outdoors in central Connecticut. Larval and adult vouchers and digital images have been deposited at the University of Connecticut. Results. On 6 and 11 July, 2007, young laivae of N. cilicoides were found feeding on the shoots of fringed loosestrife plants that were coming into flower. Laivae were located by surveying for loosestrife plants with flagging shoots. Each flagged shoot had a small entry hole and small area of cored pith, 5-10 centimeters below the shoot apex (Figs. 2, 4). On each flagged plant, we located 1 or 2 approximately 3rd instar N. cilicioides feeding externally (Fig. 3), distal from the wound, on floral buds, flowers, and new leaves. Laivae showed a preference for buds and flowers (Figs. 6, 7) over leaves. Some larvae tied together adjacent leaves and flowers, forming a weak shelter, but silk deposition was sparse and their shelters were poorly defined. Laival densities were low; only five caterpillars were found on roughly 200 plus fringed loosestrife stems that were searched in early July. In the vicinity of the colony and at other locations in New London and Tolland counties, we searched for, but did not find, laivae on either swamp loosestrife [L. terrestris (L.) Britton, Stems, & Poggenb.j or whorled loosestrife (L. quadrifolia L.). On 9 and 19 August, 2007, we returned to the same stand of fringed loosestrife in Lebanon, ostensibly to look for cocoons. While we failed to find cocoons on more than 60 checked stems, and any evidence on newly bored shoots, we discovered something surprising: young Nola cilicoides larvae, 2-3 mm in length mining leaves of fringed loosestrife. Laivae had entered through the lower epidermis, often adjacent to a vein, and then tunneled into the lamina, removing parenchymatic tissues, leaving a full depth blotch mine (Fig. 5). All but one mine had a single entrance/exit hole on the lower leaf surface; the one exception had three entry holes. Dark green to black frass collected in the Volume 63, Number 1 49 Figs. 1—9. Nola cilicoides (Grote), all Lebanon, New London Co., Connecticut except adult from Whitefield, New Hampshire. 1. Adult female. 2. Shoot damaged by single larva (arrows point to entry points). Note wilted section of stem above bore hole. 3. Middle instars feeding on young leaves. 4. Detail of 2. 5. Occupied leafmine. 6. Last instar “coring” flower bud with head fully in¬ serted. 7. Last instar feeding on remains of what was previously an open flower. 8. Last instar. While not fully visible, SD setae ex¬ tend close to all four edges of the image (see text). 9. Cocoon under construction, upper end with abdominal terminus visible. 10. Finished cocoon. Note discolored area of stem above and below cocoon where bark strips were removed. 50 Journal of the Lepidopterists’ Society mine, i.e., little had been expelled. The smallest mine located, was only a few mm in diameter and contained the cadaver of a larva 1.25 mm in length. In the wild, we found approximately 3rd instar larvae grazing on patches of parenchyma from leaf undersides — scraping away small patches of leaf tissue without feeding through to the upper leaf surface — as late as mid September, indicating that at least some N. cilicoides larvae vacate their mines to feed externally, before the onset of winter. Beginning as early as late August, our captive (lab- reared larvae) spun small, circular to oval, cocoons between overlapping leaves. Cocooned larvae were foreshortened, only 2. 6-2. 8 mm in length, and evidently had entered a non-feeding diapause in which they would pass the winter. [Meganola spodia Franclemont, an oak-feeding noline, also overwinters as a partially grown larva (Butler 1989).] The full grown larva is 10-12 mm in length (n=8). It is typical for a noline: the prolegs are absent on A3 and secondary setae are present, grouped onto five (dorsal, subdorsal, lateral, subventral, and ventral) warts. Both the dorsal and subdorsal warts have an incomplete black ring around their bases (Figs. 6, 7). The ground color is dirty yellow-orange above, more yellow laterad, and mostly unpigmented below the subspiracular stripe; the creamy to pale orange middorsal stripe is weak over Tl. Snbdorsal warts on T1-T3 and A7-A9 bear one exceptionally long seta that extends well beyond the body (see below). The head, prominently marked with black (Fig. 7), is drawn into the prothorax at rest. |nly-collected, lab-reared larvae formed their pupal cocoons along a stem (Figs. 9, 10), but extensive searches of plants in the wild, including those with wilted tips, did not yield any cocoons. The central chamber of cocoons measured roughly 3 mm x 10 mm. The lower end of the cocoons had an additional 5 nun- long ‘‘tail’' that tapered downward; the upper end was higher in profile (n=3) (Figs. 9, 10). Like other nolines, numerous strips of stem epidermis "bark" were collected and then woven into the wall of the cocoon. Our caterpillars positioned more than 40 narrow strips of bark, removed from adjacent areas of the stem, into their cocoon walls. Eclosion occurred 1-2 weeks after pupation. Discussion. The greatly elongated setae that extend from the SD warts at either end of the body of Nola larvae (Fig. 8 and caption) presumably sen e as an "early warning" system for the larvae. Those issuing from T3 are the longest: in one individual, the longest SD seta extended forward 5.8 mm on a larva only 10 mm in overall length. The rearward-projecting SD seta on A9 of this same caterpillar measured 3.6 mm. In addition. the subdorsal warts on these segments are oriented in such a way (in living larvae) that the hypertrophied setae are directed well forward on the thoracic and rearward on the abdominal segments. Analogous long dorsal or subdorsal setae also have been reported from ehoreutids (e.g., Brenthia ), pyralids (e.g., Monoloxis), as well as other noctuids (e.g., Aboblemma Walker) (Aiello & Solis 2003; Rota 2003, 2008; Wagner et al. 2008; Rota & Wagner 2008). Although Lijsimcichia ciliata has not been investigated, the tissues of other Lijsimachia species have been found to contain a variety of bioactive compounds including flavonoids, triterpenoid saponins, and phenolic acids (Chang et al. 2004). To what degree these are transportable or serve in defense remains unstudied. Our thesis presupposes that N. cilicoides larvae tunnel out the pith of their hostplant to incapacitate its vascular tissues — we acknowledge that the pithing behaviors reported here may have little to do with circumventing the plant’s ability to translocate antiherbivory substances. Perhaps pithing and the subsequent flagging help the larva to construct its leaf shelter, prevent subsequent growth and maturation of the shoot, enhance survivorship from attacks by natural enemies, or in some other way improve or enhance larval performance or survival. Regardless, the stereotypical behavior is commonly expressed, and stem boring and bud coring represent unique traits for Nearetic nolines, which tend to be rather unspecialized leaf feeders (Forbes I960; Wagner 2005; Wagner et al. 2010). As noted above, larvae preferentially sought out flower buds and flowers over leaf tissues. A preference for reproductive tissues has been reported for other nolines. Nola sorghiella Riley feeds on the seed heads of sorghum and various other grasses and appears to be completely tied to reproductive tissues (our wild- collected larvae failed when offered only leaves). In the Old World, spiny bollworms of the genus Earias are major pests of cotton flowers (Pinhey 1975; Kitching & Rawlins 1998). More smprising was the finding that N. cilicioides larvae engage in leaf mining. Leafmining is rare among noctuids. In the East only a handful of apaineines mine, and most of these only for the first