Volume 59 Number 3 28 October 2005 fle ISSN 0024-0966 ENT Journal of the Lepidopterists Society Published quarterly by The Lepidopterists’ Society THE LEPIDOPTERISTS’ SOCIETY Executive CouNcIL Feuix A. H. Speriinc, President Jens Rotanp, Vice President James K. Apams, Immediate Past President Ernest H. Wittiams, Secretary Curtis J. Cattacuan, Vice President Keiy M. Ricrers, Treasurer Timotrny L. McCase, Vice President Members at large: Akito Kawahara Robert M. Pyle Richard A. Anderson Jane M. Ruffin John A. Shuey John V. Calhoun Erik B. Runquist Andrew D. Warren Amanda Roe Eprrorrat Boarp Caria M. Penz (Chairman) Micuact E. Tottver ( Journal) Lawrence F. Gait (Memoirs) Puitup J. Scuarrert (News) Joun A. Sxyper (Website) Joun W. Brown (at large) Honorary Lire MEMBERS OF THE SOCIETY Cuartes L. RemMincton (1966), E. G. Munror (1973), [an F. B. Common (1987), Lincotn P. Brower (1990), Freperick H. Rinpce (1997), Ronatp W. 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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, % Natural History Winesnna, 900 Exposition Blvd., Los Angeles, CA 90007-4057. Cover illustration: Monarch, Danaus plexippus (Linnaeus), and milkweed aphids, feeding on Swamp Milkweed, Asclepias incarnata L., September, 2005, central Illinois, USA. Photo by Peg Toliver VOLUME 59, NUMBER 3 JouRNAL OF Tue LeEepripopteRistTs’ SOCIETY Volume 59 2005 Number 3 Journal of the Lepidopterists’ Society 59(3), 2005, 121-133 SOME EUCOSMINI (TORTRICIDAE) ASSOCIATED WITH EUCOSMA EMACIATANA (WALSINGHAM) AND EUCOSMA TOTANA KEARFOTT; FOUR NEW SPECIES, A NEW COMBINATION, AND A NEW SYNONYMY DONALD J. WRIGHT 3349 Morrison Ave., Cincinnati, Ohio 45220-1430, USA email: wrightdj@fuse.net ABSTRACT. Eucosma emaciatana (Walsingham) is transferred to Pelochrista Lederer, and Pelochrista perpropinqua (Heinrich) is recognized as a junior synonym of P. emaciatana. Three species considered by previous authors to be superficially similar to ema- ciatana are reviewed: Eucosma larana (Walsingham), Eucosma totana Kearfott and Pelochrista popana (Kearfott). Four previously unrecognized species are described: Eucosma piperata, new species, Eucosma nordini, new species, Eucosma taosana, new species, and Pelochrista powelli, new species. Lectotypes are designated for emaciatana and larana. Adults and genitalia of these species are illustrated, and new distributional records are presented. Additional key words: Oletreutinae, Pelochrista, Nearctic. The Rocky Mountain and Great Basin regions of western United States are home to many camilan looking species of Eucosmini, some of which were named by Walsingham in the last quarter of the nineteenth century. Lacking access to the Walsingham types, early North American tortricid specialists often had difficulty applying those names, and in some instances the confusion that resulted has persisted to the present day. One such case involves Eucosma emaciatana (Walsingham), Eucosma larana (Walsingham), and Pelochrista perpropinqua (Heinrich). Heinrich (1923) confused emaciatana with larana (see discussion below) and later (1929) described perpropinqua based on a series of specimens of emaciatana. I confirmed that emaciatana and perpropinqua refer to a single taxon by examining the types. Based on male genitalia, the appropriate generic assignment for this species is Pelochrista Lederer, a conclusion reached by Powell (1983) in his placement of perpropinqua, so I propose to resolve this situation by transferring emaciatana to Pelochrista and treating perpropinqua as a junior synonym. In examining specimens from various institutional and private collections I encountered two previously unrecognized species of Eucosma Hiibner that have been confused with larana. They are described below as E. piperata, new species, and E. nordini, new species. Also included are reviews of E. totana Kearfott and P. popana (Kearfott), two species considered by previous authors to be similar in appearance to larana and/or emaciatana. Finally, descriptions are provided for two additional new taxa, Eucosma taosana, new species, and Pelochrista powelli, new species. The former has previously been misidentified as totana; the latter is superficially similar to totana and taosana. Walsingham (1884) described Paedisca emaciatana from three male specimens collected by H. K. Morrison in Arizona. Fernald [1903] placed this species in Eucosma, and there it has resided ever since. The Fernald collection, acquired by the United States National Museum (USNM) in 1924-25, included two male specimens determined by Walsingham as emaciatana. Neither has an abdomen. One was collected by Morrison in Arizona in 1883 and agrees with the description of emaciatana, the other is lacking collection data and is in such poor condition that I cannot confirm the accuracy of its determination. Heinrich's review (1923) of emaciatana makes no mention of these two specimens, so I assume he did not examine them. His treatment was based on a series of specimens from Utah, and he illustrated the genitalia (Fig. 193) of a male collected by Tom Spalding at Eureka, Utah, on 27 July 1911. I examined that specimen and a number of other USNM specimens determined by Heinrich as emaciatana and concluded (see discussion below) that they represent E. larana. This explains why Heinrich, when presented with specimens of emaciatana collected in Arizona by O. C. 122 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 1-12. 1, P. emaciatana, lectotype male. 2, E. larana, lectotype male. 3, E. larana, male, Albany Co., Wyoming. 4, E. piperata, male, Oneida Co., Idaho. 5, E. nordini, holotype male. 6, E. larana, female, Oneida Co., Idaho. 7, E. totana, male, Grand Co., Colorado. 8, E. totana, male, Oneida Co., Idaho. 9, E. taosana, holotype male. 10, P. emaciatana, male, Cochise Co., Arizona. 11, P. popana, male, Larimer Co., Colorado. 12, P. powelli, holotype male. VOLUME 59, NUMBER 3 Poling, interpreted them as representing a new species, which he described (1929) as E. perpropinqua. Walsingham (1879) described E. larana from three specimens (2 ¢, 12) collected in Siskiyou County, California. The forewing (Fig. 2) of the lectotype (designated below) is white with pale brownish-orange markings; that of the female paralectotype is white with a few black specks and only a hint of brownish-orange markings. The specimens misidentified by Heinrich as emaciatana have pale yellowish-white forewings that are generously overlaid with pale brownish- -orange coloration. They also have brownish-gray markings (as in Fig. 3). Specimens I collected in Southeastern Idaho (Fig. 6) have very pale yellowish-white forewings with just a trace of brownish-orange coloration. I found no substantial differences in the male and female genitalia of these various specimens, and lacking any distinguishing biological information, I concluded that they all represent a single variable species. Curiously, Heinrich (1923) did correctly identify the male that he illustrated (Fig. 197) as larana. Two of the new species described below are similar to larana in forewing color, and each is sympatric with /arana in at least a portion of the latter species! range: piperata in Utah and southeastern Idaho, nordini in southeastern Wyoming. Kearfott (1907) reported a type series for totana consisting of five specimens collected by Tom Spalding and O. C. Poling in Stockton, Utah, and South Utah, respectively. The American Museum of Natural History (AMNH) has a male collected in So. Utah labeled LECTOTYPE, a designation Klots (1942) attributed to Heinrich (1923). I assume from Kearfott's remarks that the collector was Poling, but there is no such indication on the pin labels. I located four Spalding specimens from Stockton, Utah, that are likely to be the other syntypes. Only one is actually totana, a male in the USNM with no capture date. Both it and the lectotype bear the handwritten label “Eucosma totana Cotype Kearf.” and Kearfott's printed red label “TYPE Collection of W. D. Kearfott”. A female in the AMNH collected VIII-4-4 and bearing the red Kearfott “TYPE” label is no doubt the specimen referred to by Klots (1942) as a paralectotype, but its genitalia indicates it is not totana. The remaining two, a USNM specimen dated VIII-30-4 and an AMNH specimen dated VII-1- 4, are conspecific with a series of USNM specimens determined by Heinrich as totana but bearing a hand written label with the notation “dark var. ’ They agree with specimens I collected in New Mexico that are sufficiently distinct in maculation and genitalic details to justify separate species status. Although I was unable to locate females of this taxon, the likelihood of it being confused with totana prompted me to describe it here as E. taosana. This investigation also brought to my attention the previously unrecognized P. powelli, which can be confused with totana and taosana. Finally, Kearfott (1907) based his description of popana on 27 syntypes collected by Tom Spalding at Stockton, Utah, with capture dates between 1 June and 8 August. Klots (1942) reported thirteen specimens in the AMNH as belonging to the type series, including one labeled LECTOTYPE. The lectotype designation should be credited to Klots (1942), even though he attributes it to Heinrich (1923). I examined this material and found the lectotype to be a female, rather than a male as stated by Klots. I believe nine of the other twelve specimens are popana, but one has a capture date of 9 August, which is inconsistent with Kearfott's remarks. Of the remaining three, two are males of Epiblema sosana (Kearfott), and one is a female of uncertain identity. Listed below as paralectotypes are the eight AMNH popana specimens whose capture dates agree with Kearfott's statements and seven USNM specimens that I judge to belong to the popana type series. I am designating lectotypes for emaciatana and larana and have attached designation labels to those specimens. The specimens chosen for this purpose were originally selected by Obraztsov, but his designations were never published. MATERIALS AND METHODS This study is based on 607 adult specimens and 111 associated genitalia prepar ations. I examined the types of the five previously described species. Material was borrowed from the following institutional and private collections: AMNH, George if Balogh (GJB), Canadian National Collection (CNC), Colorado State University (CSU), Essig Museum of Entomology (EME). Clifford D. Ferris (CDF), Los Angeles County Museum of Natural History (LACM), Museum of Comparative Zoology (MCZ), John S. Nordin (JSN), The Natural History Museum, London (BMNH), USNM, Donald J. Wright (DJW), and University of Wyoming (UWY). The line drawings were based on images generated by a Ken-A-Vision microprojector ( (Model X1000-1), and each associated scale bar represents 0.5 mm. Images of the genitalia of the lectotypes of larana and emaciatana were obtained by scanning negatives of photographs taken by Obraztsov of slides he had prepared. All measurements were estimated to the nearest tenth of a millimeter with the aid of a reticule mounted in a Leica MZ95 stereomicroscope. Ratios of measurements were rounded to two decimal places. Forewing length (FWL) indicates the distance from base to apex, including fringe. Aspect ratio (AR) refers to the ratio of FWL to forewing width, the latter quantity being measured midway between base and apex. In males, the ratio of forewing costal fold length to FWL is denoted by CFR (costal fold ratio), and the ratio of valval neck width to width of basal portion of valva by NR (neck ratio). Reported values of AR, CFR and NR are averages of such values calculated for a small sample of specimens. The number of items supporting a particular statistic is indicated by n. Some species discussed here only vaguely display the putative, ancestral, fasciate forewing pattern for the Tortricidae discussed by Brown and Powell (1991) and Baixeras (2002), but their terminology is used when possible in the forewing descriptions. SPECIES ACCOUNTS Eucosma lence (Walsingham) (Figs. 2, 3, 6, 14, 15, 25, 36) Paedisca larana Tate 1879:43. Eucosma larana: Fernald [1903]:456; Barnes and McDunnough 1917:169: Heinrich 1923:110: McDunnough 1939:47; Powell 1983:34. Eucosma emaciatana: (not Walsingham 1884) Heinrich 1923:108; McDunnough 1939:46; Powell 1983:34. Types. Lectotype here designated (Figs. 2, 14): ¢ Sheep Rock, Siskiyou Co., California, W% alsingham, 3 Sept. 1871, genitalia slide 11502 BMNH. Paralectotypes: same data as lectotype (1 d, 1 2, 2 genitalia slide 11759, BMNH). Diagnosis. Darkly marked specimens of larana can be recognized by forewing pattern (Fig. 3), but pale specimens (Fig. 6) might be confused with nordini or piperata. The forewing of nordini (Fig. 5) is pale yellowish white, has a gray streak on the costal fold, and shows no indication of brownish-orange mottling. The combination of white forewing color, black speckling, and brown costal marks distinguishes piperata (Fig. 4). One can also separate larana, piperata, and nordini by the shapes of the sterigmata (Figs. 36, 35, 31) and by 13 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY subtle but consistent differences in valval shape (Figs. IUay, 1s}, SAIL), Description. Head: Very pale yellowish white, long scales of upper frons and vertex often with pi ale gray shading prece ding whiter apices; labial palpus with pale-gray lateral surface, third segment enclosed by long narrow scales of se cond se ement; antenna ye lowe white. Thorax: inercal and ventral surfaces concolorous with head, legs yellowish white to pale yellowish brown. Forewing (Figs. 2, 3, 6): 3 FWL 8.5-12 mm (mean = ee n = 13), AR = 3.16, CFR = 0.3, ¢ FWL 8.5-11.1 mm (mean = 10, n = 10), AR = 3.05; costa nearly straight, apex mildly acute, ee n i duct dorsal surface white to pale yellowish w hite and variably ov ltl td pale brownish-orange mottling, darker specimens with four brownish- gray marks, the first a triangular pretornal mark on dorsum, often ah a few black scales at its anterior extremity, the second a subbasal mark on fold that is usually connected to dorsum by brownish- -orange scaling, the third at distal end of cell, often divided longitudinally by a brownish-orange streak, the fourth anterior to ocellus; ocellus obscure, variably overlaid with brownish-orange scales and crossed longitudinally by up to four black dashes; distal one-half of costa with fon obscure, paired, white strigulae; male costal fold usually grayer than adjacent portion of wing, fringe white to yellowish white, with gray shading near apex and »rownish-orange suffusion near tornus. Hindwing: Uniformly pale brownish gray with lighter fringe. Male genitalia (Fig. 14, 15): Uncus divided medially into two variably deve eloped setose lobes with convex ateral margins; dorsolateral shoulders of tegumen well developed, often rounded and hunched, sometimes with angular corners; socii ong, flat, and densely setose; vesica with 2-7 deciduous cornuti (n = 10): valva with costal margin concave, apex and ventral angle evenly rounded, distal margin very weakly convex, ventral invagination moderate, NR = 0.61, cucullus of nearly uniform width, medial surface densely setose, sacculus sparsely setose, margin of basal opening with patch of short slender spines. Female genitalia (Fig. 25): Papillae anales facing ventrolaterally and densely setose, medial margins very weakly sinuate, surfaces finely ridged transversely, long setae on lateral mar: gins curving ventrally, remaining setae shorter with hooked apices; tergum VIII sparsely setose; lamella antevaginalis (Fig. 36) ringlike and weakly sclerotized, lamella postv: vaginalis with semitriangular posterolateral corners; membrane between sterigma and eee extremities of tergum VIII setose; sternum VII with posterior margin approximate to sterigma and roundly invaginated to depth of one-half length of sterigma; ductus bursae_ strongly constricted anterior to ostium, widening anteriorly; corpus bursae with large signum near juncture with ductus bursae and small spike-shaped signum on opposite wall. Distribution and biology. I examined 53 specimens (40 d, 13 2) from the following states and counties: CALIFORNIA: Siskiyou, Tulare; IDAHO: Oneida; UTAH: Juab; WYOMING: Albany. The flight period extends from early July to the beginning of September, and capture sites range in elevation from 5000! to 8000". FIGS. 13-14.Genitalia of lectotypes. 13, P. emaciatana, slide BMNH 11571. 14, E. larana, slide BMNH 11502. VOLUME 59, NUMBER 3 23 Fics. 15-23. Male genitalia. 15, E. larana, slide DJW 763. 16, E. totana, slide DJW 1022. 17, P. emaciatana, slide DJW 952. 18, E. piperata, slide DJW 762. 19, E. taosana, slide DJW 1035. 20, P. popana, slide DJW 1068. 21, E. nordini, slide DJW 760. 22, P. powelli, slide DJW 1027. 