AY Volume 61 Number 3 OVS 24 October 2007 Se [eo i ENT ISSN 0024-0966 Journal of the Lepidopterists Society Published quarterly by The Lepidopterists’ Society THE LEPIDOPTERISTS’ SOCIETY EXECUTIVE CoUNCIL Joun H. Acorn, President Jorn Litt, Vice President WituiaMm E. Conner, Immediate Past President Davin D. Lawrir, Secretary Anpre V.L. Freitas, Vice President Ketiy M. Ricuers, Treasurer Axiro Kawanara, Vice President Members at large: Kim Garwood Richard A. Anderson Michelle DaCosta Kenn Kaufman John V. Calhoun John H. Masters Harry Zirlin Amanda Roe Michael G. Pogue EprrortaL Boarp Joun W. Brown (Chair) Micuaer E. Tourer (Member, at large) Brian Scholtens (Journal) Lawrence F. Gaui (Memoirs) Date Criark (News) Joun A. Snyper (Website) Honorary Lire MEMBERS OF THE SOCIETY Cuarces L. RemMincTon (1966), E. G. Munroe (1973), [An F. B. Common (1987), Lincoin P. Brower (1990), Freperick H. Rinpcr (1997), Ronatp W. Hopces (2004) The object of The Lepidopterists’ Society, which was formed in May 1947 and formally constituted in December 1950, is “to pro- mote the science of lepidopterology in all its branches, . . . to issue a periodical and other publications on Lepidoptera, to facilitate the exchange of specimens and ideas by both the professional worker and the amateur in the field; to secure cooperation in all mea- sures” directed towards these aims. Membership in the Society is open to all persons interested in the study of Lepidoptera. All members receive the Journal and the News of The Lepidopterists’ Society. Prospective members should send to the Assistant Treasurer full dues for the current year, to- gether with their full name, address, and special lepidopterological interests. In alternate years a list of members of the Society is is- sued, with addresses and special interests. annual dues $45.00 within the U.S., $50.00 outside the U.S. Affiliated members—annual dues $10.00 within the U.S., $15.00 outside the U.S. Student members—annual dues $20.00 within the U.S., $25.00 outside the U.S. Sustaining members—annual dues $60.00 within the U.S., $65.00 outside the U.S. Life members—single sum $1,800.00 Institutional subscriptions—annual $60.00 Airmail postage for the News $15.00 Active members Send remittances, payable to The Lepidopterists’ Society, to: Kelly M. Richers, Treasurer, 9417 Carvalho Court, Bakersfield, CA 93311; and address changes to: Julian P. Donahue, Natural History Museum, 900 Exposition Blvd., Los Angeles, CA 90007-4057. For information about the Society, contact: Ernest H. Williams, Department of Biology, Hamilton College, Clinton, NY 13323. To order back issues of the Memoirs, write for availability and prices to Kenneth R. Bliss, 28 DuPont Avenue, Piscataway, NJ 08854. The additional cost for members outside the U.S. is to cover mailing costs. Journal of The Lepidopterists’ Society (ISSN 0024-0966) is published quarterly by The Lepidopterists’ Society, %o Los Angeles County Museum of Natural History, 900 Exposition Blvd., Los Angeles, CA 90007-4057. Periodicals postage paid at Los Angeles, CA and at additional mailing offices. POSTMASTER: Send address changes to The Lepidopterists’ Society, ‘o Natural History Museum, 900 Exposition Blvd., Los Angeles, CA 90007-4057. Cover illustration: Males (lines 1 & 3) and females (lines 2 & 4) of two closely related species in each of three genera of neotropical skippers: Polyctor (left), Cobalus (center), and Neoxeniades (right). Reared specimens from northwestern Costa Rica, photographed by Daniel H. Janzen. See paper by John M. Burns et al., page 138. JouRNAL OF Tue Lepiporterists’ S OCIETY Volume 61 Journal of the Lepidopterists’ Society 61(3), 2007,117-124 2007 Number 3 TAXONOMY OF FOUR SPECIES OF EUCOSMINI (TORTRICIDAE) ASSOCIATED WITH PELOCHRISTA COROSANA (WALSINGHAM) INCLUDING A NEW SYNONYMY AND DESCRIPTION OF A NEW SPECIES DONALD J. WRIGHT 3349 Morrison Ave., Cincinnati, Ohio 45220-1430, USA, e-mail: wrightdj@fuse net ABSTRACT. Pelochrista gelattana, new species, is described from Alberta, Utah, and Wyoming. Reviews are included of P. corosana (Walsingham) and Par, genteana (Walsingham), its two closest congeners based on male genitalia. Pelochrista idahoana (Kearfott) is recognized as a junior synonym of P. argenteana. Eucosma nuntia Heinrich, a species that i is similar in many respects to P. corosana, is also reviewed. Illustrations detailing the considerable intraspecific variation in wing color and/or male genitalia i in the aforementioned taxa are provided, and current distributional information is summarized. A lectotype is designated for P. argenteana. Additional key words: Olethreutinae, Eucosma, argenteana, idahoana, nuntia. In Nearctic Olethreutinae, the lineage represented by the genera Eucosma Hiibner and Pelochrista Lederer is comprised of approximately 180 recognized species, with the greatest diversity occurring in the arid and mountainous regions of western North America. There is a long history of confusion regarding species boundaries in the western the primary contributing factors being the common occurrence of similar looking taxa and the frequent presence of intraspecific variation in both wing color and genitalic characters. Recent progress in resolving some of these problems is reported in Wright (2005, 2007). This paper is primarily a study of the variation occurring in Pelochrista corosana (Walsingham) and P. argenteana (Walsingham), western species that share a distinctive form of male genitalia. It was prompted by the recent discovery in southeastern Wyoming of a previously unrecognized member of this group, described below as Pelochrista gelattana, new species. A fourth name, P. idahoana (Kearfott), which refers to a taxon long considered to be a very close relative of argenteana, is recognized here as a junior synonym of argenteanda. Heinrich (1929) noted that some phenotypes of P. corosana could easily be mistaken for Eucosma nuntia Heinrich, based on forewing color and maculation, so a review of the latter species is included. These two taxa are similar not only in forewing pattern but even more strongly so in female genitalia. The male genitialia of fauna, nuntia lacks the large spiniform seta at the anal angle of the cucullus associated with members of Pelochrista, but in other respects they are quite similar to those of the other species considered here. They also exhibit a range of variation in valval shape that is comparable to that in argenteana. MATERIALS AND METHODS This study was based on 497 adult specimens and 78 genitalia preparations from the following collections: American Museum of Natural History (AMNH), George J. Balogh, Canadian National Collection (CNC), University of California Berkeley (UCB), Loran D. Gibson, Mississippi Entomological Museum (MEM), John S. Nordin (JSN), United States Museum of Natural History (USNM), and Donald J. Wright (DJW). Forewing length (FWL) refers to the distance from base to apex (including fringe) and was measured to the nearest one tenth of a millimeter with a graticule mounted in a Leica MZ95 stereomicroscope. Aspect ratio (AR), defined as FWL divided by medial forewing width, is reported as the average of such values calculated for a small sample of specimens. The number of measurements supporting a particular statistic is indicated by n. Forewing pattern terminology follows Brown & Powell (1991) as modified by Baixeras (2002). The digital photographs were edited in Adobe Photoshop CS. For stability of nomenclature I am designating as lectotype for argenteana a specimen selected for that 118 purpose by Obraztsov. I did not see the specimen itself, but I did examine a 35 mm slide of the adult and a negative of the genitalia, both made by Obraztsov. The included images of the lectotype were obtained by scanning those photographs. SPECIES ACCOUNTS Pelochrista corosana (Walsingham) (Figs. 1-6, 19, 27) Paedisca corosana Walsingham 1884:139, pl. IV, Fig. 6. Eucosma corosana: Fernald [1903]:460; Barnes and McDunnough 1917:171; Heinrich 1923:127, Fig. 219; McDunnough 1939:47. Pelochrista corosana: Powell 1983: 35; Miller 1987:55. Type. Holotype: ¢, Montana, Yellowstone R., Morrison, genitalia slide 11570, BMNH. Discussion. In most specimens of corosana the forewing color is olive gray to olive brown (Figs. 3-6). One phenotype from Arizona has a reddish appearance (Fig. 1), and traces of this reddish hue are found in some specimens from New Mexico (Fig. 2). The principal forewing markings are dark olive gray to blackish brown and include an incomplete subbasal fascia running obliquely outward from dorsum to radius and a median fascia from iid costa to the pretornal portion of the dorsum. Occasionally, extensive suffusion of the forewing with dark coloration renders the fasciae barely discernable (Fig. 6). The median fascia is bordered distally by a narrow white line, a feanive that is present even in the darkest specimens. Between the white line and the apex there generally is a semitriangular patch of dark scales that connects to a dark apical mark. Most individuals have a streak of white at the base of the fringe along the anterior two thirds of the termen. The ocellus, which is anes ure but usually recognizable, is crossed by 3 to 4 black longitudinal streaks (often reduced to black dots) and is bordered basally and distally by variably expressed, silvery-gray, transverse bars. Forewing statistics: ¢ FWL 7.5-10.6 mm (mean = 9.4, n = 31), AR = 3.01, 2 FWL 7.7-10.5 mm (mean = 9.3, n = 19), AR = 2.94. Male genitalia (Fig. 19): Valva with neck long, apex nearly right- angled, ventral angle produced into triangular projection supporting one stout spine, oneal invagination baw! and shallow, proximal margin of medial surface with well dev eloped pulvinus; aedeagus long etna narrow, vesica with 5-12 deciduous cornuti (n = 15). Female genitalia (Fig. 27): Papillae anales facing laterally and moderately setose; lamella postvaginalis well dev elope d, with posterolateral comers semitriangular; sternum VII with strongly sclerotized and sometimes mildly raised posterior and lateral margins; ductus bursae with sclerotized patch incorporating a distinct fold posterior to juncture with ductus seminalis; corpus bursae with two large signa situated opposite one another on lateral surfaces of the membrane and projecting anteriorly into bursa, adjacent membrane variably wrinkled. Distribution and biology. This moth is widely distributed in western North America and is quite common in the high plains east of the Rocky Mountains. I examined 203 adult specimens (155 ¢, 48 °) from Arizona, Colorado, Iowa, Kansas, Montana, Nebraska, New Mexico, North Dakota, Oklahoma, Texas, and Utah. Capture dates range from June through September. No larval food plant has been reported. 1880, Pelochrista argenteana (Walsingham) (Figs. 9-12, 17, 18, 20, 22, 26) Paedisca argenteana Walsingham 1895:504, pl. XII, Fig. 13. Eucosma argenteana: Fernald {1903]:460; Barnes and McDunnough JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 1917:169; Heinrich 1923:86, Fig. 216; McDunnough 1939:45. Pelochrista argenteana: Powell 1983:35. Eucosma idahoana Kearfott 1907:90; Barnes and McDunnough 1917:169; Heinrich 1923:86, Fig. 217; McDunnough 1939:45, new synonymy. Pelochrista idahoana: Powell 1983:35. Ee Paedisca argenteana. Lectotype here designated (Figs. 17, 18): d, Loveland, C olomdor July 1891, Smith, ani slide 11566, BMNH. Paralectotypes: same data as lectotype (3 2), BMNH. Eucosma idahoana. Holotype ¢, Blackfoot, Idaho, ae 3, Arthur J. Snyder, genitalia slide CH 2 Dec. 1919, AMNH. Discussion. The description of idahoana was based on a single male. Kearfott apparently was unaware that this taxon is nearly identical in appearance to argenteana. Heinrich (1923) recognized the similarity but treated aahoana as a distinct species based on what he considered to be a more olivaceous forewing color and a somewhat differently shaped cucullus. I compared photographs of the argenteana lectotype and the idahoana holotype and found no significant differences in forewing color. I examined a photograph of the slide prepared by Obraztsov of the argenteana lectotype, Heinrich’s slide of the idahoana holotype, and an additional 18 slides prepared from specimens with the argenteana-idahoana forewing pattern. The observed range of variation in the shape of the cucullus is illustrated in Fig. 22. Figure 22b resembles the idahoana holotype, Fig. 22e the argenteana lectotype. T The full range of | shapes was found in each of two local populations. I also examined 5 female genitalia preparations and found no significant variation. On these grounds I conclude that idahoana should be treated as a junior synonym of argenteana. Differences in argenteana forewing appearance, which can be attributed to varying intensities and shades of color, are illustrated in Figs. 9-12. The pattern of maculation is stable and includes silvery- white longitudinal streaking on a straw-yellow background as follows: a narrow Camel streak from distal end of fold to apex, a wider streak anterior to cubital vein from base to distal margin of discal cell, a thin line along distal one half of CuA2 terminating at tornus, a narrow streak along 1A+2A from base to tornus, and usually a narrow edging along the dorsal margin. The distal one half of the second streak expands to fill the titel cell and is divided longitudinally by a straw yellow bar that is variably overlaid with dark- brown to black scaling. In some specimens (Fig. 12) the white streaks are suffused with straw yellow to the point BG being barely distinguishable. Between the cell and the termen there is variably expressed white streaking on the veins, with straw-yellow to brown scaling between the veins. The white streak on R4 is usually joined basally and distally to the costa by short white dashes. The termen is lined with a narrow white band, the distal scales of which have black to brown apices, producing a thin dark line along the base of the fringe. Fringe scales are white basally, shading to pale straw yellow distally. The brown longitudinal streaking varies considerably, but the dark terminal line is always present. Hindwing color varies from dark gray to grayish white (Figs. 9-11). In females it is nearly always gre ayish white; males exhibit the full range of variation. Forewing statistics: 6 FWL 7.7—-11.4 mm (mean = 9.9, n = 88), AR = 3.53, 2 FWL 7.6-10.2 mm (mean = 8.7, n = 22), AR = 3.33. The male genitalia (Figs. 20, 22) are quite similar in general structure to those of corosana (Fig. 19), the primary differences being in the shape of the cucullus (at least toward the lower end of the spectrum in Fig, 22) and in the basal projection on the medial edge of the valval opening. The latter is only a small bulge in argenteana vs. a fully developed pulvinus in corosana. There appears to be a connection between cucullus shape and hindwing color: males with the idahoana shape have dark gray hindwings, those with the argenteana shape grayish-white. The vesica he 5-11 deciduous cornuti (n = 19). The most prominent feature of the female genitalia (Fig. 26) is the extensive sclerotization of the ductus bursae, which extends from ostium to corpus bursae and expands anteriorly along the membrane of the bursa into two triangular projections flanking the two signa. The papillae anales face laterally and are sparsely setose, the lamella postvaginalis has acute posterolateral corners, and the concavely VOLUME 61, NUMBER 3 119 1s Fics. 1-16. Adults. 1-6, P corosana. 1, 3, Coconino Co., Arizona. 2, 3, Lincoln Co., New Mexico. 3, °, Morton Co., Kansas. 4, ' Morgan Co., Colorado. 5, 3, Weld Co., Galorenta, 6, 2, Gove Co., Kansas. 7-8, P. gelattana. 7, ce) holotype. 8, 3, Albany Co., Wyoming. 9-12, P. argenteana. 9, ¢, ApbeDy, Co., Wyoming. 10, 3, Chaffee Co., Colorado. 11, 2, Weston Co., Wyoming. 12, ¢, Al- bany Co., Wyoming. 13-16, iE See 13,6 _ Grand Co., Colorado. 14, J, Sweetwater Co., Wyoming. 15, 3, Walla Walla Co., Wash- ington. 16, é, Douglas Co., Nevada. Fics. 17-18. Lectotype, P. argenteana. 120 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics: 19-22. Male genitalia. 19, PR. corosana, slide DJW1233. 20, P. argenteana, slide DJW1270. 21, P. gelattana, slide DJW1287. 22 a-f, P. argenteana, slides DJW1271, 1245, 861, 1250, 1270, 1251. Scale bar = 0.5 mm. VOLUME 61, NUMBER 3 emarginated posterior margin of sternum VII has a ventral bulge that weakly shields the ostium. Distribution and biology. I examined 196 specimens (171 ¢, 25 2) from Alberta, British Columbia, California, Colorado, Iowa, Montana, New Mexico, North Dakota, Oregon, South Dakota, Saskatchewan, Texas, Utah, and Wyoming. With the exception of one September record from Sioux City, Iowa, all were collected between 30 May and 28 July. No larval host has been reported. Pelochrista gelattana, new species (Figs. 7, 8, 21, 29) Diagnosis. The shape of the male valva distinguishes gelattana from all Nearctic species of Pelochrista except argenteana and corosana. In most cases these three species are easily separated on the basis of forewing color and maculation. The argenteana phenotype with muted white forewing streaking (Fig. 12) might be confused with the non-speckled form of gelattana (Fig. 7), but gelattana lacks the black terminal line and has several small brown marks on the distal one half of the costa. The female genitalia of the three species are quite distinct (Figs. 26, 27, 29). Description. Head: Frons and vertex pale yellowish tan; labial Fics.. 23-25. palpus with basal segment white, medial surface and dorsal margin white, lateral surface tan, third segment concealed by long narrow scales on ventral margin of second segment; dorsal and lateral surfaces of antenna with pale tan scaling. Thorax: Dorsal surface concolorous with head; ventral surface creamy white, legs tan to creamy white. distal ends of tarsal segments ringed with white scales, the latter usually inconspicuous on hindleg. Forewing (Figs. 7, 8): ¢ FWL 10.2-13.4 mm (mean = 11.9, n = 44), AR = 3.4; 2 EWL 12.7 mm, AR = 3.34 (n = 1); costa nearly straight, vertex acute, termen straight; dorsal surface uniformly pale yellowish brown, sometimes with very pale reddish-brown overtones, frequently with some blackish-brown tipped scales producing a sparsely speckled effect; male costal fold usually slightly darker than adjacent wing area; ocellus obscure; costal stieulas weakly defined by habe brown to blackish-brown marks: fringe scales creamy white basally with very pale tan to brown apices. Hindwing: Gray brown to blackish brown, fringe white. Male genitalia (Fig. 21): Uncus semitriangular and dorsally setose; dorsolateral shoulders of tegumen moderately developed; socii of medium length, tapering distally, and moderately setose; gnathos a narrow band; aedeagus long and narrow, vesica with 3-8 deciduous cornuti (n = 4); valva with costal margin weakly concave, distal margin convex, ventral invagination broad and shallow, ventral angle produced into moderately long projection supporting one stout spine, cucullus with medial surface densely setose, margin of basal opening with weakly developed pulvinus supporting patch of short stout setae. Female petal (Fig. 29): Papillne anales large, facing ventrally, finely ridged transversely, and densely setose, setae on lateral margins of lobes long and curving ventrally, those near anal opening with hooked apes, apophyses posteriores short, length ca. 0.33 x that of apophyses anteriores; tergum VIII with long setae along posterior margin: lamella postvaginalis strongly developed, with semirectangular lateral extensions and a shallow central trough from ostium to medial invagination of posterior margin, long setae on lateral extentions and on membrane between sterigma and ventral extremities of tergum VI; posterior margin of stemum VII with medial, triangular, posteriorly directed projection overlapping ostium; ductus bursae with small sclerotized patch midway between ostium and juncture with ductus seminalis; corpus bursae small, wrinkled, with one small spike- ) PPO — ; Male E. nuntia genitalia. 23, slide DJW248. 24, slide DJW1663. 25 a-h. slides USNM70479, 70478, 70472. DJW806, 1747, 1748, USNM70474, DJW1665. Scale bar = 0.5 mm. shaped signum and one larger thimble-shaped signum. Holotype ¢: Wyoming, “Albany Co., TI5N R75W $29, W. side Gelatt tia nl 'S. Nordin, 6 June 2004, 7250 ft., USNM. Range Sta. D. F. Paratypes: ALBERTA: Mannyberries, Dom. Hardvick. 13 July 1951 (1 ¢, genitalia slide DANY 1358). UTAH: Daggett Co., 4 mi S. of Manile G. J. Balogh, 20 July 1994 (1 9, genitalia slide DJW 711). WYOMING: Albany Co., TISN R75W 829, W. side Gelatt Lake, J. S. Nordin, (1250 ft., 3 June 2004 (2 ¢, genitalia slide DJW 1112), 6 June 2004 (3 3, genitalia slides DJW 1286, 1287), 6 Ne 2006 (5 d), 12 ye 2006 (3 4), 16 June 2005 (1 ¢), 17 [Rite 2005 (4d), 18 June. 205 (4d), 19 Je, 2008 (1d), 20 june 2005 (2d), 21 une 2005 (5d), 27 une 2003 (2d), 29 June 3006 (5 , 30 June 2005 (2.3), 1 July 2005 (3 3), 6 July 2005 34), 7 July 2008 ); 1 mi. E. of ieee 2217 shy View Lane, J. S. Nordin, 8 June. 1990 (1 d). Paratype depositories: AMNH, “BMNH, CNC, Colorado State University, UCB, MEM, JSN, USNM, University of Wyoming, DJW. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Etymology. The specific name derives from the type locality, Gelatt Lake, which is located ca. 15 miles WSW of the city of Laramie in Albany Co., Wyoming. Distribution and biology. The type series consists of 46 males from southeastern Wyoming, one male from the southeast commer of Alberta, and one female from northeastern Utah. These specimens document a flight period extending from early June to mid July. The larval host is unknowns The type locality is noted for its alkali soil. The area surrounding Gelatt Lake has large stands of Atriplex (Saltbush), but the traps producing the Fics. 26-27. Female genitalia. 26, P. argenteana, slide DJW1246. 27, P. corosana, slide DJW1283. Scale bar = 0.5 mm. VOLUME 61, NUMBER 3 gelattana specimens were placed near patches of Gutierrezia (Snakeweed). Discussion. Three fourths of the gelattana specimens exhibited the nearly immaculate forewing illustrated in Fig. 7. The others presented varying amounts of dark speckling, the blackest of which is represented in Fig. 8. The one female specimen reported here is tentatively determined as gelattana. It was not collected in association with any males but is identical in appearance and general proportions to the gelattana holotype. Despite persistent efforts over two FIcs.. 28-29. years, only males have been collected at the gelattana type locality. Eucosma nuntia Heinrich (Figs. 13-16, 23-25, 28) Eucosma nuntia Heinrich 1929:10, Fig. 15; McDunnough 1939:47: Powell 1983:35. Types. Holotype: ¢, Callao, Juab County, Utah, 17 June 1922, genitalia slide USNM 72811, USNM. Paratypes: same locality as holotype, 17 June 1922 (1 4), 23 June 1922 (1 ¢), USNM. [Heinrich (1929) reported one male and one female paratype, but the two specimens mentioned above are males, and both bear Heinrich’s PARATYPE label. Heinrich erroneously reported the capture dates of Female genitalia. 28, E. nuntia, slide DJW1664. 29, P. gelattana, slide DJW711. Scale bar = 0.5 mm 124 the holotype and the first paratype as “17-IV-22”.| Discussion. In forewing pattern, nuntia (Figs. 13-16) resembles corosana, but the color tends toward yellow brown rather than olive brown or olive gray. In many of the specimens from the Great Basin the fasciae are very weakly expressed. Similarities with corosana include a whitish streak along the distal margin of the median fascia and often some white scaling along the termen. Forewing statisticts: ¢ FWL 6.3-10.2 mm (mean = 8.4, n = 29), AR = 2.72; 2 FWL 8.5-9.7 mm (mean = 9.1, n = 5), AR = 2.93. I examined 21 genitalia preparations (18 ¢, 3 2). Figures 23-25 show the variation in the shape of the valva: apex evenly rounded to acutely angular, neck wide to narrow, comer of sacculus broadly obtuse to nearly right angled, distal margin of cucullus with rounded to angular medial bend, and anal angle variably rounded with 2-3 moderately stout setae. The vesica has 9-15 deciduous cornuti. Figure 25g most closely resembles Heinrich’s (1929, Fig. 15) illustration of the holotype. Figures 25a, b, e, f, g illustrate males from a single population at Walla Walla, Washington. The female genitalia are similar to those of corosana, but the sclerotization of the ductus bursae does not include a distinct fold. Distribution and biology. I examined 39 adult specimens (34 ¢, 5 2) from Colorado, Nevada, Utah, Washington, and Wyoming. Capture dates range from 22 May to 15 August, with roughly two thirds of the records between mid June and mid July. No larval host has been reported. ACKNOWLEDGEMENTS I thank J. W. Brown, R. L. Brown, J.