one or two instars (■Wagner et al. 2010; Eric Quinter pers. com¬ munication). It is exceptional for a "leafminer" to bear secondary setae; more commonly, setae tend to be reduced or absent among leafmining clades (cf. Stehr 1987). We did not observe leafmining behaviors prior to our August visits. Interestingly, life histories with stem mining in spring generations and leafmining in late summer generations have been reported for a number Volume 63, Number 1 51 of European Momphidae (Koster 2002). In North America, Mompha solomonella Adamski & Brown, tunnels in shoots of Cephalanthus accident alio L. (Rubiaceae) in the spring, but second and subsequent generations mine leaves (Wagner et al. 2004). Much remains to be elucidated about the biology Nola cilicoides. As noted, the selective advantages of pithing and ultimate explanation for this exceptional behavior have yet to be determined. Both observational and controlled studies are needed to better understand the roles of the hypertrophied tactile setae of Nola and other lepidopterans. We are also in a quandary about the phenology and broodism of N. cilicoides in New England. We were only able document the existence of a single brood — one that produced its pithing larvae in early July. Either we missed a small spring brood, or the partially grown larvae that spun overwintering cocoons in August, waited until early July of the following season to break their diapause. Acknowledgments Tim McCabe, as so many times before, guided DLW’s initial efforts. Deane Bowers, Dave Dussourd, Laura McLoud, Jessica McShane, April Rodd, and Bo Sullivan read drafts of the paper. An anonymous reviewer offered many helpful suggestions. An¬ drea Farr assembled the plates. Both authors were supported on a grant from the CT-Department of Environmental Protec¬ tion Wildlife Division through the State Wildlife Grants Pro¬ gram; additional support was provided by Northeast Utilities. Literature Cited Aiello, A. & M. A Solis. 2003. Defense mechanisms in Pyralidae and Choreutidae: fecal stalactites and escape holes, with remarks about cocoons, camouflage, and aposematism. J. Lepid. Soc. 57: 168-175. Albanese, G., M. W. Nelson, P. D. Vickery, & P. R. Sievert. 2007. Larval feeding behavior and ant association in frosted elfin, Cal- lophrys irus (Lycaenidae). J. Lepid. Soc. 61: 61-66. Butler, L. 1989. Observations on Meganola spodia Franclemont (Lepidoptera: Noctuidae) with a description of the mature larva. Proc. Entomol. Soc. Wash. 91: 615-619. Chang.-H.T., W. L. Kong, & P. F. Tu. 2004. [Chemical and pharma¬ cological advances of study on Lijsimachia). Zhongguo Zhong Yao Za Zhi. 29: 295-298. Covell, C. V., Jr. 2005. A field guide to the moths of eastern North America. 2nd Ed. Virginia Museum of Natural History, Mar¬ tinsville, VA. 496 pp. Dussourd, D. E. 1993. Foraging with finesse: caterpillar adaptations for circumventing plant defenses. Pp. 92-131. In N. E. Stamp & T. M. Casey (eds.), Caterpillars: Ecological and evolutionary con¬ straints on foraging. Chapman and Hall, New York. - & T. Eisner. 1987. Vein-cutting behavior: Insect counterploy to the latex defense of plants. Science 237: 898-901. Dyar, H G. 1899. On the larvae of North American Nolidae with de¬ scriptions of new species. Canad. Entomol. 31: 61-64. Ferge, L. A. & G. J. Balogh. 2000. Checklist of Wisconsin moths. Superfamilies Drepanoidea, Geometroidea, Mimallonoidea, Bombycoidea, Sphingoidea, and Noctuoidea. Milwaukee Public Museum Contributions in Biology and Geology. Number 93. Milwaukee, Wisconsin. 55 pp. Forbes, W. T. M. 1960. The Lepidoptera of New York and neighbor¬ ing states. IV. Agaristidae through Nymphalidae, including but¬ terflies. Memoir 371. Cornell University Agricultural Experiment Station, Ithaca, New York. 188 pp. Forcella, F. 1982. Why twig-girdling beetles girdle twigs. Naturwis- senschaften 69: 398-400. Handheld, L. 1999. Le guide des papillons du Quebec. Broquet Inc., Boucherville, Quebec. 536 pp + 123 plates. Hanks, L. M. 1999. Influence of the larval host plant on reproductive strategies of cerambycid beetles. Ann. Rev. Entomol. 44: 483-505. Kitching, I. J. & J. E. Rawlins. 1998. The Noctuoidea. Pp. 355-401. In Kristensen, N. P. (ed.), Lepidoptera, moths and butterflies. Volume 1: Evolution, systematics, and biogeography. Handbook for zoology. Volume IV. Arthropoda: Insecta. Walter de Gruyter, Berlin, Germany. Koster, J. C. 2002. Momphidae. Pp. 224-254. In Emmet, A. M. & J. R. Langmaid (eds.). The moths and butterflies of Great Britain and Ireland. Harley, Martins, England. Pinhey, E. C. G. 1975. Moths of southern Africa. Tafelberg Publ.. Cape Town, South Africa. 274 pp. Rings, R. W., E. H. Metzler, F. J. Arnold. & D. H. Harris. 1992. The owlet moths of Ohio (Order Lepidoptera: Family Noctu¬ idae). Ohio Biological Survey Bulletin. New Series. Vol. 9, Col¬ lege of Biological Sciences, Ohio State University, Columbus, Ohio. 219 pp. Rota, J. 2003. Choreutidae (Lepidoptera) of La Selva Biological Sta¬ tion (Costa Rica): Taxonomy, ultrastructure of immature stages, and life history. Master’s Thesis, University' of Connecticut, Storrs, Connecticut. 159 pp. - . 2008. Immature stages of metalmark moths from the genus Brenthia Clemens (Lepidoptera: Choreutidae): moqrhology and Life History' Notes. J. Lepid. Soc. 62: 121-129. - & D. L. Wagner. 2008. Wormholes, sensory nets, and hyper¬ trophied tactile setae: the extraordinary defence strategies of Brenthia caterpillars. Animal Behaviour. In press. Stehr, F.W., ed. 1987. Immature Insects. Volume 1. Kendall/Hunt Publ. Co., Dubuque, Iowa. 754 pp. Wagner, D. L. 2005. Caterpillars of eastern North America: A guide to identification and natural history. Princeton University' Press, Princeton, New Jersey. 512 pp. - . D. Adamski, & R. L. Brown. 2004. A new species of Mompha (Lepidoptera: Coleophoridae: Momphinae) from buttonbush (Cephalanthus occidentals L.) with descriptions of the early stages. Proc. Entomol. Soc. Wash. 106: 1-18. - , J. Rota, & T. L. McCabe. 2008. Larva of Abablemma Nye (Lep¬ idoptera: Noctuidae: Hvpenodinae) with notes on lichenivory and algivory in Macrolepidoptera. Ann. Entomol. Soc. Amer. 101: 40-52.' - . D. F. Schweitzer, J. B. Sullivan & R.C. Reardon. 2010. Owlet cateqiillars of eastern North American (Lepidoptera: Noc¬ tuidae). Princeton Univ. Press. In review. Wood, D. L. 1982. The role of pheromones, kairomones, and al¬ lomones in the host selection and colonization behavior of bark beetles. Ann. Rev. Entomol. 27: 411-446. David L. Wagner, Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, Connecticut 06269, USA, e-mail: clavid. ivagner @uconn.edu and Bryan Connolly, Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, Connecticut 06269, USA. Received for publication 6 Nov 2007; revised and accepted 28 May 2008 52 Obituary Journal of the Lepidopterists’ Society Journal of the Lepidopterists’ Society 63(1), 2009, 52-57 EUGENE G. MUNROE (8 September 1919-31 May 2008) I recently wrote an obituary for Eugene M unroe for the Bulletin of the Entomogical Society of Canada [2008. 