23, P. powelli, slides DJW 1032 and 705 This moth has been collected in open sage brush habitat in Idaho and Wyoming. No larval host has been reported. Comments. The variation in forewing color appears to have a geographic component. Specimens from northern California and southeastern Idaho have very pale yellowish-white forewings with pale to nearly obsolescent brownish-orange markings, those from Wyoming tend to be darker, with brownish-gray markings and extensive brownish-orange mottling, and those from Utah and central California appear to be intermediate. The medial division of the uncus varies from an inconspicuous line to the pronounced indentation illustrated in Figure 15. Eucosma totana Kearfott (Figs. 7, 8, 16, 28, 32) Barnes and 1923:108; Eucosma totana Kearfott 1907:32; McDunnough 1917:169: Heinrich McDunnough 1939:46; Powell 1983:34. Eucosma spodias: Meyrick 1912:35. Types. Lectotype designated by Heinrich (1923): ¢, South Utah, July 1900, genitalia slide CH, 2 Dec 1919, AMNH. Paralectotype ¢: Stockton, Utah, Tom Spalding, USNM. Diagnosis. This species can be confused with taosana, popana and powelli, but the following combination of dark brown forewing markings usually suffices for diagnosis: a subbasal mark on fold, a thin line along fold from subbasal mark to tornus, a pretornal triangular mark based on fold, and a chevron shaped mark at distal end of cell. Some specimens do not show the line on the fold. Superficially, totana is most similar to taosana (Fig. 9), but the latter species does not have 126 a dark line on the fold or a chevron shaped mark at the end of the cell. Male genitalic characters separating totana and taosana include: subtle differences in shape of cucullus (Figs. 16, 19), distinctly different shape of uncus, and number of cornuti in vesica (10 for totana vs. 5 for taosana). Eucosma totana is easily separated from popana and powelli by the presence in the latter two species of a stout spine at the ventral angle of the cucullus. Females of taosana are not known, but the sterigmata (Figs. 32, 37, 34) of totana, popana and JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY powelli are easily distinguished from one another. Description. Head: Scales of frons and vertex white, sparsely marked with pale gray; labial palpus porrect, length ca. 3x eye diameter, second segment with medial surface and dorsal margin white, lateral surface pale brownish gray, scales of ventral and dorsal margins long and slender, concealing third segment; antenna white. Thorax: Scales of dorsal surface and tegulae white basally and apically, brownish gray medially, producing a speckled effect; ventral surface and hindlegs white, fore and midlegs white posteriorly, brown to pale brown anteriorly, with white annular markings on tarsus and tibia. Forewing (Figs. 7, 8): 0 FWL 8.7-12 mm (mean = 10.1, n = 68), AR = 3.2, CFR = 0.27, ° FWL 7.4-10.5 mm (mean = 8.9, n = 18), AR = 3.11; costa nearly straight, apex acute, termen weakly convex; dorsal FIGS. 24-25. Female genitalia. 24, P. emaciatana, slide DJW 989. 25, E. larana, slide DJW 789. VOLUME 59, NUMBER 3 26 27 Fics. 26-27. Female genitalia. 26, E. piperata, slide DJW 1061. 27, E. nordini, slide DJW 1073 surface white with brown to brownish-black irrorations on basal two- thirds and five brownish-black marks, the first a subbasal mark on fold, sometimes extending weakly to dorsum, often bordered distally by a thin line of black scales, the second a very thin line along fold from subbasal mark to tornus, the third a triangular mark based on fold and projecting anteriorly along basal margin of ocellus, the fourth a longitudinally elongate mark anterior to ocellus, narrowing basally and extending into cell, the fifth a chevron on distal margin of cell, sometimes connecting anteriorly to mid costa and posteriorly to line on fold, often divided medially by aforementioned longitudinal mark, ocellus bordered on basal, distal and tornal margins with lustrous, pale, yellow-brown to yellow-gray bars, white central field crossed by up to four, black, longitudinal dashes; costal margin brownish black, crossed by numerous paired white strigulae, numbers four through nine usually sharply delineated; dorsal margin with 10-12, small, evenly spaced, brownish-black marks; scales along terminal margin white with subapical black markings, fringe usually whiter with more extensive dark markings between M1 and M3. Abdomen: Scales on posterior margin of eighth segment in females brownish black with white apices. Hindwing: Uniformly pale brownish gray with lighter fringe. Male genitalia (Fig. 16): Uncus triangular, dorsal surface setose, posterior surface developed into medial wedge-shaped ridge, dorsolateral shoulders of tegumen well developed; socii long, flat. and densely setose; aedeagus tapered distally, vesica with ca. 10 deciduous cornuti (n = 13); valva with costal margin concave, apex and ventral angle evenly rounded, distal margin convex, invagination of ventral 125 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY FIGS, 28-29. Female genitalia. 28, E. totana, slide DJW 1028. 29, P. popana, lectotype, slide DJW 1085. margin moderate, NR = 0.6, cucullus with densely setose medial surface, sacculus and margin of basal opening moderately setose. Female genitalia (Fig. 28): Papillae anales facing laterally and densely setose, surfaces finely ridged transversely, setae on medial margins with hooked apices, those on lateral margins twice as long and curving ventrally; posterior margin of tergum VIII with 3-4 rows of setae; lamella antevaginalis (Fig. 32) ringlike and very weakly sclerotized, lamella postvaginalis well developed, width of posterior margin ca. 2x ostium diameter, posterolateral corners acute, a depressed trough from mid posterior margin to ostium; membrane between sterigma and ventral extremities of tergum VIII setose; sternum VII with length of posterior margin ca. 3x ostium diameter, roundly invaginated to depth of one-third length of sterigma, approximate to sterigma medially; ductus bursae constricted anterior to ostium; corpus bursae with two signa. Distribution and biology. My study sample included 128 specimens (105 4, 23 2) from the following states and counties: ARIZONA: Coconino; COLORADO: Chaffee, Fremont, E] Paso, Grand: IDAHO: Lincoln, Oneida; MONTANA: Jefferson; NEW MEXICO: Santa Fe; OREGON: Harney; UTAH: Juab, Sanpete; WYOMING: Albany. I have occasionally found this species to be abundant in sagebrush habitat at elevations between 5000' and 8000'. Brown et. al. (1983) reported Chrysothamnus nauseosus (Pall.) Britt. (Asteraceae) as a larval host in Idaho. Comments. The forewing markings are stable, but the overall appearance of totana varies from very pale tan to medium brown. In lighter specimens the brown irrorations are restricted to the basal one-third of the wing, the median area is mostly pale yellowish white to white, and the markings are orangish brown. Darker VOLUME 59, NUMBER 3 specimens are much more densely irrorated and have dark brownish-black markings. Eucosma piperata Wright, new species (Figs. 4, 18, 26, 35) Diagnosis. Reasonably fresh specimens of this species are readily identified by forewing g pattern: white, peppered with minute black specks, rath brown marks on distal one-half of costa. Description. Head: Frons and vertex white, a small patch of pale- brown scales anterior to eye; labial palpus with medial surface white, lateral surface pale brown; antenna white, scape sometimes pale brown dorsally. Thorax: Dorsal and ventral surfaces white; legs with anterior surfaces pale brown, posterior surfaces white, and distal ends of tarsal segments lightly ringed with white. Forewing (Fig. 4): d FWL 10.8-11.5 mm (mean = 11.2, n = 6), AR = 3.11, CFR = 0.32, 2 FWL 11.5-13.5 mm (mean = 12.6, n = 8), AR = 2.99: distal three- fourths of costa straight, apex mildly acute, termen weakly convex; dorsal surface white, sparsely speckled with black scales between radial vein and dorsum, costal strigulae delimited by narrow, brown, costal marks but otherwise not distinguishable from ground color, a conspicuous, oblique, brown mark at apex; ocellus obscure, variably marked on basal and distal margins by a few black and brown scales, respectively, ca. three, black, weakly expressed, longitudinal dashes in central field; costal fold of male pale grayish brown along costal margin; fringe scales white basally, very pale orange brown distally. Hindwing: White, a shade grayer than forewing, fringe white. Male genitalia (Fig. 18): Uncus a semicircular dorsally setose lobe, supported laterally by well developed and mildly hunched shoulders; socii densely setose, tapered distally, with lateral margins variably serrate; gnathos a narrow band; vesica with ca. 14 deciduous cormuti (n = ays valva with costal margin concave, apex rounded but moderately acute, distal margin convex, ventral angle gently rounded, ventral invagination shallow, NR = 0.7, cucullus with medial surface densely setose, sacculus sparsely setose, a patch of setae on margin of basal opening. Female genitalia (Fig. 26): Papillae anales facing laterally and densely setose, medial margins weakly sinuate, surfaces finely ridged transversely, setae toward lateral margins strongly curved ventrally, those near anal opening with hooked apices; posterior one- half of tergum VIII with ca. four rows of setae; lamella antevaginalis (Fig. 35) ringlike; lamella postvaginalis widening posteriorly to ca. 2x ostium diameter, with triangular, mildly setose, posterolateral corners; stemum VII with posterior margin roundly invaginated to three- fourths length of sterigma and approximate to sterigma; ductus bursae weakly constricted anterior to ostium, gradually widening anteriorly; corpus bursae with two similarly shaped signa, membrane finely wrinkled near signa, interior surface of bursa minutely microtrichiate. Holotype. ¢, Vineyard, Utah, 9 July 1912, Tom Spalding, genitalia slide USNM 70391, USNM. Paratypes. ARIZONA: Lupton, A. K. Wyatt, 3 July 1951 (1 genitalia slide DJW 1061). CALIFORNIA: Inyo Ca 9 mi. W. ae Pine, P. D. Hurd & J. A. Powell, 19 July 1961 (1 °); Westguard Pass, White Mts., P. Opler & J. A. Powell, 19 July 1968 (1 2). COLORADO: Mesa Co., Colo. N. Monument, Head of Red Canyon, J. Moore, 3 July 2001 (3 6, 12, 6 genitalia slide DJW 1132, ° genitalia slide DJW1133). IDAHO: Oneida Co., Curlew NG, 4 mi ENE of Holbrook, 5050', D. J. Wright, 18 July 2001 (1 ¢, genitalia slide DJW 762). NEVADA: Nye, Co., Currant Cr. Cpgd., J. Scott, 20 July 1968 (1 2), P. Opler & J. A. Powell, 20 July 1968 (2 2, genitalia slides T. Gilligan 265, DJW 1288); Lincoln Co., Cathedral Gorge St. Pk., J. Doyen, 12/13 July 1971 (12). OREGON: Baker Co., Burnt River Cyn., 3200!, 44E 33.08! N, 117E 39.75' W, C. D. Ferris (1 2, genitalia slide DJW 1134). UTAH: Vineyard, 4 July 1912 (1 2); Vineyard, Tom Spalding, 14 July 1912 (14, genitalia slide DJW 1058); Juab Co., Eureka, ae Spalding, 20 July 1911 (1 2); Sevier Co., Richfield, 15 June 1930 (1d), 15 July 1930 (1 genitalia slide DJW 1060). Paratype dhvactiaaes CDF, CSU, EME. LACM, USNM, DJW. Etymology. The specific epithet, deriving from the Latin word for pepper, refers to the minute black speckling on an otherwise white forewing. Distribution and biology. The 20 specimens (9 ¢, 11 2) reported above suggest that the range of this moth may be restricted to the Great Basin. The flight period extends from mid June through July. The larval host is unknown. Comments. In some specimens the black speckling is barely discernable, but the brown costal marks, particularly the apical one, are usually conspicuous. The wrinkling of the membrane of the corpus bursae in the vicinity of the signa is variable. Eucosma nordini Wright, new species (Figs. 5, 21, 27, 31) Diagnosis. The immaculate pale yellowish-white forewing is diagnostic for this species. Males have a blackish-gray streak along the anterior edge of the costal fold. Description. Head: Upper frons and vertex very pale yellowish white, labial palpus white, lateral surface of second segment with pale gray shading; antenna white. Thorax: Dorsal surface pale yellowish white, ventral surface white, legs pale yellowish white, anterior surfaces sometimes darker. Forewing (Fig. 5): ¢ FWL 10-13.7 mm (mean = 11.9, n = 14), AR = 3.35, CFR = 0.29, 2 FWL 10.8-13 mm (mean = 11.5, n = 5), AR = 3.13; costa straight, apex acute, terminal margin weakly convex; dorsal surface very pale yellowish white, without mar kings, fringe white, males with blackish-gray streak along costal edge of fold. Hindwing: Pale brownish gray, fringe white. Male genitalia (Fig. 21): Uncus semitriangular, apex rounded, lateral margins weakly convex, dorsal surface setose, shoulders of tegumen well “developed: socii long, flat, tapering distally, and densely setose; gnathos a narrow band; aedeagus tapering distally, vesica with 4-10 deciduous cornuti (n = 5); valva with costal margin concave except for slight, elongate, convex protrusion on cucullus, apex semirectangular, distal margin convex, ventral angle gently rounded, ventral invagination shallow, NR = 0.75, cucullus with medial surface densely setose, sacculus and margin of basal opening moderately setose. Female genitalia (Fig. 27): Papillae anales facing laterally and densely setose, surfaces finely ridged transversely, medial margins mildly sinuate, long setae on lateral margins curving ventrally, setae near anal opening shorter with hooked apices: three to four rows of setae on posterior one-third of tergum VIII; lamella antevaginalis (Fig. 31) ringlike; lamella postvaginalis with triangular, _ setose, posterolateral, corners, posterior margin weakly invaginated medially; sternum VII with posterior margin slightly wider than sterigma. roundly invaginated to one-third length of sterigma and approximate thereto; ductus bursae strongly constricted anterior to ostium, gradually widening toward corpus bursae, corpus bursae with large signum near juncture with ductus bursae and smaller spike-like signum on opposite wall posterior to mid bursa, inner surface minutely microtrichiate. Holotype. ¢, Wyoming, Albany Co., Medicine Bow NF, 11.5 mi SE Laramie, Jctn. Forest Rds 707 and 705, 4 August 2001, D. J. Wright, 8220', genitalia slide DJW 760, deposited in USNM. Type locality at 41° 11.75' N, 105° 23.7' W. Paratypes. COLORADO: Chaffee Co., Salida, G. M. and Jel: Sperry, 24 August 1938 (1 2, genitalia slide DJW1165). WYOMING: Albany Co., TI5N 873W Sec. 1, 7450', C. D. Ferris, 25 July 2000 (1 3), 28 July 2000 (1 ¢, genitalia slide USNN 91928), 30 July 2003 is 3), 5 August 2003 (2 3), 9 August 2002 (1 ¢), 9 August 2003 (1 3), 10 August 2002 (3 4, genitalia slide DJW oma 10 August 2003 (1 3), 11 August 2003 (1 3), 13 August 2002 (1 ¢), 14 August 2002 (1 5 15 August 2002 (3 d), 15 August 2003 (2 3), 16 August 2002 (5 ¢), 17 August 2002 (1d, 1), 18 August 2002 (1 ), 18 August 2003 (2 3). 1S 130 August 2002 (5 ¢, 1°), 19 August 2003 (1 22 August 1999 (1 °), 22 August 2002 (1 24 August 2002 (1 ¢), 25 August 2002 (1 °, genitalia slide DJW 1074); Albany Co., TI5N S73W Sec. 1, 2217 Sky View Ln., 7468', J. S. Nordin, 27 July 1994 (1 °, genitalia slide DJW 312), 11 August 1995 (1 ‘, genitalia slide DJW 271), 14 August 2001 (1 '), 16 August 1999 (1 S$), 17 August 2002 (1 6), 19 August 1995 (1 S). 5% August 1998 (1 ), 21 August 1998 (1 ¢, bate slide J. W. Brown 1173), 25 August 1998 1 3); Albany Co., TI5N S71W Sec. 14, E of Pilot Hill Road, 8600’, J. ’, genitalia slide DJW 1078), , genitalia slide DJW 1073), S. Nordin, 25 August 1998 (2 ¢); Albany Co., Upper Blair PG, N. of Rd. 705, J. S. Nordin, 8200', 12 August 2003 (1 2); Albany Co., NE of Pole Mtn., S. of Happy Jack Rd., 8320', J. S. Nordin, 12 August 2001 (1 d); Albany Co., 1.5 mi NW wood Landing, Fox Creek, J. S. Nordin, 7600', 31 July 2002 (1 ¢); Albany Co., 8 mi. NE Laramie, Rogers Canyon, M. Pogue, 22 eae 1980 (3 9, genitalia slide DJW 1135); Albany Co., Medicine Bow NF, 10.5 mi SE Laramie, 8300', D. J. Wright, 4 August 2001 (2 4); Albany Co., Medicine Bow NF, 11.5 mi SE Laramie, $220', D. J. aah 4 August 2001 (1 ¢); Teton Co., Grand Teton NP, Teton Sciences School, P. A. Opler, 3 August 2001 (1 3); Washakie Co., Tensleep Preserve, T47N RS6W S32, 6400', § August 1999, C. D. Ferris (3 3). eee de »positories: AMNH, BMNH, CNC, CDF, CSU, EME, JSN, LACM, USNM, DJW, UWY. Etymology. It is a pleasure to name this species after John S. Nordin, whose extensive collecting around Laramie, Wyoming, has made a significant contribution to our knowledge of the lepidopter ran fauna of that region. Distribution and biology. I examined 115 specimens (104 ¢, 11 2) from the following states and counties: COLORADO: Chaffee; WYOMING: Albany, Teton, Washakie. Capture sites range in elevation from 6400! to 8300'. Flight occurs from late July to the end of August. The larval host is unknown. Eucosma taosana Wright, new species (Figs. 9, 19) Diagnosis. The forewing of taosana has a conspicuous band of orange- brow n scales along the costa and a line of similarly colored scales along 1A+2A. The male genitalia of taosana is similar to that of totana (Figs. 16, 19), but the apex of the cucullus is more angular and the uncus lacks a wedge shaped posterior projection. Description. Head: Lower frons white, scales of vertex brownish gray medially, lighter toward base and apex; labial palpus with medial surface white, lateral surface brown; antenna brown. Thorax: Dorsal surface brown, scales on apex of tegulae brownish black with white ue es, ventral surface pale tan, legs with anterior surfaces dark gray- brown, posterior surfaces pale tan, distal e xtremities of tarsal segments with pale tan. Forewing (Fig. 9): 0 FWL 7.5-9.5 mm (mean = 12), AR = 3.19, CFR = 0.31; costa and termen nearly straight, surface brown with brownish-black markings, a -brown coloration from base to apex between costa and radial vein, a narrow similarly colored band from base to tornus along 1A+2A, a brownish-black, outwardly oblique, subbasal mark on dorsum extending forward into cell, a triz angular, brownish-black, subtornal mark on dorsum projecting ‘ anteriorly along basal margin of ocellus, both marks divided by orange-brown line along 1A+2A, a narrow elongate patch of white- -tipped, dark grayish-brown scales anterior to ocellus, extending and tapering basz ally to middle of cell, mildly constricted at distal end of cell: ocellus with basal, distal