-F. Landry, J. S. Nordin, J. A. Powell, and R. T. Schuh for the loan of specimens under their care. Clifford Ferris first called my attention to the argenteana specimens from southeast Wyoming with the idahoana form of cucullus. I particularly appreciate the collecting efforts of John Nordin, who supplied me with many specimens of argenteana and nearly all the specimens of gelattana. LITERATURE CITED BaIxeERrAS, J. 2002. An overview of genus level taxonomic problems surrounding Argyroploce Hiibner (Lepidoptera: Tortricidae), with description of a new species. Ann. Entomol. Soc. Am. 95:422-431. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Barnes, W. & J. MCDUNNouGH. 1917. Checklist of the Lepidoptera of Boreal America. Herald Press, Decatur, Illinois. 392 pp. Brown, R. L. & J. A. POWELL. 1991. Description of a new species of Epiblema (Lepidoptera: Tortricidae: Olethreutinae) from coastal redwood forests in California with an analysis of the forewing pat- tern. Pan-Pacific Entomol. 67:107—114. FERNALD, C. H. [1903]. In Dyar, H. G., A list of North American Lep- idoptera, U.S. Nat. Mus. Bull. 52:1-723. HEINRICH, C. 1923. Revision of the North American moths of the sub- family Eucosminae of the family Olethreutidae. U.S. Nat. Mus Bull. 123:1-298. . 1929. Notes on some North American moths of the subfamily Eucosminae. Proc. U.S. Nat. Mus. 75:1—-23. Kearrotr, W. D, 1907. New North American Tortricidae. Trans. Amer. Entomol. Soc. 33:1-98. McDunnoucu, J. 1939. Check List of the Lepidoptera of Canada and the United States of America. Part II Microlepidoptera. Mem. South. Calif. Acad. Sci. 2:3-171. MILLER, W. E. 1987. Guide to the Olethreutine moths of midland North America (Tortricidae). U.S.D.A. For. Serv. Agric. Hand- book 660:1—104. POWELL, J. A. 1983. Tortricidae, pp. 31-41. In Hodges, R. W. et al. (eds.), Check list of the Lepidoptera of America north of Mexico. E. W. Classey & Wedge Entomol. Res. Foundation. London, England. WALSINGHAM, T. de Grey, Sixth Earl. 1884. North American Tortrici- dae. Trans. Entomol. Soc. Lond. 1884:121-147 + 1 pl. . 1895. New Species of North American Tortricidae. Trans. En- tomol. Soc. Lond. 1895:495-518 + 1 pl. Wricut, D. J. 2005. Some Eucosmini (Tortricidae) associated with Eucosma emaciatana (Walsingham) and Eucosma totana Kear- fott: four new species, a new combination, and a new synonymy. J. Lepid. Soc. 59:121-133, . 2007. Notes on nearctic Eucosma Hiibner: a new species, a resurrected species, and three new synonymies (Tortricidae). J. Lepid. Soc. 61:38-49. Received for publication 21 February; revised and accepted 7 June 2007. VOLUME 61, NUMBER 3 Journal of the Lepidopterists’ Society 61(3), 2007, 125-137 THE BUTTERFLY DRAWINGS BY JOHN ABBOT IN THE HARGRETT RARE BOOK AND MANUSCRIPT LIBRARY, UNIVERSITY OF GEORGIA. jonn V. CALHOUN! 977 Wicks Dr., Palm Harbor, FL 34684 ABSTRACT. Artist-naturalist John Abbot completed 105 drawings of insects that are now deposited in the Hargrett Rare Book and Manu- script Library, University of Georgia. The provenance of these drawings i is unknown, but available evidence dates them to ca. 1820-1825. The adults in the 32 butterfly drawings are identified and the figures of larvae and pupae are assessed for accuracy. The illustrated plants are also identified and their status as valid hosts is examined. Abbot's accompanying notes are transcribed and analyzed. Erroneous figures of larvae, pupae, and hostplants are discussed using examples from the Hargrett Library. At least four of the butterfly species portrayed in the drawings were probably more widespread in eastemm Georgia during Abbot's lifetime. Additional key words: Larva, Lepidoptera, pupa, watercolors In 1776, the English artist-naturalist John Abbot (1751-ca.1840) arrived in Georgia, where he documented species of animals and plants for the next six decades. Living in Burke, Bullock, Chatham, and Screven Counties of eastern Georgia, he explored a region roughly bound by the cities of Augusta and Savannah, between the Oconee, Altamaha, and Savannah Rivers. Abbot longed to expand his travels, writing in 1819, “I had thoughts of taking a Trip to the back State of Tennessee to collect insects and Birds, but I think when Florida is taken possession of and settled by the United States, it will afford an ample field for collecting if Life and health permits” (William Swainson correspondence, Linnean Society of London). Abbot never fulfilled his desire to explore Tennessee and Florida, yet he worked tirelessly in Georgia for the remainder of his life. Abbot was the first to record thousands of New World species. His drawings and specimens formed the basis of numerous new taxa that were described by prominent American and European naturalists. His drawings continue to serve as an important source of information about the flora and fauna of southeastern North America. However, Abbot's illustrations and written observations often contradict our current understanding of many species (Calhoun 2007). It is helpful to examine his artwork more closely and over the course of his long career in America. I have previously analyzed Abbot's entomological contributions in Calhoun (2003, 2004, 2005, 2006a, 2006b, 2007). As another installment in this study, I present a review of Abbot's butterfly drawings that are preserved in the Hargrett Rare Books and Manuscript Library, University of Georgia. ‘Research Associate, Florida State Collection of Arthropods, DPI, FDACS, Gainesville, Florida 32614, USA METHODS I visited the Hargrett Rare Book and Manuscript Library (University of Georgia) in April, 2005. Digital photographs were taken of John Abbot's butterfly drawings and their accompanying notes. The adult butterflies were identified and the figures compared with those in other sets of Abbot's drawings that are deposited elsewhere (e.g. the Houghton Library, Harvard University, and the Alexander Turnbull Library, Wellington, New Zealand). Figures of butterfly larvae and pupae were analyzed for accuracy using written descriptions, line drawings, and photographs of living specimens. Botanist Mark A. Garland provided identifications of the depicted plants, which were then evaluated as hosts. RESULTS Analysis. The insect drawings by John Abbot in the Hargrett Library are contained in two volumes, each bound in full contemporary brown leather. The gilt spine titles read “MOTHS OF GEORGIA” with an indication of volume number. The board covers are adorned with elaborate gilt borders and blind tooling. Both volumes include yellow marbled endpapers. The volumes are enclosed in a modern rigid black case with a gilt spine title that reads “MOTHS OF GEORGIA/ JOHN ABBOT/WATER-COLOUR MANUSCRIPT” This set of 105 insect drawings includes Coleoptera (3), Hemiptera (1), Hymenoptera (1), Lepidoptera (99), and Orthoptera (1). The drawings are rendered on cream-colored wove paper, measuring 25.1 x 34.3 cm (9.5 x 13.5 in). Some sheets of paper possess undated watermarks of “T G & C” (T G & Co). This paper was manufactured by Thomas and Joshua Gilpin, whose mill was located north of Wilmington, Delaware from 1787 until 1837 (Gravell & Miller 1979). Other sheets with the watermark of “W B” came from the mills of William 126 Barber (Barbour), who produced paper in Berks County, Pennsylvania as early as 1S08 (Gravell & Miller 1979). Although Abbot employed English papers earlier in his career, he was now primarily using American papers. The Lepidoptera drawings are intended to portray the life history of each species and include figures of the larva, pupa, and a supposed hostplant (Figs. 1-4). Thirty-two of these drawings portray butterflies (Table 1). Drawing no. 9 was figured by Sotheby's (1985a). The accompanying eleven pages of manuscript notes are included separately and written in Abbot's hand on wove paper measuring 20.7 x 31.8 cm (8.13 x 12.5 in). They are entitled “Notes to the Drawings of Insects” and include numbered entries that correspond to the drawings. Abbot identified the insects and plants in his drawings using either English or Latin names. When he completed these illustrations he was using Latin names more regularly, even though their spelling and application were inexact (C Calhoun 2007). The insect and plant names that Abbot used in his notes are inscribed in pencil on many of the drawings in an unknown hand. Based on the calligraphic style, these inscriptions were probably added by an early owner and likely date to the early or mid- nineteenth century. For most butterflies, Abbot recorded the dates that each species “tyed up” (larva suspended prior to pupation), “changed” (pupated), and “bred” (eclosed as an adult). For skipper butterflies of the family Hesperiidae, he recorded when the larva “spun up” or “spun up in the leaves” (pupated). Virtually nothing is known about the provenance of these drawings. Sotheby's bookseller Maggs Brothers (Sotheby's 1985b, Leab & Leab 1986). In 1985, they were purchased at a auction in London for £11,000 by the London 1985a, Sotheby's Sotheby's (1985a) mistakenly believed that they were not the work of Abbot, but instead were “undoubtedly executed by a pupil or imitator.” This opinion was based on the higher quality of Abbot's earlier drawings that are preserved in The Natural History Museum, London. Sotheby's (1985a) hesitantly dated the drawings to ca. 1820. Not long after this auction, the volumes were purchased by the New York City bookseller Donald A. Heald, who sold them in 1998 to the University of Georgia (M. E. Brooks pers comm.). Also included in this sale were a set of Abbot's spider drawings and a unique copy of Smith & Abbot (1797) that contains plates printed on vellum (Calhoun 2006a). Owing to the British spelling of “Water-Colour” on the spine, the black case was added by either Sotheby's or Maggs Brothers. The name “J. McDougal” is inscribed in moder blue ink on a flyleaf of each volume. This is possibly the signature JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY of the consignee for the Sotheby's auction. Sotheby's (1985a) did not identify the prior owner. Completion. Because nothing is known about the early history of these drawings, existing evidence was used to determine their age. Aspects of Abbot's artistic methods and written observations were discussed in Calhoun (2006a, 2007). The Hargrett Library drawings are arranged in simple numerical order and are not haphazardly numbered like drawings that Abbot had completed ca. 1800-1810. Many of the butterfly compositions are duplicates of Abbot's life history drawings, ca. 1810-1815, that were ee for color plates in Boisduval & Le Conte (1829-[1837]) (Calhoun 2004). Entries in the accompanying notes for those drawings are also similar to the Hargrett Library notes. A large number of the Hargrett Library drawings are duplicated within a set of 103 watercolors that Abbot completed between 1816 and 1818 for the English naturalist William Swainson (Calhoun 2007). The associated notes and watermarks of these drawings are likewise very similar. Although Abbot duplicated compositions for many years, the: diminished quality of the Hargrett Library drawings i is reminiscent of his later work. It seems that Abbot abandoned life history drawings during the late 1820s in favor of less complex geometric patterns of adult insects. Abbot employed a greater number of Latin names for the Hargrett Library drawings than those for William Swainson. Over the years, Abbot less often recorded when butterfly larvae ‘ ‘tyed up.” The notes for the Swainson drawings include 17 such references, while those at the Hargrett Library include only three. Some of the Hargrett Library notes incorporate additional comments that are not found in the Swainson set. The binding of the Hargrett Library volumes is very similar to copies of Smith & Abbot (1797) that were bound in Britain around 1825 (Calhoun 2006a). Based on this evidence, the Hargrett Library drawings were probably completed ca. 1820-1825. They may represent some of Abbot's last drawings of this type. After residing in London for over 160 years, the drawings were rete to within 270 km (168 mi) of their origin in Bullock Co., Georgia, where Abbot lived from 1818 until his death. DISCUSSION Erroneous associations. Abbot's _ life ; drawings frequently deviate from reality and those in the Hargrett Library are no exception. His figures of larvae and pupae are sometimes inconsistent with the associated adults. Others are too imprecise to identify, clearly fabricated, or “borrowed” from his illustrations of other species. The depicted hostplants are often untenable or require confirmation (Calhoun 2006a, history VOLUME 61, NUMBER 3 127 Fics. 1-4. John Abbot butterfly drawings in the Hargrett Library. 1, Zerene cesonia (no. 17). 2, Limenitis archippus (no. 10). 3, Autochton cellus (no. 21). 4, Atlides halesus (no. 31) (erroneous larva and hostplant). 128 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TaBLe 1. Adult butterflies, early stages, and plants depicted in John Abbot drawings in the Hargrett Rare Book and Manuscript Library. Abbot’s original manuscript entries are included for each (Abbot's grammar and spelling are preserved). Insect nomencla- ture follows Opler & Warren (2003). Adult insect figures: D=dorsal, V=ve tral, m=male, f=female. Early stages: L=larva, P=pupa, a=acceptable, u=unacceptable. Status of figured hostplants (in brackets): C=confirmed, NC=needs confirmation, E=erroneous. Drawing Figured adults Plant species and Manuscript entry by J. Abbot No. and early stages host status 6 Papilio glaucus Linnaeus Styrax americanus Lam. (Styracaceae)[C] 6. Papilio Glaucus. The Caterpillar feeds on the plant, figured, Styrax laevigata, Swamp Ash and Hiccory, Tyed itself up by the tail 11" Oct’ Changed the 13" into Chrysalis. The Butterfly was bred 24 April. It also breeds again in the Summer. The Caterpillar is very rare, and the Butterfly not common. Df, Vf, La, Pa “Styrax laevigata” is a synonym of S. americanus. “Swamp Ash” (probably Fraxinus pennsylvanica Marsh.) (Oleaceae) and “Hiccory” [hickory] (Carya sp.) (Juglandaceae) are also confirmed hostplants. NOTES: only the dark form of the female is portrayed. Duplicate figures by Abbot were reproduced for Plates 8 and 9 of Boisduval & Le Conte (1829-[1837]). It is interesting that Abbot followed contemporary wisdom in treating this form as a separate species from the nea in drawing no. 8, especially since he reared both and noted that each fed on “Swamp Ash”. Moreover, he portray ayed slightly different immatures in these drawings. Abbot possibly knew the truth about this form, but was hesitant to refute more “learned” naturalists who were also paying customers. 0 Papilio cresphontes Cramer Zanthoxylum clava-herculis L. (Rutaceae) [C] 7. Papilio Thoas. Feeds on the Xanthoxylum Clava sterenlus, and the “Xanthoxylum Clava sterenlus” is a misspelled orange tree. Tyed up the 6" May; changed 7 a aip: Dm, Vm, La, Pa reference to Z. clava-herculis (Rutaceae). the 7th bred 27" another that changed the “Orange tree” (Citrus sp.) (Rutaceae) is also a 15" May, was bred 34 June, and another confirmed hostplant. that changed the 30" June, bred 19" July. It frequents in, and in the neighbourhood of Savannah, but is not to be met with a few miles inland. NOTES: duplicate figures by Abbot were reproduced for Plates 12 and 13 of Boisduval & Le Conte (1829-{1837]). The cultivation of orange trees was probably responsible for the occurrence of this butterfly “in the neighbourhood of Savannah.” Specimens of P. cresphontes were generally identified as Papilio thoas Linnaeus until they were recognized as a different species. 8 Papilio glaucus Linnaeus Ptelea trifoliata L. (Rutaceae) [C] 8. Papilio Eq. Gr. Turnus. Feeds on the Ptelia trifoliata, and Swamp Ash, changed “Swamp Ash” (probably Fraxinus the 20" June, bred 4" July. May be met Dm, Vm, La, Pa 5 eaceae he pennsylvanica Marsh.) (Oleaceae) is alsoa with thinly scattered in most parts of the confirmed hostplant. Country NOTES: see drawing no. 6. Duplicate figures by Abbot were reproduced for Plates 6 and 7 of Boisduval & Le Conte (1829-[1837]). Abbot's name for the butterfly was derived from the Linnaean classification system; “Eq.” refers to the group Equites (Eques) and “Gr.,” a mistake for “Tr.” refers to the subgroup Trojani (Troés). This subgroup is also in error, as the name Papilio turnus L. was originally placed by Linnaeus into the subgroup Achivi. 9 Papilio palamedes Drury Magnolia virginiana L. (Magnoliaceae) [E] 9. Papilio Chalcas. Feeds on the Magnolia Glauca, changed the 31% May, bred 14" Dm, Vm, La, Pa “Magnolia Glauca” is a synonym of M. June another that changed tie 18" Sep" virginiana. bred the 24'" March. Continues to breed all the Summer, and is frequent all over the Country NOTES: a duplicate drawing by Abbot was figured in Calhoun (2007). Duplicate figures of the larva and pupa by Abbot were reproduced for Plate 5 of Boisduval & Le Conte (1829-[1837]). The erroneous association of this butterfly with Magnolia virginiana was discussed in Calhoun (2007). The pupa is too colorful, but conceptually accurate. “Papilio Chalcas” (i.e. Papilio chalcas Fabricius) is now considered to be a junior synonym of P. palamedes. VOLUME 61, NUMBER 3 129 TABLE 1. Continued Drawing Figured adults Plant species and Manuscript entry by J. Abbot No. and early stages host status 10 Limenitis archippus (Cramer) Licania michauxii Prance 10. Black veined orange Butterfly. Feeds (Chrysobalanacae) [NC] on the species of Papaw figured, but is most frequent on Willow changed the 31" “Papaw” apparently refers to the depicted July, bred 7" August, neither the plant (commonly known as gopher apple), Caterpillar or Butterfly is common. though Abbot validly used this name for species of Asimina (Annonaceae). “Willow” (Salix sp.) (Salicaceae) is a valid hostplant. Dm, Df, Vf, La, Pa NOTES: see Fig. 1. Portions of a duplicate drawing by Abbot were reproduced for Plate 55 of Boisduval & Le Conte (1829-[1837]). The depicted plant may be the result of an association error or an aesthetic substitution. Based on a penciled inscription on a duplicate drawing at Harvard University, Scudder (1888-1889) identified the plant as Chrysobalanus oblongifolius Michx., which is now considered to be a synonym of L. michauxii. 11 Astererocampa clyton (Boisduval & Le — Vaccinium stamineum L. (Ericaceae) [E] 11. Orange coloured Butterfly. Feeds on Conte) the wild Gooseberry, changed 21* May, “Sugarberry” refers to Celtis. bred 9" June, is very rare. Dm, Df, Vf, La, Pu NOTES: Calhoun (2007) figured a duplicate drawing by Abbot and discussed the erroneous larva, pupa, and hostplant in this composition (see text). Scudder (1888-1889) identified the larva as P. interrogationis and the pupa as Polygonia comma (Harris). However, I have found no evidence that Abbot ever encountered P. comma in Georgia, nor does the larva resemble that species. 12 Chlosyne gorgone (Hiibner) Helianthus divaricatus L. (Asteraceae) [C] 12. Cross wort Frittilary Butterfly. Feeds on the Cross wort, and sunflower, t “Cross wort” apparently refers to H. changed 17" May, bred 26". Frequents Dm, Df, Vm, La, Pa divericatus (see Calhoun 2003). This is the Oak Woods of Burke County but is possibly a misapplication ofacommon __ not in the lower parts of the Country. name for the British yellow-flowered herb, Cruciata laevipes Opiz (Rubiaceae). “Sunflower” probably indicates another species of Helianthus. NOTES: : this drawing was figured in Parkinson & Rogers-Price (1984) and Calhoun (2003). Duplicate figures by Abbot were reproduced for Plate 46 of Boisduval & Le Conte (1829-[1837]) to accompany the original description of the enigmatic taxon Melitaea ismeria (Calhoun 2003, 2004, 2005). It is believed that Abbot's mention of “the lower parts of the Country” refers to bottomland habitats. This phrase, not included in the accompanying notes for three other known duplicates of this drawing, offers further evidence that Abbot did not consider C. gorgone to be the same species as Chlosyne nycteis (Doubleday) as suggested by Gatrelle (2003). The larva in this composition is conceptually consistent with C. gorgone. “Frittilary” is a misspelling of the British name “Fritillary.” 13 Phyciodes phaon (Edwards) Chrysopsis mariana (L..)Elliott 13. Small Frittilary Butterfly. Feeds on (Asteraceae) {[NC/E] the flower figured, changed the 10" June, Dm bred 21". The Caterpillar is rare, but the Butterfly is frequent in all parts of the Phyciodes tharos (Drury) Country, the whole Summer. Df, Vf, La, Pa NOTES: Abbot obviously did not distinguish between these two species of butterflies. The larva and pupa are most consistent with P. tharos. The plant may be a possible natural host of P. tharos, but not of P. phaon, which is known to feed almost exclusively on species of Phyla (Verbenaceae) (see text). 14 Asterocampa celtis (Boisduval & Le Celtis cf. tenuifolia Nutt. (Celtaceae) [C] 14. Papilio Portlandia. Feeds on the Conte) Sugar berry, changed 7" May, bred 20". “Sugarberry” refers to the figured Celtis. Is very rare Dm, Df, Vm, Lu, Pa NOTES: portions of a duplicate drawing by Abbot were reproduced on Plate 57 of Boisduval &Le Conte (1$29-[1837]) to accompany the original description of this species. The larva, and possibly also the pupa, is A. clyton (see drawing no, 11). Boisduval & Le Conte (1829-[1837]) and Scudder (1888-1889) identified the depicted plant as Celtis occidentalis L. (Celtaceae). Abbot repeatedly misapplied the name “Papilio Portlandia” (i.e. Papilio portlandia Fabricius) to this species (Calhoun 2007). 130 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TABLE 1. Continued Drawing Figured adults Plant species and Manuscript entry by J. Abbot No. and early stages host status 15 Libytheana carinenta (Cramer) Celtis cf. tenuifolia Nutt. (Celtaceae) [C] 15. Snout Butterfly. Feeds on the Sugarberry, or Hackberry,changed 29" “Sugarberry” and “Hackberry” refers to April, bred 8" May, is rare. 7. D c : ¢ Dm, Vm, La, Pa Celtis NOTES: a duplicate drawing by Abbot was figured in Calhoun (2004). With the exception of the adult figures, most of another duplicate drawing by Abbot was reproduced for Plate 64 of Boisduval & Le Conte (1829-[1837]). Scudder (1888-18889) identified the plant in duplicate drawings as Celtis occidentalis L. (Celtaceae). Species of Libytheidae have long been called “Snout” butterflies in Britain. 16 Ascia monuste (Linnaeus) Cleome gynandra L. (Capparaceae) [C] 16. White Butterfly Vanessa. Feeds on the Cleome pentaphillas. changed 17" July, Dm, Df(2), Vf, La, Pa “Cleome pentaphillas,” a misspelling of — bred 23", many of the female Butterflies C. pentaphylla L., is a synonym of C. varies being of a dingy black as figured, gynandra. They are some Years in plenty in, and about Savannah. NOTES: duplicate drawings by Abbot were figured in Gilbert (1998) and Calhoun (2004). Duplicate figures by Abbot were reproduced for Plate 16 of Boisduval & Le Conte (1829-[1837]) (Calhoun 2004). Abbot’s notes aptly describe the irregular migratory presence of the subspecies A. m. phileta (Fabricius) in coastal Georgia (Calhoun 2004). Abbot's name for this species, “Vanessa, is misapplied. In an 1813 letter, the botanist William Baldwin noted that the figured hostplant, C. gynandra, grew “spontaneously about the suburbs of Savannah” (Darlington 1843). 17 Zerene cesonia (Stoll) Dalea pinnata (J. F.Gmelin)Bameby 17. Clouded yellow Butterfly. P. Philippi. (Fabaceae) [C] Feeds on the plant figured, changed 19" : April, bred 2° May, continues to breed all Dm, Df, Vm, La, Pa the Summer and Autumn. Is most common in the Pine woods. often settles several together to suck the moist places in roads, and other places. NOTES: see Fig. 1. The late season form of this species is portrayed. Duplicate figures of the larva and pupa by Abbot were reproduced for Plate 22 of Boisduval & Le Conte (1829-[1837]). Boisduval & Le Conte misidentified the depicted plant as Tagetes papposa Vent., a synonym of Dyssodia papposa (Vent.)Hitche. (Asteraceae). Abbot’s Latin name, “P. Philippi,’ is a misspelling of Papilio phillipa Fabricius, a junior synonym of Papilio cesonia. 18 Cercyonis pegala (Fabricius) Panicum sp, possibly P. dichotomiflorum 18. Great meadow brown Butterfly. Feeds Michx. or P. rigidulum Nees (Poaceae) on the grass figured, and other grasses, Dm, Df, Vf, La, Pa [NC] changed 20" June, bred 5" July. Frequents the Pine woods, is not common. NOTES: portions of a duplicate drawing by Abbot were reproduced for Plate 59 of Boisduval & Le Conte (1829-[1837]) (the figure of the larva was reversed). Additional comments about this composition were included in Calhoun (2007). Abbot’s English name for this butterfly was derived from its superficial resemblance to the common European butterfly, Maniola jurtina (L.), known in Britain as the meadow brown since the early eighteenth century. 