40 (1): 142-146], Unfortunately there was not enough space for his list of publications. In 2000, I wrote a detailed account of Genes life and professional career for a Festschrift in Tropical Lepidoptera [11: 1-6]; the list of publications was incomplete. Gene’s impressive publication list is published in its entirety here. See also the 2008 Pyraloid Planet [2:2] for reminisces by colleagues. Michael Shaffer, The Natural History Museum, London, generously reviewed this version of Gene’s publications. To learn more about Gene’s life, accomplishments, enthusiasm for research, and love of Lepidoptera, refer to the publications mentioned above. Here, I would like to briefly share some historical information regarding the Lepidopterists’ Society and Gene. He was a Charter and Honorary Life Member of the Lepidopterists1 Society. In preparation for the First Annual Lepidopterists’ Society Meeting, Dec. 29th-30th, 1950, held at the American Museum of Natural History, New York, the President, ]. H. McDunnough, appointed three committees. Program, Local Arrangements and Nominating. Gene was appointed member of the Nominating Committee. Also, at the first meeting Gene participated (1951e) in a symposium entitled “Geographic subspeciation in the Lepidoptera” where Charles Remington also presented the introduction. Gene was involved in the first annual Season Summary as a coordinator (1947b, 1949b, c, 1951b) for Area 7, the Northeast, and later as editor- (1951d) of the Season Summary. He served as President of the society in 1958. In 2004 Gene received the Karl Jordan medal from the Lepidopterists’ Society. Gene Munroe passed away 31 May 2008 at the age of 89 in Ontario, Canada. List of Publications Munroe, E. G. 1942. The Caribbean races of Anartia jatrophae Jo¬ hansson (Lepidoptera: Nymphalidae). American Museum Novi- tates 1179: 1-4. _ . 1947a. Four new Pieridae from the West Indies. American Museum Novitates 1362: 1-5. _ (coordinator). 1947b. The field summary of Lepidoptera - 1947 season - 7, Northeast-Maryland north to Quebec and Newfound¬ land. Lepidopterists' News 1: 95. _ . 1948a. The geographical distribution of butterflies in the West Indies. Ph.D. thesis. Cornell University, Ithaca, New York. _ . 1948b. Further North American records of Acentropus niveus (Lepidoptera: Pyralidae). Canadian Entomologist 79: 120. _ . 1949a. Some remarks on the genus concept in Rhopalocera. Lepidopterists’ News 3: 3-4. _ (coordinator). 1949b. The field season summary of North American Lepidoptera for 1949, Area 7, Northeast-Maryland north to southern Quebec. Lepidopterists' News 3: 97-101. _ (coordinator). 1949c. Field season summary of North American Lepidoptera for 1948, Area 7, Northeast. Lepidopterists’ News 8 (supplement): 9-11. _ . 1949d. An unnoticed character in the Satumioidea. Entomo¬ logical News 60: 60-65. _ . 1949e. Some remarks on the orithija group of the genus Junonia (Lepidoptera: Nymphalidae). Entomologist 82: 157-158. _ . 1949f. A new genus of Nymphalidae and its affinities (Lepi¬ doptera: Rhopalocera). Journal of the New York Entomological Society 57: 67-78. _ . 1949g. Notes on fish of the interior of the Labrador Peninsula. Arctic 2: 165-173. _ . 1950a. The occurrence of a butterfly in the Pribilof Islands. Lepidopterists’ News 4: 44. _ . 1950b. The generic positions of some North American Lepi¬ doptera commonly referred to Pyrausta Schrank. (Lepidoptera: Pyralidae). Canadian Entomologist 82: 217-231. _ . 1950c. Subventral tubercles of Saturnoid larvae — a supple¬ mentary note. Entomological News 61: 39-41. _ . 1950d. The dina group of the genus Eurema in the West Indies (Lepidoptera: Pieridae). Journal of the New York Entomological Society 58: 172-191. _ . 1950e. The systematic^ of Calisto (Lepidoptera: Satyrinae), with remarks on the evolutionary and zoogeographic significance of the genus. Journal of the New York Entomological Society 58: 211-240. _ . 1951a. Field notes on the butterflies of Knob Lake, northern Quebec. Lepidopterists’ News 5: 7-9. _ (coordinator). 1951b. Field season summary of North American Lepidoptera for 1950, Area 7, Northeast-Delaware & Pennsylva¬ nia north to southern Quebec. Lepidopterists’ News 4: 102-106. _ . 1951c. The De Rabie paintings of Lepidoptera in the Blacker Library of Zoology, McGill University, with notes on the butter¬ flies represented therein. Lepidopterists' News 5: 55-57. _ (ed. ). 195 Id. The field season summary of North American Lepidoptera for 1951. Lepidopterists’ News 5: 81-110. _ . 1951e. V. Subspeciation in the Microlepidoptera. Lepidopter¬ ists’ News 5: 29-31. _ . 1951f. A previously unrecognized species of Nymphula (Lepi¬ doptera: Pyralidae). Canadian Entomologist 83: 20-23. _ . 1951g. The identity and generic position of Chauliodes dis- junctus Walker (Megaloptera: Corydalidae). Canadian Entomol¬ ogist 83: 33-36. _ . 1 95 1 h . Pest Trichoptera at Fort Erie, Ontario. Canadian Ento¬ mologist 83: 69-72. _ . 1 95 1 i . North American Pvraustinae: notes and descriptions (Lepidoptera: Pyralidae). Canadian Entomologist 83: 161-169. _ . 195lj. The geographic variations of Dasyuris polata (Duponchel) in North America. (Lepidoptera: Geometridae). Canadian Entomologist 83: 290-294. _ . 1951k. The genus Junonia in the West Indies (Lepidoptera: Nymphalidae). American Museum Novitates 1498: 1—16. Miller, M. J. & E. G. Munroe. 1951. Schistosome dermatitis in Quebec. Canadian Medical Association Journal 65: 571. Munroe, E. G. 1952a. The illibalis group of the genus Palpita Hiib- ner (Lepidoptera: Pyralidae). Canadian Entomologist 84: 43-55. Volume 63, Number 1 53 _ . 1952b. Support for the proposals submitted by Mr John G. Franclemont in regard to the generic name “Phalaena” Linnaeus, 1758, and the names published by Linnaeus for groups of that genus (Class Insecta, Order Lepidoptera). Bulletin of Zoological Nomenclature 6: 317. _ . 1952c. Support for the proposed use of the plenary powers to validate the trivial name “adippe” (as published in the combina¬ tion “ Papilio adippe”) as from Denis and Schiffermiiller, 1775 (Class Insecta. Order Lepidoptera). Bulletin of Zoological Nomenclature 9: 136. _ . 1952d. Comment on Mr. C. P. dos Passos proposal relating to the trivial name “ plexippus ” Linnaeus, 1758, as published in the combination “ Papilio pexippus” (Class Insecta, Order Lepi¬ doptera). Bulletin of Zoological Nomenclature 9: 139-140. _ . 1953a. The Karanasa butterflies — a critical review. Lepi- dopterists’ News 7: 16- 23. _ (ed.). 1953b. Field season summary of North American Lepi¬ doptera for 1953. Lepidopterists' News 7: 66-118. _ . 1953b. Lines of communication. Division of Entomology Newsletter 34(4): 1-2. _ . 1953c. Chaidiodes disjunctus Walker: a correction with de¬ scriptions of a new species and new genus (Megaloptera: Cory- dalidae). Canadian Entomologist 85: 190-192. _ . 1953d. Report on field work at Knob Lake, Quebec, 1948. In Entomological Research in Northern Canada. Progress Report, 1948. Part 1. The Northern Insect Survey. Appendix D. Ottawa: Canada Dept. Natl. Def.. Def. Res. Bd. (Rep. No. DR18): 1-34. _ . 1953e. Some aspects of butterfly distribution in the West In¬ dies. (Abstract of invitation paper). Proceedings of the Entomo¬ logical Society of Washington 55: 112. _. 1953f. The size of island faunas. (Abstract). Proceedings of the 7th Pacific Science Congress, Zoology 4: 52-53. _ . 1953g. The phylogeny of the Papilionidae. Proceedings of the 7th Pacific Science Congress, Zoology 4: 83-87. _ . 1954a. Visit to Europe, 1953. Entomology Division Newsletter 32(3): 4. _ . 1954b. The eastern North American subspecies of the bar¬ berry geometrid (Lepidoptera). Canadian Entomologist 86: 282-283. _. 1954c. A new Phlyctaenia, with notes on two related species (Lepidoptera: Pvralidae). Canadian Entomologist 86: 428-431. _ . 1954d. A new pyralid of Palaearc-tic affinities from Colorado (Lepidoptera). Canadian Entomologist 86: 499-500. _ . 1955a. The genus Epipagis Hiibner, nec Hampson, in North America. (Lepidoptera: Pvralidae). Canadian Entomologist 87: 249-252. _. 1955b. Figures of type specimens. Lepidopterists’ News 9: 140-141. _ . 1955c. Karl Jordan, the lepidopterist. Transactions Royal En¬ tomological Society London 107 (Jordan Jubilee volume): 69-75. _. 1956a. The genus Orenaia in North America, with the descrip¬ tion of a new species (Lepidoptera: Pvralidae). Canadian Ento¬ mologist 88: 74-78. _ . 