and tornal margins pale yellowish brown to yellowish gray, central field white to pale brown, crossed longitudina ly by 3-4 brownish-black ape) acute Brea band of orang dashes, the latter often connected in zig-zag pattern; distal one-half of costa usually with four, sharply deaneds paired, white strigulae, costal fold on male brownish black; termen with band of white- tipped JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY brownish-black scales from apex to tornus, fringe scales similarly marked near apex, lighter and more uniformly brownish gray toward tornus. Male genitalia (Fig. 19): Uncus triangular and dorsally setose, divided medially by shallow indentation; dorsolateral shoulders of tegumen well developed and hunched; aedeagus long, slender, and tapering distally, vesica with 4-5 deciduous cornuti (n = 9); gnathos a narrow band; valva with costal margin concave, apex rounded but moderately acute, distal margin convex, with ca. 8 stout setae along ventral two-thirds, ventral angle rounded, ventral invagination moderate, NR = 0.61, cucullus with distal one-half of medial surface densely setose, sacculus moderately setose, margin of basal opening with setose medial projection. Holotype. ¢, New Mexico, Taos Co., S. Side US 64, 10 mi. SE Tres Piedras, 7550', 11 August 1999, D. J. Wright, genitalia slide DJW 1035, deposited in USNM. Type locality at 36° 34.5! N, 105° 48.2' W. Paratypes. NEW MEXICO: Same data as holotype (17 ¢ genitalia slides DJW 528, 1034); Luma Co., Deming, 16-23 August (2 ', genitalia slides USNM 70399, DJW 1070); Fort Wingate, 24-30 June (1 ¢, genitalia slide USNM 70396), 24-31 July (3 d, genitalia slide USNM 70398). UTAH: Tooele Co., Stockton, Tom Spalding, 30 August 1904 (1 4, genitalia slide DJW 1064). Paratype depositories: AMNH, BMNH, CNC, CSU, EME, LACM, USNM, DJW. Etymology. The specific epithet refers to Taos County, New Mexico. Distribution and biology. Of the 29 specimens examined, one is from central Utah and the rest are from New Mexico. Capture dates range from late June through August. The type locality is open sagebrush habitat at an altitude of 7550 feet. The larval host is unknown. Pelochrista emaciatana (Walsingham), new combination (Figs. 1, 10, 13, 17, 24, 33) Paedisca emaciatana Walsingham 1884:137, pl. IV, Fig. 7 Eucosma emaciatana: Fernald [1903]:460; Barnes and McDunnough 1917:171; McDunnough 1939:46; Powell 1983:3 Eucosma aeraroptngoies Heinrich 1929:8; McDunnough 1939:47, new synonymy. Pelochrista perpropinqua: Powell 1983:35. Types. Paedisca emaciatana. Lectotype here designated (Figs. 1, 13): ¢, Arizona, Morrison, 1882, genitalia slide 11571, BMNH. Paralectotypes: same data as lectotype, (2 6, BMNH). Eucosma perpropinqua. Holotype: ¢, Arizona, Pima Co., Indian Oasis, Sells Post Office, 15-30 April 1923, O. C. Poling, genitalia slide 72797, USNM. Paratypes: same site and collector as holotype, 1-15 April 1923 (3 °, genitalia slides DJW 809, 955, USNM; 1 2, CNC), 15-30 April 1923 (1 9, USNM:;: 12°, AMNH). Diagnosis. Reasonably fresh specimens can be identified on the basis of forewing g pattern (Fig. 10), but dissection is recommended for positive determination. Males are distinguished by the general shape of the valva and the size and position of the ventral spike (Fig. 17), females by the sclerotized plate on the dorsolateral surface of the corpus bursae, the presence of only one VOLUME 59, NUMBER 3 signum, and the shape of the sterigma (Figs. 24, 33). Description. Head: Frons and vertex white, scales anterior to eye light brown; labial palpus elongate, lateral profile triangular, length more than 2x eye diameter, first segment and medial surface of second segment white, scales on lateral surface and dorsal margin of second segment pale brown with white apices, scales of asearnl segment concealing third segment; antenna white. Thorax: Dorsal surface white with pale- brown shading, scales of tegulae light brown with white apices, ventral surface w hite, legs light Drees with white tarsal annulations. Forewing (Figs. 1, 10): $ FWL 10.8-13 mm (mean = 11.5,n=7), AR=3.1, CFR= 0.2 27, 29 FW. LZ. 9-9.9 mm (mean = 9.4, n = 6), AR = 3.3; costa straight, vertex acute; dorsal surface white with brown markings, appearing streaked longitudinally, a dark-brown, outwardly oblique, subbasal mark on fold, a brown pretornal mark on dorsum, often one or more variably expressed, disjunct, brown marks between mid costa and pretornal mark, and an elongate patch of pale brown scales with white apices anterior to ocellus en connected by oblique spur of similar scaling to brown apical mark; ocellus obscure, white to pale brown, crossed longitudinally by three light-brown streaks, the latter often marked medially by a few bros black scales; termen with several rows of pale brown scales with white apices, the latter preceded basally by a streak of white scales from distal margin of ocellus to apex; distal one-half of costa with four indistinct, white, paired strigulae, merging into light post-costal streak from mid costa to apical mark; male costal fold darker than adjacent forewing scaling. Hindwing: Uniformly brownish gray with pale white fringe. Male genitalia (Figs. 13, 17): Uncus a ‘dorsally setose convex lobe, shoulders of tegumen moderately dev eloped; socii curving dorsally and moderately setose; aedeagus tapered distally, vesica with no indication of cornuti (n = 10): walkin with costal margin concave, apex evenly rounded, outer margin convex, ventral angle with well dev eloped projection supporting a stout spine, usually a spine of similar size and several smaller spines on distal margin of cucullus, ventral invagination moderate, NR = 0.7, cucullus with distal two-thirds of medial surface densely setose, sacculus moderately setose. Female genitalia (Fig. 24): Papillae anales facing laterally and densely setose, medial margins mildly sinuate, surfaces finely ridged transversely, long setae on lateral margins strongly curved ventrally: lamella ‘antevaginalis (Fig. 33) ringlike and very weakly sclerotized; lamella postv aginalis well dev eloped, depressed medially, with variably inv aginated posterior margin and sharply acute anterolateral projections; sternum VII with posterior margin weakly invaginated and closely approximate to sterigma; ductus bursae uniformly narrow, constricted anterior to ostium; corpus bursae with large sclerotized patch on dorsolateral surface at juncture with ductus bursae, a large signum on ventral surface, and a faint indication of a reduced signum at center of anterior margin of sclerotized patch. Distribution and biology. I examined 96 specimens (66 ¢, 30 2): one each from San Bernadino Co., California, Clark Co., Nevada, and Kimble Co., Texas: the rest (to the extent determinable by specimen data) from Cochise, Pima, Pinal, and Santa Cruz Counties in Arizona. Three specimens had capture dates in September or October; the others were collected between early March and mid June. Ninety percent of the records were dated between | April and 31 May. No larval host information has been reported. Pelochrista popana (Kearfott) (Figs. 11, 20, 29, 37) Eucosma popana kearfott 1907:31; McDunnough —_1917:169; — Heinrich McDunnough 1939:47. Eucosma carcharias: Meyrick 1912:35 Barnes & 1923:109; 13] Pelochrista popana: Powell 1983:35. Types. Lectotype designated by Klots (1942): 2, Stockton, Utah, Tom Spalding, 3 July 1904, genitalia slide DJW 1085, AMNH. Paralectotypes: UTAH: Stockton, Tom Spalding, 1 June 1904 (3 ¢, AMNH; 1 ¢. USNM), 8 June 1904 (2 6, USNM), 10 june 1904 (2 ¢ USNM), 14 June 1904 (2 6, AMNH: 1 6, USNM), 15 June 1904 (1 6, USNM), 28 June 1904 (12, AMNH), 3 July 1904 (2 2, AMNH). Diagnosis. The forewing maculation of popana is grayish-black, as opposed to brown in totana, taosana, and powelli. The dark mark anterior to the ocellus nearly always connects to an apical dash of the same color, often connects to costa, and frequently extends along distal edge of ocellus toward tornus, forming a distinctive Y-shaped mark. The shapes of the valva (Fig. 20) and sterigma (Fig. 37) separate popana from the other species considered here. Description. Head: Lower frons white, scales of upper frons and vertex long, gray to brownish gray medially, with lighter apices: labial palpus wah ‘Gace segment and medial qnttres of second segment white, lateral surface Oh second segment gray to brownish gray, third segment concealed by scales of second | segment; antenna grayish white, often darker distally. Thorax: Dorsal surface and tegulae with brownish-gray, white-tipped scales, ventral surface white, _ legs with anterior Perce. brownish gray, posterior surfaces white, tarsal segments with white distal annulations. Forewing (Fig. 11): ¢ FWL 8- 9. 8 mm (mean = 8.6, n = 12), AR = 3.21, CFR = 0. aii 2 FWL 6.7- 8.3 mm (mean = 7.4, n = 19), AR = 3.21; costa weakly convex, apex acute, termen weakly convex; dorsal surface white with brownish- black to brownish-gray markings, basal and median areas white and variably irrorated with Ibxomaalh gray, an outwardly oblique subbasal mark extending from dorsum to cell, a broken median fascia consisting of free marks, the first at mid costa, the second at distal end of Cell, the third semitriangular and projecting anteriorly from pretornal portion of dorsum along basal margin of ocellus, the first two median marks often connected, the latter two usually separated by narrow band of white scales, an elongate patch of w! hite- -tipped, black to brownish-gray scales anterior to ocellus, usually connected to apex by oblique dash of similar coloration: ocellus with basal, distal and tornal margins pale pinkish brown, central field a narrow, vertical. light brown streak, crossed longitudinally by ca. 4 black dashes; distal one half of costa with four paired white strigulae, costal fold of male dark gray, fringe scales white basally and apically, black to brownish- gray medially, Abdomen: Females with dark gray scales on posterior margin of eighth segment. Hindwing: Uniformly brownish gray, fringe lighter. Male genitalia (Fig. 20): Uncus dorsally setose and semitriangular, apex sometimes weakly indented; socii long and setose, curving dorsally; aedeagus long, tapering distally, vesica with 3- 6 deciduous Soran (n = 4); y val a with costal margin concave, apex nearly right angled but rounded, distal margin convex with 2-3 stout spines, ventral angle developed into triangular lobe supporting 1 or 2 stout spines, ventral invagination broad and shallow, NR = 0.7, cucullus with medial surface moderately setose, sacculus sparsely setose, margin of basal opening with weakly developed setose projection. emale genitialia (Fig. 29): Papillae anales facing ventrolaterally and densely setose, surfaces finely ridged transv ersely, medial margins sinuate, setae on lateral margins long, curving ventrally, those near anal opening shorter, with hooked apices: sterigma (Fig. 37) with anterior margin very weakly sclerotized, lamella postvaginalis extending laterally to width of ca. 3x ostium diameter, widening posteriorly, length ca. 0.5x width, posterior margin with scalloped appearance due to sharply acute posterolateral comers and concave medial invagination, a very shallow trough from mid posterior margin to ostium, surface finely microtrichiate; sternum VII with posterior and lateral margins strongly sclerotized, posterior margin with medial triangular projection overlapping ostium; ductus bursae of nearly uniform width, sclerotized from constriction anterior to ostium to ductus seminalis; corpus bursae with two similarly sized signa, membrane variably crinkled around signa, interior surface minutely microtrichiate. Distribution and biology. I examined 167 specimens (146 ¢, 21 2) from the following states and counties: COLORADO: Chaffee, Grand, Larimer, Mesa; IDAHO: Blaine; MONTANA; NEVADA: Lander, White Pine; NEW MEXICO: Taos; UTAH: Cache, Garfield, San Juan, Tooele, Uintah; WYOMING: Albany, Carbon, Fremont, Park, Sublette, Teton. They document a flight period from early June to late August. No larval host has been reported. Pelochrista powelli Wright, new species (Figs. 12, 22, 23, 30, 34) Diagnosis. This moth has a pale brown appearance. By contrast, popana is brownish-black to gray, with considerable contrast between markings and ground color. The forewing pattern of powelli lacks the chevron shaped mark at the end of the cell in totana and the orange-brown scaling along the costa and along [A+2A in taosana. In females of powelli, the scales on the posterior margin of the eighth abdominal segment are brown and inconspicuous, they are brownish-black to gray in popana. The v-shaped posterior margin of the sterigma separates powelli from the other species considered here. Description. Head: Lower frons pale tan, scales of vertex white distally, shading to tan basally; labial palpus tan to pale brown; antenna pale tan. Thorax: Dorsal surface pale brown, ventral surface pale tan, legs with anterior surfaces pale brown, posterior surfaces white to tan, distal ends of tarsal segments ringed with pale tan. Forewing (Fig. 12): d FWL 7.1-10.5 mm (mean = 8.7, n = 10), AR = 3.3, CFR = 0.26, 2° FWL 8.2-9.2 mm (mean = 8.8, n = 5), AR = 3.11; costa weakly convex, apex acute, termen straight to weakly convex; dorsal surface pale tan with brown markings, a brownish- black subbasal mark on fold, a thin brown line along fold from subbasal mark to tornus, a narrow, triangulate, brown mark based on fold and projecting toward apex along basal margin of ocellus, an elongate patch of white-tipped brownish-black eealles anterior to ocellus, extending basally through distal one-half of cell, usually constricted and darker at distal end of cell; ocellus obscure, variably bordered on basal, distal and tornal margins with pale pinkish-tan bars, central field white, crossed by 4-6 ibrar black dashes that are often joined in zig-zag pattern; distal two-thirds of costa with numerous white strigulae, delineated by brown costal marks and thin brown striae, male costal fold brownish black; termen with band of white-tipped brownish-black scales extending from apex to tornus, fringe scales lighter with pale-brown medial markings. Hindwing: Pale gray-brown with paler fringe. Male genitalia (Figs. 22, 23): Uncus se mitriangular with waning apex, dorsal surface setose; tegumen long, dorsolateral shoulders well developed and hunched; socii long, flat, tapering distally, and moderately setose; gnathos a narrow ahead aedeagus long, tapered distally, vesica with ‘4-9 deciduous cornuti (n = 7); wala sath costal margin weakly concave, apex rounded to angular, distal margin convex, ventral angle with triangular projection supporting stout spine, neck long and narrow, NR = 0.5, ventral inv agination broad and moderate, nani with densely setose medial senrbye and 3-5 stout spines on distal margin, sacculus moderately setose, margin of basal JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fic. 30. Female genitalia. 30, P. powelli, slide DJW 1024. opening with weakly developed setose projection. Female genitalia (Fig. 30): Papillae anales facing ventrolaterally and densely setose, medial margins sinuate, surfaces very finely ridged tre ansversely, setae on lateral margins long and curving ventrally, those near anal opening shorter with hooked apices; posterior one- -half of tergum VIII with 4- 5 rows of setae; sterigma (Fig. 34) with anterior margin very weakly sclerotized, lamella “postvaginalis developed posterolaterally into triangular projections, posterior margin with v-shaped medial invagination, surface finely microtrichiate; stemum VII with posterior margin concavely invé aginated except for mild, convex, medial, projection overlapping ostium; ductus bursae narrow, constricted anterior to ostium; corpus bursae with two signa, inner surface minutely microtrichiate. Holotype. ¢, Idaho, Oneida Co., Curlew NG, 4 mi. ENE of Holbrook, Jctn. Forest Rds. 056 and 057, 5050', 7 July 2001, D. J. Wright, genitalia slide DJW 1032, deposited in USNM. Type locality at 49° 11.35! N, 112° 34.92' W. Paratypes. IDAHO: Oneida Co., Curlew NG, 4 mi. ENE of Holbrook, 5050', D. J. Viper 25 July 2003 (2 ¢, genitalia slide DJW1027), 26 July 2003 (2 4, 1 9); Oneida Co., Curlew NG, T14S R32E $30, D. J. Wright, 28 July 2003 (4 9, genitalia slides DJW1024, 1029). UTAH: Daggett Co., 4 mi. S. of Manila, G. J. Balogh, 20 July VOLUME 59, NUMBER 3 FIGS. 31-37. Sterigmata of specimens illustrated in Figs. 24-30. 31, E. piperata. 36, E. larana. 37, P. popana. 1994 (5 ¢, genitalia slides DJW 697, 705, 707); Juab Co., Eureka, Tom Spalding, 18 August 1911 (1 d, genitalia slide DJW1069); Garfield Co., 3 mi. W. Bryce Jct., 2300m, J. A. Powell, 28/29 June 1992 (1 d, genitalia slide EME 5755); Garfield Co., Kings Cr. campgr., 15 km SW Bryce Jct., 2300m, J. A. Powell, 18 July 1993 (12 d, genitalia slide EME 5756). Paratype depositories: BMNH, CNC, EME, GJB, USNM, DJW. Etymology. This species is named after J. A. Powell, who collected nearly half of the specimens in the type series. Distribution and biology. The 29 specimens (24 ¢, 5 2) reported above were collected in southeastern Idaho and Utah, suggesting a Great Basin distribution for this insect. The type locality is open sage brush habitat. The larval host is unknown. Comments. The shape of the male valva is variable (Figs. 22, 23), the cucullus illustrated in Fig. 22 being the most angular of the nine I examined. Forewing color also varies from very light tan in the specimens from Idaho to a pale brown in those from Utah. ACKNOWLEDGEMENTS I thank B. Brown, J. W. Brown, J. D. Lafontaine, P. A. Opler, P. D. Perkins, J. A. Powell, E. Quinter, S. Shaw and K. R. Tuck for the loan of specimens under their care. I particularly appreciate the efforts of J. S. Nordin and C. D. Ferris, who supplied me with many study specimens from southeastem Wyoming. Finally, thanks to T. Gilligan for his help in producing the electronic files for the illustrations and to the reviewers, R. L. Brown and W. E. Miller, for many helpful suggestions. LITERATURE CITED Barxeras, 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(4):422-431]. Barnes, W & J. McDunNoucu. 