19 Cyllopsis gemma (Hiibner) Panicum sp. (Poaceae) [NC] 19. Swamp brown Butterfly. Feeds on the grass figured and other grasses, Tyed up F This species feeds on grasses, but natural 10" April, changed the 11" bred 24". Dm, Df, Vm, Lu, Pu hostplants are poorly known. Frequents Swamps and hammocks, is not common. NOTES: most of a duplicate drawing by Abbot was reproduced for Plate 62 of Boisduval & Le Conte (1829-[1837]). The head of the depicted larva lacks the distinctive homs of this species. This same figure of the larva is found in at least two of Abbot's drawings of Hermeupychia sosybius (drawing no. 20), suggesting that he “borrowed” it to illustrate the life history of C. gemma. Moreover, the pupa in this drawing also lacks horns and is consistent with H. sosybius. VOLUME 61, NUMBER 3 13] TABLE 1. Continued Drawing Figured adults Plant species and Manuscript entry by J. Abbot No. and early stages host status 20 Hermeupychia sosybius (Fabricius) Carex sp., possibly C. hyalinolepis 20. Small Ringlet Butterfly. Feeds on the Steudel (Cyperaceae) [NC] Grass figured, and other grasses, changed 24" August, bred 11" Sep’. Frequents the Swamps and fields, is frequent in most parts. Dm, Df, Vm, Lu, Pa NOTES: this butterfly is known to feed only on grasses (Poaceae). Abbot may have collected the wrong plant for his illustration, possibly confusing it with the host of Neonympha areolatus (J. E. Smith), which feeds on sedges (Cyperaceae). To Abbot, sedges were simply “grasses.” The depicted larva lacks the pair of posterior appendages that are found in this species. Abbot included a more accurate larva in at least two other drawings of this species, but later applied it to C. gemma (see drawing no. 19 and text). The name “Ringlet” was derived from its remote similarity to the widespread European species, Aphantopus hyperantus (L.), which has been known as the ringlet in Britain since the mid-eighteenth century. 21 Autochton cellus (Boisduval & Le Conte) Stylisma aquatica (Walter)Raf. 21. Barrd Skipper Butterfly. Feeds on (Convolvulaceae) [E] the Convolvulus figured, spun up 4" , This plant was previously placed in the — April, bred 25‘. Frequents the sides of Dm, Vm, La, Pa genus Convolvulus L. as indicated by Swamps, is rare. Abbot. NOTES: see Fig. 3. Most of a duplicate drawing by Abbot was reproduced for Plate 73 of Boisduval & Le Conte (1829-[1837]), representing the “original description” of this species. This skipper feeds almost exclusively on Amphicarpaea bracteata (L.)Fern.(Fabaceae) in eastern North America (Burns 1984). Abbot's depiction is either an aesthetic substitution or he misidentified the plant when he collected samples for this composition. Scudder (1888-1889) identified the plant on the published plate in Boisduval & Le Conte (1829—[1837]) as Breweria aquatica (Walter)A. Gray, which is now considered to be a synonym of S. aquatica. The pupa is conceptually accurate, but should be stouter in shape. 22 Problema bulenta (Boisduval & Le Panicum sp., possibly P. dichotomiflorum 22. Broad grass Skipper Butterfly. Feeds Conte) Michx. or P. virgatum L. (Poaceae) [NC] on the broad grass, folding itself in the leaf, changed 25" July, bred 6" August. Dm, Df, Vm, La, Pa “Broad grass” refers to this or a similar _ Frequents Rice fields, ditches, and the species of grass. sides of ponds in the lower parts of Georgia. Is not common. NOTES: duplicate figures by Abbot were reproduced for Plate 67 of Boisduval & Le Conte (1829-[1837]), representing the “original description” of this species. Rather than Panicum, Abbot possibly found his larvae on Zizania aquatica L. (Poaceae) or Zizaniopsis miliacea (Michx.) Déll & Asch. (Poaceae) (Calhoun 2007). 23 Euphyes arpa (Boisduval & Le Conte) — Rhynchospora latifolia (Baldwin) W.W. 23. Georgia Skipper Butterfly. Feeds on Thomas (Cyperaceae) [NC] the Grass figured, and other grasses, Dm, Df, La, Pa spun up 25" March, bred 12" April. Frequents the sides of ponds in the pine woods, is rare. NOTES: most of a duplicate drawing by Abbot was reproduced for Plate 68 of Boisduval & Le Conte (1829-[1837]), representing the “original description” of this species. This skipper normally feeds on Serenoa repens (Bartram)Small (Palmae), but Minno (1994) reared it on a species of Cyperaceae, suggesting that Abbot could have successfully reared it on this sedge (Calhoun 2004). The larva is conceptually accurate. 24 Thorybes bathyllus (J. E. Smith) Desmodium sp., possibly D. paniculatum 24. Brown Skipper. Feeds on the Beggers (L.)DC. (Fabaceae) [C] lice (figured) spun up in the leaves 1S": Dm, Df, Vf, La, Pa Oct’ bred 20" April, is not very common “Begger's lice” refers to the figured Desmodium. NOTES: a duplicate of this drawing was reproduced in ATLET (1983). It was also figured by Reynolds (1983) and Rogers-Price (1983). Portions of another duplicate drawing by Abbot were reproduced for Plate 74 of Boisduval & Le Conte (1829-[1837]). Although I have tentatively identified the figures in this composition as T. bathyllus (also see Calhoun 2007), some characters are reminiscent of Thorybes confusis Bell, making it difficult to determine the species with certainty. The plant was identified in ATLET (1983) as Desmodium fernaldii B.G.Schub. (Fabaceae). TABLE 1. Continued Drawing Figured adults No. and early stages 25 Pyrgus communis (Grote) Dm, Df, Vf, La, Pa JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Plant species and host status Sida acuta Burm. f. (Malvaceae) [C] Manuscript entry by J. Abbot 25. Black and white Skipper. Feeds on the plant figured, spun up in the leaves 25" June, bred 7" July. Is to be met with in the Oak woods and fields, is not common. NOTES: a duplicate drawing by Abbot was figured by Calhoun (2007). The skippers portrayed in this drawing are almost certainly P. communis, as there is no evidence that the similar Pyrgus albescens Plétz occurred in Georgia during Abbot's lifetime (Calhoun 2007). 26 Ancyloxypha numitor (Fabricius) Dm, Df, Vm, La, Pa Justicia ovata (Walter)Lindau (Acanthaceae) [E] 26. Least yellow Skipper. Feeds on the plant figured, changed 12" Sepr bred 22" Ts frequent in Rice fields, and meadowy parts of branches. NOTES: this species is a grass-feeder. Justicia grows in the wet habitats where this skipper occurs, thus Abbot may have confused the host. However, an earlier composition of A. numitor by Abbot includes another erroneous host, Asclepias verticillata L. (Apocynaceae), which occurs in dry soils. Abbot ambiguously referred to both plants as “the plant figured,” suggesting that he did not recall the proper host or inserted these more colorful plants to enhance his compositions. 27 Erynnis martialis (Scudder) Dm, Df, Vf, La, Pa Indigofera caroliniana Mill. (Fabaceae) [E] 27. Least dingy Skipper. Feeds on the In this case, “Wild Indigo” apparently refers to Indigofera, not a species of Baptisia (Fabaceae). “Red root, or red shank” refers to Ceanothus americanus L. (Rhamnaceae), a confirmed hostplant Wild Indigo, and Red root, or red shank, Spun up in the leaves 25t" June, bred 8'" July, Frequents the Oak woods, is much less frequent than the other Dingy Skippers. NOTES: this species is known to feed only on Ceonothus americanus L. (Rhamnaceae) in eastern North America. In fact, Abbot illustrated this skipper with C. americanus for an earlier composition, calling the plant “Red shank or red Root” (see text). Abbot’s mistaken recollection of an alternate host may have resulted in the inclusion of I. caroliniana. He called all species of the genus Erynnis “Dingy Skippers,” after the European Erynnis tages (L.), which has long been called the dingy skipper in Britain. 28 Pholisora catullus (Fabricius) Dm, Df, Vf, La, Pa Monarda punctata L. (Lamiaceae) [E] “Rignum’” is an old name for M. punctata. “Horse mint” is also a name for this plant. “Careless” (Amaranthus sp.) (Amaranthaceae) and “lambs quarter” 28. Black Skipper Catullus. Feeds on the plant figured called here Rignum and horse mint, common and red careless, and lambs quarter. Spun up 18" June. bred 26" another that spun up 29" July, was bred the 5" August, The Butterfly is (Chenopodium sp.) (Amaranthaceae) are frequent in Corn fields and plantations in confirmed hostplants. Burke County. NOTES: another drawing of this species by Abbot, which included duplicate figures of larva and pupa, was reproduced for Plate 24 of Smith & Abbot (1797) (Calhoun 2006a). Abbot's notes for that earlier drawing include some of the same life history data as for this illustration. Abbot created at least three compositions of this species and all depict Monarda, probably because it is more visually appealing than the true hosts. 29 Amblyscirtes alternata (Grote & Rrobinson) Dm, Df, La, Pa Sorghastrum secundum (Elliott)Nash (Poaceae) [NC] “Wild Oats” apparently refers to the depicted grass. Hostplants of this rare skipper are poorly known. 29. Little brown Skipper. Feeds on the wild Oats, spun up in the leaves 31° May, bred 14" June. The Caterpillar is very rare. The Butterfly frequents the pine woods, but is not common. NOTES: Abbot portrayed fresh specimens of A. alternata, which can be boldly marked as in this drawing, especially ventrally. Duplicate figures were misidentified by Scudder (1872, 1888-1889) and Beime (1950) as Amblyscirtes hegon (Scudder) (see text). VOLUME 61, NUMBER 3 TABLE 1. Continued Plant species and host status 133 Manuscript entry by J. Abbot Drawing Figured adults No. and early stages 30 Feniseca tarquinius (Fabricius) Dm, Df, Lu, Pa Vaccinium arboreum Marshall (Ericaceae) [E] “Winter Huckleberry” apparently refers to the depicted plant. “Alder” undoubtedly refers to Alnus serrulata (Aiton)Willd. (Betulaceae), which is also an erroneous hostplant. 30. Little orange Butterfly. Feeds on the winter Huckleberry, but is most frequent on Alder, it is partly covered with a white loose down, changed the 14" of April. bred the 25". The Butterfly frequents Swamps, but is rare. NOTES: duplicate figures by Abbot were reproduced for Plate 37 of Boisduval & Le Conte (1829-[1837]). This butterfly feeds only on species of aphids (see text). The depicted larva is a slightly altered version of Abbot's larva of Callophrys niphon (Hiibner), which is very unlike the spiny and more mottled larva of F. tarquinius (see text). 31 Atlides halesus (Cramer) Dm, Df, Vm, Lu, Pa Quercus phellos L. (Fagaceae) [E] “Willow Oak” is a common name for Q. phellos, which Abbot misspelled as “phillos.” 31. Great Purple hair Streak Butterfly. Feeds on the Willow Oak, Quercus phillos, changed the 20" bred 6" Sep’ is not common. NOTES: see Fig. 4. Duplicate figures by Abbot were reproduced for Plate 25 of Boisduval & Le Conte (1829-[1837]). This species feeds on mistletoes (Phoradendron sp.) (Viscaceae), which are common hemi-parasites of oaks. The depicted larva is a duplicate of the larva that he figured in drawings of Satyrium favonius (J. E. Smith) (see text). It is inconsistent with the “swollen” larva of A. halesus. 32 Parrhasius m-album (Boisduval & Le Conte) Dm, Df, Vm, La, Pa Astragalus michauxii (Kuntze)F. J. Herm. (Fabaceae) [E?] “Oaks” (Quercus sp.) (Fagaceae) are confirmed hostplants. 32. Small purple hair Streak Butterfly. Feeds on the Astragalus, and Oaks, changed 20" Aug' bred 5" Sep" may be met with in different parts of the County. but is rare in all. NOTES: duplicate figures by Abbot were reproduced for Plate 27 of Boisduval & Le Conte (1829-[1837]). The figured plant may be an erroneous host for this oak-feeder, which has also been dubiously reported to feed on other legumes. Scudder (1888-1889) identified the plant in a duplicate drawing as Astragalus canadensis L. (Fabaceae). 38 Calycopis cecrops (Fabricius) Dm, Df, Vm, Lu, Pa Vaccinium corymbosum L. (Ericaceae) [NC] “Large black Huckleberry” apparently refers to the depicted plant, but the same plant is portrayed in drawing 36 under a different name. 33. Least purple hair Streak Butterfly. Feeds on the large black Huckleberry, changed 30" April, bred 20" May is frequent in most parts of the Country. NOTES: a duplicate drawing was reproduced in ATLET (1983). The plant in a duplicate drawing was identified in ATLET (1983) as Gaylussacia frondosa (L.) Torrey & A. Gray ex Torrey) (Ericaceae). The larva of this species is not green as illustrated, but brown or pinkish-brown. 34 Strymon melinus (Hiibner) Dm, Df, Vm, La, Pa Hypericum myrtifolium Lam. (Clusiaceae) [C] “Pines” (Pinus sp.) (Pinaceae) and “snap beans” (prob. Phaseolus vulgaris L.) (Fabaceae) are confirmed hostplants. 34. Red spotted hair streak Butterfly. Feeds on the flower figured, Pines, snap beans &c. changed 30" April bred 14" May. is not very common. NOTES: duplicate figures by Abbot were reproduced for Plate 28 of Boisduval & Le Conte (1829-[1837]). These authors identified the plant in their duplicate drawing as a Hypericum, hence the name they proposed for this butterfly, Thecla hyperici. 134 TABLE 1. Continued Drawing Figured adults No. and early stages Satyrium liparops (Le Conte) Dm, Df, La, Pa JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Plant species and host status Manuscript entry by J. Abbot Crataegus sp., possibly C. viridis L. (Rosaceae) [C] “Parsley Haw” refers to Crataegus. 35. Ogechee brown hair steak Butterfly. Feeds on the Parsley Haw, changed 18" April, bred 5"" May, This species frequents the Oak woods on the sides of Ogechee river swamp, but is very rare. NOTES: unlike his other butterfly compositions, the ventral surface of the adult is not portrayed. Another drawing by Abbot was reproduced on Plate 31 of Boisduval & Le Conte (1829- [1837]) to accompany the original description of this species, which aaron poorly understood for over a century (See Calhoun 2004, 2005). Ogeechee River of eastern Georgia. 36 Callophrys henrici (Grote & Robinson) Dm, Df, Vm, La, Pa NOTES: duplicate figures by Abbot of the larva and pupa were reproduced for Plate 31 of Boisduval & Le Conte (1829-[1837]). Vaccinium corymbosum L. (Ericaceae) [C] “Swamp Huckleberry” probably refers to the figured Vaccinium, but the same plant is portrayed i in drawing 33 under a different name. “Judas tree” refers to redbud (Cercis canadensis L.) anes The adults that Abbot saw ee ree the blossoms of redbud may have included ovipositing females, as this tree is also a confirmed hostplant. “Wild plum” (Prunus sp.) (Rosaceae) is also a confirmed hostplant. ‘Ogechee” is a misspelled reference to the occurrence of this species in the vicinity of the 36. Swamp brown hair streak Butterfly. Feeds on the Swamp Huckleberry, changed 20" April, bred 6" May. frequents the blossoms of the Judas tree, and wild plums, on the sides of swamps, is far from common. Pupae of this species typically overwinter, but Abbot's notes suggest that his larva developed into an adult during the same season. ~l Callophrys irus (Godart) Dm, Df, Vf, Lu, Pa Cyrilla racemiflora L. (Cyrillaceae) (E) 37. Little brown hair streak Butterfly. Feeds on the plant figured &c. changed 22" June, bred 20" March is very rare. NOTES: most of a duplicate drawing by Abbot was reproduced for Plate 32 of Boisduval & Le Conte (1829-[1837[) to accompany the original description of Thecla arsace Boisduval & Le Conte, now considered a subspecies of C. irus. Scott (1986) incorrectly attributed the hostplant association in this composition to C. henrici. Gatrelle (1999) claimed that Scudder (1888-1889) unsuccessfully attempted to rear C. henrici or C. irus on C. racemiflora as figured, but Scudder actually referred to Leucothoe racemosa (L.)A. Gray (Ericaecea). Nonetheless, it is probable that neither of these butterflies would accept C. racemiflora, particularly C. irus. The illustrated phenotype of this butterfly feeds primarily on species of Baptisia (Fabaceae). Although Gatrelle (1999) considered the duplicate figure of the larva in Boisduval & Le Conte (1829-[1837[) to be C. irus, its shape and coloration are more consistent with C. henrici. VOLUME 61, NUMBER 3 2007). Using unpublished and published references, including Allen et al. (2005), Minno et al. (2005), Robinson et al. (2002), and Wagner (2005), I have attempted to evaluate the validity of the associated figures in Abbot's drawings in the Hargrett Library (Table 1). Larvae and pupae were considered “acceptable” if they exhibit fundamental characteristics of the given species. At least ten of the 32 butterfly drawings in the pee Library (nos. 9, 11, 13, 21, 26, 27, 28, 30, 31, 37) portray erroneous hostplants (Table 1). Two of these emphasize the dubious nature of some of Abbot's compositions. Drawing no. 30 portrays North America’s only carnivorous butterfly, Feniseca tarquinius (Fabricius). Abbot repeatedly illustrated the larva of this species resting on partially eaten leaves of Vaccinium arborium Marshall (Ericaceae) (as in the Hargrett composition), Viburnum dentatum L. (Adoxaceae), and Crataegus sp. (Rosaceae). He wrote in his accompanying notes that this butterfly most often feeds on alder, Alnus serrulata (Aiton)Willd. (Betulaceae). Alder is a common host of wooly alder aphids (Prociphilus tessellatus (Fitch)), upon which F. tarquinius larvae often feed. Abbot even observed that the larvae were “partly covered with a white loose down,” actually a waxy secretion of wooly alder aphids. Entomologists were misled by these drawings until the biology of SN species was fully revealed oe Edwards (1 886). Drawing no. 31 portrays Atlides halesus (Cramer) with a sprig of willow oak, Quercus phellos (L.) (Fagaceae) (Fig. 4). Although Abbot also referred to this plant in his notes (Table 1), larvae of A. halesus feed only on mistletoe (Phoradendron serotinum (Raf.)M. C. Johnston) (Viscaceae), which is a common hemi-parasite of oaks in Georgia. Mature larvae of A. halesus wander from mistletoe to pupate (Wagner 2005). Abbot probably found larvae of F: tarquinius and A. halesus on the figured plants and simply assumed that they were the hosts. In addition, the larvae in both of these dr: awings are inaccurate, suggesting that Abbot illustrated these life histories at a later date. He may have been unable to relocate larvae for his compositions and rendered facsimiles based on the larvae of other species. The larva that he figured in his compositions of F. tarquinius is a slightly altered version of the larva that he included in drawings of Callophrys niphon (Hiibner). His larva of A. halesus is a duplicate of the larva that he used for later drawings of Satyrium favonius (J. E. Smith). Abbot must have perceived some connection between these butterflies and assumed that their larvae were analogous. Abbot similarly “borrowed” his larva of Hermeuptychia sosybius (Fabricius) (no. 20) for his life history 135 illustrations of Cyllopsis gemma (Hiibner) (no 19). He subsequently rendered a new, though inaccurate, replacement larva for H. sosybius. Abbot's illustration of the butterfly Asterocampa clyton (Boisduval & Le Conte) (no. 11) is one of the most remarkable examples of his proclivity to invent figures (see Calhoun 2007, Fig. 2). He mistakenly applied the larva and pupa of A. clyton to the closely related Asterocampa celtis (Boisduval & Le Conte). He fabricated figures for A. clyton, modeling them after Polygonia interrogationis (Fabricius), another orange butterfly that he found feeding on the same Celtis trees (Celtaceae). Abbot occasionally confused species of Lepidoptera, resulting in erroneous host associations (Calhoun 2006a, 2007). Drawing no. 13 in the Hargrett Library portrays two species of similar butterflies as male and female of the same species. The male is Phyciodes phaon (Edwards), while the females are Phyciodes tharos (Drury) (Table 1). The larva and pupa are most consistent with P. tharos. Larvae of P. tharos feed on Asteraceae, thus the depicted plant, Chrysopsis mariana (L.)Elliott (Asteraceae), could conceivably serve as a host. On the other hand, P. phaon feeds almost exclusively on species of Phyla (Verbenaceae). Duplication. Abbot duplicated most of his butterfly life history compositions for 20-25 years (C Calhoun 2007). At least 31 of the 32 butterfly drawings in the Hargrett Library are duplicated in other sets of Abbot's illustrations: including the life history drawings that were copied for plates in Boisduval & Le Conte (1829-[1837]) (Table 1). Those original drawings are believed lost, but their notes are deposited in the Houghton Library, Harvard University (Calhoun 2004). I previously attempted to match the entries in the Houghton Library notes with Abbot's drawings that appeared in Boisduval & Le Conte (1829- [1837 ]) (Calhoun 2004). Lacking other evidence, I tentatively attributed six of these entries solely on the basis of the limited information in the notes. Duplicate drawings and notes that I subsequently discovered in the Hargrett Library were instrumental in confirming my identifications of three of these entries. The Hargrett Library set also shares duplicate figures of adults, larvae, and pupae with other plates in Boisduval & Le Conte (1829-[1837]). These drawings, begun in 1813, are now deposited in the Thomas Cooper Library, University of South Carolina (Calhoun 2004). Abbot’s observations. The illustrations and notes of John Abbot represent a valuable window through which we can explore the natural history of Georgia before it was substantially altered by human dev elopment. Changes were already affecting the local flora and fauna 136 during the early nineteenth century (Calhoun 2007). Several of the butterflies portrayed in the Hargrett Library now appear to be less widespread in eastern Georgia than during Abbot's lifetime. These include Autochton cellus (Boisduval & Le Conte) (no. 21), Problema bulenta (Boisduval & Le Conte) (no. 22), Euphyes arpa (Boisduval & Le Conte) (no. 23), Pyrgus communis (Grote) (no. 25), and Erynnis martialis (Scudder) (no. 27). In fact, P. bulenta was suspected of being an imaginary species until it was rediscovered in 1925. Pyrgus communis is possibly being displaced in eastern Georgia by the closely related Pyrgus albescens (Plotz) (Calhoun 2007). Abbot was the first to document the life histories of virtually all the species that he illustrated. Many of his drawings were the only available source of this information for over a century. The larva and pupa of A. cellus were not observed again until 1934 (Clark 1936). Abbot illustrated FE. martialis with Ceonothus americanus (L.) (Rhamnaceae) over 150 years before Burns (1964) confirmed this association. Although the Hargrett Library drawing of E. martialis portrays an erroneous —hostplant, Abbot still referred to C. americanus in his accompanying notes, calling it “Red root, or red shank.” Until very recently, researchers knew nothing about the life history of Ambyscirtes alternata Cue & Robinson) (aio: 29). Abbot's unpublished drawings of this species were overlooked because they had been misidentified by Scudder (1872, 1888-1889) and Beirne (1950) as Amblyscirtes hegon (Scudder). Although the figured hostplants need confirmation, the early stages in these drawings are consistent with A. alternata. Despite his artistic indiscretions, Abbot's illustrations continue to offer precious insight into the natural history of an early Georgia. ACKNOWLEDGEMENTS Thanks are extended to Mary Ellen Brooks (Director of the Hargrett Rare Book and Manuscript Library) for providing ac- cess to the drawings in her care and serving as a gracious host during my visit. ‘Nelson Morgan photographed the figured drawings and granted permission to publish them. Mark A. Gar- land, former State Botanist of Florida, kindly identified the plants portrayed in the butterfly drawings. Beverly Pope (Divi- sion of Plant Industry Library, (enineeviiles Florida) helped me to obtain necessary literature. James K. Adams, Irving L. Finkelstein, and Eric H. Metzler critically reviewed the manu- script and offered helpful suggestions. LITERATURE CITED ALLEN, T. J., J. P. Brock, & J. GLASsBERG. 2005. Caterpillars in the field and garden: a field guide to the butterfly caterpillars of North America. Oxford Univ. Pr., New York, New York. 232 pp. ATLET (Alexander Turnbull Library Endowment Trust). 1983. John JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Abbot's insects of Georgia. Reproduced from the original water- colours in the Alexander Turnbull Library, Wellington, New Zealand. Fascicle one. Alex. Turnbull Lib. Endowment Trust, Wellington, New Zealand. [18] pp., 6 pl. BEIRNE, B. P. 1950. Some original paintings by John Abbot. Lepid. News 4(3):25-26. BoIsDUVAL, J. B. A. D. DE & J. E. LE Conte. 1829-[1837]. Histoire générale et iconographie des Lépidoptéres et des chenilles de Amérique septentrionale. Librairie Encyclopédique de Roret, Paris. 228 pp., 78 pl. Burns, J. M. 1964. Evolution in skipper butterflies of the genus Erynnis. Univ. California Publ. Entomol. 37:1-216. 1984. Evolutionary differentiation. Differentiating gold- banded skippers—Autochton cellus and more (Lepidoptera: Hes- periidae: Pyrginae). Smiths. Contrib. Zoology, No. 405. 38 pp. CALHOUN, J. V. 2003. The history and true identity of Melitaea isme- ria (Boisduval & Le Conte): a remarkable tale of duplication, mis- interpretation and presumption. J. Lepid. Soc. 57:204-219. . 2004. Histoire générale et iconographie des Lépidoptéres et des chenilles de ! Amérique septentrionale by Boisduval & Le Conte (1829-[1837]): original drawings for the engraved plates and the true identities of four figured taxa. J. Lepid. Soc. 58:143-168. . 2005. An early drawing of Chlosyne gorgone (Hiibner) (Nymphalidae) ) by John Abbot. J. Lepid. Soc. 59:121-122. . 