1956b. Geshna primordialis Dyar. with descriptions of two new genera and two new subspecies (Lepidoptera: Pvralidae). Canadian Entomologist 88: 126-128. _ . 1956c. Restriction and revision of the genus Diastictis Hiibner (Lepidoptera: Pvralidae). Canadian Entomologist 88: 208-228. _ . 1956d. Canada as an environment for insect life. Canadian En¬ tomologist 88: 372-476. _ . 1956e. The North American species of Diatlirausta Lederer (Lepidoptera: Pvralidae). Canadian Entomologist 88: 579-583. _. 1956f. Pvralides recoltes a l'Tle de la Guadeloupe par M. L. Berland. Revue Franyaise d'Entomologie 23: 121-127. _ . 1957a. North American subspecies of Pyrausta unifascialis (Packard) (Lepidoptera: Pyralidae). Canadian Entomologist 89: 91-94. _ . 1957b. North American components of the genus Diasemia of authors (Lepidoptera: Pvralidae). Canadian Entomologist 89: 164-170. _ . 1957c. Comparison of closely related faunas. Science 126: 437-439. _ . 1957d. Comment on Dr. Elwood C. Zimmerman's proposal to add to the "Official list of generic names in zoology" the names “ Oeobia ” Hiibner, [1825], and “Hellula” Guenee, 1845 (Class In¬ secta, Order Lepidoptera). Bulletin of Zoological Nomenclature 13: 183-185. _ . 1957e. Proposed suppression under the plenary powers of the generic name Oeobia Hiibner, [1825] (Class Insecta, Order Lep¬ idoptera) for the purposes of the law of priority but not for those of the law of homonymy. Bulletin of Zoological Nomenclature 13: 186-189. _ . 19571. Northward extension in recent years of the range of the lettered sphinx moth. Canadian Field Naturalist 71: 37. _ . 1958a. Collecting insects in Pacific islands and Southeast Asia. Division of Entomology Newsletter 36(5): 1-2. _ . 1958b. A new subspecies of Sparagmia gigantalis Guenee (Lepidoptera: Pvralidae). Canadian Entomologist 90: 127-128. _ . 1958c. Pycnarmon jaguaralis (Guenee) and an allied species (Lepidoptera: Pvralidae). Canadian Entomologist 90: 234—235. _ . 1958d. Far-eastern Pvralidae (Lepidoptera). Canadian Ento¬ mologist 90: 249-254. _ . 195Se. Some species of Polygrammodes Guenee (Lepidoptera: Pyralidae). Canadian Entomologist 90: 257-274. _ . 1958f. A revision of the genus Epicorsia Hiibner (Lepidoptera: Pyralidae). Canadian Entomologist 90: 293-301. _. 1958g. Catalogue of Lederer types of Pvralidae in the British Museum (Natural History) wfith selections of Iectotvpes (Lepi¬ doptera). Canadian Entomologist 90: 510-517. _. 1958h. Hampson's Schoenobiinae (Lepidoptera: Pyralidae). Proceedings of the 10th International Congress of Entomology' 1: 301-302. _ . 1958i. Systematics as a tool in ecology. Proceedings of the 10th International Congress of Entomology, Ecology 2: 663-667. _ . 1958j. The geographic distribution of the Scopariinae (Lepi¬ doptera: Pyralidae). Proceedings of the 10th International Con¬ gress of Entomology, Geographical Distribution 1: 831-837. _ . 1958k. Ranges of representative insects, ticks and spiders. At¬ las of Canada, Plate 37, 24 maps. _, A. Diakonoff, & E. L. Martin. 1958. Catalogue of Snellen's types of Pyralidae, with selections of Iectotvpes. Tijdschrift voor Entomologie 101: 65-88. _ . 1959a. Canadian species of Dioryctria Zeller (Lepidoptera: Pyralidae). Canadian Entomologist 91: 65-72. _ . 1959b. New Pyralidae from the Papuan Region (Lepidoptera). Canadian Entomologist 91: 102-112. _ . 1959c. Pvralidae from the collection of the California Academy of Sciences (Lepidoptera). Canadian Entomologist 91: 161-167. _ . 1959d. The phlogosaria complex of the genus Plagodis (Lepi¬ doptera: Geometridae). Canadian Entomologist 91: 193-208. _ . 1959e. New genera and species of Pyralidae (Lepidoptera). Canadian Entomologist 91: 359-371. _ . 1959f. Four new species of Evergestis (Lepidoptera: Pyrali¬ dae). Canadian Entomologist 91: 406-411. _. 1959g. Revision of the genus Linosta Moschler (Lepidoptera: Pyralidae) with characterization of the subfamily Linostinae and a new subfamily. Canadian Entomologist 91: 485—488. _ . 1959h. New species and new subspecies of Palpita (Lepi¬ doptera: Pvralidae). Canadian Entomologist 91: 641-650. _ . 1959i. Pyralidae collected in lower California by Mr. J. Powell (Lepidoptera). Canadian Entomologist 91: 725-727. _. 1959j. The pyralid fauna of the Western United States. Presi¬ dential address to the Sixth Pacific Slope Meetings of the Lepi¬ dopterists’ Society, August 1959. Journal of the Lepidopterists’ Society 13: 241-244. _ . 1960a. New tropical Pyraustinae (Lepidoptera: Pyralidae). Canadian Entomologist 92: 164-173. _ . 1960b. A new genus of Pyralidae and its species (Lepidoptera). Canadian Entomologist 92: 188-192. _ . 1960c. New species of Polygrammodes and a related new 54 Journal of the Lepidopterists’ Society genus (Lepidoptera: Pvralidae). Canadian Entomologist 92: 279-284. _ . 1960d. The Meyrick types of Scopariinae (Lepidoptera: Pyral- idae) in the British Museum (Natural History), exclusive of Hawaiian species. Canadian Entomologist 92: 891-897. _ . 1960e. Liocoris , Lygus and ethics. Bulletin of the Brooklyn Entomological Society 55: 104-108. _ . 1960f. An assessment of the contribution of experimental tax¬ onomy to the classification of insects. Revue Canadienne de Bi- ologie 19: 293-319. _ . 1960g. Professional and amateur research in Lepidoptera. (President's Annual Address, 1959). Journal of the Lepidopterists’ Society 14: 1-4. _ & P. R. Ehrlich. 1960. Harmonization of concepts of higher classification of the Papilionidae. Journal of the Lepidopterists’ Society 14: 169-175. _ . 1961a. Geography and systematic research. Proceedings of the 9th Pacific Scientific Congress 19: 72-74. _ . 1961b. The classification of the Papilionidae (Lepidoptera). Memoirs of the Entomological Society of Canada 17: 1—51 . _ . 1961c. Synopsis of the North American Odontiinae, with de¬ scriptions of new genera and species (Lepidoptera: Pyralidae). Memoirs of the Entomological Society Canada 24: 1—93. _ . 1962. A new species of Linosta (Lepidoptera: Pyralidae). Cana¬ dian Entomologist 94: 922-923. _ . 1963a. The g ilvarius group of Aspilates Treitschke (Lepi¬ doptera: Geometridae). Canadian Entomologist 95: 259-287. _ . 1963b. Perspectives in biogeography. Canadian Entomologist 95: 299-308. _ . 1963c. A new species of Nepytia (Lepidoptera: Geometridae), of economic importance to Douglas fir in British Colombia. Canadian Entomologist 95: 407— 113. _ . 1963d. A new species of Tetralopha (Lepidoptera: Pyralidae) from Vaccinium. Canadian Entomologist 95: 677-679. _ . 1963e. New Neotropical Pyraustinae (Lepidoptera: Pyralidae). Canadian Entomologist 95: 704-712. _ . 1963f. Collecting insects in the Territory of Papua and New Guinea, 1957. Proceedings of the 9th Pacific Science Congress, Entomology 4: 32 — 41 . _ . 1963g. Systematic Entomology Survey Program of the Canada Dept, of Agriculture. Proceedings of the 9th Pacific Science Con¬ gress, Entomology 4: 42-43. _ . 1963h. Entomological research in Canada. Proceedings of the 9th Pacific Science Congress, Entomology 4: 132-135. _ . 1963i. Characteristics and history of the North American fauna: Lepidoptera. Proceedings of the 16th International Con¬ gress of Zoology 4: 21-27. _ . 1964a. Problems and trends in systematics. Canadian Ento¬ mologist 96: 367-377. _ . 1964b. Four new species of Cliniodes Guenee, with notes on the genus and some relatives and segregates (Lepidoptera: Pyral¬ idae). Canadian Entomologist 96: 529-538. _ . 1964c. New genera and species of Glaphyriinae (Lepidoptera: Pyralidae). Canadian Entomologist 96: 1261-1307. _ . 1964d. Some Neotropical genera resembling Epicorsia Hiib- ner (Lepidoptera: Pyralidae). Memoirs of the Entomological So¬ ciety of Canada 33: 3-76. _ . 1964e. Insects of Campbell Island. Lepidoptera: Pyralidae. Pa¬ cific Insects Monographs 7: 260-271. _ . 