1917. Checklist of the Lepidoptera of Boreal America. Herald Press, Decatur, Illinois. 392 pp. 133 nordini. 32, E. totana. 33, P. emaciatana. 34, P. powelli. 35. E. Brown, R. L., J. F. G. CLarke, & D. H. HaBeck. 1983. New host records for Olethreutinae (Tortricidae). J. Lepid. Soc. 37:224- Pear 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- term. 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. U.S. Nat. Mus. Proc. 75:1-23. Kearrotr, W. D. 1907. New North American Tortricidae. Trans. Amer. Entomol. Soc. 33:1-98. Kiots, A. B. 1942. Type material of North American microlepi- doptera other than Aegeriidae in The American Museum of Nat- ural History. Bull. Amer. Mus. of Nat. Hist. 79:391-424. 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. Meyrick, E. 1912. On some impossible specific names in micro-lepi- doptera. Entomol. Monthly Mag., Ser. 2, 48:32-36. 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, Lorp. 1879. Illustrations of typical specimens of Lepi- doptera Heterocera in the collection of the British Museum. Part IV. North American Tortricidae. 84 pp. + 17 pls. Dept. Zoology, British Museum, London. —. 1884. North American Tortricidae. Trans. Entomol. Soc. Lond. 1884:121-147 + 1 pl. Received for publication 20 October 2004, revised and accepted 5 May 2005 Journal of the Lepidopterists’ Society 59(3), 2005, 134-142 REDISCOVERY OF ACTINOTE ZIKANI (D'ALMEIDA) (NYMPHALIDAE, JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY HELICONIINAE, ACRAEINI): NATURAL HISTORY, POPULATION BIOLOGY AND CONSERVATION OF AN ENDANGERED BUTTERELY IN SE BRAZIL RONALDO BASTOS FRANCINI, ANDRE VICTOR LUCCI FREITAS?* AND KEITH SPALDING BROWN JR.” ABSTRACT. Actinote zikani (D'Almeida) (Nymphalidae, Heliconiinae, Acraeini) was rediscovered in 1991 in Paranapiacaba, Sao Paulo, 40 years after its original description (based on specimens from the Boracéia Ecological Station, Salesépolis, Sao Paulo); 49 years after its last collection, and studied during three years. The adults go through two yearly generations, one in March (earlier than the other species of Actinote in the region) and the other in November (bivoltinism). than one week, and the sex ratio in the field is male biased. The mean residence of the adults is less The only known host plant for the species is Mikania obsoleta (Aster- aceae), and the immature stages are similar to those known for other species of Actinote. The range of A. zikani is within one of the most densely human populated regions in Brazil, making urgent the creation of effective preserved areas where colonies of this species are known. Additional key words: Atlantic forest, Bivoltinism, Mikania, Neotropical In 1941 and 1942, Romualdo Ferreira D'Almeida collected 10 individuals of a dark Actinote Hiibner at the Estacao Biolégica de Boracéia in Sales6polis, Sao Paulo, which were misidentified as Actinote morio (D'Almeida, 1943). Later, notified by J. F. Zikan (a field naturalist who lived near Itatiaia, RJ) D'Almeida corrected himself, recognizing the status of this species and describing it as Actinote zikani (D'Almeida, 1951). This species was described based on material collected by D'Almeida from Boracéia (Sales6polis) and one male collected by Roberto Spitz from Alto da Serra de Santos, SP in 1941. A survey revealed that the 11 specimens cited by D'Almeida are now in the Museu de Zoologia da Universidade de Sao Paulo (MZUSP; Lamas 1973) and the Museu de Zoologia da Universidade Federal do Parana (Mielke & Casagrande 1986). Additionally, KB saw another 18 specimens collected by R. Spitz from the Alto da Serra de Santos, in the collection of the Natural History Museum (London). From 1985 to 1990, A. zikani was searched for intensively by RBF and AVLF in the type-locality in April-May and November-December, and also in other sites sith the same environmental characteristics (Francini 1992), but none were seen. The only new information was from KB who saw a possible male of 'Universidade Catélica de Santos, Campus D. Idilio José Soares, Av. Cons. Nébias, 300, Prédio dos Laboratérios, Sala 213, CEP 11015- 200, Santos, SP, Brazil. email: francini@unisantos.br 2 Museu de Historia Natural and Departamento de Zoologia, Instituto de Biologia, Universidade Estadual ae Campinas, C P6109, Campinas, SP 13083- 970, Brazil. email: ksbrown@unicamp.br ° Corresponding author: email: Rin@anteiabe this species on the wing in April 1981 on the edge of the road from Tapirai to Sorocaba, in southern Sao Paulo state, about 1000 m altitude in a very wet forest. Because of the difficulty in finding extant colonies of this species, KB proposed the inclusion of A. zikani on the list of Brazilian species possibly threatened with extinction (Bernardes et al. 1990; Brown 1991), and since then, A. zikani has been classified as critically endangered (SP-SMA 1998, MMA 2003). With intensive searching, finally on 16 March 1991 (1100 h), on a routine trip, RBF and AVLF found a male flying at the summit of the Serra do Mar, 20 km northeast of the city of Santos, Sao Paulo. With data from this locality, the present paper describes the natural history and population biology of A. zikani, information important to the conservation of the species and its habitat. STUDY SITE AND METHODS The population studied was located in the Santo André municipality, Sao Paulo State, near the village of Paranapiacaba. Most field work was carried out in a place east of Paranapiacaba, along a road 2500m long (SW-NE following the orientation of the mountain chain), paved with concrete blocks, connecting two groups of towers (television and microwaves) on peaks of Serra do Mar reaching 1200 m (Fig. 1). Because of the high rainfall, the road on the summit was built with a good drainage network; there are also four small creeks running across the road. Sometimes the road and drains were cleared, but the trees next to them were VOLUME 59, NUMBER 3 Tropic of | Capricorn ‘Tapirai e electric poles —->~ creeks and drainage ditches t TV towers r Radio towers creek 3 creek 1 drainage ditch 1 i Sfeteihinietete ene cl ‘Salesopolis Paranapiacaba; drainage ditch 2 ——| 500 m Paranapiacaba road (details) Fic. 1. Study area in Southeastern Brazil (modified from IGGSP 1972a). In the regional map, open circles show the known present and past collecting sites of A. zikani; the question mark indicates a doubtful record and the triangle is a locality with a possible undescribed subspecies of A. zikani. Black areas indicate altitude above 1000m. always maintained. The road was mapped with the aid of a tape measure and a compass using a 1:50,000 topographic chart (IGGSP 1972). All 52 electric poles along the road were numbered, permitting the location of each butterfly to be recorded to the nearest 50 m. The study area is in the rainiest part of Brazil outside of the upper Amazon. The mean annual rainfall between 1870 and 1939 was over 3500 mm, with a minimum of 2355 mm in 1874 and a maximum of 5563 mm in 1872 (NOAA 1998). Data from SIGRH (2003) show that the mean annual rainfall between 1936 and 1996 was 3164 mm, with an extraordinary minimum of 826 mm in 1990 and a maximum of 4739 mm in 1947. The rains roughly occur 15% in winter (June- September), 25% in spring, 35% in summer and 25% in autumn (Santos 1965). Fog is frequent in the study site, and a sunny day could suddenly change to misty and rainy. The original vegetation is montane rain forest (Ururahy et al. 1984). On the edge of the road above 1000m, there are many patches of bamboos, and "manaca-da-serra", "quaresmeira" (Tibouchina spp: Melastomataceae), and the vine Mikania hirsutissima (Asteraceae) are abundant. The trunks and stems of most plants are covered by various epiphytic mosses and ferns. In this area 16 species of Asteraceae were found 136 which could be potentially used as foodplants by larvae of Actinote spp. (RBF unpublished results). After the discovery of the population of A. zikani near Paranapiacaba in 1991, 48 trips were made to the study area up through June 2004 (161 hours of field work); 24 days from January to December 1991 (1-180 days interval), 14 days from July to November 1993 (1- 40 days interval) and 10 days in March-June 1994 (1-20 days Butterflies binoculars and various aspects of interval). were observed with behavior were photographed; some individuals were collected for morphological study. All material including the reared specimens was deposited in the collection of the MZUSP. This population was studied by a mark-release- recapture method (MRR). Each captured individual received a small numbered circle of impermeable paper glued to the ventral base of the left hindwing. This marking technique permits rapid marking and data retrieval; it was previously tested by RBF and used in a population study of Actinote pellenea pellenea Hiibner in 1988, Actinote mamita mitama (Schaus) in 1990, and Stalachtis phlegia susanna (Fabricius) (Riodinidae) in 1992 (RBF, unpublished data). butterfly, sex, "age" (based on wing wear), forewing length, location and the time of ‘day were recorded (as in Freitas 1993, 1996). The relative daily abundance was obtained dividing the total number of males sighted by minutes of observation effort, later transformed to butterflies per hour (based only on days with weather conditions favorable for the flight of the butterflies). The MRR data for the summer 1993 generation (12 field days, 1-7 days interval) was analyzed by the Lincoln-Petersen-Bailey method (Southwood 1971) for estimating population parameters (software developed by RBF, UNISANTOS). In most cases, only males were analyzed because of the low recorded. For each marked number of females Daily results were tabulated as individuals present per day” (NIPD), following Ramos & Freitas (1999). To estimate the NIPD, recaptured individuals were considered to be present in the population on all previous days since the day of first capture. Numbers of reared lots are sequential in the RBF data bank. RESULTS Geographic distribution. All known present and past colonies of A. zikani are found in a limited area between Salesépolis and Paranapiacaba (Sao Paulo): Alto da Serra de Santos (a partly unknown spot that could include part of Paranapiacaba), Paranapiacaba and the Estagao Biolégica de Boracéia (Salesépolis) “number of individuals captured per day” (NICD), and “number of JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY (Fig. 1). There is also a mention of this species occurring in southern Minas Gerais (D'Almeida 1951). In November 1991, a male Actinote with whitish color pattern was collected by the authors in Penedo, Rezende (Rio de Janeiro), in the foothills of the Serra do Itatiaia. Although the genital armature of this individual is similar to that of A. zikani, more material will be necessary to confirm the status of the specimen from Penedo. Flight activity. The flight activity of A. zikani is variable and dependent on weather conditions. Butterflies usually start to fly at approximately 0800 h, when direct sunlight reaches the forest (the summits to the southeast are higher and shade the study area before this time); they open and shut their wings in direct sunlight, basking i in a cyclic process until the beginning of flight. On warm but moist days, the flight activity diminished shortly after the sun was covered by clouds. On 20 March 1991, the peak of activity was around 0830 - 0900 h, which was the warmest period of the day (Fig. 2). Males usually fly more than 2 m above the ground along the road, and in forested areas they usually fly above the canopy. Males attack any flying insect that gets less than 1 m from them, when they quickly go up to almost 10 m above the ground or rapidly cover a linear transect of more than 100 m. The relatively small males of A. zikani were seen to persecute butterflies as large as Morpho hercules (Dalman) (Nymphalidae: 105 84 et. Se 2 —@®— males 4 | - O- females >is ct = = as ve o 2 [= 25 0 08:30 09:00 09:30 10:00 10:30 11:00 11:30 12:00 Time Fic. 2. Flight activity of A. zikani in Paranapiacaba, SP, in March 20, 1991. in the area between poles 50 and 51. Morphinae). Mating. Mating in A. zikani occurs without any apparent courtship display. The patrolling male follows a female; after flying 2 to 5 m in a straight line about 2 m above ground they make a spiraled flight to the ground, at the end of which the male grasps her abdomen with his valves, forcing copulation on the ground (n = 5). There is always formation of a plug (sphragis) in the female (as in other Actinote species), but in A. zikani it is mostly internal and inconspicuous, similar to that VOLUME 59, NUMBER 3 known in A. discrepans D'Almeida. Attempts to copulate are sometimes unsuccessful, but once it occurs, it can last more than 30 minutes. On 17 November 1993 a mating pair observed at 0830 h continued “in copula” until at least 0908 h when RBF left the area. Foraging activity. Feeding activity of adults occurs in the coolest hours of the day, soon after sunrise or before sunset. In the March-April generation of 1991 and 1994, the males were observed drinking nectar from the inflorescences of Mikania triphylla (Fig. 3), Mikania micrantha, Eupatorium gaudichaudianum (Asteraceae), Mitracarpus hirtus and Borreria verticillata (Rubiaceae). All these food resources were very scarce during the study period, especially in 1994. In the November generation the main nectar sources were two species of myrtaceous trees (Myrcinia) that were common along the road. The presence of butterflies was directly related to food resources, with more butterflies present in places with more flowers. On some occasions (cloudy days) females were observed on the ground, or on petals of “manaca-da-serra” (Tibouchina sp.. Melastomataceae) or large flowers of the exotic “lirio-do-brejo” (Hedychium coronarium, Zingiberaceae). In these situations, they appeared to be drinking the accumulated water. Larval foodplant. The larval foodplant of A. zikani is Mikania obsoleta (Vell.) G. M. Barroso, discovered after observation of two ovipositing females on 1 April 1991. This is the only hostplant of A. zikani known in the study site (from a total of 13 species of Mikania present there). M. obsoleta was not observed being used by any additional species of Actinote. This plant is a climber with halberd- shaped smooth leaves (Fig. 3). It grows around tree trunks climbing to 6 m Heine Most individuals of M. obsoleta grow near small creeks, in places with wet soil in open canopy areas. The growth of M. obsoleta (measured by the number of new leaves) was relatively slow compared with that of other Mikania species in the area (e.g. M. hirsutissima) (RBF unpublished results). Flowering occurs from October to November and the flowers were not observed attracting any butterflies. A program of monitoring the hostplants revealed that many individuals of M. obsoleta tagged in November 1993 had disappeared by April 1994. More than 20 plants disappeared after cleaning of the rivulets (area between poles 16-17) and near creek 1, but some plants inside the forest on the borders of creek 3 also disappeared without any sign of human action. Oviposition behavior. Females of A. zikani had an oviposition behavior similar to that observed in other Actinote species (Francini 1989). The female flies Fic. 3. Above - flowers of Mikania triphylla. Below - Close-up view of plant of Mikania obsoleta showing details of the halberd shaped leaf and the inflorescence. a male Actinote zikani drinking nectar from around the foodplant landing briefly on some leaves, and after choosing a leaf it lands on its ventral surface. After a period of | inactivity (1-5 min) the female starts ovipositing, continuing for up to one hour. On 1 April 1991, five ovipositions were observed in the study area, including one leaf with a double oviposition (two differents females observed ovipositing together, lots F- 2337 and F-2338). Oviposition in the laboratory was also obtained with a female in a glass jar with a piece of foodplant under a 150W incandescent light bulb (following Francini 1989 and Freitas 1991) (oviposition lot F-2361). In this case, the entire process lasted five hours with the female constantly vibrating her wings. In 1993 only one oviposition was found on a plant inside the forest near a creek, and in 1994 no oviposition was observed in the study area. Immature development and behavior. Detailed descriptions of the life cycle of A. zikani will be presented in a further paper (RBF in prep.). Eggs of a double oviposition collected in the field on 1 April 1991 (F-2337 and F-2338) hatched on 15 April 1991 in laboratory conditions. First instar larvae of A. zikani left the egg after eating the lateral walls of the chorion, leaving the remainder of the egg intact. Feeding activity started after 3 ventral epidermis. The fecal pellets were glued onto the leaf by silk, not falling to the ground. There was no significant difference between the mean cursgen of the larval periods for males (72.5 days, SD = 2.76, n = 16) and females (73.6 days, SD = 2.68, n = 12) (t = -1.0414, -5 hours, and eral larvae ate only the P = 0.30, DF= 26), nor between the mean duration of the pupal periods for males (mean = 17.2 days, SD = 1.41; n=16) and females (mean = 16.9 days, SD = 0.94, n = 12) (t = 0.4473; P = 0.66, DF= 26). In laboratory conditions, the larvae hatched on 15 April 1991 reached the last instar at the beginning of June; pupation occurred at the end of June and adults emerged in August (almost two months earlier than the flight pened in fie field). The mean total duration of the life- -cycle in the laboratory (egg to adults) was 105.7 days (SD = 3.92, n = 28) or roughly three and half months. Tri ips to the field during all these months showed absence of adults before November. The same life- -cycle pattern was observed in 1993 and 1994. Chemical protection and predation. Qualitative tests for cyanogenesis (following Francini 1989) were done with one male, five eggs, two first instar, and one last instar, and all were positive as for other known species of Neotropical Acraeinae (Brown & Francini 1990). In the field, one oviposition (F-2342) was observed being partially eaten by ants of the genus Pheidole. Additionally, a dead male was observed in a web of Nephila clavipes (Arachnida: Araneida). No predation on larvae and pupae were observed in the field. Population biology. Four generations of A. zikani were followed between March 1991 and April 1994. Adults of A. zikani are bivoltine, with flight periods of about one month; the first generation occurs in March/April (autumn generation) and the second in November (summer generation). In 1991, the autumn generation flew from 16 March to 17 April, and the summer generation from 7 November to 24 November. In 1993 the summer generation flew from 1 November o 28 November, and in 1994 the autumn generation flew from 19 March to 4 April. The number of butterflies sighted per hour (BSH) varied between 3 and 8 (mean = 5.05, SD = 2.39) in the March-April generation of 1991, between 1 and 17 (mean = 5.33, SD = 4.81) in the November generation of 1993 (Fig. 4) and between 13 and 20 (mean = 14.85, SD = 2.98) in the March-April generation of 1994. Of the 190 males captured and marked in November, 1993 only 10 (5.3%) were recaptured. Eight individuals were recaptured once, and two individuals were JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY recaptured twice. In March-April, 1994, due to the unfavorable weather conditions, 35 males were marked and only two recaptured (5.7%). In November 1993, males started to fly on 1 November and females only on 6 November. Both sexes reached highest numbers from 12 - 18 November, after which the population diminished to low numbers until disappearing before the beginning of December (Fig. 4). The number of individuals present per day (NIPD) in the summer 1993 generation varied from 1 to 48 (mean = 18.3, SD = 15.5, = 11 days) (Fig. 4). The estimated popnlacen size nem on Lincoln-Petersen-Bailey in November 1993 showed that population peaks can include more than 1000 males in the study site (Table 1). 505 404 | —e—Males | --0-- females | w--- BSH | Number of individuals - SLES SE NN SL Ga 10 12 14 16 18 20 22 24 26 28 Days (november/1993) Fic. 4 - Number of individuals present per day (NIPD) for males (solid circles) females (open circles) and number of butterflies sighted per hour for males (BSH solid squares) in November 1993 in Paranapiacaba, SP. Sex ratio. The sex ratio in the field was male biased in all study periods (Table 2), with lowest male:female ratios in November 1993. In the laboratory, the sex ratio was not different from 1:1 (Table 2). In the November 1993 generation, males were the dominant sex in all but one day with more than 15 individuals (Table 1). Age structure and residence time in 1993. Most of the first captures of both sexes were individuals of “intermediate” age (59% of males and 63% of females). The age structure in November 1993 shows a clear pattern of individuals becoming older from the beginning to the end of flight period (Fig. 5). Residence time for males ranged from two to six days, with six males lasting two days, three males lasting three days and a single male lasting six days (mean = 2.7 days, SD = 1.25;n = 10). A single female was recaptured during the study, with a residence of 10 days. Vagility. The average distance traveled by males in 1993 was 232.3 meters (SD = 287.9; n = 17), not significantly different of that of females (mean = 136.7 VOLUME 59, NUMBER 3 139 TABLE 1 - Summary of population data for the summer 1993 generation of Actinote zikani in Paranapiacaba, SE Brazil. NICD = number of individuals captured per day, M = males, F = females; NIPD = number of individuals present per day (males only), BSH = butterflies sighted per hour (males only) rounded off to nearest integer, LPB - number of butterflies estimated by Lincoln-Petersen-Bailey, SE - standard error. An asterisk indicates a male biased sex ratio (chi square test [x7], p < 0.05). Date NICD Sex ratio NIPD BSH LPB SE M F 1993 OU/NOV 8 0 —_— 8 P) 8 — 03/NOV 1 0 — 1 1 1 04/NOV 1 (0) — 1 1 16 21 06/NOV 14 2 Wale 15 4 142 142 09/NOV 18 3 6:1° 18 4 270 295 12/NOV 29 1 29:1° 29 11 157 98 14/NOV 31 10 Oils 38 444 481 15/NOV 22 15 ES 24 1104 1514 17/NOV 45 27 (she 48 16 1034 366 18/NOV 20 5 4:1° 21 1 63 8 21/NOV 0 1 — 0 0 — — 28/NOV 2, 1 — 1 1 — a TABLE 2 - Sex ratio of marked and reared Actinote zikani from Paranapiacaba, SE Brazil. An asterisk indicates a male biased sex ra- 7 tio (chi square test [x°], p < 0.001). 6 Males Females Sex ratio a 3 5 Field captures i 4 Autumn 1991 50 ll 4.5:1 24.9° a Summer 1993 190 65 29:1 61.3° 2 Autumn 1994 35 2 17.5:1 23.5 < 2 Reared material 3 ; Lot 2337 3 3 1:1 — Lot 2338 13 9 14:1 0.727 Percentage of age classes 11 13 15 17 19 21 Days (november/1993) 23° 25 27 Fic. 5. Age structure of Actinote zikani in Paranapiacaba, Santo André, SP, in November 1993. Black = fresh individuals, gray = intermediate individuals, white = old individuals as % of each day’s captures. 21-25 26-50 51-100 101-200 201-400 400-800 ->800 Distance traveled (m) FIG. 6. Maximum distances traveled by Actinote zikani, using MRR data from generations of November, 1993 and March, 1994 in Paranapiacaba, SP. meters; SD = 195.0: n = 3) (t = -0.5474, P = 0.59, DF = 18). Individual butterflies were recaptured up to 1000 m from their marking point. On 6 April 1991 a female of A. zikani was collected flying in a straight line in a westward direction along the road to Paranapiacaba, about 4 km from the study area. Fig. 6 shows the vagility of A. zikani based on data from summer 1993 and autumn 1994. Adult size. Based on sampled individuals from March-April 1991, the forewing length of females (mean = 37.14 mm, SD = 2.575, n = 64) was greater than that of males (mean = 32.88 mm, SD = 2.539, n = 197) (t = 12.281, p < 0.05, DF = 259). In the same period, the dry weight of males varied from 0.30 to 0.48g (mean = 0.41g, SD = 0,052, n = 10), and the dry 140 weight of the females varied from 0.65 to 0.S9g (mean = 0.74g, SD = 0.098, n = 5), showing that even though females weighed almost twice as much than males, the length of their forewing was only 1.1 times greater. Adults of A. zikani are large, compared with other species in the genus Actinote (RBF unpublished data). DISCUSSION Natural History and Population Biology. In most aspects of population biology and natural history, A. zikani is similar to the other known species of Actinote from SE Brazil. most known species of Actinote, except A. pellenea and A. brylla Oberthiir, that can be multivoltine in warm places, and most red species in the “red mimicry complex” that are univoltine (Francini 1989, 1992, Penz & Francini 1996). The adult permanence in the population of less than one week is low if compared with most neotropical butterflies (Ramos & Freitas 1999), but it is similar to the values obtained for most species of Actinote (Francini 1989 and unpublished data). These low values are suggested as a combination of short lifespan and high dispersal rates in these butterflies (Francini 1989). Even if flight periods and time intervals (about one month) are similar to the duration of generations of other Actinote species (Francini 1989), there is an asynchrony of A. zikani with relation to other Actinote species of about one month. During the present study, other species of Actinote like A. canutia (Hopffer), A. carycina Jordan, A. parapheles Jordan, A. melanisans Oberthiir and A. genitrix D'Almeida began to fly in Paranapiacaba only in the middle of April in the autumn generation (A started in early March). It is interesting to note that the labels of Museum specimens (April 1941 and December 1931) contributed to the delay in finding the species, since much time was spent in the field 20-30 days after the flight period of A. zikani. The male biased sex ratios recorded for A. zikani in the field are similar to those of the other 13 species of Actinote from SE Brazil (Francini 1989). Male biased sex ratios are . zikani usually observed in butterflies in the field even if laboratory broods are 1:1 (Brussard & Ehrlich 1970, Freitas 1993, 1996, Ramos & Freitas 1999). The recapture rate of about 5% recorded for A. zikani is low even if compared with those of other species of Actinote (Francini 1989). Francini (1989) recorded recapture rates of 12% and 8% for A. pellenea pellenea and A. brylla Oberthiir, 1917 respectively in the coastal plain of Sao Paulo. General features of the immatures conform to those of other species of Actinote (Francini 1989, 1992). The host plant agrees with the suggestion of D'Almeida The bivoltinism is characteristic of JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY (1951) who proposed that the host plant of A. zikani should be a species of Mikania. Immatures of A. zikani are gregarious like all other known Neotropical Acraeinae (Francini 1989), and the presence of a double oviposition (two different ovipositions in the same leaf) is also observed in other species that apparently have gregarious ovipositing (Francini & Freitas unpublished data) . The pattern of lateral eclosion differs from all other known Actinote, whose larvae exit through the Maybe the most fast See of ’ three and half months without periods of diapause or slow growing. micropylar region (Francini 1989). remarkable immatures in laboratory conditions; only feature is the Even though the larvae were reared under temperature conditions (25°C + 2°C) different from those in the field (that can be near 0°C on some winter nights), the duration of the larval stage in A. zikani was exceptionally short for an Actinote species, except for some broods of Actinote pellenea pellenea that were reared in summer at sea level (about two months, Francini 1989 and unpublished data). This capacity to grow quickly at high temperatures could be an indication that any Actinote could be multivoltine if conditions are adequate. Conservation of A. zikani and its habitats. The conservation status of A. zikani was defined as critically endangered in the most recent evaluation of the Brazilian list of endangered species (MMA 2003), based on a combination of: restricted area of occupancy, few known and declining populations and sites, small total population size and extreme fluctuations in number of mature individuals (categories B2 bii, iii, iv c Cb - criteria from IUCN 2001). Only one colony is known at present (this paper); based on museum specimens there are at least two other possible sites where A. zikani has occurred (see Fig. 1). Undoubtedly, there is an urgent need for discovering more colonies of this species in the region. The analysis of topographic charts (1:50,000) between Paranapiacaba and Boracéia revealed 10 areas with altitude from 1100 to 1200 m, within 10 km from the Serra do Mar break, all potentially suitable for a population of A. zikani. Visits to some of these areas should be an immediate priority. Further areas SW of Paranapiacaba should also be visited; although winter temperatures are lower towards the south (Nascimento & Pereira 1988: Nimer 1972, 1989) little is known about the tolerances of this species and the potential occurrence on both NW- and SE-facing summits. In any case, if there is still a population present in the Estagao Biolégica de Boracéia, or in any additional location northward, it will be partially isolated from the population studied in Paranapiacaba, since much of the original forest throughout this region has been replaced by eucalyptus trees. VOLUME 59, NUMBER 3 Even the colony of Paranapiacaba is not completely protected. The area is constantly visited for maintenance of the towers, resulting in clearing of the roads and of the drainage network, leading to vigorous growth of more aggressive Mikania species that overcome M. obsoleta. The heavy ecotourism in the region contributes to environmental degradation and accumulation of garbage in the initial part of the area. The entire ridge of the Serra do Mar area including the region of Paranapiacaba is discontinuous to the southwest, where two major superhighways, two railroads, many oil ducts and service roads, several cleared tracks for power line maintenance, and increasing urbanization of the slopes of the mountains (below 400 m) create a mosaic of unsuitable habitats. Perhaps due to the extensive and continuous modification of the vegetation in the entire region, recent trips to the area (1997 to 2005) showed no trace of A. zikani and few plants of M. obsoleta. This microsystem appears to be composed of fugitive species (cf. Horn & MacArthur 1972) which compete poorly with relatives better adapted to this patchy environment (see the example of Heliconius nattereri Felder & Felder in Santa Teresa, Espirito Santo in Brown 1972). Nowadays the area seems apparently less impacted (with a single new tower constructed there), but the food plants (M. obsoleta) continue to disappear. The history of environmental conservation in the area of the Serra do Mar in the State of Sao Paulo starts at the beginning of twentieth century, when Herman von Ihering, director of the Museu Paulista (now Museu de Zoologia da Universidade de Sao Paulo) demonstrated the need to create federal legislation to regulate bird hunting (Ihering 1902), and later emphasized the conservation of forests (Ihering 1911). At the end of 1909 he set up on his own property a sanctuary, the "Estagao Biol6gica do Alto da Serra", now under responsibility of the Instituto de Botanica da Secretaria da Agricultura do Estado de Sao Paulo (Kirizawa et al. 2004). This protected area is located SW_ of Paranapiacaba, and unfortunately has been heavily polluted with fluoride and other chemical contaminants from the Cubatao industrial area at the base of these mountains (Klump et al. 1996; Kirizawa et al. 2004). Future perspectives. The observations made here are an initial step to a better understanding of this fugitive pair of species A. zikani and M. obsoleta. To help answer the open questions, the following actions are needed: more accurate estimates of population parameters (mean residency time and home-range), verification of the impact of the ants that prey on eggs, better estimates of population parameters of the foodplant, and laboratory testing of larval acceptance of 14] other Mikania species. In spite of the study site being close to a contaminated area (see above), the population of A. zikani is relatively protected against air pollution by the summits of a nearby mountain. range, that deflect the winds toward the west. The reasons for the observed disappearance of the population in the study area (see above) were not investigated. Perhaps there is a natural cycle of this pair of species, becoming common in a few years and scarce in most (as observed for some species of Actinote by the authors). ACKNOWLEDGEMENTS We would like to thank Paulo Vanzolini, Cleide Costa, Ubira- jara Martins, M. Marques and Francisca C. do Val for permis- sion to study the material in the collection of MZUSP. and A. P. A. F. Moraes, Carlo L. B. Francini, Ronaldo B. Francini Filho. A. S. Gongalves and Renato R. Ramos for their help in field- work. Carla Penz and Fernando Frieiro-Costa provided essen- tial literature and Marcio Uehara-Prado, Danilo Bandini Ribeiro, Carla Penz and Gerardo Lamas gave valuable sugges- tions for the manuscript. This study was funded by Fapesp (grants 84/0432-3, 86/0618-5, 88/3069-8 and 93/0097-9 to RBF. and 00/01484-1 and 04/05269-9 to AVLF) and is part of the pro- ject “Lepidoptera of Sao Paulo State” (BIOTA-FAPESP pro- gram - grant 98/05101-8). André V. L. Freitas thanks also the National Science Foundation (DEB-0316505). LITERATURE CITED BERNARDES, A. T., A. B. M. MACHADO & A. B. RyLanps. 1990. Fauna brasileira ameagada de extingao. Fundacao Biodiversitas/IBAMA, Belo Horizonte, 60 pp. Brown, K. S., Jr. 1972. The Heliconius of Brazil (Lepidoptera: Nymphalidae). Part III. Ecology and biology of Heliconius nattereri, a key primitive species near extinction, and comments on the evolutionary development of Heliconius and Eueides. Zoologica, 57: 41-69. ——. 1991. Conservation of Neotropical environments: insects as in- dicators, pp. 349-404. In Collins, N. M. & J. A. Thomas (eds.). The Conservation of Insects and their Habitats, Academic Press. London. Brown, K. S., JR. & R. B. FRANCINI. 1990. Evolutionary strategies of chemical defense in aposematic butterflies: cyanogenesis in Asteraceae-feeding American Acraeinae. Chemoecology, 1: 52- 56. BrussarD, P & P. R. ERLICH. 