2006a, A glimpse into a “flora et entomologia”: The Natural History of the Rarer Lepidopterous Insects of Georgia by J. E. Smith and J. Abbot (1797). J. Lepid. Soc. 60:1-37. . 2006b. John Abbot's “lost” drawings for John E. Le Conte in the American Philosophical Society Library, Philadelphia. J. Lepid. Soc. 60:211-217 . 2007. John Abbot's butterfly drawings for William Swainson, including general comments about Abbot's artistic methods and written observations. J. Lepid. Soc. 61:1-20. Ciark, A. H. 1936. The gold-banded skipper (Rhaboides cellus) (with eight plates). Smiths. Misc. Coll. 95(7 ):1-50, § pl. DARLINGTON, W. 1843. Reliquiae Baldwinianae: selections from the correspondence of the late William Baldwin, M. D. surgeon in the U. S. Navy, with occasional notes, and a short biographical memoir. Kimber & Sharpless, Philadelphia. 346 pp. Epwarps, W. H. 1886. On the history and the preparatory stages of Feniseca tarquinius, Fabr. Can. Entomol. 18:141-153. GATRELLE, R. R. 1999. An evolutionary subspecific assessment of De- ciduphagus henrici (Lycaenidae) based its utilization of Ilex and non-Ilex hosts: description of a third Hex associated subspecies. Designation of a neotype and type locality for Deciduphagus irus. Taxon. Rpt. 1(6):1-10. . 2003. A taxonomic review of Chlosyne ismeria with descrip- tion of a new subspecies from the southern Appalachian moun- tains. Taxon. Rpt. 4(4):1-15. GILBERT, P. 1998. John Abbot: birds, butterflies and other wonders. Merrell Holberton, London. 128 pp. GRaVELL, T. L. & G. MILLER. 1979. A catalogue of American water- marks 1690-1835. Garland Publ., New York, New York. 230 pp. Leas, K. Y. & D. J. Leas (eds). 1986. American book prices current. Vol. 91: the auction season September 1984—August 1985. Ban- croft-Parkman, Inc. New York, New York. 1131 pp MINNO, M.C. 1994. Immature stages of the skipper butterflies (Lep- idoptera: Hesperiidae) of the United States: biology, morphology, and descriptions. Unpubl. Ph.D. dissertation. Univ. Florida, Gainesville. 509 pp MINNO, M. C., J. F. BuTLer, & D. W. HALL. 2005. Florida butterfly caterpillars and their host plants. Univ. Pr. Florida, Gainesville. 360 pp. OpLeR, P. A. & A. D. WARREN. 2003. Butterflies of North America. 2. Scientific names list for butterfly species of North America, north of Mexico (May 15, 2003, with additions). Contrib. C. P. Gillette Mus. of Arth. Biodiv., Colorado State Univ., Ft. Collins, Colorado. 83 pp. PARKINSON, P. G. & V. ROGERS-PRICE. 1984. Pioneer naturalist re- VOLUME 61, NUMBER 3 vived. John Abbot emerges from cocoon of neglect. Australian Antique Coll. 28:55-58. REYNOLDS, E. P. 1983. John Abbot, pioneer naturalist. Georgia Re- view 37:816-S24. RocERs-PRrIcE, V. 1983. John Abbot in Georgia: the vision of a natu- ralist artist (1751—ca.1840). Madison-Morgan Cult. Ctr., Madison, Georgia. 149 pp. ROBINSON, G. S., P. R. ACKERY, I. J. KITCHING, G. W. BECCALONI, & L. M. HERNENDEZ. 2002. Hostplants of the moth and butterfly caterpillars of America north of Mexico. Mem. Amer. Entomol. Inst. 69:1-824. Scorr, J. A. 1986. The butterflies of North America: a natural history and field guide. Stanford Univ. Pr., Stanford, California. 583 pp, 64 pl. suey S. H. 1872. Abbott’s [sic] notes on Georgian butterflies. Can. Entomol. 4:73-77, 84-87. 1888-1889. Butterflies of the eastern United States and Canada. 3 vols. Cambridge, Massachusetts. 1958 pp., 89 pl. 137 SMITH, J. E. & J. ABBOT. 1797. The natural history of the rarer lepi- dopterous insects of Georgia, including their systematic charac- ters, the particulars of their several metamorphoses, and the plants on which they feed, collected from the observations of Mr. John Abbot, many years resident in that country. 2 vols. J. Ed- wards, Cadell & Davies, and J. White, London. 214 pp., 104 pl. SOTHEBY'S (firm). 1985a. Natural History, scientific and medical books and books from the collection of the late Irving Davis. Sotheby's, London. [158] pp. . 1985b. Price list. Sale of printed books, Tuesday/Wednesday, 2nd/3rd April, 1985. Sotheby's, London. [4] pp. Wacner, D. L. 2005. Caterpillars of eastern North America: a guide to identification and natural history. Princeton Univ. Pr., Prince- ton, New Jersey. 512 pp. Received for publication 6 February 2007; revised and accepted 17 June 2007. 138 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Journal of the Lepidopterists’ Society 61(3), 2007, 138-153 DNA BARCODES OF CLOSELY RELATED (BUT MORPHOLOGICALLY AND ECOLOGICALLY DISTINCT) SPECIES OF SKIPPER BUTTERFLIES (HESPERIIDAE) CAN DIFFER BY ONLY ONE TO THREE NUCLEOTIDES JOHN M. Burns Department of Entomology, National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, MRC 127, room E-515, Washington, DC 20013-7012, USA, email: burnsj@si.edu DANIEL H. JANZEN Department of Biology, University of Pennsylvania, Philadelphia, PA 19104, USA, email: djanzen@sas.upenn.edu MEHRDAD HaJjIBABAEI Department of Integrative Biology, University of Guelph, Guelph, ON, Canada N1G 2W1, email: mhajibab@uoguelph.ca WINNIE HALLWACHS Department of Biology, University of Pennsylvania, Philadelphia, PA 19104, USA, email: whallwac@sas.upenn.edu I g) 5 S P I AND PauL D. N. HEBERT Department of Integrative Biology, University of Guelph, Guelph, ON, Canada N1G 2W1, email: phebert@uoguelph.ca ABSTRACT. Unlike most species of Lepidoptera whose DNA barcodes have been examined, closely related taxa in each of three pairs of hesperiids (Polyctor cleta and P. polyctor, Cobalus virbius and C. fidicula, Neoxeniades luda and N. pluviasilva Burns, new species) seem in- distinguishable by their barcodes; but that is when some of the cytochrome c oxidase I (COI) sequences are short and sample sizes are small. These skipper butterflies are unquestionably distinct species, as evidenced by genitalic and facies differences and by ecologic segregation, i.e., one species of each pair in dry forest, the other in adjacent rain forest in Area de Conservacién Guanacaste in northwestern Costa Rica. This national park is the source of the specimens used in this study, all of which were reared. Larval foodplants are of no or problematic value in dis- tinguishing these species. Large samples of individuals whose barcodes are acceptably long reveal slight interspecific differentiation (involving just one to three nucleotides) in all three pairs of skippers. Clearly, the chronic practice of various taxonomists of setting arbitrary levels of dif- ferentiation for delimiting species is unrealistic. Additional key words: Burns, n. sp. Area de Conservacion Guanacaste, Costa Rica, dry forest, rain forest, foodplants, genitalia, Neoxeniades pluviasilva A DNA barcode is the base pair (bp) sequence of a short (~650 bp), standard segment of the genome (Hebert et al. 2003). In animals, this is part of the mitochondrial gene cytochrome c oxidase I (COI). Because the COI gene generally mutates at evolutionarily rapid rates, comparison of barcodes in a sample of individuals best reveals differentiation at low taxonomic levels. Hence barcodes can be extremely useful in distinguishing and identifying species. Coupling this concept with the idea of always comparing the same short length of COI across a wide diversity of taxonomic groups—and doing so with demonstrable success—is what led to the catchy name “DNA barcodes” (Hebert et al. 2003). Even though COI had been used effectively in various evolutionary and taxonomic studies at and around the species level well before this epithet appeared, in the few years since its introduction, barcodes have been used for their specific purpose with notable results and with rapidly increasing frequency. The rearing of myriad wild-caught caterpillars in Area de Conservacion Guanacaste (ACG) in northwestern Costa Rica is now approaching its thirtieth year (for information about both site and rearing process, see Miller et al. 2006, Janzen & Hallwachs 2006, Burns & Janzen 2001). DNA barcodes (of a total of 4,260 reared adults) have been able to distinguish among almost 98% of 521 previously known species of the lepidopteran families Hesperiidae (skipper butterflies), Sphingidae (sphinx moths), and Saturniidae (wild silk moths) VOLUME 61, NUMBER 3 (Hajibabaei et al. 2006, Janzen et al. 2005). Rare cases where barcodes failed, which always involved closely related congeners, are worth examining in more detail. In this paper we treat three such pairs of congeneric skipper species (noted in Hajibabaei et al. 2006:table 1). We map the ecologic separation of the species in each pair in and very near ACG. We document the species status of each member of a pair (and describe one as new) on morphologic grounds. We discuss the various degrees to which larval diets, although specialized, are unreliable for species discrimination. And we show, by scrutinizing sequence length and composition, that DNA barcodes separate the species after all. Despite the immense value of DNA barcodes and the fact that they have often indicated overlooked species, it is important to consider characters besides the barcodes themselves—a point made repeatedly in the revelation of 10 cryptic species hiding under the one name Astraptes fulgerator (Walch) in ACG (Hebert et al. 2004). 139 THE SPECIES PAIRS IN QUESTION Ecologic separation (Figs. 1-3). Each pair ae a dry-forest species and a rainforest species. Parapatry of this kind is a recurrent distribution pattern among closely related lepidopteran species in ACG. In each pair of the following list, the dry-forest species comes first: Polyctor allt Evans and P. polyctor (Prittwitz), Cobalus virbius (Cramer) and C. fidicula (Hewitson), luda (Hewitson) and N. pluviasilvua Burns (a new species described below). t pyrgine and Cobalus Neoxeniades are hesperiine genera. From our distribution data, parapatry in both pairs of hesperiine species appears to be complete (Figs. 2, 3) whereas that in the pyrgine pair does not (Fig. 1). Neoxeniades Polyctor is a genus, and Out of 211 reared individuals (wild-caught as caterpillars) of the rainforest species P. polyctor, four came from dry forest. The genitalia of these apparent strays have been KOH- dissected and thoroughly studied to be sure of Area de Conservacion Guanacaste - World Heritage Site 86°00' 85°50" 85°40" 85°30" 85°20' 85°10' W Pacis 11°10" N AMB. gp ABLE, # "2, a) bs = p aN Ad 5 D. %, < ~MINAE 44°00" TP 11°00° ~ Maritza ©) Punta Santa Elena Murciélago | c Te Ban | e ° Santa Ro: 40°50" ( area Wey —_S Nancite | Playa ORGY Naneite all 1S) = Dry Forest Playa Naranjo * aS wate = SS | Rain Forest of pt AREA, . ~ 7) BD Cloud Forest Pacific Ocean = gaaa <‘- @ Polyctor cleta & g Oo 2 4 = Poll ’ = olyctor polyctor %, e} = © Stations on s 10°40" 1k Towns 40°40’ SFNac ee + International Airport an aa 0 ASS international Border SPananays) Interamerican Highway LY (VU nilewe: H PAY Roads f aor [a H (5) Protected Area ) ‘ « 86°00" 85°50" 85°40' 85°30" 85°20' 85°10" Scale: i) 10 20 30 40 50 60 Kilometers Produced by Waldy Medina : Area de Conservacion Guanacaste - 2006 Fic. 1. Spatial distribution of Polyctor cleta and P. polyctor in and near ACG. 140 their specific determination. this essentially parapatric pair of Polyctor eat the same three species of foodplants (Table 3), and because one of these plants occurs in both rain and dry forest, female wz andering from rain forest can find an attractive foodplant in dry forest and oviposit on it. The flight of these skippers is far stronger than necessary to travel the distance involved. Of the four P. polyctor caterpillars found in dry forest, three were eating the species of foodplant most often eaten by this skipper in rain forest (and because two of those were found on the very same plant, they are probably offspring of a single female); the fourth caterpillar was eating an exceedingly common, but strictly dry-forest, species that is by far the preferred foodplant of P. cleta. A small number of P. cleta caterpillars found in disturbed ecotone between dry and rain forest, and less than 2 km from the latter, were eating the main foodplant of P. polyctor. KOH-dissection and examination of the genitalia of the four adults reared Because both species of JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY from this group gave no hint of hybridization. Morphologic differences (Figs. 4-41; Tables 1, 2). In all three pairs, the brown ground color of the adult averages paler in the dry-forest species than it does in its rainforest counterpart (Figs. 4-27). This is especially evident when comparing long series of more or less recently reared (therefore unfaded) specimens. In both sexes of Polyctor, a spot spanning the distal end of the forewing cell is hyaline in P. cleta but opaque in P. polyctor. A male secondary sex character in these species of Polyctor comprises a tuft of long hairlike scales arising near the base of the dorsal hindwing costa, as well as an elongate patch of pale specialized scales embraced by the swollen beginning of vein 7 and a similarly swollen, closely adjacent length of vein 6; in both veins, sw velling extends out to the end of the cell: and the hairlike scales are long enough to overlie the special patch. These presumably pheromone- disseminating hairs are mostly to entirely dark in P. cleta but pale (often orangish) i in P. polyctor (cf. Figs. 4 and 6). Area de Conservacion Guanacaste - World Heritage Site 86°00' 85°50' 85°40" 11°10" 11°00" Punta Santa Eleng 10°50" [4 Dry Forest Rain Forest Cloud Forest @ Cobalus virbius Cobalus fidicula ©) Stations 10°40" |e Towns +4 International Airport ANA international Border Interamerican Highway avd Roads é [2] Protected Area 86°00" 85°50' 85°40’ 0 Scale: 85°30" 85°20' 85°10' W 85°30' 85°20' 85°10° 60 Kilometers Produced by Waldy Medina Area de Conservacién Guanacaste - 2006 Fic. 2. Spatial distribution of Cobalus virbius and C. fidicula in and near ACG. VOLUME 61, NUMBER 3 Area de Conservacion Guanacaste - World Heritage Site 86°00" 85°50" 85°40' 85°30" 85°20' 85°10' W 11°10' See Punta Santa Elena Murciélago oo eee nnn nn, . Pu [) Dry Forest Rain Forest GA sCloud Forest Playa Naranjp Pacific Ocean @ Neoxeniades luda Neoxeniades pluviasilva (©) Stations J 10°40' %& Towns SN Plana rated “a + International Airport ee, aural AA international Border J fimo at Ne Ta | rene Interamerican Highway S-taopnn Sie a Roads aa) 4 Ke Protected Area 86°00' it) Scale: 85°20' 85°10' Produced by Waldy Medina __60 Kilometers Area de Conservacién Guanacaste - 2006 Fic. 3. Spatial distribution of Neoxeniades luda and N. pluviasilva in and near ACG. Though clearly variations on a theme, the male genitalia of these two Polyctor species differ in striking ways. Despite substantial individual variation, almost every genitalic part differs interspecifically to at least some extent; but it is the highly asymmetric valvae that differ most (see Table 1 and cf. Figs. : 28-33). Even the less elaborate female genitalia are notably distinct in the two species (Table 2). Both species of Cobalus, which are predominantly brown, have a conspicuous white patch both dorsally and ventrally on a distal area of the hindwing. In ACG specimens, this patch is restricted to males of C. fidicula but expressed by both sexes of C. virbius (Figs. 8-11, 20-23), except for two females in which it is barely perceptible. Both species express it more fully vent: ally 7 than dorsally. In C. fidicula the patch stops before the outer margin so as to leave a narrow strip of dark brown ground color, ventrally the patch extends from mid space Ic to vein 6, and the white of the patch looks creamy on the ventral surface. In C. virbius the patch reaches the outer margin, ventrally extends from the tornus to vein 6, and looks pure white on both wing surfaces. Lateral orange scaling—broad on the outer side of the palpus and narrow behind the eye—is bright in C. fidicula but just dully suggested, and only on the palpus, in C. virbius. Cobalus fidicula is a little larger than C. virbius, and its forewing hyaline spots are likewise larger. The os genitalia (which are symmetric) differ in two obvious respects. The ventral distal division of the valva is longer and dorsally dentate in C. fidicula (cf. Figs. 35 anal 37). The very broad uncus in dorsal view shows a pair of prominent lateral sw ellings in C. virbius (cf. Figs. 34 and 36). Neoxeniades pluviasilva Burns, new species (Figs. 3, 14, 15, 26, 27, 40, 41, 45, 46; Table 3) Etymology. The species name, a noun in apposition, comes from the Latin pluvia for rain and silva for forest. Diagnosis. This is a rainforest species whereas TABLE 1. Major differences between male genitalia of Polyctor cleta and P. polyctor. Polyctor cleta Polyctor polyctor LEFT VALVA: Curved flap extending inward from dorsal margin Narrower at base Wider at base Distal dorsal division Not expanded; teeth finer Expanded; teeth coarser Distal ventral division Bent sharply dorsad_ Evenly curved dorsad RIGHT VALVA: Lower at anterior end Higher at anterior end Distal dorsal division: Dentation on edge of | More distal; finer — More dorsal; coarser SACCUS More delicate; shorter to about gone Heavier; longer UNCUS, dorsally nr. origin Pair of longitudinal humps No longitudinal humps N. luda is a species of the dry forest (Fig. 3). Ata glance, N. pluviasilva is darker than N. luda and does not express a large, pale, outer marginal area on the ventral side of the hindwing nearly as well (cf. Figs. 26, 27 with 24, 25). In females of N. pluviasilva, the double hyaline cell spot of the forewing extensively overlaps the spot in space 2 whereas in N. luda females, this forewing cell spot overlaps the spot in space 2 little or not at all (cf. Figs. 15, 27 with 13, 25). Description. Member, with N. luda, of mainly South American N. scipio species complex—treated by Evans (1955) as a polytypic species. This complex closely related to type species of Neoxeniades, N. musarion Hayward of Rio de Janeiro and Petrépolis, Brazil. Facies: Brown ground color dark. Ventral brown ground color sexually dimorphic: warmer and rustier in male (Fig. 26); colder, with purplish gray overscaling, in female (Fig. 27). Sexual dimorphism also pronounced in size of forewing hyaline spots in space 2 and in cell: much larger in female than in male, with spots overlapping one another (cf. Figs. 15, 27 with Figs. 14, 26) [no such pronounced sexual dimorphisms in N. luda (Figs. 12, 13, 24, 25)]. Large, outer marginal, pale patch on ventral hindwing inconspicuous in male (Fig. 26) to all but nonexistent in female (Fig. 27) [patch always obvious in both sexes of N. luda (Figs. 24, 25)]. Dorsally, male always showing hyaline spot in space 1b just above mid vein 1, but spot almost always small (Fig. 14) to tiny. Female, more often than not, lacking this spot dorsally; but spot, when showing, usually tiny (Fig. 15). [In N. luda, male always with this spot, and spot usually well-expressed (Fig. 12); about two-thirds of females with this spot which, when present, more often small (Fig. 13) than tiny. ] Male genitalia: Symmetric; short anterior edge of slightly raised, dorsally dentate, distal end of valva curved somewhat cephalad in lateral view (Fig. 41) [about straight to curved somewhat caudad in lateral view (Fig. 39) in N. luda]. Female genitalia: Immediately anterior to papillae anales, sclerotized transverse plate of lamella postvaginalis with broad, rounded midventral elevation [narrower, more pointed in N. luda}. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Type specimens: Holotype: male (Figs. 14, 26, 45, 46 [see arrow]), voucher code 06-SRNP-31674 (Janzen & Hallwachs 2006), Sendero Memos, Sector Pitilla, Area de Conservaci6n Guanacaste, Costa Rica, 740 m, _ latitudel0.98171, longitude -85.42785. Deposition: National Museum of Natural History, Smithsonian Institution (USNM). Labelled (yellow): LEGS AWAY/FOR DNA. DNA barcode (658 bp) of holotype (coded MHAHH575-06106- SRNP-31674|Neoxeniades pluviasilva): AACTTTATACTTTATTTTTGGAATTTGAGCAGGAATATTAGG AACTTCATTAAGTTTATTAATCCGTACAGAATTGGGAAATCCAG GATTTTTAATTGGAAATGATCAAATTTACAATACTATTGTTACAG CTCATGCATTTATTATAATTTTTTTITATAGTTATACCTATTATAAT TGGAGGATTTGGAAATTGATTAGTACCTTTAATATTAGGAGCT CCAGATATAGCTTTCCCTCGATTAAATAATATAAGATTTTGATTA TTACCTCCTTCTTTAATACTTTTAATTTCAAGAAGAATTGTAGA AAATGGAGCTGGCACTGGATGAACTGTTTATCCCCCTCTITC CTCTAACATTGCTCATCAAGGATCATCTGTAGATTTAGCAATCT TCTCACTCCATCTAGCTGGAATTTCATCTATTTTAGGAGCTATT AATTTTATTACCACAATTATTAATATGCGAATTAAAAATTTATCTT TTGATCAAATATCTTTATTCGTGTGATCTGTTGGTATTACTGCT TIACTTTTACTCTTATCTCTACCAGTCTTAGCTGGAGCTATTAC AATATTACTTACTGACCGAAATCTTAATACTTCTTTTITTCGACC CAGCAGGAGGAGGAGATCCTATTTTATATCAACATTTATTT Paratypes: 14 males, 19 females, ACG, Costa Rica (USNM). The lone synonym of N. luda is Proteides hundurensis Mabille. Mabille’s (1891) original description of one female from Honduras clearly applies to N. luda. Foodplants: Larval foodplants of this rainforest skipper at least eight species in five genera (one introduced) of Bromeliaceae (Table 3) [no known overlap in foodplant use at species level with dry-forest, sister species N. luda; but one overlap at genus level]. Larval foodplants (Table 3). Foodplants do not distinguish sister species of the pyrgine genus Polyctor: both of these skippers eat one and the same plant species in each of three genera (Allenanthus, Coutarea, and Exostema) of the family Rubiaceae. Nevertheless, P. cleta and P. polyctor use these plants at very different frequencies. Although this may reflect different preferences of the skippers, it more likely stems from the different ecologic distributions of the foodplants: Coutarea hexandra occurs at low frequency in both rain and dry forest, A. erythrocarpus is a rainforest plant TABLE 2. Major differences between female genitalia of Polyctor cleta and P. polyctor. Polyctor cleta Polyctor polyctor Sclerotized sterigmal Smaller Larger area Overall asymmetry Not extreme Pronounced Sclerotized flap on Absent or small Very large right side of sterigma Sclerotized ostium Conspicuously Relatively flush with bursae projecting (tubelike) from surface of sterigma surface of sterigma VOLUME 61, NUMBER 3 TABLE 3. Larval foodplants of two species of Polyctor, Cobalus, and Neoxeniades in Area de Conservacion Guanacaste, northwestern Costa Rica; the number of rearing records for each plant species is given (source, Janzen & Hallwachs 2006). Polyctor cleta Rubiaceae Allenanthus erythrocarpus 9 Coutarea hexandra 7 Exostema caribaeum 1 Exostema mexicanum 124 Polyctor polyctor Rubiaceae Allenanthus erythrocarpus 44 Coutarea hexandra 166 Exostema mexicanum 1 Cobalus virbius Arecaceae Acrocomia aculeata ite Bactris guineensis 5) Cobalus fidicula Arecaceae Astrocaryum alatum 25 Bactris gasipaes (introduced) 5 Bactris gracilior 6 Bactris hondurensis 10 Chamaedorea dammeriana 1 Chamaedorea pinnatifrons 1 Cryosophila warscewiczii 1 Prestoea decurrens 1 Neoxeniades luda Bromeliaceae Aechmea bracteata 4 Aechmea magdalenae 30 Bromelia pinguin 138 Neoxeniades pluviasilva Bromeliaceae Aechmea pubescens 12 Ananas comosus (introduced) 3 Guzmania desautelsii 3 Guzmania donnellsmithii 12 Guzmania nicaraguensis 1 Pitcairnia arcuata 1 Pitcairnia atrorubens 4 Vriesea gladioliflora 19 (used by P. cleta at the very edge of the dry forest/rain forest ecotone), and E. mexicanum is a common dry- forest plant (used once by a P. polyctor that apparently strayed some 15 km from rain forest). Conversely, foodplants do seem to distinguish the species of the species pairs in the hesperiine genera Cobalus and Neoxeniades. In each pair, the rainforest skipper feeds on more species than its dry-forest counterpart. Cobalus virbius eats two plant species in two genera, and C. fidicula eight plant species in five genera, of the family Arecaceae (palms). Neoxeniades luda eats three plant species in two genera, and N. pluviasilva eight plant species in five genera, of the family Bromeliaceae (bromeliads). The species in each skipper pair share no foodplant species and just one genus. However, these considerable differences in diet may have little taxonomic significance. Palms and bromeliads are far more diverse in rain forest than they are in dry forest, so that the rainforest skippers have a wider choice. Were either species of a pair to invade the other's ecosystem, it might well find its relative’s foodplants acceptable. This conjecture is somewhat weakened by the fact that the rainforest skippers, though polyphagous, still restrict their diet to fewer species of palms and bromeliads than are available to them; and even one dry-forest species is somewhat choosy. To illustrate, the ACG dry forest provides only two species of palms for C. virbius, both of which it eats, whereas the rain forest offers >10 species of palms beyond the eight so far recorded for C. fidicula. The dry-forest skipper N. luda eats the three terrestrial bromeliad species, but not the epiphytic ones, available to it, whereas N. pluviasilvua eats both kinds of bromeliads, but seems nevertheless to ignore many of the epiphytic species at hand. DNA barcodes (Figs. 42-46). For our barcoding methods, see Hajibabaei et al. (2006:971). We determined sequence divergences among individuals in each species pair (as well as in a related outgroup species) by means of the Kimura-2-Parameter (K2P) distance model (Kimura 1980), and then showed these divergences in neighbor-joining (NJ) trees (Saitou & Nei 1987). A paper in preparation will deposit in GenBank all of the sequences used here (along with thousands more for hundreds of species in various lepidopteran families, including Hesperiidae). All voucher specimens have been deposited in the National Museum of Natural History, Smithsonian Institution. Since we could not discriminate between the species in each pair during our early barcoding efforts, we greatly increased sample sizes; but we still included some COI sequences that were too short to qualify as legitimate barcodes. Two of the resulting NJ trees 144 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 4-15. Reared adults in dorsal view of Polyctor, Cobalus, and Neoxeniades from ACG, Costa Rica (specimens in USNM). Males even-numbered, females odd-numbered. Wingspan and voucher code given for each specimen. 