1964f. Biosystematics and dynamic ecology. Royal Ontario Mu¬ seum, University of Toronto Life Sciences Contributions 59: 1-17. _ . 1965. Zoogeography of insects and allied groups. Annual Re¬ view of Entomology 10: 325-344. _ . 1966. Revision of North American species of Udea Guenee (Lepidoptera: Pyralidae). Memoirs of the Entomological Society of Canada 49: 1-57. Wier, J. R. & E. G. Munroe. 1966. The interdisciplinary research. Agricultural Institute Revue. November-December: 9-13. Munroe, E. G. 1967a. Four new species of Udea from South America (Lepidoptera: Pyralidae). Canadian Entomologist 99: 392—397. _ . 1967b. A new species of Thliptoceras from Thailand, with notes on generic and specific synonymy and placement and with designations of lectotypes (Lepidoptera: Pyralidae). Canadian Entomologist 99: 721-727. _ . 1967c. A new genus and a new species of Pyraustinae (Lepi¬ doptera: Pyralidae). Canadian Entomologist 99: 1104-1110. _ . 1967d. Parapilocrocis citribasalis, a new genus and species of Pyraustinae (Lepidoptera: Pyralidae). Canadian Entomologist 99: 1156-1159. _ . 1967e. Titles. Nature 214: 1064. _ . 1968a. New Pvralidae (Lepidoptera) from the Papuan Region. II. Canadian Entomologist 100: 394—401. _ . 1968b. New species of Apilocrocis [Amsel], with a key to known species (Lepidoptera: Pyralidae). Canadian Entomologist 100: 441^448. _ . 1968c. Insects of Ontario: geographical distribution and post¬ glacial origin. Proceedings of the Entomological Society of On¬ tario 99: 43-50. _ & A. MUTUURA. 1968a. Contributions to a study of the Pyrausti¬ nae (Lepidoptera: Pyralidae) of temperate East Asia. I. Canadian Entomologist 100: 847-861. _ & _ . 1968b. Contributions to a study of the Pyraustinae (Lepidoptera: Pyralidae) of temperate East Asia. II. Canadian Entomologist 100: 861-868. _ & _ . 1968c. Contributions to a study of the Pyraustinae (Lepidoptera: Pyralidae) of temperate East Asia. III. Canadian Entomologist 100: 974-985. _ & _ . 1968d. Contributions to a study of the Pvraustinae (Lepidoptera: Pyralidae) of temperate East Asia. IV. Canadian Entomologist 100: 986-1001. _ . 1969a. Transfer of Cacographis colombiana [Schaus] from Midilinae to Nymphulinae, with description of Midilambia n. gen. (Lepidoptera: Pyralidae). Canadian Entomologist 101: 994-996. _ . 1969b. Scoparia macroplianes Meyrick, 1888 (Insects, Lepi¬ doptera, Pyralidae): request for suppression of neotype designa¬ tion following rediscovery of holotvpe. Bulletin of Zoological Nomenclature 26: 167-170. _ . 1969c. Insects of Ontario: geographical distribution and post¬ glacial origin. Proceedings of the Entomological Society of On¬ tario 99: 43-50. _ . 1969c!. Book review - Ecology and biogeography of high alti¬ tude insects, by M. S. Mani. Canadian Entomologist 101: 1007-1008. _ & A. Mutuura. 1969a. Contributions to a study of the Pyrausti¬ nae (Lepidoptera: Pyralidae) of temperate East Asia. V. Canadian Entomologist 101: 299-305. _ & _ . 1969b. Contributions to a study of the Pyraustinae (Lepidoptera: Pyralidae) of temperate East Asia. VI. Canadian Entomologist 101: 897-906. _ & _ . 1969c. Contributions to a study of the Pyraustinae (Lepidoptera: Pyralidae) of temperate East Asia. VII. Canadian Entomologist 101:1069-1077. _ & _ . 1969d. Contributions to a study of the Pyraustinae (Lepidoptera: Pvralidae) of temperate East Asia. VIII. Canadian Entomologist 101: 1239-1248. _ & P. D. McTaggart-Cowan. 1969. The science council of Canada. Agricultural Institute Review March-April 1969: 1820. Mutuura, A., E. G. Munroe, & D. A. Ross. 1969a. American species of Dioryctria Zeller (Lepidoptera: Pyralidae). I. Western Cana¬ dian species of the zimmermani group. Canadian Entomologist 101: 1009-1023. _ , _ , & _ . 1969b. American species of Dioryctria Zeller (Lepidoptera: Pvralidae). II. Western Canadian species of the baumhoferi and ponderosae groups. Canadian Entomologist 101: 1042-1047. Munroe, E. G. 1970a. Book review- A short history of the browntail moth, by William Curtis. Canadian Entomologist 102: 170. _ . 1970b. A new genus and three new species of Chrysauginae (Lepidoptera: Pvralidae). Canadian Entomologist 102: 414^120. Volume 63, Number 1 55 _ . 1970c. A short history of the brown-tail moth. Canadian Ento¬ mologist 102: 640. _ . 1970d. Types of Nymplnilinae, Scopariinae, Cybalomiinae, Odontiinae, Evergestinae, and Pyraustinae (Lepidoptera: Pyrali- dae) in the Zoological Institute, Academy of Sciences of the U.S.S.R., Leningrad, with selections of lectotvpes. Canadian En¬ tomologist 102: 1025-1035. _ . 1970e. Revision of the subfamily Midilinae (Lepidoptera: Pvralidae). Memoirs of the Entomological Society of Canada 74: 1-94. _ . 1970f. New species of Syllepis Poey ([Lepidoptera]: Pvralidae: Pyraustinae), with a key to known species. Journal of the Lepi- dopterists’ Society 24: 287-291. _ & A. MUTUURA. 1970a. Contributions to a study of the Pyrausti¬ nae (Lepidoptera: Pvralidae) of Temperate East Asia. IX. Cana¬ dian Entomologist 102: 294-304. _ & _ . 1970b. Contributions to a study of the Pyraustinae (Lepidoptera: Pvralidae) of Temperate East Asia. X. Canadian Entomologist 102: 1489-1507. Mutuura, A. & E. G. Munroe. 1970. Taxonomy and distribution of the European coni borer and allied species: genus Ostrinia [Hiibner] (Lepidoptera: Pvralidae). Memoirs of the Entomologi¬ cal Society of Canada 71: 1-112. Munroe, E. G. 1971a. Eudonia borealis (Tengstrom) and E. alpina (Curtis) (Lep., Pvralidae). Entomologist's Gazette 22: 96. _ . 1971b. A new name for Papilio zetes (Westwood, 1847), nec. Linnaeus, 1758 (Papilionidae). Journal of the Lepidopterists’ So¬ ciety 25: 142. _ . 1971c. Distribution and geographical differentiation of Marpe- sia eleuehea Hiibner (Nvmphalidae), with descriptions of two new subspecies. Journal of the Lepidopterists’ Society 25: 185-193. _ . 1971d. Status and potential of biological control in Canada. Part IV, Review of biological control in Canada, 1959-1968. C.A.B., London 4: 213-255. _ & A. Mutuura. 1971a. Contributions to a study of the Pyrausti¬ nae (Lepidoptera: Pyralidae) of Temperate East Asia. XI. Cana¬ dian Entomologist 103: 173-181. _ & _ . 1971b. Contributions to a study of the Pyraustinae (Lepidoptera: Pvralidae) of Temperate East Asia. XII. Canadian Entomologist 103: 503-506. _ & _ . 1971c. Geographical distribution of Pyraustinae (Lep¬ idoptera: Pyralidae) of temperate East Asia. Transactions of the Lepidopterists’ Society of Japan 22: 1-6. Coulson, R. N., A. Mutuura, & E. G. Munroe. 1972. The Dioryc- tria species of loblolly pine in East Texas, with comments on the occurrence of two new species. Journal of Economic Entomology 65: 868-870. Munroe, E .G. 1972a. A new species of Midila Walker (Lepidoptera: Pyralidae: Midilinae) from Venezuela. Canadian Entomologist 104: 711-713. _ . 1972b. An entomologist in Brazil. Tableau: 2. _ . 1972c. Centime de lepidopteres de file de Cuba, by Ph. Poey. Reprinted by E. W. Classey Ltd. Journal of the Lepidopterists’ Society 26: 123. _ . 1972d. Book review — Jamaica and its butterflies, by F.M. Brown and B. Heineman. Journal of the Lepidopterists’ Society 26: 264-267. _ . 1972e. Pvraloidea. In The moths of America north of Mexico. Fasc. 13. 1A. Subfamilies Scopariinae, Nymphulinae. R. B. Do¬ minick, C. R. Edwards, D. C. Ferguson, J. G. Franclemont, R. W. Hodges, and E. G. Munroe. E. W. Classey, (eds.), London: 1-134. _ . 1972f. Pyraloidea. In The moths of America north of Mexico. Fasc. 13. IB. Subfamilies Odontiinae, Glaphyriinae. R. B. Do¬ minick, C. R. Edwards, D. C. Ferguson, J. G. Franclemont, R. W. Hodges, and E. G. Munroe. E. W. Classey, (eds.), London: 137-250. Mutuura, A. & E. G. Munroe. 