1970. The population structure of Erebia epipsodea (Lepidoptera: Satyrinae). Ecology, 51: 119-129. D'ALMEIDA, R. F. 1943. Algumas observacoes sobre 0 Actinote morio Oberthiir, 1917 (Lep. Heliconidae, Acraeinae). Papéis Avulsos do Departamento de Zoologia, Sao Paulo, 3: 107-110. ——. 1951. Umanova espécie de Actinote do Sul do Brasil (lep. He- liconiidae, Acraeinae). Arquivos do Museu Nacional,Rio de Janeiro, 42: 1-5. FRANCINI, R. B. 1989. Biologia e ecologia das borboletas Actinote (Lepidoptera, Nymphalidae, Acraeinae) na transicao subtropical no sudeste do Brasil. Dissertagao de Mestrado em ecologia, Uni- versidade Estadual de Campinas. 236 pp. 1992. Ecologia das taxocenoses de Actinote (Lepidoptera: Nymphalidae) em Asteraceae (Angiosperma: Magnoliatae) no sudeste do Brasil: subsidios para conservagao. Tese de doutora- mento em ecologia, Instituto de Biologia, Universidade Estadual de Campinas, SP, 194 pp. Freitas, A. V. L. 1991. Variagao morfolégica, ciclo de vida e sis- tematica de Tegosa claudina (Eschscholtz) (Lepidoptera, Nymphalidae, Melitaeinae) no estado de Sao Paulo, Brasil. 142 Revta. bras. Ent. 35: 301-306. 1993. Biology and population dynamics of Placidula euryanassa, a relict ithomiine butterfly (Nymphalidae: Ithomiinae). J. Lepid. Soc, 47: 87-105. ——. 1996. Population biology of Heterosais edessa (Nymphalidae) and its associated Atlantic Forest Ithomiinae community. J. Lepid. Soc. 50: 273-289. Horn, H. S. & R. H. MacArtuur. 1972. Competition among fugitive species in a harlequin environment. Ecology, 53: 749-752. IGGSP (editor) 1972. Folha SANTOS, SF-23-Y-D-IV-4. Mapa topografico em escala 1:50.000. Instituto Geografico e Geolégico do Estado de Sao Paulo, Sao Paulo. IHERING, H. 1902. Necessidades de uma lei federal de caga e protecgao de aves. Revista do Museu Paulista, 5: 238-260. —. 1911. Devastagao e conservagao das mattas. Revista do Museu Paulista, 8: 485-500. IUCN 2001. IUCN Red List Categories and Criteria version 3.1. IUCN, Gland and Cambridge. Krrizawa, M., M. SuctyaAMa, E. A. Lopes & A. CUSTODIO-FILHO. 2004. Flora da Reserva Bioldgica do Alto da Serra de Paranapia- caba Santo André, Sao Paulo, Brasil. http://www.ibot.sp.gov.br/PESQUISA/paranapiacaba/paranapi_re- sultados.htm Kiumep, A., M. DomINcos & G. KLuMpp. 1996. Assessment of the vegetation risk by fluoride emissions from fertiliser industries at Cubatao, Brazil. The Science of Total Environment, 192: 219- 228. Lamas, G. 1973. The type material of Lepidoptera Rhopalocera con- tained in the collections of the Museu de Zoologia da Universi- dade de Sao Paulo. Papéis Avulsos de Zoologia, Sao Paulo, 26: 179-185. MIELKE, O. H. H. & M. M. CasaGRANDE. 1986. Sobre os tipos de Lepidoptera depositados em museus brasileiros. II. Nymphali- dae (Danainae, Brassolinae, Morphinae, Satyrinae e Acraeinae), descritos por R.F.D'Almeida. Revista Brasileira de Entomologia, 30: 141-152. MMA, 2003. Anexo a Instrucao Normativa n° 3, de 27 de maio de 2003, do Ministério do Meio Ambiente. Lista das Espécies da JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fauna Brasileira Ameagadas de Extingao. http:/Avww.ibama. gov. br/fauna/downloads/lista%20spp.pdf NASCIMENTO, C. M. & M. A. M. G. PEREIRA, 1988. Atlas clima- tolégico do Estado de Sao Paulo. Fundagao Cargill, Campinas. NiMeER, E, 1972. Climatologia da regiaio sudeste do Brasil. Intro- dugao a climatologia dinamica. Revista Brasileira de Geografia, Rio de Janeiro, 34; 3-48. ——. 1989. Climatologia do Brasil. Fundagao IBGE, Rio de Janeiro, 421 pp. NOAA 1998. The Global Historical Climatology Network (GHCN),. http:/Awww.nede.noaa.gov/oa/pub/data/ghen/v2/ghenftp_zipd.html PENz, C. M. & R. B. FRANCINI. 1996. New species of Actinote Hiib- ner (Nymphalidae: Acraeinae) from Southeastern Brazil. Journal of the Lepidopterists' Society, 50: 309-320. Ramos, R. R. & A. V. L. FreITAS. 1999. Population biology and wing color variation in Heliconius erato phyllis (Nymphalidae). J. Lepid. Soc. 53: 11-21. Santos, E. O. 1965. Caracteristicas climaticas, pp. 95-150. In Azevedo, A. (ed.). A baixada Santista, aspectos geograficos, vol. 1. Cia. Editora Nacional/EDUSP, Sao Paulo. SIGRH. 2003. Sistema Integrado de gerenciamento de recursos hfdricos do Estado de Sao Paulo: Banco de dados Pluviométricos do Estado de Sao Paulo. http://www.sigrh.sp.gov.br/sigrh/basecon/bancodedados/plw/plu.htm SouTHWOOD, T. R. E. 1971. Ecological Methods with Particular Ref- erence to the Study of Insect Populations. Chapman & Hall, London, 524 pp. SP-SMA. 1998. Fauna ameagada no estado de Sao Paulo. Secretaria do Meio Ambiente, SMA/CED, Sao Paulo. Ururany, J. C. C., J. E. R. CoLLareE S, M. M. Santos & R. A. A. BarRETTO. 1984. 4. Vegetac&o, pp. 553-611. In Projeto RADAMBRASIL, levantamento de Recursos Naturais. Vol. 32. fls. Sf 23-24 (Rio de Janeiro e Vitéria). Fundagao IBGE, Rio de Janeiro VOLUME 59, NUMBER 3 143 Journal of the Lepidopterists’ Society 59(3), 2005, 143-160 EXTRINSIC EFFECTS ON FECUNDITY-MATERNAL WEIGHT RELATIONS IN CAPITAL-BREEDING LEPIDOPTERA WILLIAM E. MILLER Department of Entomology, University of Minnesota, St. Paul, Minnesota 55108 USA email: mille0]4@umn.edu ABSTRACT. Capital-breeding Lepidoptera depend for reproduction on metabolic resources assembled either entirely or primarily by their larvae, the former termed 'perfect' the latter 'imperfect'. Empirical evidence suggests that maternal size determines capi- tal-breeder fecundity. The fecundity-maternal size relation is usually formulated as F = bW + a, where F is fecundity, W is final ma- ternal size in units such as weight of newly transformed pupae, b is the slope, and a the intercept. Exhaustive search yielded 71 fe- cundity-maternal pupal weight relations for 41 capital breeders in 15 families, 58 of which, including 2 previously unpublished, were based on individual specimens, and 13 on grouped specimens. In 22 individual-specimen relations, cohorts divided into 2 or more subgroups were reared simultaneously at different temperatures, on different diets, or exposed to other extrinsic factors. These 22 ‘multiform! relations were compared with 36 ‘uniform! relations, and where possible cohort subgroups were compared. Pupal weights of cohort subgroups were affected much oftener than underlying slopes and intercepts. Individual-specimen slopes based on transformed data ranged 0.52-2.09 with a mean and standard error of 1.13+0.04, and slopes did not differ significantly among perfect, imperfect, multiform, and uniform categories. Despite the evident similarity, one relation does not apply to 5 all capital breed- ers. Tradeoffs sometimes occur between fecundity, F, and mean egg weight, E. Reaction norms of fecundity and pupal weight across extrinsic-factor ranges were overwhelmingly congruent, which supports axiomatic status for the dependence of fecundity on capital- breeder maternal size. Cooler rearing temperatures usually produced heavier female pupae and greater fecundities, a phenomenon of population dynamics interest. The two sides of practically all fecundity-maternal weight regressions are not statistically indepen- dent, in effect stating F = b(W + [F x E]) + @, which artificially inflates test statistics. Where desirable, the fully independent rela- tion R = b(W - [F x E]) + a can be used, where R is reproductive bulk, the mathematical product of F x E. Additional key words: temperature, diet quality, population dynamics 'Capital-breeding' describes Lepidoptera that depend for reproduction entirely or primarily on metabolic resources assembled by their larvae, in contrast to ‘income-breeding', which describes those that depend for reproduction primarily or entirely on resources assembled by their adults (Boggs 1992, Miller 1996, Tammaru and Haukioja 1996). The gypsy moth, Lymantria dispar (L.) (Lymantriidae), is a capital breeder; the monarch butterfly, Danaus plexippus (L.) (Nymphalidae), an income breeder. In four butterfly income breeders in two families, income contributed = 80% to fecundity, and capital < 20% (Boggs 1997, Fischer and Fiedler 200la). Based on sizes of superfamilies (Kristensen and Skalski 1999) and the extent to which income breeding is phylogenetically limited, probably =90% of extant Lepidoptera are capital breeders. Most outbreak Lepidoptera also are capital breeders (Miller 1996, Tammaru and Haukioja 1996). Capital breeders have an ovigeny index, OI, of 1 or >> 0, referring to the proportion of lifetime potential fecundity that consists of mature eggs at eclosion, whereas income breeders have an OI of 0 or <<1 (Jervis and Ferns 2004). Capital breeders with nonfeeding adults and Ols of 1 are here termed 'perfect', whereas those with OIs of >>0 whose adults may feed, but do so less than income breeders, are termed 'imperfect'. Maternal size is widely believed to determine fecundity in capital breeders (Leather 1988, Honek 1993). This belief derives not from experimentation but from long empirical observation. Direct fecundity-size relations occur in the lepidopteran phy logenetic sequence at least as early as Tineidae, the basal-most lineage of Ditrysia (Titschack 1922, Kristensen and Skalski 1999) and are probably part of the ground plan of Ditrysia, if not all Lepidoptera. This densndenes implies that whatever influences maternal size may influence fecundity and its associated quality attributes, and thus population fluctuations. Fecundity can be a proxy for net reproductive rate (Carey 1993, Huey and Berrigan 2001) and has been implicated in capital- breeder population fluctuations, as in Bupalus piniaria L. (Geometridae) (Klomp 1966), Bucculatrix pyrivorella Kuroko (Bucculatricidae) (Fujiie 1980), Leucoptera spartifoliella (Hiibner) (Lyonetiidae) (Agwu 1974), and in capital-breeding Noctuidae (Spitzer et al. 1984). Traditionally, the relation between fecundity, F, and maternal weight, W, usually has been defined by linear regression as F = bW + a, where W refers to newly transformed pupae or newly eclosed adults, b is the slope, and a is the intercept or scaling parameter. Honek (1993) devised a fecundity- Tmatemall weight relation for insects generally, as well as one for Lepidoptera, but he did not segregate capital breeders for special study nor exhaustively seek examples. Honek noted that w eight appears on both sides of fecundity-maternal weight regressions, but that statistically independent measures of fecundity and maternal weight are practically 144 nonexistent. In effect, such relations state that F = b(W + [F x E]) + a, where E is mean egg weight. The resulting nonindependence inflates test statistics and minimizes variation between response and explanatory variables. The practical usefulness of the traditional regressions is not necessarily impaired, but their statistics should not be used where strict independence between the variables is assumed. As discussed further on, a fully independent alternative relation emerged from this study. In any capital-breeder reared under homogeneous conditions, intrinsic effects alone will produce a direct relation between fecundity and maternal size. If a cohort of eggs or hatchlings is divided into subgroups, and each subgroup reared at a different level of an extrinsic factor, such as a different temperature, or on a different diet, then extrinsic effects are likely to be added to the intrinsic ones. Here I examine extrinsic effects on fecundity-maternal pupal weight relations during rearing of capital breeders. I focus on effects produced by different temperatures—as might occur during anomalous weather, or between microhabitats, or between generations or seasons—and by differing diet quality—as might occur on variably stressed or different kinds of foodplants, or on different kinds or amounts of adult nourishment. MATERIALS AND METHODS I assembled as many statistical fecundity-maternal weight relations as possible from a personal reference collection, electronic databases including Biosis, Biological Abstracts, and the Zoological Record, and from citations in references. Most velutions were based on observations of specimens individually, a few on means of grouped specimens. Individual-specimen relations were admitted if based on samples numbering > 20, grouped-specimen relations if based on groups numbering =5. No relations were excluded because of non-English text. In the 58 assembled individual-specimen relations, weights and fecundities were available in numerical form for three published and two unpublished ones (Table 1); weights and fecundities for the remainder were transcribed from enlarged photocopies of published scatterplots. Because transcription creates error—when one point covers another, for instance—I tested slopes of transcribed relations against corresponding slopes given in sources. A few departures were statistically significant, but most were not (F-tests, P = 0.99-0.009; median P = 0.76; n = 35). If P was < 0.25, I retranscribed, but in no case did retranscription change the outcome appreciably. I accepted scatterplots at face value despite minor inconsistencies, except that JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY for Philosamia ricini Hutt. (Singh and Prasad 1987), which seemed too anomalous. In the 13 grouped- specimen relations, most weights and fecundities were available in numerical form (Table 2). Study relations consisted of perfect and imperfect groups and uniform and multiform subsets. 'Uniform! denotes homogeneous conditions of development expected to produce only intrinsic effects, and 'multiform! denotes heterogeneous conditions expected to produce extrinsic as well as intrinsic effects. I examined relations for extrinsic effects first by meta- analysis (Gates 2002) and second by comparing cohort subgroup relations provided in sources or obtained by deconstruction. A standardized maternal weight was desirable, and I chose fresh pupal weight. By ce pupal stage metabolic resources for ovigenesis are in place. Moreover, pupal weight has been most often used in describing fecundity-size relations (42 of 58 relations in Table 1, 12 of 13 in Table 2), and explanatory variables based on weight outnumber those based on lineal dimensions such as forewing length and pupal diameter. I maximized the number of relations for study by converting female adult fresh weight, W,, to fresh pupal weight, W_, where Ww, = W, x 1.85, a fete based on four observations: (1) Arst= day female pupae of Malacosoma disstria (Hbn.) (Lasiocampidae) in a previously unpublished study averaged 1.98 times heavier than first-day adults (n = 30 weighings, paired); (2) a corresponding value of 1.81 for Epiphyas postvittana (Walker) (Tortricidae) (n >130 weighings, unpaired) (Danthanarayana 1975); (3) a corresponding value of 1.74 for Streblote panda (Hbn.) (Calvo and Molina 2005); and (4) a corresponding value of 1.67 for Cnephasia jactatana (Walker) (Tortricidae) (Ochieng'- Odero 1990). Fecundity had been estimated in sources by various methods, all internally consistent and all accepted here. Methods included counting unlaid eggs in dissections of newly eclosed females, counting only eggs actually laid, and combining eggs laid with residual eggs in ovaries after death. For meta-analysis, I transformed fecundities and pupal weights of each relation to percentages of their midranges (Honek 1993) ([individual value/midrange value] x 100, where midrange = 0.5 x [minimum value + maximum value]). This transformation enabled comparison of relations for different species and groups on a single scale, as between large saturniids and small tortricids. Statistics were generated by SYSTAT (1992) software. Student's t used pooled variances except where noted otherwise. In analyzing and comparing cohort subgroup VOLUME 59, NUMBER 3 relations, I used nontransformed transcribed data when tabulated data were unavailable. In tests of differences among fecundity-maternal weight regressions of cohort srnineasaTeT Lincludeda categorical erallaeniony variable, as well as a maternal ore X group interaction term, the latter enabling tests of differences among slopes and the former enabling tests of differences among heights of regression lines. Height tests were used here as tests of intercept differences. Where subgroup relations within multiform relations were not provided in sources and deconstruction was not possible, the fact is noted. Wherever possible, I report whether reaction norms of fecundity and maternal pupal weight are congruent. A reaction norm is the trajectory of response values across the range of an extrinsic factor (Schlichting and Pigliucci 1998). Congruency signifies that fecundity and female pupal relent peak at the same value of an extrinsic factor. Bescon norms to rearing temperatures are illustrated in Fig. la, and those to different diets in Fig. 2a. In sources where it was not possible to evaluate congruency, it is noted as indeterminate. RESULTS The 71 assembled fecundity-maternal pupal weight relations represent 41 capital breeders in 15 families (Tables 1 and 2). The 58 individual-specimen relations (Table 1) consist of 35 in perfect and 23 in imperfect groups, 36 of the 58 uniform and 22 multiform. Perfect- uniform and perfect-multiform categories number 23 and 12, and imperfect-uniform and imperfect- multiform categories each number 13. The 13 grouped- insect relations (Table 2) consist of 10 of the perfect kind and 3 of the imperfect, with 5 uniform and 8 multiform. Individual-specimen relations are unaffected by aggregation bias and thus receive more attention here than grouped-specimen relations. In all discussions, attributes and their numerical values appear in parallel sequences. 