4, 5, P. cleta: 32 mm, 02- SRNP-32285; 40 mm, 06-SRNP-869. 6, 7, P. polyctor: 35 mm, 05-SRNP-42248; 34 mm, 03-SRNP-9639. 8, 9, C. virbius: 32 mm, 92-SRNP-6215.1; 32 mm, 92-SRNP-46. 10, 11, C. fidicula: 39 mm, 05-SRNP-23068; 36 mm, 04-SRNP-32408. 12, 13, N. luda: 49 mm, 01-SRNP-11651; 53 mm, 03-SRNP-38341. 14, 15, N. pluviasilva: 43 mm, 06-SRNP-31674; 52 mm, 05-SRNP-22927. VOLUME 61, NUMBER 3 145 Fics. 16-27. Reared adults in ventral view of Polyctor, Cobalus, and Neoxeniades from ACG, Costa Rica (specimens in USNM). Males even-numbered, females odd-numbered. Same specimens in same sequence as in Figs. 4-15. 16, 17, P. cleta. 18, 19, P. polyctor. 20, 21, C. virbius. 22, 23, C. fidicula. 24, 25, N. luda. 26, 27, N. pluviasilva. 146 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 28, 29. Asymmetric male genitalia in left lateral view of two species of Polyctor from ACG, Costa Rica (USNM), scale = 1.0 mm. 28, P. cleta, genitalia dissection code X-6165, voucher code 03-SRNP-30880. 29, P polyctor, X-6159, 02-SRNP-7128. nearly separated the two species of Polyctor (Fig. 42) and those of Cobalus (Fig. 44). However, the third tree appreciably intermixed the two species of Neoxeniades (Fig. 45). The short sequences lacked diagnostic sites and therefore compromised the NJ analysis. Subsequent exclusion of short sequences resulted in clear species separation (Figs. 43, 46). Close examination of the barcode nucleotides showed that the two species of Polyctor consistently differ at three nucleotide positions (610, 616, and 625), and the two species of Neoxeniades, at one (115). Similarly, the two species of Cobalus differ in one nucleotide (at position 181), except for two females of C. virbius (04- SRNP-21798 and 06-SRNP-22664) whose “diagnostic” nucleotide is the same as that of the C. fidicula specimens. The two females of C. virbius whose barcodes match those of C. fidicula are not the two females of C. virbius (noted above, under “Morphologic differences” [with voucher codes 92-SRNP-340 and 06- SRNP-13344]) whose hindwing facies approaches that of C. fidicula females. Because, in each pair of skipper species, the interspecific nucleotide differences are very few (compared with many species of skippers previously examined), full-length, high- quality barcode sequences (~650 bp) are critical for distinguishing the species in each pair. Of course, characters of this kind—like many others—may vary geographically. The levels at which these species are distinguished is so low that, in many other circumstances, their differences could easily qualify as nothing more than individual variation. It follows that the dlewisrnettiora of some percentage or degree of divergence as a point VOLUME 61, NUMBER 3 147 Fics. 30, 31. Asymmetric male genitalia in right lateral view of two species of Polyctor from ACG, Costa Rica (USNM),. scale = 1.0mm. 30, P. cleta, X-6165, 03-SRNP-30880. 31, P. polyctor, X-6159, 02-SRNP-7128. 32 Fics. 32,33. Asymmetric male genitalia in dorsal view of two species of Polyctor from ACG, Costa Rica (USNM), scale = 1.0 mm. 32, P. cleta, X-6165, 03-SRNP-30880. 33, P. polyctor, X-6159, 02-SRNP-7128. 148 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 34-37. Male genitalia of two species of Cobalus from ACG, Costa Rica (USNM), scale = 1.0 mm. 34, 35, C. virbius, X- 5873, 99-SRNP-6213. 36, 37, C. fidicula, X-5876, 99-SRNP-5803. 34, 36, Tegumen, uncus, and gnathos in dorsal view. 35, 37, Complete genitalia in left lateral view. VOLUME 61, NUMBER 3 149 ws Fics. 38-41. Male genitalia of two species of Neoxeniades from ACG, Costa Rica (USNM), scale = 1.0 mm. 38, 39, N. luda. X- 6372, 95-SRNP-10736. 40, 41, N. pluviasilva, X-5799, 00-SRNP-2211. 38, 40, Tegumen, uncus, and gnathos in dorsal view. 39, 41, Complete genitalia in left lateral view. 150 42 ®@ Polyctor polyctor © Polyctor cleta Polyctor enops 1% © 03-SRNP-37549|629bp © 95-SRNP-9502|630bp © 03-SRNP-30883|630bp © 04-SRNP-16175|658bp © 04-SRNP-16111|658bp © 04-SRNP-15455|658bp lO 04-SRNP-15360|658bp 0 93-SRNP-5853|612bp IO 95-SRNP-9500|614bp lO 05-SRNP-61212|615bp © 03-SRNP-37550|614bp © 03-SRNP-30882|614bp © 03-SRNP-30880|614bp 0 02-SRNP-32285|614bp 0 02-SRNP-31414|614bp © 00-SRNP-2150|614bp lO 00-SRNP-20253|614bp © 92-SRNP-4986|602bp @ 02-SRNP-27779|618bp © 98-SRNP-4183)517bp @ 05-SRNP-41712/658bp @ 80-SRNP-364|573bp @ 03-SRNP-21456|576bp @ 03-SRNP-29770)51 1bp @ 03-SRNP-21493|509bp @ 03-SRNP-9643|578bp @ 06-SRNP-31806|658bp @ 05-SRNP-42247|658bp @ 05-SRNP-41517|658bp @ 04-SRNP-61026|658bp @ 04-SRNP-61027|658bp @ 05-SRNP-42248|657bp @ 06-SRNP-31800|657bp @ 04-SRNP-61128|637bp @ 04-SRNP-56096|634bp @ 05-SRNP-34721|621bp @ 04-SRNP-60861|658bp @ 04-SRNP-60859|658bp @ 04-SRNP-21386|658bp @ 04-SRNP-14594|658bp @ 04-SRNP-14595|658bp @ 04-SRNP-23706|658bp @ 04-SRNP-61064/658bp @ 02-SRNP-31973|628bp @ 03-SRNP-9641|630bp @ 05-SRNP-6247|657bp @ 02-SRNP-7132|633 @ 05-SRNP-1141|611bp @ 05-SRNP-33391|621bp @ 05-SRNP-6792|202bp @ 04-SRNP-21385|658bp @ 04-SRNP-42603|658bp @ 05-SRNP-41516|658bp @ 02-SRNP-7127|630bp @ 03-SRNP-9701|630bp @ 95-SRNP-9527|630bp @ 03-SRNP-9706|578bp @ 03-SRNP-20028/624bp @ 03-SRNP-9639|616bp @ 02-SRNP-7128|614bp @ 03-SRNP-20449|614bp @ 03-SRNP-29891|601bp }@ 00-SRNP-14180|606bp 1 05-SRNP-24532|621bp 1 05-SRNP-24533|658bp JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 43 © 95-SRNP-9502|630bp lo 03-SRNP-37549|629bp lo 03-SRNP-30883|630bp lo 04-SRNP-15455|658bp lo 04-SRNP-15360|658bp lo 04-SRNP-16175|658bp lo 04-SRNP-16111|658bp lo 93-SRNP-5853|612bp lo 95-SRNP-9500|614bp lo 03-SRNP-30880|614bp lo 05-SRNP-61212|615bp lo 03-SRNP-37550|614bp lo 03-SRNP-30882|614bp lo 02-SRNP-31414|614bp lo 02-SRNP-32285|614bp lo 00-SRNP-2150|614bp lo 00-SRNP-20253|614bp © 05-SRNP-41712|658bp © 02-SRNP-27779|618bp @ 03-SRNP-20449|614bp le 02-SRNP-7128|614bp je 03-SRNP-9639|616bp @ 05-SRNP-33391 621 bp Je 05-SRNP-1141|611bp le 03-SRNP-20028|624bp le 04-SRNP-14594|658bp le 06-SRNP-31806|658bp le 05-SRNP-42247|658bp le 05-SRNP-41517|658bp le 05-SRNP-41516|658bp le 04-SRNP-61064|658bp le 04-SRNP-61027|658bp le 04-SRNP-61026|658bp le 04-SRNP-60861|658bp le 04-SRNP-60859|658bp le 04-SRNP-42603|658bp le 04-SRNP-23706|658bp le 04-SRNP-21386|658bp lo 04-SRNP-21385|658bp le 04-SRNP-14595|658bp le 95-SRNP-9527|630bp le 03-SRNP-9701|630bp le 03-SRNP-9641|630bp le 02-SRNP-7127|630bp le 02-SRNP-31973|628bp le 05-SRNP-42248|657bp le 06-SRNP-31800|657bp le 05-SRNP-6247|657bp le 02-SRNP-7132|633 le 04-SRNP-61128|637bp le 05-SRNP-34721|621bp le 04-SRNP-56096|634bp 11 05-SRNP-24532|621bp 1 05-SRNP-24533|658bp 1 05-SRNP-24534|658bp 0 02-SRNP-30383|645bp 1 05-SRNP-24534|658bp DD 02-SRNP-30386|644bp 0 02-SRNP-30383|645bp 0 02-SRNP-30386|644bp Fics. 42, 43. Neighbor-joining (NJ) trees based on Kimura-2-Parameter (K2P) distances for cytochrome c oxidase I (COI) of Polyctor from ACG, Costa Rica. Outgroup, P. enops. Rearing voucher code and sequence length given for each individual. To find the larval foodplant of any individual, enter its voucher code in the ACG database (Janzen & allwache 2006). 42, 19 P. cleta and 44 P. polyctor with various COI sequence lengths. 43, 17 P. cleta and 35 P. polyctor with COI barcodes of gulieiot lengths (ap- proaching or exceeding 650 base pairs) for distinguishing these species Ol VOLUME 61, NUMBER 3 ] @ 02-SRNP-28956|624bp @ 02-SRNP-4281|626bp @ 03-SRNP-1656|630bp @ 04-SRNP-30067|630bp @ 99-SRNP-12115|631bp @ 99-SRNP-5803|632bp @ 04-SRNP-20859|647bp @ 00-SRNP-1518|646bp @ 02-SRNP-29526|653bp @ 04-SRNP-42650|658bp @ 04-SRNP-42942|658bp @ 04-SRNP-61415|658bp @ 05-SRNP-20957|658bp @ 05-SRNP-20848|658bp @ 05-SRNP-201|658bp @ 06-SRNP-1014|658bp @ 05-SRNP-43578|658bp @ 05-SRNP-70497|658bp @ 05-SRNP-23068|658bp @ 04-SRNP-60363|658bp @ 03-SRNP-20043|643bp @ 03-SRNP-31144|657bp @ 99-SRNP-5802|658bp @ 06-SRNP-22121|658bp @ 02-SRNP-33898|658bp @ 02-SRNP-4283|658bp @ 02-SRNP-600|658bp @ 06-SRNP-22543|658bp @ 06-SRNP-9326|658bp @ 04-SRNP-32408|522bp @ 06-SRNP-40104|658bp @ 06-SRNP-1576|658bp @ 00-SRNP-4589|658bp @ 02-SRNP-4284|658bp @ 05-SRNP-86|620bp @ 05-SRNP-2180|618bp @ 03-SRNP-1606|621bp @ 99-SRNP-5801|579bp O 04-SRNP-21798|657bp O 06-SRNP-22664|658bp @ 04-SRNP-42312|615bp O 06-SRNP-13344|658bp O 92-SRNP-6215.1|609bp O 99-SRNP-6212|636bp O 92-SRNP-46|516bp O 92-SRNP-340|549bp O 99-SRNP-6213|600bp (1 04-SRNP-41736|658bp 1 04-SRNP-31491|658bp (1 04-SRNP-42853|658bp C0 04-SRNP-34841|658bp (1 04-SRNP-34350|658bp @ Cobalus fidicula © Cobalus virbius Aides brino 1% Fic. 44. NJ tree based on K2P distances for COI of 8 Cobalus virbius and 39 C. fidicula from ACG, Costa Rica. Outgroup, Aides brino. Rearing voucher code and sequence length given for each individual. @ 00-SRNP-2440|593b @ 03-SRNP-12943.1|598bp @ 05-SRNP-4183|596bp @ 06-SRNP-31674|658bp @ 05-SRNP-22927|614bp @ 04-SRNP-4952|600bp @ 05-SRNP-2697|596bp @ 05-SRNP-2089|591bp @ 06-SRNP-925|544bp O 95-SRNP-287|379bp @ 05-SRNP-7791|381bp @ 05-SRNP-43047|381bp @ 04-SRNP-42436|379bp @ 04-SRNP-42073|596bp @ 03-SRNP-37401|611bp @ 04-SRNP-42888|380bp @ 05-SRNP-42875|381bp @ 04-SRNP-5000|381bp @ 04-SRNP-42896|380bp O 03-SRNP-1072|387bp @ 06-SRNP-926|391bp @ 06-SRNP-928/391bp. O 93-SRNP-8348|362bp O 93-SRNP-8426|357bp O 06-SRNP-500|391 bp @ 04-SRNP-40975|381bp @ 04-SRNP-4951|381bp @ 04-SRNP-4907|380bp O 00-SRNP-2151|378bp O 00-SRNP-2217|567bp O 00-SRNP-4479|597bp O 04-SRNP-24531|591bp O 03-SRNP-38341|592bp O 06-SRNP-12188|658bp O 06-SRNP-12189|658bp O 06-SRNP-12187|658bp 45 @ Neoxeniades pluviasilva © Neoxeniades luda Neoxeniades Burns04 1 04-SRNP-31794|6: Fics. 45, 46. NJ trees based on K2P distances for COI of Neoxer described species). Rearing voucher code and sequence length given for each individual. Arrow indicates holotype. O 06-SRNP-32563|658bp 0 00-SRNP-2591|574bp JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Pp @ 03-SRNP-37355|538bp 46 <— @ 00-SRNP-2440|593bp @ 03-SRNP-37355|538bp @ 03-SRNP-12943.1|598bp @ 05-SRNP-22927|614bp @ 05-SRNP-4183|596bp @ 06-SRNP-31674|658bp <— @ 04-SRNP-4952|600bp @ 04-SRNP-42073|/596bp @ 05-SRNP-2089|591bp @ 05-SRNP-2697|596bp @ 06-SRNP-925|544bp @ 03-SRNP-37401|611bp O 00-SRNP-4479|597bp O 03-SRNP-38341|592bp O 04-SRNP-24531|591bp O 00-SRNP-2217|567bp O 06-SRNP-12188|658bp 1% © 06-SRNP-12189|658bp O 06-SRNP-12187|658bp 1 06-SRNP-32563|658bp 1 00-SRNP-2591|574bp 1 04-SRNP-31794/658bp 58bp tiades from ACG, Costa Rica. Outgroup, N. BURNS04 (an un- 45,13 N. luda and 24 N. pluviasilva with various COI sequence lengths. 46, 7 N. luda and 12 N. pluviasilva with COI barcodes of sufficient lengths for distinguishing these species. considered many be though below which individuals should conspecific is unrealistic (even taxonomists have done so, in various contexts, for a great many years). Speciation is not tidy. ACKNOWLEDGEMENTS We thank Waldy Medina for making distribution maps, Don- ald Harvey for dissecting genitalia, Young Sohn for drawing some of them, Karie Darrow for reducing “eral numbering the drawings, the ACG parataxonomists (see Burns & Janzen 2005a, 2005b, Miller et al. 2006) for finding and rearing caterpillars, Tanya Dapkey for plucking and shipping legs from reared adults for DNA analysis (i.e., “LEGS AW. ‘AY/FOR DNA”), Stephanie Kirk and Rebecca Cowling for molecular work, Janet Topan and Angela Holliss for DNA sequence analysis, and John Shuey and Felix Sperling for helpful comments. Support for this study came from the National Museum of Natural History Small Grants Program (J. M. B.); National Science Foundation grants BSR 9024770 and DEB 9306296, 9400829, 9705072, 00727 30, and 0515699 (D. H. J.); and Genome Canada through the On- tario Genomics Institute, and the Gordon and Betty Moore Foundation (P. D. N. H.). VOLUME 61, NUMBER 3 LITERATURE CITED Burns, J. M. & D. H. JANZEN. 2001. Biodiversity of pyrrhopygine skipper butterflies (Hesperiidae) in the Area de Conservacién Guanacaste, Costa Rica. J. Lepid. Soc. 55:15-43. . 2005a. Pan-neotropical genus Venada (Hesperiidae: Pyrginae) is not monotypic: Four new species occur on one volcano in the Area de Conservacién Guanacaste, Costa Rica. J. Lepid. Soc. 59:19-34. —. 2005b. What's in a name? Lepidoptera: Hesperiidae: Pyrginae: Telemiades Hiibner 1819 [Pyrdalus Mabille 1903]: new combina- tions Telemiades corbulo (Stoll) and Telemiades oiclus (Mabille)— and more. Proc. Entomol. Soc. Wash.107:770-781. Evans, W. H. 1955. A catalogue of the American Hesperiidae indi- cating the classification and nomenclature adopted in the British Museum (Natural History). Part IV. Hesperiinae and Mega- thyminae. British Museum, London. 499 pp., pls. 54-88. HayipaBakl, M., D. H. JANZEN, J. M. BurNs, W. HALLwacus & P. D. N. HEBERT. 2006. DNA barcodes distinguish species of tropical Lepidoptera. Proc. Natl. Acad. Sci. USA 103:968-971. HEBERT, P. D. N., A. Cywinska, S. L. BALL & J. R. DEWaARD. 2003. Biological identifications through DNA barcodes. Proc. Roy. Soc. Lond. B 270:313-321. Hebert, P. D. N., E. H. PENTON, J. M. Burns, D. H. JANZEN & W. HaLuwacus. 2004. Ten species in one: DNA barcoding reveals cryptic species in the neotropical skipper butterfly Astraptes ful- gerator. Proc. Natl. Acad. Sci. USA 101:14812-14817. JANZEN, D. H., M. Hajipabacl, J. M. Burns, W. HaLnwacus, E. REMIGIO & P. D.N. HEBERT. 2005. Wedding biodiversity inventory of a large and complex Lepidoptera fauna with DNA barcoding. Phil. Trans. Roy. Soc. B 360:1835-1845. (doi:10.1098/rstb.2005.1715) JANZEN, D. H. & W. Hatiwacus. 2006. Event-based database of caterpillars, their host plants, and their parasitoids in Area de Conservacién Guanacaste, northwestern Costa Rica. (http://janzen.sas.upenn.edu). Kimura, M. 1980. A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences. J. Mol. Evol. 16:111-120. MABILLE, P. 1891. Description dhespérides nouvelles. Ann. Soc. en- tom. Belgique 35:LIX—LXXXVIII. MILLER, J. C., D. H. JANZEN & W. Hatuwacus. 2006. 100 Caterpil- lars. Belknap Press of Harvard Univ. Press, Cambridge, Mass. 264 pp. Saitou, N. & M. NEL 1987. The neighbor-joining method: a new method for reconstructing phylogenetic trees. Mol. Biol. Evol. 4:406-425. Received for publication 27 March; revised and accepted 25 May 2007. 154 Journal of the Lepidopterists’ Society 61(3), 2007, 154-164 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY THE INFLUENCE OF HURRICANE AND TROPICAL STORM ACTIVITY ON RESIDENT BUTTERFLIES IN THE LOWER FLORIDA KEYS Mark H. SAtvaTo! AND HOLLy L. SALVATO 1765 17th Avenue, Southwest, Vero Beach, Florida 32962, USA, email: anaea_99@yahoo.com ABSTRACT. Butterfly populations were monitored at two coastal locations in the lower Florida Keys to observe and evaluate their re- sponse to hurricane and tropical storm activity. Four major hurricanes—Dennis, Katrina, Rita and Wilma—occurred within the vicinity of the Florida Keys during 2005. The ocean-facing exterior portions of both study areas were heavily damaged by hurricane and tropical storm force winds, salt spray and storm surge, resulting in greatly reduced butterfly abundance and species richness. More interior portions of the study areas, while inundated with floodwaters, retained the majority of their vegetation throughout the storm season allowing for an assemblage of butterflies similar in richness, albeit reduced in abundance, to pre-storm conditions. At each study area butterfly recovery time appeared re- lated to availability of appropriate host and nectar plant species. However, the decline, disappearance or slow recovery of certain butterflies suggests that storm activity had a deleterious influence on the natural histories of select butterflies. Additional key words: adverse weather, population dynamics, Cyclargus. Butterflies of the Florida Keys have adapted over time to the influence of tropical storms and other forms of adverse weather conditions (Covell 1976, Minno and Emmel 1993, Smith et al. 1994). merely mentioning the threat that hurricanes may pose to localized aornullsiieies of endangered species in the region (Minno and Emmel 1993, 1994, USFWS 1999, Calhoun et al. 2000), there is a scarcity of published data on the effects of tropical storms on butterfly populations However, aside from and their natural histories. During the active storm season of 2005, we closely monitored butterflies at two locations in the lower Florida Keys evaluate their response to hurricane and tropical storm activity. Surveys of the study areas conducted by the authors during 2004, to observe and in which no substantial storm activity occurred in the lower keys, provided a baseline for comparison. METHODS The survey areas and pre-storm butterfly diversity. Cactus Hammock, located on southeastern Big Pine Key within the National Key Deer Refuge (NKDR), contained a variety of plant communities, including coastal scrub, mangroves, salt marsh and tropical hardwood hammocks that allowed for a diversity of butterfly species. Brephidium isophthalma pseudofea ( (Morrison) (Lycaenidae) is prolific in coastal within the lower keys, including Cactus Hammock, where the species hostplants, Salicornia bigelovii L. (Chenopodiaceae) and Batis maritima L. (Bataceae) are abundant (Minno and Emmel 1993, Salvato 1998). Other species frequently encountered within Cactus Hammock historically included Junonia areas ‘Biologist, United States Fish and Wildlife Service, Vero Beach, Florida 32960, USA evarete (Cramer) (Nymphalidae), Strymon martialis (Herrich-Schiiffer) (Lycaenidae), Ascia monuste phileta Fabricius (Pieridae), Heliconius charithonia tuckeri Comstock and Brown (Nymphalidae), and Panoquina panoquinoides (Skinner) (Hesperiidae) (Salvato and Salvato, unpublished data). A 5-hectare portion of Cactus Hammock was monitored during this study to observe the possible influence of tropical storm conditions on butterfly species richness and abundance. Bahia Honda State Park (Bahia Honda), located approximately 8 km (5 miles) east of Big Pine Key, also had a variety of natural habitats, including tropical hammocks, mangroves, coastal scrub and beacon as well as an old berm that historically served as a railroad bridge. The areas surrounding this berm were heavily vegetated on both the south (Atlantic Ocean) and north (Gulf of Mexico) sides by a variety of native plant species. Our surveys at Bahia Honda were limited to an area approximately 1-hectare in size along the old railroad berm. Vegetation within the study site included Coccoloba uvifera L. (Polygonaceae), Swriana maritima (Surianaceae) and Caesalpinia bonduc Roxburgh (Fabaceae), the latter of which is a hostplant of the endangered Cyclargus —thomasi —_ bethunebakeri Comstock and Huntington (Lycaenidae) (Pyle 1981, Calhoun et al. 2000). Other butterfly species occurring on Bahia Honda prior to the survey period included Phoebus agarithe maxima (Neumoegen) (Pieridae), Agraulis vanillae nigrior Michener (Nymphalidae), H. c. tuckeri, Hemiargus ceraunus antibubastus Hiibner (Lycaenidae), Leptotes cassius theonus (Lucas) (Lycaenidae), Hylephila phyleus phyleus (Drury) (Hesperiidae) and Cymaenes tripunctus tripunctus (Herrich-Schiffer) (Hesperiidae) (Minno and Emmel 1993, Smith et al. 1994, Calhoun et al. 2000, Salvato and Salvato, unpublished data). VOLUME 61, NUMBER 3 The study areas were monitored monthly from June 2005 to February 2006 to determine butterfly abundance and species richness. A standard walking route was established at each location that allowed two researchers to observe and record butterfly activity within the varied environs of each survey location. Both locations were visited on each survey date, except during November 2005, when Cactus Hammock and Bahia Honda were surveyed on 5 November and 11 November, respectively (Bahia Honda was closed to the public due to storm damages on 5 November). Surveys were conducted on warm, clear days under conditions that were considered sufficient for butterflies to be flying. Each sampling date included approximately 6-8 hours of field time (between 8:00-16:00 h). On each sampling date approximately 4 to 5 and 2 to 3 hours was spent monitoring at Cactus Hammock and Bahia Honda, respectively. Butterfly diversity was determined on each sampling date by visually observing and recording the individuals and species encountered. Both researchers traversed the same survey route in unison, with one researcher (MHS) counting (with a hand counter) the most numerous species flying on a given sampling date while HLS tallied the remaining less abundant species. The storms. Four major hurricanes—Dennis, Katrina, Rita and Wilma—occurred within the vicinity of the Florida Keys during 2005 (Fig. 1 indicates the relative paths for each storm). Storm accounts discuss data gathered and summarized by the National Oceanic \— 4 ae 2 * cactus Hammock q> ) 6 8 10 Kilometers. . . x ” as? a Pr gussnesnent® sane and Atmospheric Administration (NOAA) for Key West. Florida (www.nhe.noaa.gov/archive/2005), which was located 48.3-56.3 km (30-35 miles) to the southwest of the study areas on Big Pine and Bahia Honda Keys. Dennis, had been a strong Category 4 hurricane (winds in excess of 241 km [150 mph]) prior to landfall in south-central Cuba, but weakened as it entered the southern Gulf of Mexico. Dennis passed approximately 137 km (85 miles) to the west of Key West on 9 July 2005 generating hurricane and tropical storm force winds in the extreme lower keys and a storm surge of up to 1.8 m (6 feet) above normal high tide levels within our coastal survey areas. Katrina, which crossed southern peninsular Florida and entered the Gulf as a weak hurricane, passed 28 km (45 miles) to the northwest of Key West on 26 August 2005 generating tropical storm force winds and storm surge on the northern side of the keys. Katrina had little influence on our study areas on southern Big Pine and Bahia Honda Keys. Rita passed within $1 km (50 miles) of Key West on 20 September 2005 and rapidly intensified as it traversed the entire stretch of the Straits of Florida. Rita's relatively close proximity generated hurricane and tropical storm force winds of greater duration and intensity than Dennis, with a storm surge up to 2.1 m (7 feet) above normal high tide levels within our study area. Wilma, which passed rapidly to the northeast of the lower keys on 24 October 2005, generated hurricane and tropical storm force winds and a storm surge up to 2.7 m (9 feet) that resulted in extensive damages on the FENLARGED AREA . A AREA a , ENLARGED oo 3 Se SN \ a! Bahia Honda Key ES) * N oO 25 50 75 100 Kilometers Fic. 1. The paths of Hurricane Dennis, Katrina, Rita and Wilma in relation to south Florida and the study areas in the lower keys. 156 Fic. 2. Exterior of Cactus Hammock, as shown on 24 Sep- tember 2005, shortly after Rita, was severely damage from hur- ricane and tropical storm force winds, salt spray and storm surge (Photo Credit; H.L. Salvato). northern side of the lower keys. However, as with Katrina, this storm had little direct impact to our survey areas on the southern sides of the islands. RESULTS Cactus Hammock-post storms. A survey conducted on 16 July 2005, one week following Dennis, found that the majority of hammock and coastal scrub habitat along the coastal exterior of Cactus Hammock had been either severely damaged or destroyed by tropical storm force winds, salt spray and storm surge. Aside from migratory species such as, A. m. phileta and P. a. maxima, no other butterfly activity was observed within the heavily damaged exterior portions of Cactus Hammock. The interior of Cactus Hammock, while inundated with floodwaters, was otherwise undamaged. These more inland portions of the survey area maintained a richness of butterfly species similar to what was recorded prior to the storm; however, overall butterfly abundance was greatly reduced, particularly for Lycaenids, such as B. i. pseudofea, L. c. theonus and H. c. antibubastus. Papilio cresphontes Cramer (Papilionidae), P. panoquinoides and H. c. tuckeri, species frequently encountered within Cactus Hammock prior to Dennis, were absent following the storm. Additional post-Dennis surveys of Cactus Hammock indicated that several butterfly species, such as J. evarete, A. m. phileta and Anartia jatrophae guantanamo Munroe (Nymphalidae), had dispersed within the survey area and were observed ovipositing on available hostplants. However, butterfly abundance and species richness within Cactus Hammock continued to decline post-Dennis, while select species remained absent. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fic. 3. Floodwaters intruded deep into Cactus Hammock fol- lowing storm activity as a result of coastal dune erosion along the southeastern portion of the study area, covering extensive areas of Salicornia (Photo Credit; H.L. Salvato). Rita passed directly to the south of the lower keys, generating storm surge, hurricane and tropical storm force winds, further adding to the damages initially caused by Dennis. Although quantitative measures on vegetation were not conducted as part of this study, we visually estimated that 60-80% of the hammock vegetation had been damaged or destroyed during Dennis and Rita (Fig. 2). Floodwaters that had receded during the interim between storms intruded deeper into Cactus Hammock following Rita as a result of coastal dune erosion along the southeastern portion of the study area (Fig. 3). Natural rehabilitation of coastal and hammock vegetation that had begun shortly after Dennis was hindered due to the influence of Rita. The abundance of select butterflies continued to decline, Fic. 4. Storm surge and salt spray from Dennis and Rita greatly damaged vegetation on the ocean side of the Bahia Honda survey area, such as Caesalpinia bonduc, host plant for the endangered Cyclargus thomasi bethunebakeri (Photo Credit; H.L. Salvato). VOLUME 61, NUMBER 3 while other species either decreased or remained absent. Overall butterfly richness within the area following Rita and into the fall of 2005 was much lower from what was noted during the previous year. Butterflies that were observed in Cactus Hammock during the late summer and fall months consisted largely of migratory species such as Phoebus sennae eubule (L.) (Pieridae), Urbanus proteus proteus (L.) (Hesperiidae) and Danaus plexippus (IES) (Nymphalidae), which were found nectaring on Stachytarpheta jamaicensis (L.) Vahl (Verbenaceae). Danaus plexippus was observed roosting on damaged and defoliated hammock trees along the coastal portions of the study site. Wilma resulted in extensive damage on the Gulf side of the keys. However, a survey of Cactus Hammock on 5 November 2005, two weeks after Wilma, found little indication of major damage from this storm. New foliage that was sprouting on trees and shrubs that had been damaged or defoliated by Hurricanes Dennis and Rita appeared unaffected by Wilma. However, butterfly species richness and abundance continued to decline. In mid- eee 2005 and into early 2006, with the exception of B. i. pseudofea, which appeared to be returning to Beseonnl abundance, overall species richness and abundance in Cactus Hammock remained lower from what had been recorded during the previous year and prior to the first storm activity of 2005. Figures 5 and 6 indicate monthly ‘butterfly species richness and abundance observed in Cactus Hammock from June 2005 to February 2006 during and following an active storm season. Also included is similar data on butterflies in the survey area collected during the same period of the prior year (June 2004 to February 2005), in which no substantial storm activity occurred within in the lower keys. Appendix 1 indicates which species were encountered on each sampling date during the survey periods in 2004-2006 and their abundance. Bahia Honda-post storms. On Bahia Honda, the ocean side of the old railroad berm was heavily damaged by storm surge and salt spray from Dennis (Fig. 4). One week after Dennis (16 July 2005), with the exception of migrating A. m. phileta, P. a. maxima and Kricogonia lyside (Godart) (Pieridae), no butterfly activity was observed on the ocean side of the survey area. We visually estimated that as much as 50% of the vegetation on the southern side of the island had been heavily damaged, including large stands of C. bonduc, S. maritima and other host and nectar plants of importance to butterflies on Bahia Honda. However, Cactus Hammock species richness 255 —— 2005-06 active season | —#- 2004-05 inactive season 20 4 ry 2 @ 154 Qa a - i} (-} iS 8 Katrina 2 104 26 Aug 2005 £ °o = § Dennis 9 July 2005 Rita 54 20 Sept 2005 6 Wilma 24 Oct 2005 0 T T T , T T T Jun Jul Aug Sep Oct Nov Dec Jan Feb months Fic. 5. Indicates monthly butterfly species richness observed in Cactus Hammock from June 2005 to February 2006 during and following an active storm season. Also indicated is similar data on butterflies in Cactus Hammock collected during the same pe- riod the prior year (June 2004 to February 2005), in which no substantial storm activity occurred in the lower keys. 158 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 800 ; 700 | Cactus Hammock abundance —e— 2005-06 active season 600 + —m— 2004-05 inactive season cy os s 2 500 5 2 a] & = © 400 4 [-) S 8 § 300 c s 2 a 200 | 100 Rita 20 Sept 2005 26 Aug 2005 iH 0+— r — c , + - + ———~ Jun Jul Aug Sep Oct Nov Dec Jan Feb months Fic. 6. Indicates monthly butterfly species abundance observed in Cactus Hammock from June 2005 to February 2006 during and following an active storm season. Also indicated is similar data on butterflies in Cactus Hammock collected during the same period the prior year (June 2004 to February 2005), in which no substantial storm activity occurred in the lower keys. the remaining portions of the survey area, including the areas on top and to the northern side of the old railroad berm itself, appeared relatively undisturbed from storm surge and salt spray. Whereas butterfly activity was larg gely absent on the ocean side of the survey area, the Gulf side retained its vegetation and numerous plants remained in bloom. Heliotropium angiospermum Murray (Boraginaceae), Bidens alba L. (Asteraceae) and Melanthera nivea Small (Asteraceae), were heavily visited by surviving butterflies. Unlike Cactus Hammock, where a butterfly abundance was noted following Dennis, the level of butterfly activity increased on Bahia Honda from densities recorded prior to the storm. Similar to Cactus Hammock, however, decline in was increased butterfly activity within areas less impacted from storm activities, which retained ample nectar sources and appropriate hostplants. The increased abundance in the select parts of this study area may have been the result of butterfly dispersal from storm- -damaged areas of Bahia Honda and surrounding islands. As a result of natural regeneration and park maintenance, much of the vegetation was observed recovering during continued post-Dennis surveys. However, populations of several butterfly species, such as C. t. bethunebakeri and H. c. tuckeri, declined steadily. Although both of these species lost large quantities of their respective hostplants on the ocean side of Bahia Honda as a result of Dennis, this reduction did not appear so substantial as to trigger such sharp declines. As with Cactus Hammock, storm surge and salt spray from Rita greatly damaged vegetation on the ocean side of the old railroad berm. The impacts to the ocean side of the island appeared more severe, perhaps due to the fact that large amounts of coastal vegetation had been damaged or thinned by Dennis. This vegetation may have served as a protective barrier for the study area during the first storm. Overall species richness and abundance in late September and into November decreased following Rita and was reduced from levels observed during the previous year or earlier in the 2005 survey period. Wilma caused extensive damage to the northeastern side of Bahia Honda; however, we observed little storm impact on our study area on the southwestern portion of the island during a survey on 11 November 2005, about three weeks after the storm. Surveys of Bahia Honda in mid-December 2005 and into early 2006 found butterfly richness and abundance similar to that observed a year prior and earlier in the survey period, indicating that butterflies were returning to relatively normal seasonal densities. VOLUME 61, NUMBER 3 159 30 Bahia Honda species richness —e— 2005-06 active season $ Katrina —m— 2004-05 inactive season 25 4 26 Aug 2005 — 204 a 2 o o a a r) - 154 2 oo 8 24 Oct 2005 = 2 10- 20 Sept 2005 5 Jun Jul Aug Sep Oct Nov Dec Jan Feb months Fic. 7. Indicates monthly butterfly species richness observed on Bahia Honda from June 2005 to February 2006 during and fol- ine a an active storm season. Also indicated is similar data on butterflies on Bahia Honda collected during the same period the prior year (June 2004 to February 2005), in which no substantial storm activity occurred in the lower keys. 1200 - Bahia Honda abundance —e— 2005-06 active season —a— 2004-05 inactive season 1000 + cy sg 800 + 2 2 I & | = | ° 6 600 - = o o = s z § 400 3 .) Katrina 26 Aug 2005 200 | a aad 20 Sept 2005 @ 24 oct 2005 0 + : - 7 : r + T Jun Jul Aug Sep Oct Nov Dec Jan Feb Fic. 8. Indicates monthly butterfly species abundance observed on Bahia Honda from June 2005 to February 2006 during and fol- lowe an active storm season. Also indicated is similar data on butterflies on Bahia Honda collected during the same period the prior year (June 2004 to February 2005), in which no substantial storm activity occurred in the lower keys. 160 Figures 7 and § indicate monthly butterfly species richness and abundance observed in Bahia Honda from June 2005 to February 2006 during and following an active storm season. Also included is similar data on butterflies in the survey area collected during the same period of the prior year (June 2004 to February 2005), in which no substantial storm activity occurred within in the lower keys. Appendix 2 indicates the species that were encountered on each sampling date during the survey periods in 2004—2006 and their abundance. DISCUSSION Butterflies that readily re-established themselves within the survey areas after the storms were those species most closely associated with salt marsh, mangrove and hammock vegetation that had rebounded rapidly after Dennis and Rita. In Cactus Hammock, A. m. phileta, P. a. maxima, B. i. pseudofea, Strymon istapa modestus Maynard (Lycaenidae), J. evarete, A. j. guantanamo, Phoicides pigmalion — okeechobee (Worthington) (Hesperiidae) and Polygonus leo savigny (Latreille) (Hesperiidae) (all dependant on plant species that quickly returned after the storms) were species that were most often re-encountered following the storms. The decline, disappearance or slow recovery of select species within the remainder of Cactus Hammock (such as H. c. tuckeri), suggests that storm activity had a deleterious influence on the natural histories of select butterflies. Species that had occurred locally within Cactus Hammock prior to the storms (such as S. martialis and P. panoquinoides), but not after them, will require ongoing monitoring to determine their status. Although recovery was rapid for many of the butterfly species within the Bahia Honda survey area following the storms, a number of species remained in decline or absent in post-storm surveys. Heliconius c. tuckeri was abundant immediately after the initial storm, but was not recorded again in the Bahia Honda survey area for the remainder of the study. Leptotes c. theonus and S. martialis disappeared from Cactus Hammock shortly JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY after Dennis, but both quickly re-established themselves on Bahia Honda and elsewhere in the lower keys. This suggests that they may rely on a more storm- impacted host plant within Cactus Hammock, resulting in the differing recovery outcomes. Despite the noted decline of the endangered C. t. bethunebakeri throughout the survey period further monitoring by the authors found that the species had returned to pre- storm abundance by the summer months of 2007. ACKNOWLEDGEMENTS The authors thank J. V. Calhoun, M. C: Minno, H. Pavulaan, B. Scholtens and M. Toliver for their insights and critical reviews that improved the quality of this paper. Barry Wood and T. A. Dean assisted with figure preparation. LITERATURE CITED CALHOUN J. V., J. R. SLOTTEN & M. H. SatvaTo. 2000. The rise and fall of tropical blues in Florida: Cyclargus ammon and Cyclargus thomasi bethunebakeri (Lepidoptera: Ly- caenidae). Holarctic Lepid. 7:13-20. CovELL, C.V., Jr. 1976. The Schaus swallowtail: a threatened subspecies? Insect World Digest 3(59):21-26. MINNO, M. C. & T.C. EMMEL. 1993. Butterflies of the Florida Keys. Scientific Publishers, Gainesville, Florida. 168 pp. MINNO, M. C. & T.C. EMMEL. 1994. Order Lepidoptera. In M. Deyrup and R. Franz (eds.), Rare and Endangered Biota of Florida. Vol. 4. Invertebrates, 571-575. University Press, Gainesville, Florida. 798 pp. PyLE, R. M. 1981. The Audubon Society Field Guide to North American Butterflies. Alred A. Knopf. New York. 916 pp. SmituH, D. S., L. D. MILLER & J. Y. MILLER. 1994. The Butter- flies of the West Indies and South Florida. Oxford Univer- sity Press, New York. 264 pp. 32 pl U.S. Fish and Wildlife Service. 1999. South Florida multi- species recovery plan. Atlanta, Georgia. Received for publication 31 March 2006; revised and accepted 20 June 2007. Please see the next 4 pages for Appendices. Continued VOLUME 61, NUMBER 3 161 APPENDIX l. The butterfly species encountered in Cactus Hammock during each sampling date during the survey periods in 2004-2006 and their abundance. 2004 2005 Species Jun Jul Aug Sep Oct Nov Dec Jan Feb Papilio cresphontes 4 0 1 0 1 1 0 0 0 Ascia m. phileta 225 195 36 12 0 6 8 14 10 Phoebus s. eubale 0 0 0 12 2 3 0 0 0 P. a. maxima 22 12 17 13 1 2 0 0 2 Kricogonia lyside 0 0 0 0 0 0 0 0 0 Leptotes c. theonus 87 55 37 23 27 20 25 23 20 Hemiargus c. antibubastus 42 25 22, 23 13 15 12 10 6 Brephidium i. pseudofea 245 325 317 313 280 225 267 395 451 Strymon i. modestus 2 0 3 1 0 0 6 4 0 S. martialis 4 0 2 1 0 2 4 3 1 Agraulis v. nigrior 12 15 12 15 25 23 18 25 12 Heliconius c. tuckeri 10 5 12 9 6 14 5 12 14 Junonia genoveva 2 0 1 1 3 2 2 0 0 J. evarete 12 1 2 12 25 22 12 14 3 J. coenia 0 0 0 0 0 0 0 0 0 Anartia j. guantanamo 4 1 2 2 0 2 22 8 2 Danaus plexippus 0 0 0 0 0 12 0 0 0 D. gilippus 0 0 0 0 0 0 0 0 0 Phycoides phaon 0 0 0 0 0 0 0 0 0 Urbanus p. proteus 0 0 0 1 1 2 0 0 0 Phocides p. okeechobee 6 0 2 2 0 2 5 6 3 Polygonus I. savigny 2 0 1 2 2 2 2 s 4 Pyrgus o. oileus 4 0 3 2 1 2 0 0 0 Hylephila p. phyleus 0 0 0 0 0 0 0 0 0 Wallengrenia otho 13 2 8 4 12 3 12 10 2 Panoquina panoquinoides 3 5 2 1 0 1 1 0 0 162 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY APPENDIX 1. continued 2005 2006 Species Jun Jul Aug Sep Oct Nov Dec Jan Feb Papilio cresphontes i) 2 0 0 0 0 0) 0 0 Ascia m. phileta 157 18 15 8 6 1 2 (| 2 Phoebus s. eubale 0 0 0 4 0 0 0 0 0 P. a. maxima 28 4 12 6 2 0 1 0 2 Kricogonia lyside 13 4 1] 0 0 0 0 0 0 Leptotes c. theonus 125 1 17 0) 2 0 0 0 0 Hemiargus c. antibubastus 48 0 4 (0) 0 0 0 0 0 Brephidium i. pseudofea 312 161 68 12 0 1 157 215 431 Strymon i. modestus 0 0 0 0 0) 0 3 2 0 S. martialis 3 0 0 0 0 0 0 0 0 Agraulis v. nigrior 12 3 8 2 3 0 5 12 1 Heliconius c. tuckeri 14 0 0 0 0 0 0) 0 0 Junonia genoveva 0 0 0 0 0 0 0 l 0 J. evarete 14 0 7 5 6 0 1 3 2 J. coenia 0 (0) 0 0 0 0 1 0 0 Anartia j. guantanamo 3 0 4 5 2 0 0 0 0 Danaus plexippus 0 0) 0 0) 4 6 12 0 0 D. gilippus (0) 0 0 0 0 0 8 0 0 Phycoides phaon 0 0 0 0 0 1 0 0 0 Urbanus p. proteus 0 0 0 0 0 0 3 0 0 Phocides p. okeechobee 2, 0 1 0 0 0 0 1 1 Polygonus I. savigny 3 2 2 0 0 0 0 5 3 Pyrgus o. oileus 0 0 0 0 0 0 0 0 0 Hylephila p. phyleus 12 2 2 0 0 1 16 5 2 Wallengrenia otho 2, 0 0 0 0 0 12 0 0 Panoquina panoquinoides 0 0 0 0 0 0) 0 0 0 VOLUME 61, NUMBER 3 163 APPENDIX 2. The butterfly species encountered on Bahia Honda Key during each sampling date during the survey periods in 2004-2006 and their abundance. 2004 2005 Species Jun Jul Aug Sep Oct Nov Dec Jan Feb Papilio cresphontes 2 1 0 0 0 0 0 0 0 Ascia m. phileta 197 212 25 15 26 14 5 6 2 Phoebus s. eubale 0 0 12 12 15 3 0 0 0 P. a. maxima 21 13 4 6 10 8 6 2 4 P. p. philea 1 0 2 1 2 0 0 0 0 Nathalis iole 0 15 0 2 5 0 0 0 0 Eurema l. lisa 0 0 0 5 5 0 2 1 0 E. daira 0 0 0 1 3 0 0 0 0 Kricogonia lyside 0 12 5 12 2 2 0 0 0 Leptotes c. theonus 34 27 45 41 25 37 35 142 174 Hemiargus c. antibubastus 18 20 25 15 10 12 18 15 23 Cyclargus t. bethunebakeri 28 35 15 55 75 46 27 53 69 Electrostrymon a. angelia 2 1 0 1 0 0 0 0 0 Strymon i. modestus it 3 10 12 12 16 12 7 15 S. martialis 6 3 1 1 10 23 13 15 12 Agraulis v. nigrior 73 35 255 202 175 82 95 275 200 Heliconius c. tuckeri 85 5 150 97 105 61 75 110 65 Dryas i. largo 0 0 0 0 0 0 0 0 0 Anartia j. guantanamo 2 0 10 12 12 13 25 209 24 Vanessa cardui 1 0 0 0 0 0 0 0 0 Junonia evarete 0 0 0 0 0 0 0 0 0 J. coenia 0 0 0 0 0 0 0 0 0 Danaus plexippus 0 0 0 0 0 22 0 0 0 D. gilippus 0 0 0 2 8 8 0 0 0 Urbanus p. proteus 0 0 0 1 0 2 4 2 2 Phocides p. okeechobee 2 0 15 13 15 3 2 2 0 Polygonus 1. savigny 1 0 0 2 6 7 4 0 2 Pyrgus o. oileus 1 0 1 0 0 0 0 0 2 Hylephila p. phyleus 4 1 10 15 20 16 20 14 15 Polites v. vibex 2 1 10 8 6 6 0 0 2 Wallengrenia otho 3 0 10 12 20 12 15 8 2 Cymaenes t. tripunctus 1 0) 0 0 1 0 0 0 0 164 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY APPENDIX 2. Continued. 2005 2006 Species Jun Jul Aug Sep Oct Nov Dec Jan Feb Papilio cresphontes 0 2 1 0 0 0 0 0 0 Ascia m. phileta 125 116 87 15 5 0 1 0 0) Phoebus s. eubale 0 0 0 12 6 1 0 0 0 P. a. maxima 25 47 32 14 1 0 1 1 2 Pp. philea 2 0 2 0 0 0 0 0 0 Nathalis iole 0 0 0 0 0 0 0 0 0 Eurema l. lisa 2 0 1 1 2 0 1 0 0 E. daira 2 0 1 0 0 0 0 1 0 Kricogonia lyside 28 32 27 uf 0 0 0 0 0 Leptotes c. theonus 27 26 29 24 5 25 155 325 152 Hemiargus c. antibubastus 39 38 35 28 23 10 32 10 12 Cyclargus t. bethunebakeri 57 82 30 10 6 17 2 1 0 Electrostrymon a. angelia 6 2 1 0 0 0 0 0 0 Strymon i. modestus 16 62 20 4 1 0 3 2 42 S. martialis q 6 12 4 0 1 28 2 5 12 Agraulis v. nigrior 224 268 132 187 60 35 88 371 190 Heliconius c. tuckeri 114 197 18 0 0 0 0 0 0 Dryas i. largo 0 0 0 0 1 0 0 3 0 Anartia j. guantanamo 2 0 5 8 0 1 12 354 5 Vanessa cardui 0 0 2 0 0 0 0 0 0 Junonia evarete 1 0 0 0 0 0 0 0 0 J. coenia 0 0 0 0 2 0 0 0 0 Danaus plexippus 0 0 0 0 0 30 0 0 1 D. gilippus 0 0 0 0 0 0 (0) 0 0 Urbanus p. proteus 0 1 0 0 0 2 2 0 2 Phocides p. okeechobee 0 0 0 0 0 0 3 0 0 Polygonus l. savigny l 1 0 0 0 0 0 0 0 Pyrgus o. oileus 1 0 1 0 0 0 0 0 0 Hylephila p. phyleus 7 5 5 2 2 0 4 6 8 Polites v. vibex 2 1 1 0 0 0 0 0 0 Wallengrenia otho 3 3 2 0 1 0 2 1 0 Cymaenes t. tripunctus 3 3 1 0 0) 0 0 0 0 VOLUME 61, NUMBER 3 Journal of the Lepidopterists’ Society 61(3), 2007, 165-171 FIRST REPORT OF OECOPHORA BRACTELLA (L 165 .) (OECOPHORIDAE) IN NORTH AMERICA MERRILL A. PETERSON (MAP) Biology Department, Western Washington University, Bellingham, WA 98225, USA (e-mail: peterson@biol.wwu.edu) Eric H. LAGASA (EHL) Washington State Department of Agriculture, Olympia, WA 98504, USA (e-mail: elagasa@agr.wa.gov) STEVEN PASSOA (SP) USDA, APHIS, PPQ, The Ohio State University, Museum of Biodiversity, 1315 Kinnear Rd., Columbus, OH 43212, USA (e-mail: Steven.C.Passoa@usda.gov) GADEN S. ROBINSON (GSR) Department of Entomology, Natural History Museum, Cromwell Road, London SW7 5BD, UK (e-mail: G-Robinson@nhm.ac.uk) AND Davip HOLDEN (DH) Canadian Food Inspection Agency, 400 - 4321 Still Creek Drive, Bummaby, BC V5C 687 Canada (e-mail: holdend@inspection.ge.ca ABSTRACT. The first report of Oecophora bractella (L. .) from North America is given, based on collection records from Washington and British Columbia. This species is found throughout Europe, but is generally rare to uncommon through most of its range. Larvae occur on rot- ting wood in association with certain fungi and are not expected to reach pest status. Adults were captured at nine locations, with the first record from Seattle, WA in 1998. This first record and some of the subsequent records are from sites immediately adjacent to or associated with in- ternational shipping routes. Other sites with O. bractella are from residential areas, two of which have established breeding populations. The introduction of this moth is surprising, particularly given the low interception rate at United States ports and its specialized feeding niche. A di- agnosis and photographs of adults are provided to enable North American researchers to identify this species and to monitor its spread. Additional key words: Lepidoptera, Red Data Book, specialist, exotic, introduced, Pacific Northwest We report in this paper the surprising discovery of Oecophora bractella (Linnaeus, 1758) in Washington and British Columbia, apparently reflecting an introduction of this species to North America from Europe. Oe¢ecophora bractella is widely known in Europe, including the UK, as a strikingly beautiful, but infrequently seen, microlepidopteran occurring from the British Isles (but not Ireland) and the southern half of Scandinavia south to the Mediterranean and east to western Estonia and the Ukraine (Karsholt & Razowski 1996; Jiirivete et al. 2000; Lvovsky 2003). Oecophora bractella has a provisional listing as rare ('‘pRDB3') in the UK's Red Data Book (Kimber 2007), has similar Red Data Book status in both Estonia (Commission for Nature Conservation of the Estonian Academy of Sciences, 2002) and the Austrian state of Carinthia (Wieser & Hiimer 1999), and is considered ‘regionally threatened’ in continental Finland (K. Silvonen, pers. comm.). In many other parts of its range, O. bractella does not have a formal conservation listing, but it is generally characterized as being scarce or having a localized distribution (Lindsey 2006; de Prins & Steeman 2007). The exceptions to this general pattern are Sweden, where O. bractella is considered locally common (N. Ryrholm pers. comm.), and eastern Denmark, where it is generally common (Palm 1989). The larvae of O. bi Peel feed in the bark of dec: aying wood, often in close association with the my celia of fungi, especially honey fungus (Armillaria mellea (Vahl:Fr.) Kummer) (Sterling, 1984). It remains unclear whether the larvae eat these fungi (Harper et al. 2002). Larvae can be found in the bark of a variety of different tree species, including Quercus, Betula, Fraxinus, Corylus, Prunus, Larix, Pinus, Picea, and Tsuga, feeding from January to May in the UK. Larvae typically live under a loose layer of silk and frass (Harper et al. 2002). Between April and June, they pupate in their feeding sites, forming a cocoon also lined with silk and frass. Adults fly from late May to the end of July in the UK where O. bractella is considered to be univ eline (Harper et al. 2002). In Denmark, the flight season is slightly longer (mid-May to mid-August), suggesting possible bivoltinism (Palm 1989). Adults are primarily crepuscular, being most active from the early morning 166 light to sunrise, as well as in the late afternoon and evening (Harper et al. 2002; Palm 1989). They are seldom found at lights (Kimber 2007) and spend most of the daylight hours in seclusion (de Prins & Steeman 2007). The reliance of O. bractella on dead wood, coupled with its apparent association with specific fungi, may explain why this species is generally limited to ancient woodlands in the UK (Harper et al. 2002; Kimber 2007). Its scarcity in the UK may also be exacerbated by the common forestry practice of removing trees that are infected with Armillaria mellea, a policy motivated by the fact that the fungus can kill stressed Vancouvel = Lette, Ik JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY trees (Sterling 1984). The specialized life history (Sakai et al. 2001; Suarez et al. 2005), and scarcity of interception records for this species at U.S. ports (USDA, APHIS, PPQ Pest Interception Database (PestID), Riverdale, Maryland) suggests that O. bractella should be an unlikely species for inadvertent introduction. Nonetheless, we report herein its establishment in western North America. In addition, we describe the species and provide photographs of living and pinned specimens, both to facilitate recognition of this species, and to enable North American entomologists to monitor its spread. The introduction of this species WASHINGTON Bellingham km el 0 50 100 Distribution of collection records for O. bractella in Washington and British Columbia. Open circles and site numbers correspond to the sites listed in Table 1. Closed circles indicate the location of cities. This map was generated with Online Map Creator (http:/Avww.aquarius.geomar.de/make_map.html). VOLUME 61, NUMBER 3 adds to a growing list of wood-associated oecophorids of European origin that have been introduced to this continent (Powell 1964, 1968; Hodges 1974). Oecophora bractella in North America. The first specimen of O. bractella found in North America was a male taken at a blacklight trap in the industrial area of Seattle, Washington on June 30, 1998. Two and a half weeks later, a second male was taken in this same trap. Subsequent to this collection, an additional eight specimens were collected in pheromone traps at five other south Seattle locations, most of which were in