1972. American species of Dionjctria Zeller (Lepidoptera: Pvralidae). III. Grouping of species: species of the auranticella group, including the Asian species, with the description of a new species. Canadian Entomologist 104: 609-625. Munroe, E. G. 1973a. A supposedly cosmopolitan insect: the celery webworm and allies, genus Nomopliila Hiibner (Lepidoptera: Pyralidae: Pyraustinae). Canadian Entomologist 105: 177-216. _ . 1973b. A new genus and species of Odontiinae from Arkansas (Lepidoptera: Pyralidae). Canadian Entomologist 105: 669-671. _ . 1973c. A new species of Trischistognatha from Venezuela (Lepidoptera: Pyralidae: Evergestinae). Canadian Entomologist 105: 1081-1083.' _ . 1973d. A new genus for Noctueliopsis radialis (Lepidoptera: Pvralidae: Odontiinae). Canadian Entomologist 105: 1361-1362. _ . 1973e. Transfer of Cymoriza abrotalis Walker, 1859, from Nytnphula Schrank to Disimidila Dvar (Pyralidae: Nymphulinae, Midilinae). Journal of the Lepidopterists’ Society 27: 290-201. _ . 1973f. Book review — Die tagfalter Europas und Nordwest Afrikas, ein Taschenbuch fur Biologen, by L. G. Higgins and N. D. Riley, illustrated by B. Hargreaves, and translated by W. Forster. Bulletin of the Entomological Society of Canada 5(3): 30. _ . 1973g. Book review — An index to the described life histories, early stages and hosts of the macrolepidoptera of the continental United States and Canada, by H. M. Tietz. Bulletin of the Ento¬ mological Society of Canada 5(3): 94-95. _ . 1973h. Book review — Emperor moths of south and central Africa, by E. Pinhey. Bulletin of the Entomological Society of Canada 5(3): 96-97.' Mutuura, A. & E. G. Munroe. 1973. American species of Dionjctria [Zeller] (Lepidoptera: Pvralidae). IV. The schuetzeeUa group and the taxonomic status of the spruce cone moth. Canadian Ento¬ mologist 105: 653-668. Munroe, E. G. 1974a. Pyraloidea. In The moths of America north of Mexico. Fasc. 13. 1C. Evergestinae. R. B. Dominick, D. C. Fer¬ guson, J. G. Franclemont, R. W. Hodges, and E. G. Munroe. E. W. Classey, (eds.), London: 253-304. _ . 1974b. Three new genera and four new species of Pyralidae (Lepidoptera) from South America. Canadian Entomologist 106: 11-20. _ . 1974c. New Pvralidae (Lepidoptera) from the Papuan Region. III. Canadian Entomologist 106: 21-30. _ . 1974d. Pyralidae of the Third Archbold Expedition. I. Sub¬ family Odontiinae (Lepidoptera). Tijdschrift voor Entomologie 117: 27-37. _ . 1974e. New genera, species, and generic transfers of Neotrop¬ ical Odontiinae (Lepidoptera: Pvralidae). Canadian Entomologist 106: 127-137. _ . 1974f. Transfer of Cybalomia angustalis Dognin to Udea Guenee, with description of a related new species (Lepidoptera: Pyralidae). Canadian Entomologist 106: 139-142. _ . 1974g. A new Noctueliopsis from Texas (Lepidoptera: Pyrali¬ dae: Odontiinae). Canadian Entomologist 106: 655-658. _ . 1974h. Hard decisions ahead for Canada’s biological scientists. Science Forum 37 7(1). Mutuura, A. & E. G. Munroe. 1974. A new genus related to Dioryc- tria Zeller (Lepidoptera: Pyralidae: Phycitinae), with definition of an additional species group in Dioryctria. Canadian Entomologist 106: 937-940. Munroe, E. G. 1975a. Biogeography of North American Odontiinae. Canadian Entomologist 107: 129-154. _ . 1975b. Butterflies and moths in winter. Nature Canada 4(1): 26-29. _ . 1975c. Entomology in the N.C.C. Symposium on the ecology of the Mer Bleue. 2pp. _ . 1975d. Science policy committee. Bulletin of the Entomologi¬ cal Society of Canada 7: 77. _ . 1975e. Moths of importance to crops, forests and environmen¬ tal quality in North America, with particular reference to un¬ solved taxonomic problems. The Wedge Entomological Research Foundation, New York, N.Y. 31pp. Dominick, R. B. & E. G. Munroe. 1975. Why should individuals, or- 56 Journal of the Lepidopterists’ Society ganizations or groups support the Wedge Entomological Re¬ search Foundation? The Wedge Entomological Research Foun¬ dation, New York, N.Y. 13pp. MUNROE, E. G. 1976a. Pvraloidea. In The moths of America north of Mexico. Ease. 13. 2A , Pvralidae, Subfamily Pvraustinae, Tribe Pyraustini. B. Pyralidae, (conclusion). R. B. Dominick, T. Do¬ minick, D. C. Ferguson, J. G. Franclemont, R. W. Hodges, and E. G. Munroe. E. W. Classey, (eds.), London: 1-78. _ . 1976b. Pvraloidea. In The moths of America north of Mexico, Fasc. 13. 2B. Pyralidae, Subfamily Pvraustinae, Tribe Pyraustini (conclusion). R. B. Dominick, D. C. Ferguson, J. G. Fran¬ clemont, R. W. Hodges, and E. G. Munroe. (eds.), E.W. Classey, London: 81-150. _ . 1976c. Hapalia nigristriatalis a synonym of Udea angustalis (Pyalidae: Pvraustinae). Journal of the Lepidopterists' Societv30: 11. _ . 1976d. New genera and species of Pyraustinae (Lepidoptera: Pyralidae), mainly from the collection of the British Museum (Natural History). Canadian Entomologist 108: 873-884. _ . 1976e. Book Review — Macrolepidoptera of Fiji and Roturna: a taxonomic and hiogeographic study, by Gaden S. Robinson. Bul¬ letin of the Entomological Society of Canada 8: 13-14. _ . 1976f. Obituary — Richard Bayard Dominick, 1919-1976. Bul¬ letin of the Entomological Society of Canada 8(4): 19-20. _ . 1977a. Lepidoptera. In Biota Acuatica de Sudamerica Austral. S. II. Hurlbert (ed.), San Diego State University, San Diego, Cal¬ ifornia 14: 254-255. _ . 1977b. Two species of Palpita Hiibner described by Janse, with descriptions of new species (Lepidoptera: Pyralidae: Pvrausti¬ nae). Canadian Entomologist 109: 67-76. _ . 1977c. Synopsis of Heortia Lederer, with descriptions of a new species and a new subspecies (Lepidoptera: Pyralidae: Odonti- inae). Canadian Entomologist 109: 429-441. _ . 1977d. A new genus and two new species of Neotropical Pyraustinae (Lepidoptera: Pyralidae). Canadian Entomologist 109: 481^185. _ . 1977e. A new genus and species of Odontiinae, with remarks on Balaenifrons Hampson and Taurometopa Meyrick (Lepi¬ doptera: Pyralidae). Canadian Entomologist 109: 865-869. _ . 1977f. New Indo-Malayan Odontiinae (Lepidoptera: Pyrali¬ dae), with taxonomic and synonymic notes. Canadian Entomolo¬ gist 109: 1341-1358. _ . 1977g. Additional material of Scoparia huachucalis Munroe, with description of male genitalia (Pyralidae, Scopariinae). Jour¬ nal of the Lepidopterists' Society 31: 63-66. _ . 1977h. Canada’s endangered species and habitats: Insects. Why protect insects? Canadian Nature Federation: 144-147. _ . 1977i. Introduction for Monographic des Phycitides et Gal- leriides par E. L. llagonot. Facsimile edition. E.W. Classey Ltd. _ . 1977j. Book review — Insects that feed on trees and shrubs: an illustrated practical guide, by W. T Johnson & H. II. Lyon. Cana¬ dian Field-Naturalist 91: 105-106. ' _ . 1977k. Book review — Biology of insects, by David J. Horn. Canadian Field-Naturalist 91(3): 326. Carde, R. T„ W. L. Roelofs, R. G. Harrison, A. T. Vawter, P. F. Brussard, A. Mutuura, & E. G. Munroe. 1978. European Com Borer: pheromone polymorphism or sibling species? Science 199: 555-556. Munroe, E. G. 1978a. A new species related to the garden webworm. with a key to American members of the genus Achyra Guenee (Lepidoptera: Pyralidae: Pyraustinae). Canadian Entomologist 110: 499-505. _ . 1978b. Epicorsia I Iiibner and allied Neotropical genera: sup¬ plementary notes and descriptions (Lepidoptera: Pyralidae: Pyraustinae). Canadian Entomologist 110: 673-695. _ & A. Mutuura. 1978. Meyrick's record of “Mecyna fumacalis, GN” from Fiji, with a new generic assignment for Pyrausta homaloxarvtha Meyrick (Pyralidae: Pyraustinae). Journal of the Lepidopterists’ Society 32:130-134. Mutuura, A. & E. G. Munroe. 1978. An Old World leaf roller po¬ tentially injurious to Rosaceae, newly discovered in British Co¬ lumbia, Croesia holmiana (Lepidoptera: Tortricidae). Canadian Entomologist 110: 641-646. Munroe, E. G. 1979. Lepidoptera. In Canada and its insect fauna. H.V. Danks (ed). Memoirs of the Entomological Society of Canada 108: 427-481. Mutuura, A. & E. G. Munroe. 