'Tradeoffs' refer to any change in proportion between fecundity and egg size. Egg size refers to the mean weight of one egg in an eg load. Individual-specimen relations Meta-analysis. Midrange maternal pupal weight, which approximates the mean, varied from 12-9435 mg (Table 1), averaging 1401 and 63 mg for perfect and imperfect groups. The difference is highly significant (Student's t [separate variances] = 3.74; df = 34. 1;P< 0.001). Range in transformed weights of pupae (greatest % of midrange minus smallest % of midrange) varied from 34-179 (Table 1), averaging 97 and 99 for perfect and imperfect groups. Corresponding uniform and multiform ranges are 34-179 and 48-143 (Table 1), averaging 97 and 100. Slopes of fecundity-maternal pupal weight relations varied from 0.52—2.09 (Table 1, Fig. 3), with the mean and standard error, SE, 1.13+0.04. This overall mean significantly exceeds 1.00 (Student's t [one-tailed test] = 3.02; df = 57; P = 0.02). Slopes for perfect and imperfect groups average 1.14 and 1.12; those for uniform and multiform subsets also 1.14 and 1.12. The four slopes for perfect-uniform and perfect-multiform, and imperfect-uniform and imperfect multiform categories average 1.13 and 1.15, and 1.14 and 1.09. Differences among them are not significant (F = 0.08; df = 3, 54; P = 0.97). Pooled slopes are likewise unrelated to midrange pupal weight (Pearson's r = 0.06; df = 56; P = 0.68) or to range (Pearson's r = 0.04; df = 56; P = 0.77). Although a positive correlation among _ relations between range and number of observations either as n or log.n could be expected statistically, it did not miaterialize (Pearson's r = 0.16; df = 56; P = 0.24). Slope variability as SE,/b among perfect-uniform and perfect- multiform categories averaged 9.5 and 8.9, and among imperfect-uniform and imperfect-multiform, 13.0 and 15.7. The mean for the entire imperfect group, 14.8, is seemingly higher than that for the entire perfect group, 9.3, but the difference is not strictly significant (Student's t [separate variances] = 1.85; df = 27.4; P = 0.07). So-called funnel diagrams—plots of SE, vs. b— visually suggest greater scatter among imperfect than perfect groups (Fig. 3a, b). Spatial and temporal effects. Coefficients of fecundity-maternal pupal weight relations for the same species often differ geographically (Table 1), but only those obtained in the same way by the same workers can be meaningfully compared, as in the following examples. Lorimer (1979) found slopes and intercepts of two uniform relations for Malacosoma disstria from Indiana and Michigan to differ significantly. Parry et al. (2001) found fone of the six M. disstria slope comparisons for Michigan, Manitoba, and Louisiana between two years to differ significantly, as well as all of the six intercept comparisons. In M. disstria, however, a tradeoff between fecundity and egg size occurs as a geographic NW-SE cline (Parry et al. 2001). At two Quebec locations, slopes of 0.26 and 0.18 for Lymantria dispar differed significantly (F = 20.5; df= 1, 111; P < 0.001) (transcribed data) (Madrid and Stewart 1981). Egg size is notably plastic in L. dispar (Rossiter 1991). Lorimer and Bauer (1983) found that fecundity-pupal weight slopes for Choristoneura fumiferana differed between New Hampshire and Minnesota; atypically, significant correlation absent in the oe (transcribed data). 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Both fecundity and female pupal weight peaked at 22.7°, making their reaction norms congruent. Points on the pertinent source scatterplot were not temperature-coded, which precluded deconstruction. Relation 31. Calvo and Molina (2005) reared Streblote panda at four arbitrary constant temperatures from 19-28° C on two foodplants. Congruency of fecundity and maternal weight to the temperatures and foodplants is indeterminate, and the scatterplot precluded deconstruction. Relation 34. Kamata and Igarashi (1995) reared Quadricalcarifera punctatella at arbitrary constant temperatures from 10-25° C. At the lower temperatures, most larvae had four stadia, and at higher ones, most had five. The authors also field-collected pupae for comparison with the laboratory rearings. Points on their scatterplot of fecundity vs. adult female weight were coded as four-stadium, five-stadium, and field-collected, which permitted deconstruction for analysis and comparison of cohort subgroup regressions. noncoded After adult-to- pupal weight conversion, female pupae averaged 390, 520, and 741 mg for the four-stadium (cooler), five-stadium (warmer), and field-collected, all of which are inferred to differ significantly from one another (F = 391.8; df = 2, 74: P < 0.001) (transcribed data). In the three subgroup fecundity-maternal weight relations, slopes and intercepts were 0.72 and -108.4, 0.64 and -74.8, and 0.64 and -48.3. re slopes did not differ significantly (F = 0.82; df = 2, 71; P = 0.44), nor did the intercepts (F = 0.88; df = 2, 71; P = 0.42) (transcribed data). Congruency of fecundity and maternal weight is indeterminate. = (.79; white oak, F = 0.88 W - fecundity vs. pups al weight after transformation to percentages of poole »d midrange values. E quation is F, e Pignut hickory .__ Pupal weight (mg) (W) =i WA = 0.92. Data Fecundity as % of pooled midrange (Fp) 160 120 80 * White oak 4 © Pignut hickory © Northern red oak 4 White oak 40 200 250 300 350 30) 50. 70 (90!) 110 180" 150551770) Pupal weight as % of pooled midrange (Wp) Relation 35. Mizuta et al. (1969) reared Japanese and Chinese varieties of Bombyx mori, and resulting points on their scatterplot of fecundity vs. female pupal weight were coded as spring or summer and assumed to represent individual specimens. No rearing temperatures were given, but temperatures in spring were likely cooler than those in summer. Mean spring and summer female pupal weights were 1451 and 1133 mg, and the difference, 318 mg, is highly significant (Student's t = 10.5; df = 144; P < 0.001) (transcribed data). Corresponding fecundity-maternal weight slopes and intercepts are 0.30 and 70.8, and 0.45 and - 128.3. The slopes do not strictly differ (F = 3.0; df = 1, 142; P = 0.085), nor do the intercepts (F = 3.79; df = 1, 142; P = 0.053). Occurrence of a fecundity-egg size tradeoff could not be ascertained, but given the many varieties of B. mori involved here and sown generally (Hiratsuka 1999), tradeoffs would not be surprising. Congruency of fecundity and maternal weight is indeterminate. Relation 53. This previously unpublished Choristoneura fumiferana relation is drawn from a two- part experiment. The first part used the whole sample, which focused on the response of female pupal weight to different rearing temperatures. The second part used a subsample of the whole in which fecundity as well as female pupal weight was measured. Certain comparisons in both groups are abbreviated because some data were unfortunately lost as detailed further on. Foliage harboring overwintered second instars within 3 m of the ground was collected in early May from Abies baleen (L.) Mill. and Picea glauca (Moench) Voss (both Pinaceae) near Cloquet, Minn. The second instars were light-extracted (Miller 1958) and placed at densities of 1 and 3 per cup in 25 ml VOLUME 59, NUMBER 3 Fic. 3. Funnel diagrams showing standard errors of slopes, SE,, relative to the corresponding slopes, b. a. Perfect group. b. Imperfect group. SE of slope (a) Perfect ° Uniform i @® Multiform 0.40 0.30 0.10 0.4 0.7 1.0 1.3 1.6 1.9 Slope plastic cups with artificial diet. Cups with each density were divided into three subgroups and reared as outlined by Grisdale (1970) in three growth chambers programmed for different temperatures. The first chamber simulated the natural seasonal march of daily mean day-night temperatures based on long-term averages at a weather station = 20 km from the collection site, and presumably approximated the temperature regime to which the insect was acclimated. Chamber temperatures were increased 1° C every 4-6 days during May and June until adult eclosion. At the start, day-night temperatures were 10° and 3°, and at the end, 26 and 12° C. The second and third chambers housed cups at each density, with one chamber programmed 5° cooler than presumed acclimation and the other 5° warmer, their temperatures being increased incrementally as in the first chamber. All chambers were fluorescent-illuminated on a 16:8 D:N schedule. In early June, four weeks after the first collection, fifth instars were collected at the site, placed singly in cups with artificial diet, and added to the experiment for exposure to the rearing temperatures only during the late larval stage, from the latter part of stadium 5 through final stadium 6. All pupae were weighed and sexed within 24 hr after transformation. The subsample for computing fecundity-maternal weight regressions consisted of female-male pupal pairs placed one each with a6 cm long sprig of foodplant in 0.5 liter cardboard ice cream containers in which they eclosed, mated, and oviposited. The OI of this species is = 0.4 (Outram 2.2 0.4 0.7 1.0 1.3 1.6 1.9 (b) Imperfect QO Uniform = Multiform Oo i) i) Slope 1971, Miller 1987), but no liquids were provided to adults during posteclosion ovigenesis. Laid eggs were counted daily, and unlaid chorionated eggs were added to the daily counts after females died, with chorionation evaluated by ovarian staining with methylene blue (Miller 1987). As in the ehole sample, a subsample subgroup developed under each of three temperature regimes, under long and short exposures to temperature regimes, and at two rearing densities. In the whole sample, mean weights of long-exposure female pupae at single and triple densities across all temperatures were 100.0 and 101.7 mg, indicating no real difference due to density. Long-exposure single- density female pupal weights at presumed acclimation, at 5° cooler, and at 5° warmer, were 103.8, 106.9, and 85.8 mg, and the 18.0 mg lesser weight at warmer than presumed acclimation is highly significant (Student's t = 3.75; df = 196; P < 0.001). Long-exposure triple-density weights were 99.4, 111.9, and 93.6 mg, and the 12.5 mg greater weight at cooler than presumed acclimation is significant (Student's t = 2.90; df = 248; P = 0.006). Short-exposure single-density female pupal weights were 111.3, 110.8, and 104.7 mg, and the 6.6 mg lesser weight at 5° warmer than presumed acclimation is significant (Student's t = 2.24; df = 293; P < 0.038). The foregoing reliance on t-tests rather than comprehensive F-tests was necessitated by loss of some data before analysis was completed. In the fecundity- maternal pupal weight subsample with densities and exposure lengths pooled, mean fecundities and mean more 154 female pupal weights at presumed acclimation and 5° warmer were 167.2 and 106.6, and 112.6 and 88.5, congruent temperatures in the surviving data, data from 5° cooler indicating reaction norms to rearing than acclimation having been lost. pupal weights mirrored those of the whole sample, but These subgroup the associated fecundities were depressed compared with published experiments in which females received imbibing liquids (Miller 1987, 1989). Slopes and intercepts of pooled subsample data at presumed ,and 1.91 and -56.3, but although seemingly disparate, neither acclimation and 5° warmer were 1.26 and 23.2 slopes nor intercepts differed significantly mslepesF eo 0.046; df = 1, 38; P = 0.83, intercept F = 0.12: df 38; P = 0.75). Relation 58. Hamilton and Zalucki (1991) reared Crocidosema plebejana at a number of arbitrary constant temperatures from 14-31° C. Fecundity and female adult weight were congruent. Their scatterplot of fecundity vs. female adult weight is temperature- coded for seven rearing temperatures, which enabled deconstruction for analysis of constituent regressions. I computed two fecundity- maternal weight regressions, one for 14—21° C, and the other for 25-31°. Their slopes and intercepts were 51.8 and -237.6, and 47.9 and -195.0, and neither the 7 Siopes nor gigs differed significantly (slope F ee : df = 1, 71; P = 0.42, intercept F = 0.99; df ae p= Pa, (transcribed data). In addition, the ae showed statistically that fecundity depended on female weight independently of temperature. Diet effects. Twelve multiform relations involved cohort subgroups that received different larval diets (Nos. 26, 28, 32, 33, 50, 52, 54-56, 67) or different adult diets (Nos. 49, 51). Subgroup larvae were reared separately on different foodplants or on diets differing otherwise in quality or quantity. In the adult group, different imbibing liquids were provided, or withheld: during posteclosion ovigenesis. Each relation is discussed below. Relation 32 for Ennomos subsignarius is discussed first because it typifies the effect different larval diets have on fecundity-maternal weight relations. Drooz (1965) reared this species on pignut hickory, Carya glabra (Mill.) Sweet (Juglandaceae), northern red oak, Quercus rubra L., and white oak, Q. alba L. (Fagaceae). He found that fecundity and female pupal weight were significantly higher on pignut hickory than on the oaks (transcribed surrogate values here in Fig. 2a). Fecundity and female pupal weight across foodplants were congruent. In a subsample, Drooz computed regressions of fecundity vs. female pupal weight by juvenile-mature aK JURNAL OF THE LEPIDOPTERISTS’ SOCIETY individual and pooled foodplants (transcribed surrogate values in Fig. 2b). I confirmed that differences among slopes and intercepts of these regressions do not differ significantly (slope F = 0.63; df = 2, 30; P = 0.54, intercept F = 0.51; df = 2, 30; P = 0.50) (transcribed data). The three subgroups are pooled in the summary relation (Fig. 2c, Table 1). Relation 26. (1997) reared cohort subgroups of Lymantria dispar on juvenile Cambini and Magnoler foliage alone and mixed juvenile-mature foliage of cork oak, Quercus suber L., and holm oak, Q. ilex L. They showed that female pupae were significantly heavier on cork than on holm oak, and on juvenile than on mixed foliage. Six-stadium females predominated on cork oak, and 7-stadium females predominated on holm oak. The authors also showed that neither slopes nor intercepts of fecundity-maternal pupal weight relations among the subgroups differed significantly by foodplant. Fecundity and female pupal weight were consistently congruent. Relation 28. Van der lends and Votite (1967) reared Euproctis chrysorrhoea on English oak, Quercus robur L., and buckthorn, Rhamnus sp. (Rhamnaceae), and their scatterplot of fecundity vs. pupal weight was foodplant-coded, which permitted deconstruction. Mean weights of female pupae were 186 mg on English oak and 268 on buckthorn, and the difference, 82 mg, is highly significant (Student's ¢ = 6.66; df = 54; P < 0.001) (transcribed data). Corresponding slopes and intercepts were 1.78 and -112.4, and 1.63 and -62.1, but neither slopes nor intercepts differed significantly (slope F = 0.31; df = 1, 52; P = 0.58, intercept F = 0.61; df = 1, 52; P = 0.44). Congruency of fecundity and female pupal weight is indeterminate. Relation 33. Gruys (1970) reared Bupalus piniaria at different including 1 larva/container, considered uncrowded, and 2-5 larvae/container, considered crowded. He presented uncrowded and crowded fecundity-maternal weight _ scatterplots separately. Mean weights of uncrowded and crowded female pupae were 184 and 152 mg, and the wes 32 mg, is highly significant (Student's ¢ = 15.1; df = P < 0.001) (transcribed data). Gruys speculated a increased bodily contact in the crowded group adversely affected nutrition. He found that both slopes and intercepts of fecundity-maternal pupal weight regressions differed significantly between the two groups. He also found that crowding significantly reduced mean egg weight, which indicates a fecundity- egg size tradeoff between the two densities. Fecundity and female pupal weight proved congruent. Because naturally, uncrowded and crowded densities density varies VOLUME 59, NUMBER 3 subgroups are pooled in the summary relation. REMatiOn 49. Tisdale and Sappington (2001) fed three groups of mated female adult Spodoptera exigua different liquid diets—10% honey in water, 10% sucrose in water, and plain water. They showed that fecundity of females on both carbohydrate diets significantly exceeded that of females on plain water. Their diet- specific scatterplots of fecundity vs. female pupal weight allowed deconstruction. I confirmed that pupal weights among treatment groups were statistically equal at the start (F = 0.51; df = 2, 126; P = 0.60) (transcribed data). Slopes and intercepts for the honey, sucrose, and plain water diets were 13.3 and 12.8, 12.4 and 43.6, and 16.5 and 1.78. The slopes did not differ significantly (F = 1.00; df = 2, 122; P = 0.37), nor did the intercepts (F = 2.03; df = 2, 122; P = 0.14) (transcribed data). It should be mentioned that S. exigua might be borderline between capital and income breeding, Relation 50. Bessin and Reagan (1990) field- Gollected pupae of Diatraea sacc haralis from two varieties of sugarcane, Saccharum spp.; corn, Zea mays L.; and johnsongrass, Sorghum halapense (L.) Persoon (all Gramineae). They showed that weights of female pupae differed significantly by foodplant but that differences among slopes and intercepts of the corresponding fecundity- size regressions did not differ significantly. Congruency of fecundity and pupal weight is indeterminate. Relation 51. Hagstrum and Tomblin (1975) provided drinking water to mated female adults of one group of stock-culture Cadra cautella and withheld it from a second group. They presented scatterplots of fecundity vs. female weight separately for drinkers