residential areas (Table 1). Interestingly, these moths were found in traps baited with four different pheromones. The pheromone-trap and light-trapping surveys in which these specimens were captured were annual projects funded in part by grants from USDA APHIS, as part of the national Cooperative Agricultural Pest Survey (CAPS) program. In addition to the Seattle-area specimens, numerous O. bractella specimens have been collected more recently at three locations N of Seattle, near the US/Canada border (Table 1). Four of these specimens were found in a residential neighborhood in Bellingham, one was found on a pallet upon inspection for U.S. import in Blaine, Washington, and the rest were captured in a residential neighborhood in Port Coquitlam, British Columbia and a riparian area in Burnaby, British Columbia. The locations of all known North American collection sites for O. bractella, indicated in Fig. 1, are listed in Table 1, with collection and deposition data. A USDA, APHIS, PPQ Pest Interception Database query turned up no reports of interceptions of O. bractella at ports and airports other than the Blaine, WA specimen. The initial identification of O. bractella was made by one of the authors (GSR) on the basis of a photograph of a live specimen from Bellingham, WA (Fig. 2). This identification was confirmed by another author (SP) upon comparison of two Seattle-area males with both a specimen of German origin and literature illustrations of the genitalia (Palm 1978). Several lines of evidence suggest that O. bractella was inadvertently introduced to North America via shipments of bulk wood products. First, because the larvae feed on Armillaria-infected dead tree trunks, the shipment of raw timber and/or bulk wood products could easily facilitate introduction of this species. More importantly, several of the collection locations are near international shipping terminals. Indeed, the first specimens recorded from the region were from near Seattle's shipping terminal, and the specimen from Blaine, WA was found in a shipping container upon inspection for U.S. import. The 167 container held wood pallets loaded with insulating bricks that were shipped from Denmark (Interception # APSWA061931374003, USDA, APHIS, PPQ Pest Interception Database (PestID), Riverdale, Maryland). The establishment of breeding populations of O. bractella in North America has been confirmed at two locations (Bellingham, Washington, and Burnaby, British Columbia), where immatures were found in spring 2007. At Burnaby, a half dozen unknown larvae living under the peeling bark of a dying Alnus rubra were collected for rearing in March. On May 3 an adult male O. bractella was found in the rearing chamber. Subsequent collections in this riparian habitat yielded specimens on many different A. rubra offering similar larval habitats, as well as one ornamental Acer sp. in a nearby landscaped environment. Most collections were on dying some were on recently felled standing trees, but Fic. 2. Live O. bractella female from Bellingham, Washing- ton. Photograph by MAP. Fic. 3. O. bractella female from Port Coquitlam, British Co- lumbia. Photograph by DH. 168 trees. In this same vicinity, larvae were not found under peeling bark on completely dead trees with very little moisture content. In Bellingham, larvae and pupae were found in a 9 May 2007 under the bark of 2"_4" diameter dead standing branches of Laburnum anagyroides Medik. (Fabaceae), an introduced ornamental commonly called Golden Chain Tree. Adults were subsequently reared from three of these larvae, and a fourth larva produced an unidentified tachinid. Also on 29 May 2007, adults were found on the bark of the dead branches as well as on nearby overhanging live branches of L. anagyroides and neighboring trees and bushes. From 29 May to 12 June 2007, 14-18 adults were associated w ‘ith this L. anagyroides tree, with thereafter until 17 July 2007. Larvae were present through the entire six week period. Although breeding has not been documented at other sites from which adults have been collected, it is likely that this species is established outside of Burnaby and Bellingham. Notably, adults were found in two successive years at one site south of Seattle (Site 2) . It remains unclear whether the Pacific Northwest populations of O. bractella are associated with Armillaria mellea, the fungus with which larvae are typically associated in Europe. The larvae found in Bellingham were on branches that had peeling bark under which the wood was blackened, perhaps from fungal infection. At Burnaby, the highest densities of larvae appeared to be associated with an unknown bracket fungus but fungal feeding was not evident. Larvae under bark did not appear to be in close residential area on 2 diminishing numbers association with visible fruiting bodies or mycelia and they appeared to be rasping or chewing the surface of the phloem which resulted in a colour change of the sapwood. This discoloration was a good indicator of larval presence as was their frass filled silk webbing. Although some authors suggest that Armillaria mellea occurs in the Pacific Northwest (Shaw 1973), the taxonomy of this group has been recently revised (Burdsall & Volk 1 1993). Under this revision, A. mellea is no longer considered to occur in Washington and Oregon, but other species of Armillaria are common in the region (Burdsall & Volk 1993), suggesting that the fungal associations of this species may include more than one Armillaria species. Indeed, given the recent taxonomic changes for Armillaria (Burdsall & Volk 1993), it is possible that some of the previously reported associations with A. mellea may be inaccurate. The fact that A. mellea is found in northern California and eastern North America JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY (Burdsall & Volk 1993) suggests that O. bractella could become widespread in North America. Formal nomenclature. Oecophora bractella (Linnaeus, 1758) Phalaena (Tinea) bractella Linnaeus, 1758, Systema Naturae (edn 10): 540, no. 280. Lectotype, 2, ['Europa'] labelled ‘bractella’ [by Linnaeus|/bractella 894 [by J.E. Smith] designated by Robinson & Nielsen (1983: 204) (Linnean collection! London). Phalaena bractella; Clerck, 1759; Icones Insectorum Rariorum, pl. 12, fig. 4. Phalaena (Tinea) {bracteella; Linnaeus, 1761, Fauna Svecica (edn 2): 366, no. 1426 [incorrect subsequent spelling]. Oecophora bractella (L Insectorum 180: 21. It is likely that Clerck figured Linnaeus's specimen, as there are no specimens in the Clerck collection (Robinson & Nielsen, 1983). High-resolution digital images of the lectotype should be available shortly through the Linnean typification project on the Linnean Society's website: http:/Avww.linnean.org. During the latter part of the nineteenth and the early twentieth century bractella was placed in the genus Alabonia, which was synonymized with Oecophora by Meyrick (1922). Diagnosis. The adults and immature stages are described in detail elsewhere in the literature (e.g. Meyrick 1922; Toll 1964; Fetz 1994; Harper et al. 2002; Patoéka_& M. Turédni 9005). Here, we discuss the diagnostic features of O. bractella adults (see also Figs. 2, 3) to allow identification by North American researchers. Head and thorax: The head and thorax are covered with bright yellow, appressed scales. The long, recurved labial palps are mostly dark fuscous, with pale tips and some yellow on the inner and ventral surfaces. The legs are dark fuscous with white tarsal and tibial rings. Wings: The forewings (5.5-7.5mm wing length) of O. bractella are dark brown or gray to black, patterned with bright yellow and metallic blue or purple patches. The basal 1/3 of the forewings is bright yellow, except along the costal margin, which is dark brown to black. Just distal to the basal yellow area is a band of metallic blue or purple scales, running straight from the costa to the dorsum, and separated from the yellow basal area by a thin band of dark fuscous to black. The forewing also features a single smaller anteapical yellow spot along the costal margin. Basal and medial to this spot is a short band of metallic blue or purple scales that does not reach the margin of either wing. The distal margin of the forewing features metallic blue or purple scaling. The fringe scales are generally dark fuscous to black, with those near the apex tipped with white. The .); Meyrick, 1922, Genera VOLUME 61, NUMBER 3 169 TABLE 1. Collection and deposition data for all known Oecophora bractella specimens from North America. Locality USA: Washington 1) King Co., Seattle, 47.57°N 122.34°W; industrial/port 2) King Co., 4.8km N of Burien, 47 A88°N 122.351°W; residential 3) King Co., Saltwater State Park, Des Moines, 47.375°N 122.322°W; park/residential 4) King Co., West Seattle, 47,552°N 122.398°W; residential 5) King Co., Normandy Park, 47.433°N 122.348°W; residential 6) King Co., Burien, 47.467°N, 122.361°W; urban 7) Whatcom Co., Bellingham, 48.741°N 122.474°W; residential 8) Whatcom Co., Blaine, 49.000°N122.738°W; import inspection station CANADA: British Columbia 9) Port Coquitlam, 49.244° N 122.778°W: residential 10) Burnaby, 49.15° N 123.00°W; riparian near commercial development Date Method Collector Determination 30 June 1998 Blacklight M. Allen E.H. LaGasa 17 July 1998 Blacklight M. Allen S. Passoa 8 August 2000 European Corn Borer P. Hertzog S. Passoa lure trap? 2 July 2001 Proeulia lure trap? P. Hertzog E.H. LaGasa 2 August 2001 — Proeulia lure trap? P. Hertzog E.H. LaGasa 7 August 2001 Proeulia lure trap? P. Hertzog E.H. LaGasa 8 July 2002 Leek moth lure trap? S. Williams E.H. LaGasa 17 July 2002 Plum fruit moth lure — S. Williams E.H. LaGasa trap? 17 July 2002 Plum fruit moth lure S$. Williams E.H. LaGasa 17 June 2006 29 May 2007 11 July 2006 23 June 2006 26 June 2006 1 June 2007 2 June 2007 27 June 2007 28 June 2007 2 July 2007 4 July 2007 30 May 2007 3 May 2007 to 12 June 2007 trap* At window of house Dead branches of Laburnum anagyroides Shipping container Mercury vapor lamp Mercury vapor lamp Mercury vapor lamp Mercury vapor lamp Mercury vapor lamp Mercury vapor lamp Mercury vapor lamp Mercury vapor lamp Sweep net Reared from bark of Alnus rubra & Acer sp. M. Peterson M. Peterson J. Boyer . Holden . Holden . Holden . Holden . Holden . Holden . Holden . Holden Piss Se) eo) (oi 1S) 4s) D. Holden . Holden o G. Robinson M. Peterson ZA Passoa . Pohl . Pohl . Holden . Holden . Holden . Holden . Holden . Holden Do AS Ae oS) ye) i) . Holden . Holden a Number & Deposition! 1 male; WSDA 1 male; USNM 1 male; USDA 1 sex unknown (missing abdomen & hindwings); WSDA 2 males; WSDA (only one specimen retained) 1 female; WSDA il male; specimen not kept 1 male; specimen not kept 1 male; specimen not kept 1 female; WWU 2 females, 1 male: WWU 1 female; USDA 1 female; DH 2 females; CNC 1 female; DH 2 females; DH 1 female; DH 2 females; DH 1 female; DH 1 male; DH 1 male; DH 22 males: 20 females, DH 'WSDA = Washington State Department of Agriculture, Olympia, WA; USDA = U.S. Dept. of Agriculture, APHIS-PPQ, The Ohio State University, Museum of Biodiversity, Columbus, OH; USNM = Smithsonian Institution; WWU = Biology Department, Western Washington University, Bellingham, WA; DH = David Holden, personal collection; CNC = Canadian National Collection, Ottawa, ON. *Pheromone lure traps consisted of gray rubber septa (West Co., Lionville, PA, cat. no. 1060-0275) loaded with specific pheromone lures, in a Pherocon 2 type trap. European Corn Borer lure: 0.5mg Z-11-14:AC, 0.5mg E-11-14:AC; Proeulia lure: 0.lmg E-11-14:0H; Leek Moth lure: 1 mg Z-11-16:AL, Plum Fruit Moth lure: 0.1mg Z-8-12:AC, 0.004mg E-8-12:AC, 0.025mg Z-8-14:AC, 0.005mg Z-10-14:AC, 0.2mg 14:AC. 170 hindwing is uniformly dark gray. Rectiostoma fernaldella (Riley, 1889) is the ‘only gelechioid in western North America that could perhaps be confused with O bractella. This species, occurring only in the SW U.S., is superficially similar to O. bractella, but the basal area of the forewing of R. fernaldella is dull (not bright) yellow, the forewing lacks an anteapical yellow spot, the apex of the forewing is strongly squared off (not rounded), and the hindwings are broader than in O. bractella. The somewhat dorsoventrally-flattened larvae of O. bractella are grayish brown, with darker gray thoracic segments and terminal abdominal segment, and a brown head capsule. Each abdominal segment has a subdorsal furrow. Pest Status. O. bractella is unlikely to attain pest status in North America, as its larvae feed in the bark of decaying wood. Although O. bractella is unlikely to have any economic effects, it is possible that its establishment could have minor ecological ramifications, through its impact on detritivore food webs. Perhaps of broader interest is that fact that the arrival of O. bractella underscores the ease with which species can exploit the colonization opportunities afforded by international shipping. Given the profound economic and environmental costs associated with invasive species that do attain pest status (Wilcove et al. 1998; Mack et al, 2000; Pimentel et al. 2001), the introduction of this rather uncommon and specialized species is alarming. Furthermore, this introduction underscores the difficulty in making generalities regarding which insects are most likely to he introduced ( Simberloff 1989). Relevance to Conservation Efforts. The residential and urban locales in Washington and British Columbia from which O. bractella Wie: been recorded are unlike the habitats this species exploits in its native range. Through most of its range, O. bractella is generally restricted to forests and other wooded regions (Novak & Severa 1980; Palm 1989), while in the UK, it is generally restricted to ancient forests (Harper et al. 2002; Kimber 2007), and in Estonia, it is found only in sparse juniper woodlands on limestone with sparse vegetation (M. Martin, pers. comm.). However, in some regions this species is found in managed habitats such as hedgerows (Lindsey 2006). The ability of O. bractella to occupy residential habitats in North America suggests that further research on this species in North America may provide insights into managing habitat for this species in portions of its native range in which it is of conservation concern. Furthermore, the discovery that O. bractella can be taken by pheromone traps reveals that such traps may enable better monitoring of this species in its native range. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY ACKNOWLEDGMENTS W. de Prins, O. Karsholt, J. Langmaid, M. Matin, M. Parsons, M. Ponomarenko, N. Ryrholm, K. Silvonen, and S. Sinev pro- vided helpful information on the distribution, abundance, and conservation status of O. bractella in its native range, and helped us locate the pertinent literature. M. Honey assisted with the initial identification of the Bellingham specimen, and pointed out useful references. In addition, G. Pohl provided an initial identification of the Port Coquitlam, B.C. specimens. W. Carl- son sent the Blaine, WA interception to one of us (SP). P. Mar- quez queried the USDA, APHIS, PPQ Pest Interception Data- base for previous O. bractella records. J. Brown confirmed the identification and verified the lack of previous North American records. His comments and those of two anonymous reviewers improved a previous draft of this manuscript. F. Rhoades pro- vided helpful information on the distribution and taxonomy of Pacific Northwest Armillaria. Pheromone lures were provided by the USDA APHIS Otis Methods Lab. USDA APHIS CAPS grants (#98, 00, 01, and 02-8553-0249-CA) to WSDA supported the Seattle-area light-trap and pheromone trapping. MAP was supported by a Summer Research Grant from Western Wash- ington University's Office of Research and Sponsored Programs. LITERATURE CITED BurbsALL, H.H. JR. & T.J. VOLK. 1993. The state of taxonomy of the genus Armillaria. Mcllvainea 11:4-11. CLERC K, C. 1759-1764. Icones insectorum rariorum. [xii] + 55 pls. Holmiae. COMMISSION FOR NATURE CONSERVATION OF THE ESTONIAN ACADEMY OF SCIENCES. 2002. Red data book of Estonia. http://www. zbi. ee/punane/english/index.html., De Prins, W, & C. STEEMAN. 2007. Catalogue of the Lepidoptera of Belgium. http:/Avebh01.ua.ac.be/vve/Checklists/Lepidoptera/Oe- cophoridae/Obractella.htm Fetz, R. 1994. Larvalmorphologische Beitriige zum phylogenetischen System der ehemaligen Oecophoridae (Lepidoptera, Gele- chioidea). Neue Entomologische Nachrichten 33, 290 pp. HARPER, M.W.,, J.R. LANGMaID, & A.M. EmMMET. 2002. Oecophoridae, pp. 43-177. In: A.M. Emmet & J.R. Langmaid (eds), The moths and butterflies of Great Britain and Ireland. 4(1). Harley Books, Colchester, UK. Honpces, R.W. 1974. The Moths of America North of Mexico, Fasci- cle 6.2, Gelechioidea, Oecophoridae. E.W. Classey Limited and R.B.D. Publications, Inc., London. 142 pp. JUrivere, U, J. Karrina, T. KesktLa, K. NUPPONEN, J. VIDALEPP & E. Ounap. 2000. Estonian Lepidoptera: Catalogue. Estonian Lepi- dopterologists' Society, Tallinn. 150 pp. KarSHOLT, O. & J. Razowsk1. 1996. The Lepidoptera of Europe: a dis- tributional checklist. Apollo Books, Stenstrup, Denmark. 380 pp. Kimper, I. 2007. UK Moths. 651. Oecophora bractella. http://uk- moths.org.uk/ show.php?bf=651 LINDsEy, J. K. 2006. Ecology of Commanster. http://popgen.uni- maas.nl/~jlindsey/ commanster.html LINNAEUS, C, 1758. Systema Naturae .. [iii] pp. . Editio 10. 1. iv + 824 pp., Holmiae. . 1761. Fauna Svecica ... Editio reformata. [xlviii] + 578 pp., Stockholmiae. Lvovsky, A.L. 2003. Check-list of the broad-winged moths (Oe- cophoridae s. |.) of Russia and adjacent countries. Nota lepid. 25:213-220. Mack, R.N., D. SIMBERLOFF, W.M. LONSDALE, H. Evans, M. CLOUT, & FA. Bazzaz. 2000. Biotic invasions: causes, epidemiology, global consequences, and control. Ecol. Applic. 10:689-710. Meyrick, E. 1922. Oecophoridae. In: Wytsman, P. (ed.) Genera In- sectorum, fasc. 180. Bruxelles. 222 pp. Novak, I. & F. SEvERA. 1980. Der Kosmos-Schmetterlingsfiihrer. W. Keller & Co., Stuttgart. Pa, E. 1978. De danske Oecophoridae, Lepidoptera, Saernummer 4. 100 pp. VOLUME 61, NUMBER 3 . 1989. Nordeuropas prydvinger, handbook on the Oecophorid moths of North Europe. Danmarks Dyreliv, Vol. 4. Fauna Boger, Copenhagen. 247 pp. PaTOcka,J. & M. TuRCANI 2005. Lepidoptera pupae, Central Euro- pean species. Apollo Books, Stenstrup, Denmark. 863 pp. PIMENTEL, D., S. MCNaIr, J. JANECKA, J. WIGHTMAN, C. SIMMONDS, C. O'CONNELL, E. Wonc, L. RUSSEL, J. ZERN, T. AQUINO, & T. TsoMONDO. 2001. Economic and environmental threats of alien plant, animal, and microbe invasions. Agric. Ecos. Envy. 84:1—20. POWELL, J.A. 1964. Two scavenger moths of the genus Borkhausenia introduced from Europe to the west coast of North America. Pan-Pac. Ent. 40:21§-221. . 1968. Discovery of Esperia sulphurella (F.) in California. Pan- Pac. Ent. 44:78. ROBINSON, G.S. & E.S. NIELSEN. 1983. The Microlepidoptera de- scribed by Linnaeus and Clerck. Syst. Ent. 8:191-242. Sakal, A.K., FW. ALLENDORF, J.S. Hott, D.M. LopcE, J. MOLOFsky, K.A. WiTH, S. BAUGHMAN, R.J. CaBIN, J.E. COHEN, N.C. ELL- STRAND, D.E. MCCAULEY, P. O’NEIL, I.M. PARKER, J.N. THOMP- son, & $.G. WELLER. 2001. The population biology of invasive species. Ann. Rev. Ecol. Syst. 32:305-332. SHaw, C.G. 1973. Host fungus index for the Pacific Northwest — 1. Hosts, Wash. State Ag. Exp. Sta. Bull. 764: 1-121. 171 SIMBERLOFF, D. 1989. Which insect introductions succeed and which fail?, pp. 61-75 In: J.A. DRAKE, H.A. Mooney, F. pi Castri, R.H. GRrovEs, F.J. KRUGER, M. REJMANEK, & M. WILLIAMSON (eds.), Biological invasions: A global perspective. John Wiley & Sons. NY. STERLING, P.H. 1984. Oecophora bractella in Hampshire. Ent. Rec. J. Var. 96:54-55. Suarez, A.V., D.A. Houway, & P.S. Warb. 2005. The role of opportu- nity in the unintentional introduction of nonnative ants. PNAS (USA) 102:17032-17035. TOLL, S. 1964. Lepidoptera — Oecophoridae. Klucze do Oznaczania Owad6w Polski, Ser. 43, Part 27, No. 35. 174 pp. WiESER, C. & P. HUMER. 1999. Rote Listen der Schmetterlinge Karn- tens. In: W.E. Holzinger, P. Mildner, T. Rottenburg & C. Wieser (eds.), Rote Listen gefahrdeter Tiere Karmtens. Naturschutz in Karnten 15:133-200. WILCOVE, D.S., D. ROTHSTEIN, J. DuBow, A. PHILLIPS, & E. Losos. 1998. Quantifying threats to imperiled species in the United States. BioScience 48:607-615. Received for publication 9 March 2007; revised and accepted 29 June 2007. Journal of the Lepidopterists’ Society 61(3), 2007, 172-175 GENERAL NOTES JOURNAL OF THE LEPIDOPTERISTS’ SOCLETY FIRST REPORT OF THE PALEARCTIC SPECIES CYDIA CONIFERANA (TORTRICIDAE) IN THE WESTERN UNITED STATES A single adult male of Cydia coniferana (Saxesen, 1840) was collected in Tumwater, Thurston County, Washington State, in 2000. This moth was a non-target in a United States Department of Agriculture/Animal and Plant Health Inspection Service/Cooperative Agricultural Pest Survey (USDA/APHIS/CAPS) program aimed at detecting the European com borer (ECB), Ostrinia nubilalis (Hiibner). The collection method was a pheromone-trap baited with hybrid northern/southem strain ECB lure (1:1 Z11/E11- 14Ac). This was the first record of coniferana recorded from North America since several adults were reared from the bark of red pine in New York (Schaffner 1959). The New York population apparently never established (W. Miller pers. comm.), although the species was listed from North America by Powell (1983). Cydia coniferana is native to the Palearctic Region where it ocurrs throughout Europe east to Russia, China, and Mongolia. (Bradley et al. 1979; Zhang 1994). The larvae feed in the cambium layer of many coniferous trees, including Pinus spp., Picea spp., Abies spp. and Larix sp. (Bradley et al. 1979; Karsholt and Razowski 1996; Zhang 1994). Published information on larval feeding impacts is limited and variable. In England, where coniferana infestation is associated with fungal disease, it is not a significant pest (Bradley et al. 1979). Central Europen populations of coniferana are reported to enter the cambium only after the tree is wounded (Patocka and Turéani 2005). However, Razowski (2003) reported Fic. 1. Adult males of Cydia coniferana in a pheromone trap showing variation in forewing pattern. “occasional damage”, especially to pine nurseries, and coniferana is identified as a “harmful species” in western Russia (Medvedev 1987). Adult coniferana are small (10-14 mm) dark moths (Fig. 1) with a wing pattern that is typical of many Cydia and Grapholita species (see Komai 1999; Razowski 2003). Recognition in sticky traps is difficult without genitalic dissection. Male moths can be identified by a short row of deciduous cornuti in the aedeagus and a ventral toothlike projection on the valves (Fig. 2) (Bradley et al. 1979). Some variation of adult morphology was noted in our samples. In particular, the white dorsal patch of the forewing is sometimes absent as noted by Razowski (2003). One feature of the male genitalia, the length of the ventral toothlike projection, was also variable. Descriptions or illustrations of the female genitalia (Bradley et al. 1979), pupa (Patocka and Turéani 2005), and larva (Swatschek 1958) of coniferana have also been published. In the summer of 2005, a preliminary pheromone trap delimiting survey for coniferana was conducted in westemm Washington from King County south to the Oregon border Fic. 2. Cydia coniferana male genitalia. Arrows point to the cornuti ei the aeadeagus and ventral toothlike projection of the valve. Note variation a length of the ventral tooth of valve. VOLUME 61, NUMBER 1 173 ohomish omish™\__s N Ww pe Jefferson SS Ls Kittitas BPS Yakima 2005 Cydia coniferana trap sites O Negative Sites @ Positive Sites Interstate 5 Positive counties Skamania Eric LaGasa, WSDA Pest Program, 360 902-2063 Fic. 3. Cydia coniferana collection sites in southwestern Washington State. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TABLE 1. 