1979. American species of Dioryctria (Lepidoptera: Pyralidae). 5. Three new cone-feeding species from the southeastern U.S. Journal of the Georgia Entomological Society 14: 290-304. Bolte, K. B. & E. G. Munroe. 1979. Hemithea aestivaria in British Columbia (Lepidoptera: Geometridae: Geometrinae). Canadian Entomologist 111: 1121-1126. Munroe, E. G. & M. Shaffer. 1980. A revision of Vitessidia Roth¬ schild & Jordan and Vitessa Moore (Lepidoptera: Pyralidae). Bul¬ letin of the British Museum of Natural History, Entomology 39: 241-360. Inoue, H., E. G. Munroe, & A. Mutuura. 1981. A new species of Glyphodes from Japan, with biological notes. Tinea 11: 91-97. Munroe, E. G. 1982. Lepidoptera. In Synopsis and Classification of Living Organisms. S. P. Parker, (ed.), McGraw-Hill, New York: 612-651. " _ . 1983a. Pyralidae (except Crambinae). In Check List of the Lepidoptera of America north of Mexico. R. W. Hodges, T. Do¬ minick, D. R. Davis, D. C. Ferguson, J. G. Franclemont, E. G. Munroe, and J. A. Powell. E. W. Classey and The Wedge Ento¬ mological Research Foundation (eds.), London : 67-85. _ . 1983b. Wing Patterns in Lepidoptera (Gold Medallists Ad¬ dress). Bulletin of the Entomological Society of Canada 14: 103-112. _ . 1986. Obituaiy— George Pearson Holland, 1911-1985. Trans¬ actions of the Royal Society of Canada 1: 303-305. _ . 1989. Changes in classification and names of Hawaiian Pvraloidea since the publication of Insects of Hawaii, Volume 8, by. E. C. Zimmerman (1958) (Lepidoptera). Bishop Museum Oc¬ casional Papers 29: 199-212. Shaffer, J. C. & E.G. Munroe. 1989a. Type material of four African species of Notarcha Meyrick, wfith designations of lectotypes and changes in synonymy (Lepidoptera: Crambidae: Pyraustinae). Proceedings of the Entomological Society of Washington 91: 248-256. _ & _ . 1989b. Type material of two African species of Her- petogramma and one of Pleuroptya (Lepidoptera: Crambidae: Pyraustinae). Proceedings of the Entomological Society of Wash¬ ington 91: 414-420. Smith, D. S., E. G. Munroe, & S. J. Ramos. 1989. Biogeographical affinities of the butterflies of the soutlrwest Caribbean Islands, San Andres and Providencia. Journal ol the Lepidopterists’ So¬ ciety 43(4): 274-288. Clavijo, J. A. & E. G. Munroe. 1991. Misidentification of Diaphania liyalinata (L.) (Lepidoptera: Crambidae: Pyraustinae) in the British Lepidoptera fauna. British Journal of Entomology and Natural History 4: 9-10. Munroe, E. G. 1991. Transfer of Autocodes eupselias Meyrick to Pyraustinae, with notes on the genus Marasmia Lederer and on cataclystiform wfing patterns in the family Crambidae (Lepi¬ doptera: Pyraloidea). Bishop Museum Occasional Papers 31: 122-130. Mather, B. & E. G. Munroe. 1992. Heleithia decostalis (Crambidae: Pyraustinae) new to the United States. Journal of the Lepi¬ dopterists' Society 46: 159-160. Munroe, E. G. 1992. Three nominal genera of Crambidae from “The Generic Names of Moths of the World, Vol. 5, Pvraloidea.” Jour¬ nal of the Lepidopterists’ Society 46: 75-77. _ . 1995. Usingeriessa onyxalis (Dvar) (Lepidoptera: Crambidae: Nymphulinae), a moth with presumably aquatic larvae, newly recorded from I Iawaii, with a synopsis of Hawaiian Nymphulinae. Bishop Museum Occasional Papers 42: 39—42. _ , V. O. Becker, J. C. Shaffer, M. Shaffer, & M. A. Solis, 1995. Pyraloidea. In Atlas of Neotropical Lepidoptera. Checklist: Volume 63, Number 1 57 Part 2. Hyblaeoiclea - Pyraloidea - Tortricoidea. |. B. Heppner (ed.). Association of Tropical Lepidoptera, Gainesville: 34-105, Clavijo, J. A., E. G. Munroe, & Q. Arias. 1995. The genus Diapha- nia Hiibner (Lepidoptera: Crambidae): key to species of eco¬ nomic importance. Agronomia Tropical 45: 347-358. _ & E. G. Munroe. 1996. A new species of Diaphania from Tropical America (Lepidoptera: Pyraloidea: Crambidae). Tropical Lepidoptera 7: 151-153. Munroe, E. G. 1996. Distributional patterns of Lepidoptera in the Pacific Islands. In The Origin and Evolution of Pacific Island Bio¬ tas, New Guinea to Eastern Polynesia: Patterns and Processes. A. Least and S. E. Miller (eds.). Academic Publishers, Amsterdam: 275-295. _ & M. A. Solis. 1999. Pyraloidea. In Lepidoptera, Moths and Butterflies, Vol. 1. Arthropoda. Insect, Vol. 4, Part 35. Handbook of Zoology'. N. Kristensen (ed.). Walter de Gruyter & Co., Berlin: 233-256.' Peck, S. & E. G. Munroe. 1999. Biogeography and evolutionary his¬ tory: wide-scale and long-term patterns in insects. In Ecological Entomology. C. B. Huffaker and A. P Gutierrez (eds.). John Wi¬ ley & Sons, New York: 231-261. Shaffer, J. & E. G. Munroe. 2007. Crambidae of Aldabra Atoll (Lepidoptera: Pyraloidea). Tropical Lepidoptera 14: 1-109. M. A. Solis, SEL, USD A, Smithsonian Institution, P.O. Box 37012, National Museum of Natural History, E-517, MRC 168, Washington, D.C., 20013-7012; email- alma. solis@ars.uscla.gov Received for publication 30 September 2008; revised and accepted 17 January 2009.. 58 Book Review Journal of the Lepidopteuists’ Society Journal of the Lepidopterists’ Society 63(1), 2009, 58 MOTH CATCHER: AN EVOLUTIONIST’S JOURNEY THROUGH CANYON AND PASS. By Michael M. Collins. 176 pages, 13 color plates; ISBN 978-0-87417-725-1 (hardcover), ISBN 978-0-87417- 735-0; US$49. 95 (hardcover), US$18.95 (paperback). Univ. of Nevada Press. Publication date: 2007. Two threads run through the stories of most lepidopterists that contribute to their life-long obsessive fascination with some aspect of entomology, whether they are professionals or avid amateurs. The first is some event of life-changing discovery to light the fire; the second is the timely connection with one or more knowledgeable and nurturing mentors to fuel the fire so that it burns throughout a lifetime. Such is indeed the case with Mike Collins, who shares with us the beginnings and many subsequent years of his own journey as a lepidopterist — in this case pursuing the life history, behaviors and interspecific relationships of western North American Saturniidae. Migrating from his native Iowa to the western mountains, Mike quickly passed beyond the usual goal of building a large collection of adult specimens and focused on questions of biology and evolution, prompted first by the late Walter R. Sweadners monograph on the evolution of the genus Hyalophora. Seminal to his later researches, Sweadners treatise on natural hybridization begged for substantiating research. The evolutionary relationships among the 3 Hyalophora species — cecropia, glove ri and euryalus — became the major focus of Mike’s research. Mike provided important supporting evidence for Sweadners ideas in collections and by rearing H. gloveri and H. euryalus from areas of sympatry and allopatry. The discovery and elucidation of the hybrid zone where these two species come together in Monitor Pass in eastern California proved to be a major discovery. This stoiy is the centerpiece of the book. The sibling nature of Hemileuca eglanterina, nuttalli and hero and their isolating mechanisms was another problem in Mike’s voyage of research and discoveiy. His research on these and other saturniids was woven into his undergraduate years at UC Berkeley, and PhD work under Art Shapiro at UC Davis. Friendships with several of our colleagues and stories of field adventures together spice the narrative. The account of his first meeting with Art Shapiro gave me a knowing chuckle. A short biography of Sweadner, and Mike’s meeting with his widow and daughter, was of special interest to me, as was a photo of this man who inherited W. J. Holland’s curatorial mantle at the Carnegie Museum, but for a tragically short period. Collins’ style is anecdotal, written in the first person, and easily readable. He uses it effectively when describing people and research projects without the cumbersome data found in his scientific publications. There are numerous short treatises on topics pertinent to systematic^ and ecology: western montane ecology (plant and animal), Neo-Darwinism, industrial melanism, DNA sequencing, speciation (biological species concept versus phvlogenetie species concepts), and pre-mating and post-mating barriers, to name a few. The book thus serves as a good primer for bright young people interested in entering the field of population ecology. Mike’s lifelong passion for his chosen specialty in entomology — isolating mechanisms among closely related saturniid moths in the majestic West — shines through from beginning to end. Mv minor comments concern uneven referencing: some of the major literature cited (i.e. Sweadners monograph, p.24), are mentioned generally (no complete title and other citation details), but not included in the Bibliography. The reader must scout these out from other sources. Also, some less scientifically trained readers might benefit from definitions of more of the scientific terms employed ( genome , p. 44; “relief’, p. 56). Other terms are well defined or explained. It is always fascinating to see how our colleagues began their journeys with Lepidoptera — how and from whom they drew inspiration, nurturing and training— and then see a summary of their career-long contributions to the science. Mike’s story is an enjoyable and informative addition to this lore. Charles V. Covell Jr., McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, Hull Road at SW 34th St., Gainesville, FL 32611. Email: ccovell@flmnh.ufl.edu. Received for publication 10 Dec 2008; revised and accepted 17 Jan 2009 Volume 61, Number 1 59 Journal of the Lepidopterists' Society 63(1), 2009, 59 MANUSCRIPT REVIEWERS FOR 2008 (VOLUME 62) Manuscript reviewers are anonymous contributors to the scientific rigor, clarity, and quality of text and illustrations in the papers published by the Journal of the Lepidopterists’ Society. The reviewers’ input is invaluable and always welcomed by authors, editors and readers. We hope their careful work continues to allow the Journal to increase quality and readership. On behalf of all the authors and the editorial staff of the Journal, respectful acknowledgement is given to the reviewers for contributions published in Volume 62. John Acorn, Edmonton, Alberta, Canada Matti Ahola, Koski, Finland Annette Aiello, Washington, DC Richard Arnold, Pleasant Hill, California George Balogh, Portage, Michigan Susan Rorkin, Milwaukee, Wisconsin M. Deane Bowers, Boulder, Colorado Lincoln Brower, Roseland, Virginia John Brown, Washington, DC John Bums, Washington, DC John Calhoun, Palm Harbor, Florida Steven Cary, Santa Fe, New Mexico Jeff Crolla, Toronto, Ontario, Canada Michelle DaCosta, Winston-Salem, North Carolina Robert Dana, Minneapolis, Minnesota Phillip DeVries, Metairie, Louisiana Robert Dirig, Ithaca, New York John Emmel, Hemet, California Tom Emmel, Gainesville, Florida Matthew Forister, Reno, Nevada Terry Harrison, Charleston, Illinois David Iftner, Pittsfield, Illinois Brian Inouye, Tallahassee, Florida Akito Kawahara, College Park, Maryland Ian Kitching, London, England Norbert Kondla, McBride, British Columbia, Canada David Lawrie, Edmonton, Alberta, Canada David MacDougall, Albany, New York Billy McCord, Charleston, South Carolina David McCorkle, Monmouth, Oregon William Miller, St. Paul, Minnesota Mark Minno, Gainesville, Florida Eugene Morton, Cambridge Springs, Pennsylvania Karen Oberhauser, St. Paul, Minnesota Paul Opler, Loveland, Colorado Jerry Powell, Berkeley, California Ron Priest, East Lansing, Michigan Tommaso Raeheli, Rome, Italy Robert Raguso, Ithaca, New York Jadranka, Rota, Washington, DC Ronald Royer, Minot, North Dakota Daniel Rubinoff, Honolulu, Hawaii Erik Runquist, Davis, California Ronald Rutowski, Tempe, Arizona Emily Saarinen, Gainesville, Florida James Scott, Lakewood, Colorado J. Mark Scriber, East Lansing, Michigan Paul Severns, Philomath, Oregon Jon Shepard, Nelson, British Columbia, Canada John Shuey, Indianapolis, Indiana Felix Sperling, Edmonton, Alberta, Canada Stephen Spomer, Lincoln, Nebraska Fred Stehr, East Lansing, Michigan Kieth Summerville, Des Moines, Iowa Michael Toliver, Eureka, Illinois Tonya Van Hook, Sweet Briar, Virginia Tom Vawter, Aurora, New York Andrew Warren, Castle Rock, Colorado Wayne Welding, Silver Spring, Maryland Martha Weiss, Washington, DC Donald Wright, Cincinnati, Ohio Myron Zalucki, Brisbane, Australia EDITORIAL STAFF OF THE JOURNAL Brian Scholtens, Editor Biology Department College of Charleston, 58 Coining Street, Charleston, South Carolina 29424-001 1 . 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Table 1.) and use a concise and informative heading. Type each table on a separate sheet and place after the Literature Cited section, with the approximate desired position indicated in the text. Avoid vertical lines and vertical writing. Voucher specimens: When appropriate, manuscripts must name a public repository where specimens documenting the identity of organisms can be found. Kinds of reports that require vouchering include descriptions of new taxa, life histories, host associations, immature morphology, and some experimental studies. Proofs: The edited manuscript and galley proofs will be mailed to the author for correction of printer’s errors. Excessive author’s changes will be charged to authors at the rate of $3.00 per line. A purchase order for reprints will accompany proofs. Page charges: For authors affiliated with institutions, page charges are $50 per Journal page. For authors without institutional support, page charges are $25 per Journal page. For authors who are not members of the Society page charges are $75 per Journal page. Authors unable to pay page charges for any reason should apply to the editor at the time of submission for a reduced rate. Authors of Book Reviews and Obituaries are exempt from page charges. Correspondence: Address all matters relating to the Journal to the editor. Address book reviews directly to the book review editor. PRINTED BY THE ALLEN PRESS. INC., LAWRENCE. KANSAS 6fi<)44 U.S.A. CONTENTS 3 9088 01912 3868 A New Species of Drasteria Hubner (Noctuidae: Catocalinae: Melipotini) from Arizona, with Comments on the Genus Jan Metlevski and Gregory Zolnerowich - 1 Biology and Distribution of a Recently Rediscovered Endemic Hawaiian Leafrolleii Moth, Omiodes continuatalis (Crambidae) Cynthia King, Daniel Rubinojf and William Haines — 11 A New Species of Morrisonia (Noctuidae) from Southeastern North America J. Bolling Sullivan and Janies K. Adams - 21 Tinacrucis noroesta , New Species, North America’s Largest Tortricine Moth (Tortricidae: Atteriini) Jerry A. Powell - 27 Melitara Walker (Pyralidae) in Western Canada: The Documentation of M. SUBUMBRELLA (DYAll) IN THE PRAIRIE PROVINCES DEMONSTRATES THE VALUE OF REGIONAL Collections and Species Lists. Thomas J. Simonsen, Jason J. Dombroskie and Gregory R. Pohl - 31 Not All Bilateral Gynandromoiiph Butterflies are Interspecific Hybrids: New Papilio Specimens from Lield Populations J. Mark Scriber, Rodrigo Me reader, Howard Romack and Mark Deering - 37 General Notes Pithing and Mining by a Punkie: The Unusual Leeding Strategies of Nola cilicoides (Grote, 1873) (Noctuidae: Nolinae) David L. Wagner and Bryan Connolly - 48 Obituary Eugene G. Munroe (8 September 1919-31 May 2008) M. A. Solis - 52 Book Review Moth Catcher: An Evolutionist’s Journey Through Canyon and Pass. By Michael M. Collins. Charles V. Covell Jr. - 58 List of Reviewers for Volume 62- 59 ® This paper meets the requirements of ANSI/NISO Z39. 48-1 992 (Permanance of Paper).