2005 Cydia coniferana trap site numbers and results by county. County Total Sites Positive Sites King 30 28 Pierce 50 46 Thurston 60 58 Lewis 30 26 Cowlitz 10 9 Clark 20 18 Totals 200 185 Average moths Total Moths (per Pos. Site) % Sites Positive 93.3% 885 31.6 92.0% 976 21.2 96.7% 1,453 25.1 86.7% 400 15.4 90.0% 154 W7fell 90.0% ATT 26.5 92.5% 4,345 23.5 (Fig. 3, also LaGasa and Welch 2005). Traps were placed in a total of 200 roadside or residential yard trees, mostly on Pinus spp., Abies spp., Pseudotsuga menzesii, and Picea spp. from early June until late August. Pherocon 2® type traps with lures consisting of 0.5 mg of E,$ — Dodecenyl Acetate on a hexane-rinsed red ail ibe septa were used. The lures were changed no later than every four weeks. Survey sosulte summarized by county in Table 1, and illustrated in Figure 3, clearly show that coniferana i is well established back widespread in the survey area. A total of 4,345 male coniferana were collected at 185 of 200 total sites, with an average catch of 24 per trap at positive sites. The average number of moths captured across counties surveyed was fairly uniform, and the distribution of positive sites and catch numbers did not v. ary substantially between rural/voodland areas and more urban sites. The majority of moths (80%) were captured during August with peak activity around the middle of the month. Additional monitoring is needed to determine the entire duration of adult activity and whether the species produces one or multiple generations in the Pacific Northwest. Given the prevalent and uniform occurrence of coniferana in the surveyed area, it is likely that the current distribution of this moth includes more (if not all) of western Washington and possibly adjacent areas in Oregon and British Columbia west of the Cascade Mountains. Examination (including removal of bark) of potential host tree species in the area infested by coniferana revealed some evidence of larval damage matching the description given by Bradley et al. (1979), but no larvae were found. C. coniferana was found at numerous sites where Pseudotsu ga menzesii (Douglas fir) was the only conifer present, suggesting that this species may be a potential host. Voucher specimens collected in this 2005 survey are deposited at the United States National Museum (Washington, D.C), the S. Passoa collection (Columbus, Ohio) and in the W. ashington State Department of Agriculture Insect Collection (O (O lympia, Washington). Multiple introductions of microlepidoptera can occcur on either coast of the United States (see Powell and Passoa 1991). Our data shows that a second introduction of coniferana established in the westem United States. Regulatory entomologists only consider an organism to be introduced if there is evidence of an established breeding population (Pender 1983). Our data shows this is the case for coniferana which justifies inclusion of this species in the checklist of North American Lepidoptera. We thank Dr. Wm. Miller (University of Minnesota, St. Paul) and Dr. J. Brown (Systematic Entomology Lab, Washington D.C.) for confirming the adult of coniferana, comments on the manuscript, and for valuable historical data. The USDA/APHIS/Otis Methods Development Center provided pheromone traps and lures. This survey was funded in part by a Cooperative Agricultural Pest Survey (C APS) gr rant from the USDA APHIS Westem Region (#05-8553- 0737-CA). LITERATURE CITED BRADLEY, J.D., W.G. TREMEWAN, & ARTHUR SMITH. 1979. British Tortri- coid Moths, Tortricidae: Olethreutinae. Curwen Press Ltd., London, England. viii + 219 pp. Hopces, R.W,, T. Dominick, D. R. Davis, D. C. FERGUSON, J. G. FRAN- CLEMONT, E. G. MUNROE & J. A. POWELL, 1983. Check list of the Lepidoptera of America North of Mexico including Greenland. E.W. Classey Ltd. & the Wedge Entomological Research Foundation. London, England. xxiv + 284 KarsHOLt, O. & J. Razowski (eds.), 1996. The Lepidoptera of Europe, A distributional checklist. Apollo Books. Stenstrup, Denmark. 380 pp. Komal, F. 1999. A taxonomic review of the genus Grapholita and allied genera (Lepidoptera: Tortricidae) in the Palearctic Region. Ent. Scand. 55: 1-226. LaGasa, E.H. & S. WELCH, 2005. 2005 Preliminary Western Washington Pheromone-trap Delimiting Survey for Cydia coniferana (Lepi- doptera: Tortricidae), an Old World Conifer-bark-feeding Tortricid New to Western North America. Washington State Department of Agriculture AGR PUB 805-145 (N/10/05), Olympia, Washington. 4 Dp. ME eee GS. (Editor). 1987. Keys to the Insects of the European Part of the USSR. Volume IV, Lepidoptera. Part 1. Academy of Sciences of the USSR Publication TT 81-52013. Nauka Publishers. Translated from Russian by Amerind Publishing Co., New Delhi, India. xxvi + 991 pp Parocka, J. & M. TurCANI. 2005. Lepidoptera pupae. Central European species. Apollo Books. Stenstrup, Denmark. 863 pp. PENDER, M. T. 1983. Procedures for evaluating and reacting to reports of new organism occurrences in the United States. New Pest Advisory Group. USDA/APHIS/PPQ. Unpublished report. POWELL, J. A. 1983. Tortricidae, pp. 31-41. In Hodges, R. W. et al. (eds), Checklist of the Lepidoptera of America north of Mexico. E. W. Classey & Wedge Entomol. Res. Foundation. London, England. VOLUME 61, NUMBER 1 & S. Passoa. 1991. Rapid colonization of the westem United States by the Palearctic moth, Agonopterix alstroemeriana (Oecophoridae). Journal of the Lepidopterists' Society 45(3): 234-236. Razowskl, J. 2003. Tortricidae (Lepidoptera) of Europe. Volume 2. Olethreutinae. Frantisek Slamka, Publisher. Bratislava, Slovakia. 301 eee Jr, J. V. 1959. Microlepidoptera and their parasites reared from field collections in the northeastern United States. USDA Misc. Pub. no. 767. 97 pp. SWATSCHEK, B. 1958. Die larvalsystematik der Wickler (Tortricidae und Carposinidae). Abhandlungen Zur Larvalsystematik der Insekten nr. 3. Akademie- Verlag. Berlin, Germany. 269 pp. ZHANG, B-C. 1994. Index of economically important Lepidoptera. Walling- ford, United Kingdom: CAB International. 599 pp. Journal of the Lepidopterists’ Society 61(3), 2007, 175 Eric LaGasa, Pest Program, Plant Protection Division, Washington State Department of Agriculture, P.O. Box 42560, Olympia, Washington 98504-2560 USA, email: elagasa@agr.wa.gov STEVEN Passoa, USDA/APHIS/PPQ, The Ohio State University, Museum of Biological Diversity, 1315 Kinnear Road, Columbus, Ohio 43212 USA, — email: Steven.C. Passoa@usda.gov Received for publication 22 November 2006, revised and accepted 25 May 2007. POANES MELANE (HESPERIIDAE) OVIPOSITING ON AN AUSTRALIAN GRASS NATURALIZED IN CALIFORNIA Additional key words: Rytidosperma, Danthonia, introduced species The use of introduced plants as hosts by native California butterflies has been reviewed by Shapiro (2002) and Graves and Shapiro (2003), who found that the urban and suburban fauna of that state was now largely dependent on such plants. New records of this type appear regularly, demonstrating that butterflies colonize potential host plants more or less quickly after they appear in an area. On 27 October 2006 I observed a female Poanes melane (W.H. Edwards) systematically searching for and ovipositing repeatedly on a low, tufted, apparently perennial grass I did not recognize in a parking strip in Berkeley, Alameda County, California. This grass was common in the neighborhood, occurring in lawns and waste ground in an older residential area of North Berkeley. Unable to identify it using Hickman (1993), I brought specimens to the U.C. Davis Herbarium where it was identified by Jean Shepard as Rytidosperma racemosum (R. Br.) Connor & Edgar (formerly placed in the genus Danthonia). This species was not recognized as naturalized in California when Hickman (1993) was in preparation. According to Stephen Darbyshire of Agriculture Canada, an authority on the genus, this grass was grown “experimentally” in gardens in Berkeley as early as 1941. It seems to have ecu to spread in Alameda County i in the early 1950s and is recorded as naturalized only in that county except for a 1978 record from the naval garrison on Angel Island, Marin County. According to Darbyshire and Barbara Ertter of the Jepson Herbarium at U.C. Berkeley as well as my own observations, it is now a fairly common weed in Berkeley and nearby Albany and will probably continue to spread. Rytidosperma racemosum is originally from Australia. Various members of the genus were tested in California for forage potential as early as 1911 and R. penicillatum (Labill.), more commonly known as Danthonia pilosa R.Br., is naturalized in Califormia, southern Oregon and Hawaii. It would hardly be surprising to find P. melane and other native skippers using these plants elsewhere. Although the ability of P. melane to feed and develop successfully on R. racemosum has not been demonstrated, it accepts most perennial and some annual grasses in the laboratory. Scott (1986) lists five very diverse grasses in as many genera plus one sedge (Cyperaceae). He does not provide sources for these records. Garth & Tilden (1986) record two native perennial grasses in southern California. Various other records are scattered in the literature, none of them being on Rytidosperma, which as noted above was not even recognized as being part of the California flora! Bay Area populations are now “urbanized” and routinely breed on Bermuda Grass (Cynodon dactylon (L.)Pers., while inland California populations are confined to riparian-wildland habitats (Shapiro and Manolis, 2007). I thank Jean Shepard, Barbara Ertter and Stephen Darbyshire for their help. LITERATURE CITED Garth, J.S. & J.W. Tilden. 1986. California Butterflies. University of Cali- fornia Press. 246 pp. Graves, S.D. & A.M. Shapiro.2003. Exotics as host plants of the California butterfly fauna. Biol. Cons. 110: 413-433. Hickman, J. 1993. The Jepson Manual: Higher Plants of California. Uni- versity of Califormia Press. 1400 pp. Scott, J.A. 1986. The Butterflies of North America. Stanford University Press. 583 pp. Shapiro, A.M. 2002. The Californian urban butterfly fauna is dependent on alien plants. Diversity & Distributions 8: 31-40. Shapiro, A.M. & T.D. Manolis. 2007. Field Guide to Butterflies of the San Francisco Bay and Sacramento Valley Regions. University of Califor- nia Press. 346 pp. ARTHUR M. SHAPIRO, Center for Population Biology, University of California, Davis, CA 95616, email: amshapiro@ucdavis.edu. Received for publication? November 2006, revised and accepted 12 June 2007. 176 Journal of the Lepidopterists’ Society 61(3), 2007, 176-177 HANDBOOK OF ZOOLOGY, VOLUME IV, ARTHROPODA: INSECTA, edited by M. Fisher, Part 35 Lepidoptera, Moths and Butterflies, Volume 1: Evolution, Systematics, and Biogeography, edited by N. Kristensen, 1998, 490 pp, ISBN3-11-015704-7 Volume 2: Morphology, Physiology and Dievellopanenl, edited by N. Kristensen, 2003, 564 pp. ISBN 3-11- 016210-5, de Gruyter, Berlin and New York. List price (Amazon Books): Volume 1, US$299.60; Volume 2, US$347.20. Few lepidopterists have the comprehensive knowledge or stamina to direct the production of The Handbook of Zoology — Lepidoptera, Moths and butterflies. Indeed, the two volumes that resulted from editor Kristensen’s deliberate planning are more suitably described as ‘encyclopedia’ than ‘handbook .In 40 chapters written by outstanding contributors these books cover evolution, systematics, biogeography, comparative morphology and anatomy of early stage and adult Lepidoptera. Volume 1: Evolution, Systematics, and Biogeography contains 21 chapters. The ‘Historical Introduction’ (Kristensen) enumerates critical landmarks marking the and progress of Lepidoptera studies, and functioning as a gateway for further exploration. Chapter 2, ‘Phylogeny ana Palaeontology’ (Kristensen & Skalski), is a lucid overview of superfamily-level relationships based on a deep understanding of the Lepidoptera groundplan, extant and fossil taxa. Figure 2.1 beautifully describes the groundplan, and should be of interest to all entomologists, and Fig. 2.2 gives a phylogeny for extant superfamilies of Lepidoptera. This chapter also lists known fossils by family; a useful reference guide to original studies. A chapter on ‘Classification and Keys to icher Taxa’ (Carter & Kristensen) naturally follows. While the key to adults includes 46 superfamilies and does not require illustrations (readers will recognize all characters), the key to larvae is supplied with illustrations of basic larval morphology and leads to 129 categories, including superfamilies, families, and subfamilies. Chapters 4-19 cover higher groups; “The Non-Glossatan Moths’ (Kristensen), ‘The Homoneurous Glossata’ (Kristensen), ‘The Monotrysian Heteroneura (Davis), “The Tineoidea and Gracillarioidea’ (Davis & Robinson), ‘The Yponomeutoidea’ (Dugdale, Kristensen, Robinson & Scoble), ‘The Galechioidea’ (Hodges), “The Zygaenoidea’ (Epstein, Geertsema, Naumann & Tarmann), “The Cossoid/Sesioid Assemblage’ (Edwards, Gentili, Horak, Kristensen & Nielsen), ‘The Book REVIEW JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Tortricoidea’ (Horak), “The smaller Microlepidoptera- Grade Superfamilies’ (Dugdale, Kristensen, Robinson, Scoble), “The Pyraloidea’ (Munroe & Solis), “The Axioidea and Calliduloidea’ (Minet), ‘The Butterflies: Hedyloidea, Hesperioidea, and Papilionoidea’ (Ackery, de Jong & Vane-Wright), ‘The Drepanoid/Geometroid Assemblage’ (Minet & Scoble), “The Bombycoidea and their relatives’ (Lemaire & Minet), and “The Noctuoidea’ (Kitching & Rawlins). All these chapters have a standard format (likely owing to the editor's guidance), are well illustrated, and provide detailed accounts on classification and basic morphology of adults and early stages. No other book on Lepidoptera combines such a breadth of information and extensive citations. Although relationships between groups are discussed, chapters 4-19 do not include phylogenies. I found this a sensible editorial decision. | Modern phylogenetic analyses are not homogeneously available for Lepidoptera groups, and discussions of conflicting hypotheses are better placed in the primary literature. Accordingly, chapters 4-19 rest on the solid ground of classical systematics and will remain lasting references for future generations. Chapters 20 and 21, “Evolution of Larval Food Preferences in Lepidoptera’ (Powell, Mitter & Farrell) and ‘Biogeography of the Lepidoptera’ (Holloway & Nielsen), are excellent reviews and should be required reading for general entomologists, and lepidopterists in particular. Volume 2 : Morphology, Physiology, and Development includes 19 chapters that cover all life stages. Here the chapters are slightly less homogeneous in format and coverage, and some are more thorough than others. Chapters 1 and 2 open the volume with detailed information about the ‘Integument’ (Chauvin & Kristensen) and ‘Hairs and scales’ (Kristensen & Simonsen). There one may learn, for example, that the relationship between scale size and wing length is not linear, and that several structures of the integument are still poorly understood. Chapter 3, ‘Coloration: Patterns and Morphogenesis’ (Nijhout) is a brief summary of classical and recent topics and covers both butterfly and moth color patterns. Chapters 4 and 5, ‘Skeleton and muscles: adults’ (Kristensen) and ‘Skeleton and muscles: immatures’ (Hasenfuss & Kristensen) make up one- fourth of volume 2, and these two chapters are not for the novice. They are a useful source of comparative information best appreciated by those who have wrestled with these topics for some years. Comprehensive descriptions of adult and larval muscles are provided together for the first time, and VOLUME 61, NUMBER 3 furthermore, they attempt to standardize the terminology for many skeletal structures—something that is particularly timely for genitalic morphology. Chapters 6-9 and 16 concern the “Digestive and excretory systems’ (Barbehenn & _ Kristensen), ‘Respiratory system’ (Wasserthal), ‘Circulation and thermoregulation’ (Wasserthal), and ‘Nervous system’ (Yack & Homberg) and ‘Reproductive organs’ (Kristensen). These chapters remind me of R. Chapman's original work The insects: structure and function. Although focused on Lepidoptera, these chapters are widely relevant to entomologists and provide up to date, detailed information and useful illustrations for early stages and adults. The chapters covering ‘Sensilla and proprioceptors’ (Hallberg, Hansson & Léfstedt), “Auditory and sound producing organs’ (Minet & Surlykke), and “Eyes and vision’ (Warrant, Kelber & Kristensen) provide a wealth of comparative detail that characterizes the two volumes of the Handbook. At risk of stating the obvious, it is extremely useful to be able to cross consult chapters. For example, Minet & Surlykke note that the forewing base tympanal organs are always innervated by a branch of the nerve IINIc (p 289), which is illustrated by Yack & Homberg (p 234). Chapters 13-16 explore ‘Exocrine Glands: Chemical communication and chemical defense’ (Hallberg & Poppy), ‘Labial glands, silk and saliva’ (Akai, Hakim & Kristensen), ‘Endocrine glands and hormones’ (Sridhara, Bhaskaran & Dahm), and ‘Reproductive organs’ (Kristensen). These reviews encompass processes from the cellular to ecological levels, and point to huge gaps in our knowledge of certain glands. Finally, chapters 17-19 include ‘Karyology and sex determination’ (De Prins & Saitoh), ‘Eggs’ (Fehrenbach), and “Embryology (Kobayashi, Tanaka & Ando). These three chapters are framed in a broad, order-level comparative context and present valuable baseline information. Perhaps the most important contribution of the Handbook is the scope of comparative information conveyed in the two companion volumes. Readers can browse the chapters like the stacks in a library. This encourages scholarship by providing easy access to topics of direct or potential relevance to our research on Lepidoptera. The Handbook serves as a reminder that a holistic approach to research, including familiarity with embryology, host plants, wings, systematics, genitalia, sensilla, et cetera, will lead to a more sound interpretation of our data, whatever they are, and ultimately a deeper understanding of our study organisms. More than ever, neadertic biologists in the US face a demand for increased productivity through extramurally funded research. Although productivity, is positive in principle, sacrificing quality for quantity, or depth for pace are not. The Handbook was done in the fine European tradition of bringing international contributors together with comprehensive excellence as the main goal. Some chapters combine many lifetimes of comparative research (e.g., ‘Skeleton and muscles’), and their production required dedicated work that can only be accomplished through scholarly maturity in the best sense of the word. Inasmuch as the mission of the Lepidopterists’ Society is to further our understanding of moths and butterflies in all aspects of their natural history and evolution, I feel that the Handbook is a giant step towards this goal. These two volumes will certainly stimulate us to dig deeper, look further, think laterally, and read more broadly. The cost of these books is prohibitive to students, but not beyond the reach of professionals. In closing I would like to say, Professors: share the Handbook of Zoology - Lepidoptera, Moths and butterflies with your students — they will not find such a wealth of information on the web. CarLA M. PENZ, Department of Biological Sciences, University of New Orleans, New Orleans LA 70148, USA. e-mail: cpenz@uno.edu. 178 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY COVER PHOTOS SOLICITED The Journal solicits high-quality color photographs for consideration as cover illustrations. The photographs may illustrate any aspect of Lepidopteran biology. Digital image files saved to a CD are preferred. Submit digital files with a vertical orientation of approximately 1770 pixels by 2200 pixels at a resolution of at least 300 pixels/inch ( 120 pixels/em) in *.tif format. If you wish to submit printed photographs, please ensure that they are vertical in orientation, measuring approximately 5.5. inches wide x 6.75 inches long (or 14 cm wide x 17 cm long). Date of Issue (Vol. 61, No. 3): 24 October 2007 rtey | RHR A) A allie (ig Bh iene Ais AYES deh, ie APAR Aliie ce : ee Pree | ie ee a fs eile, ob a a Si wats Me Lice) ARR ae Ca i Cie dion i t ale Usa eel A Bt haar e i Sil ie ait en A aia wil in =f ve ALLO pea Pe ee, EDITORIAL STAFF OF THE JOURNAL Brian Scuotrens, Editor Biology Department College of Charleston, 58 Coming Street, Charleston, South Carolina 29424-0011, USA scholtensb@cofc.edu Pec Toutver, Layout Editor Put DeVries, Book Review Editor Natural Imprints Department of Biological Sciences 706 Lake Road University of New Orleans, 2000 Lakeshore Dr. Eureka, Illinois 61530 New Orleans, Lousiana 70148-0001 USA naturimp@mtco.com pdevries@uno.edu Associate Editors: Gerarbo Lamas (Peru), Kenetm W. Puitie (USA), Rosert K. Rospins (USA), Fevix Spertinc (Canada), Davin L. Wacner (USA), Curister WikLunp (Sweden), Caria Penz (USA), ANDREW Warren (USA), NOTICE TO CONTRIBUTORS Contributions to the Journal may deal with any aspect of Lepidoptera study. Categories are Articles, Profiles, General Notes, Technical Comments, Book Reviews, Obituaries, Feature Photographs, and Cover Illustrations. Obituaries must be authorized by the president of the society. Requirements for Feature Photographs and Cover Illustrations are stated in Volume 44(2):111 and on the Society's web site at http:/Avww.lepsoc.org/. Send Journal submissions electronically to: scholtensb@cofc.edu or to the editor at the above address. Contributors should feel free to recommend one or two reviewers upon submission of their manuscript. Send requests to review books for the Journal and book review manuscripts to Phil DeVries at the above address (or electronically to: pdevries@uno.edu). Short manuscripts concerning new state records, current events, and notices should be sent to the News, Dale Clark, 1732 South Hampton Rd, Glenn Heights, TX 75154-8530. Before sub- mitting manuscripts or electronic files, consult instructions on the Society's web site at http:/Avww.lepsoc.org/. Although we prefer electronic submissions, authors may submit hard copy if they are unable to send material electronically. Submit manuscripts in triplicate, typewritten, entirely double-spaced, with wide margins, on one side only of white letter-sized paper. Prepare manuscript according to the following instructions, and submit them flat, not folded. Submit original illustrations only after the manuscript has been revised and accepted. Authors should provide an electronic copy of the final accepted manuscript in a standard PC or Macintosh-based word processing format. Abstract: Include an informative abstract for Articles, Profiles, and Technical Comments. Additional key words: Up to five key words or terms not in the title should accompany Articles, Profiles, General Notes, and Technical Comments. Text: Write with precision, clarity and economy, and use the active voice and the first person whenever appropriate. Make title explicit, descriptive, and as short as possible. The first mention of a plant or animal in the text should include the full scientific name with author, and family. Use metric units for measurements, and a 24-h clock (0930 h, not 9:30 AM) to express time. Underline only where italics are intended. Literature cited: References to Articles, Profiles, General Notes, and Technical Comments should be given as Sheppard (1959) or (Sheppard 1959, 1961a,b) and listed alphabetically under the heading Literature Cited, in plain text (do not capitalize author names) and without underlining as follows: Sheppard, P. M. 1959. Natural selection and heredity. 2nd ed. Hutchinson, London. 209 pp. —— 196la. 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Figures, both line drawings and photographs, should be numbered consecutively in Arabic numerals; do not use “plate.” Type figure legends double-spaced, on a separate sheet (not attached to the illustrations), headed Explanation of Figures. Use a separate paragraph for each legend. Color illustrations are encouraged; contact editor or consult our web site for submission requirements and cost. Tables: Number tables consecutively in Arabic numerals. Label them in plain text (e.g., 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 66044 U.S.A. iil ! 9088 01387 2064 It CONTENTS Taxonomy oF Four Species oF EucosmMini (TorrrictbpaAz) AssociateED WitH PELOCHRISTA COROSANA (WaLsINGHAM) INcLUDLUDING A New Synonymy AND DescripTION oF A NEW _ SPECIES Donald J. Wright -------------------------------------------------------------------------=--=--=------- LLY Tue Butrerrty Drawincs py Joun Appor IN THE Harcretr Rare Book anb Manuscripr Liprary, University or Georcia John V. Calhoun --------------------------------------------------=-------- 125 DNA Barcobes or CLOSELY RELATED (BUT MORPHOLOGICALLY AND ECOLOGICALLY DistTINcT) SPECIES oF Skipper Burrerevies (HeseerubAk) CAN Dirrer By ONLY OnE TO THreeE NUCLEOTIDES John M. Burns, Mehrdad Hajibabaei, Winnie Hallwachs and Paul D.N. Herbert ----------- 138 Tue INFLUENCE OF HurricANE AND TropicaL StorM Activity ON RestIpENT BUTTERFLIES IN THE Lower Fiorina Keys Mark H. Salvato and Holly L. Salvato ------------------------------------ 154 First Report of OgcorHora BRACTELLA (L.) (OkcopHortDAE) IN Nortu America Merrill A. Peterson, Eric H. LaGasa, Steven Passoa, Gaden S. Robinson and David Holden ---------- 165 GENERAL NOTES First Report or THE PaLeARcTIC SPECIES CypIA CONIFERANA (TORTRICIDAE) IN THE WESTERN Unitep States Eric LaGasa and Steven Passoa -------------------------------------------------=-- 2: Poanes MELANE (HesprertpaAk) OvrpostriING ON AN AUSTRALIAN Grass NaruRALIZED IN CALIFORNIA Arthur M. Shapiro --------------------------------------------------- 7-2-0222 nnn nnn nnnnnnnnnnn 175 Book Review Hanpsooxk or Zooiocy, Voutume IV, Arruroropa: Insecta, Epirep spy M. FIsHeEr, Part 35 Leprpoprera, Morus AND Butrerruirs. Carla Penz ----------------------------------=--==- 176 @ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanance of Paper).