and nondrinkers. I confirmed that weights of the two groups were statistically equal at the start iGedent Bie IL 83; df = 75: P = 0.07) (transcribed data). The authors found that drinkers laid significantly more eggs than nondrinkers, and that the corresponding fecundity- maternal weight regressions also differed significantly, Drinkers and nondrinkers are pooled in the summary relation because extent of drinking probably varies in environments where this species occurs. Relation 52. Russell et al. (1980) reared two strains of Corcyra cephalonica on grain of millet, Panicum sp., and sorghum, Sorghum sp. (both Gramineae), which had been Riested to differing moisture levels from 5-13%. Rearings were separate by strain, foodplant, and level of grain moisture. Mean weight of adult females of both strains ranged from 9.8-21.3 mg between the lowest and highest levels of grain moisture. No statistical tests were reported, but most of the weight differences are probably real. The scatterplot of fecundity vs. adult female weight was not treatment- coded so could not be deconstructed. Congruency of fecundity and maternal size is indeterminate. Relation 54. Final instars of Cnephasia jactatana were subjected to different levels of starvation to create differences in amount of food ingested (Ochieng'- Odero 1990). The author's scatterplot of fecundity vs. weight of female pupae was not treatment-coded, which precluded deconstruction. Congruency of fecundity and maternal weight is indeterminate. Relations 55, 56. Danthanarayana (1975) reared Epiphyas postvittana on four foodplants— curled dock, Rumex crispus L. (Polygonaceae); plantain, Plantago lanceolata L. (Plantaginaceae); capeweed, Arctotheca calendula L. (Asteraceae); and apple, Malus sylvestris (L.) Mill. (Rosaceae). His scatterplots of fecundity vs. female weight were based on separate samples for pupae and adults. Neither scatterplot was diet-coded, which precluded deconstruction, and congruency of fecundity and maternal weight is indeterminate. Relation 57. Torres-Vila et al. (1999) reared cohort subgroups of Lobesia botrana on Vitis sp. (Vitaceae), one on inflorescences, a second on unripe fruit, and a third on ripe fruit. These groups simulated three L. botrana generations associated with the annual march of foodplant phenology. The authors found that both maternal weight and associated fecundity differed significantly by subgroup. The diet-coded scatterplot allowed deconstruction, and slopes and intercepts for inflorescences, unripe fruit, and ripe fruit were 11.3 and 0.0, 15.5 and -15.5, and 23.8 and -35.1 (transcribed data). The slopes, although seemingly disparate, do not differ significantly (F = 1.63; df = 2, 81; P = 0.20), nor do the intercepts (F = 0:34; df = 2, 81; P = 0.71) (transcribed data). Fecundity and maternal weight proved congruent. The three generations are pooled in the summary relation. Grouped-specimen relations The 13 fecundity-maternal pupal weight relations based on grouped specimens number 10 of the perfect kind and 3 of the imperfect, and § multiform. Perfect-uniform and perfect-multiform categories number five each and imperfect-uniform and imperfect-multiform categories number zero and three (Table 2). Grouping damps variation and equally weights groups of differing sample sizes, which biases regression statistics. Presentation of statistics for fecundity-maternal weight relations is therefore limited to slopes and intercepts of summary relations (Table 2) and to cohort subgroup relations, these statistics being useful despite aggregation bias. Meta-analysis. Slopes of the summary relations vary from 0.60 to 2.06 (Table 2), with mean and SE 1.37+0.10. This mean is significantly higher than the 5 uniform, 156 1.13 for individual-specimen relations (Student's ¢ [one- tailed test] = 2.30; df = 69; P = 0.04). Slopes of the perfect-uniform category range 0.60-1.98, averaging 1.32; those of the perfect-multiform, 1.12—1.70, averaging 1.35; and those of the imperfect-multiform, 1.17—2.06, averaging 1.48. Temperature effects. One different rearing temperatures, as discussed below. Relation 13. In one experiment, Woodroffe (1951) apparently reared eee pseudospretella at two temperatures and relative humidities: 25°C-70%, 10°C-70%, and 25°C-20%, but whether larvae were reared or ovipositing adults held under these conditions is not entirely clear. Although Woodroffe reported significant differences in fecundity among some female adult weight classes, I found no differences among adult female weights oe which averaged 26, 22, and 24 mg (F = 0.31: df = 2, 18: P = 0.74) (tabulated data). I tested differences among the three corresponding fecundity-maternal pupal weight regressions whose slopes and intercepts were 12.8 and -7.4; 10.1 and -0.53; and 10.6 and -24.4, and ae slopes did not differ significantly (F = 2.54; df =2, 15; P = 0.89), nor did the intercepts (F = 0.12; df = 2, 15: P = 0.11) (tabulated data). The three presumed rearings are pooled in the summary relation. In a second rearing experiment mentioned only cursorily, fecundity and female adult weight across three temperature-humidity combinations slightly different than above proved congruent. Diet effects. Diet was involved in seven relations (Nos. 6-12 in Table 2), each of which is discussed below. Relation 6. Barah and Sengupta (1991) reared Antheraea assama on four foodplants and reported significant differences in female pupal weight by foodplant. Slopes and intercepts of fecundity- maternal pupal weight relations were 84.0 and -334.3 on Litsaea sp. No. 1, 47.3 and -95.2 on Machilus bombycina King ex Hook, 47.7 and -96.6 on Litsaea sp. No. 2, and 33.6 and -23.9 on Cinnamomum sp. (all Lauraceae), and the differences were highly significant (slope F = 7.52; df = 3, 32; P < 0.001) (transcribed data). Whether suspected tradeoffs between fecundity and egg size existed by foodplant could not be ascertained. Fecundity and maternal weight were congruent. The foodplants. occur together in nature, and data from the four are pooled in the summary relation. Relation 7. Lewitt (1934) field-collected Lymantria dispar pupae from a large area and segregated them into three groups by foodplant damage levels ranging ~5—100%. Lewitt reported that pupal weight decreased with increasing foodplant damage and attributed this to decreasing food availability. Fecundity and female pupal weight across damage levels were congruent. The data relation involved JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY could not be deconstructed for detailed analysis. Relations 8 and 9. Tsai et al. (1958) field-collected Dendrolimus punctatus pupae of the first and wintering generations from Pinus (Pinaceae) and segregated them by three levels of needle damage. Where 50, 80 and 100% of needles were damaged, mean weights of first- generation female pupae were 1210, 940, and 840 mg. Where <50, 50, 80, and 100% of needles were damaged, mean weights of wintering-generation female pupae were 1230, 1680, 1470, and 1490 mg. Although not tested, some differences among the pupal weights of both generations are probably real. Fecundity-matemal weight regressions did not differ among damage levels in either the first generation (slope F = 1.46; df = 2, 34; P = 0.25) or the wintering generation (slope F = 0.03; df = 3, 52: P = 0.99) (tabulated data). However, relations did differ significantly between generations (slope F = 135.2; df = 1, 96; P < 0.001) (tabulated data). Whether suspected tradeoffs between fecundity and egg size occurred could not be ascertained. Fecundity and maternal weight were exactly congruent in the first generation, and approximately so in the wintering generation. A summary relation is tabulated for each generation. Relation 10. Morris and Fulton (1970) reared Hyphantria cunea in different years on different but unspecified diets. Their scatterplot of fecundity vs. female pupal weight was not treatment-coded, which precluded deconstruction for detailed analysis. Congruence of fecundity and maternal weight is indeterminate. Relation 11. Henneberry and Kishaba (1966) reared 12 groups of Trichoplusia ni at 4 densities each with 3 different amounts of artificial diet. Female pupal weights declined significantly with increasing density and decreasing amount of food. Fecundity and maternal weight reaction norms across densities and amounts of food were congruent. The overall relation could not be usefully deconstructed for further analysis. Relation 12. Van Dinther and Goossens (1970) reared Diatraea saccharalis on stalks of rice, Oryza sativa L., and corn, Zea mays L. (both Gramineae), and on differing amounts of various artificial diets. Magnitudes of reported means, standard deviations, and numbers of test individuals suggest significant differences among female pupal weights. Fecundity and maternal weight were approximately congruent. It was not possible to meaningfully deconstruct the summary fecundity- maternal pupal weight regression for further analysis. DISCUSSION AND CONCLUSIONS Exhaustive search yielded 56 more relations, 27 more species, and 8 more families of capital breeders than in VOLUME 59, NUMBER 3 Honek (1993). Meta-analysis Few differences in fecundity-maternal pupal weight relations emerged at the metadata level. Individual- specimen slope means for perfect, imperfect, uniform, and multiform categories of summary relations did not differ statistically. The mean slope of 1.37 for the 13 grouped-specimen relations proved significantly higher than the 1.13 for the 58 individual-specimen relations. Also, the mean slope of 1.22 for Honek's (1993) 15 capital breeders exceeded that for the individual- specimen relations, but not significantly. Aggregation bias probably artificially elevated grouped- specimen mean slopes, including Honek's, who divided each data set into 3-8 grouped- -specimen values. That the 1.13 slope mean statistically exceeds 1.00 may indicate that fecundity is increased or decreased disproportionately by change in maternal weight. This would indirectly intensify the influence on fecundity by an extrinsic factor like rearing temperature. Fully deploy ed meta- -analysis seeks to find a single best estimate of an effect or parameter (Gates 2002), but meta-analysis here is not meant to go beyond minimizing bias in selecting data sets and enabling group and subgroup comparisons. Despite evident similarity of slopes among capital breeders, one fecundity-maternal weight relation does not apply to all. Tradeoffs between Reena and egg size sometimes occur. Also, capital breeders are taxonomically and ecologically diverse, as demonstrated by 15 families represented in this survey. Funnel diagrams of SE, plotted on b for imperfect and perfect groups exhibit much scatter, especially in the imperfect group (Fig. 3a, b), which often can indicate low study precision (Gates 2002). However, adult nutrition potential—present in the imperfect group but absent in the perfect— probably increases variation in fecundity, thereby increasing scatter, and adults of the imperfect group usually had access to fluids. Studies of individual imbibing are few, but in one, Choristoneura fumiferana females imbibed erratically, which undoubtedly increased variation in fecundity (Miller 1989). Individual-specimen midrange pupal weights averaging 63 mg for the imperfect group and 1401 mg for the perfect confirm casual observations that perfect capital breeders are typically larger-bodied than imperfect ones. Extrinsic effects Extrinsic effects on fecundity-maternal pupal weight relations were reflected predominantly in pupal weight and fecundity. In all nine individual-specimen summary relations involving different rearing temperatures (Nos. 24, 25, 29-31, 34, 35, 53 and 58 in Table 1), female pupal weight formed steeple-shaped reaction norms across the temperatures, which fecundity closely tracked, as in Fig. la. Of the six where cohort subgroups could be compared (Nos. 25, 30, 34, 35, 53, and 58). slopes may have been affected in only one (No. 35 for Bombyx mori). Hamilton and Zalucki (1991) showed statistically that in Crocidosema plebejana (Relation No. 58) fecundity was controlled directly by maternal weight and only indirectly by rearing temperature. In all face individual- -specimen summary relations concerning different larval diets where cohort subgroups could be compared (Nos. 26, 32, 33, and 57 in Table 1), female pupal weight and fecundity peaked on the same diet, and only one (No. 33 for Bupalus piniaria) exhibited an effect on slope. That maternal weight directly determines fecundity can be inferred from all of the individual-specimen relations. In the grouped-specimen multiform relations, larval diet was most often the focus, and different diets affected female pupal weight the same as in individual- specimen relations. Although fecundity and maternal weight were congruent in grouped- specimen relations, slopes sometimes differed between cohort subgroups, signaling fecundity-egg size tradeoffs. Different adult diets altered posteclosion ovigenesis in two imperfect capital breeders (Relations 49 and 51 in Table 1), but adult studies were too few to permit Broad generalizations. In a study of maternal weight and fecundity in Choristoneura fumiferana developing on normal, fourth- and fifth-year severely infested foliage—a sequence of declining diet quality— —DMiller (1957) ) found that the slopes did not differ significantly, but that the intercepts and maternal sizes declined in parallel with the declining diet-quality sequence. In a study whose surprising results need confirmation, Carisey and Bauce (2002) found that maternal size in C. fumiferana did not differ among cohort subgroups reared on three artificial diets simulating midcrown, lower crown, and old foliage of Abies balsamea, whereas fecundity and egg size declined in parallel with this sequence of declining diet quality. In several capital-breeder studies not heretofore mentioned, mean fecundity also peaked jointly with mean female pupal weight across rearing temperatures, providing additional examples of congruency. These involved Galleria mellonella (L.), Achroia grisella (F.) (both Pyralidae), Bupalus piniaria (Oldiges 1959), and Lobesia botrana (Torres-Vila 1996). Also, in a supplement to Relations 55 and 56 for Epiphyas postvittana (Table 1), Danthanarayana et al. (1995) reared E. postvittana on four foodplants, including three used in the earlier study, plus an artificial diet, at 158 six constant temperatures from 10.3-32° C. A 6- temperature X 5-diet matrix totaling 30 female pupal weights and fecundities resulted. Maternal weight and fecundity peaked exactly together at five of the six temperatures, and did so approximately at the sixth, and exactly together on three of the five diets, and approximately so on the remaining two. The authors showed that most peak fecundities and maternal weights significantly exceeded nonpeak counterparts. Rearing temperatures in most of the assembled temperature-focused studies were selected arbitrarily, but those for Choristoneura fumiferana (Relation 53 in Table 1) were selected purposefully to compare pupal weights and fecundities at warmer and cooler regimes with those at simulated natural temperatures. The natural regime was presumed to represent the regime of acclimation. Pupal weights and fecundities slumped at 5° warmer but peaked at or near 5° cooler. This occurred in both the short- and long-exposure experiments, which suggests that rapidly accelerating weight increase in the alate fifth and sixth stadia (Eidt aint Cameron 1972) made short exposure virtually equal to long exposure. Thus, brief anomalous warm weather might result in lighter, less fecund females, and brief anomalous cool weather in heavier, more fecund females. More often than not, cooler temperatures produced heavier, more fecund females even though temperatures were selected arbitrarily. Because of its population dynamics interest, this phenomenon should be sought in other capital breeders. Deconstructing some fecundity-maternal pupal weight relations to obtain cohort subgroup regressions for comparison created smaller samples with fewer degrees of freedom, which may have led to false negatives in some tests of slope differences. Possible examples are Choristoneura fumiferana, Lobesia botrana (Relations 53 and 57 in Table 1), and Hofmannophila pseudospretella (Relation 13 in Table 2). However, these few cases do not seriously challenge the conclusion that extrinsic alteration of slopes occurs infrequently, Leather (1988) cautioned against interpreting fecundity-size relations simplistically. His point that potential fecundity does not necessarily translate to field fecundity is unarguable. However, Leather's critique envisaged Lepidoptera’ as a whole — without distinguishing between the divergent life systems of capital and income breeders. Also, he did not realize that extrinsic alteration of maternal size does not necessarily alter underlying fecundity-size relations. With the tally of more than 25 exactly congruent maternal pupal weights and fecundities emerging in this survey, and with no clear counter examples, the direct JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY dependence of fecundity on maternal weight probably warrants the axiomatic status it has been accorded for capital-breeding Lepidoptera. Moreover, extrinsic influences like rearing temperature and larval diet affect the dependence through maternal weight and thus only indirectly. When slopes differed either spatially or among cohort subgroups, as they did in 10 cases, egg size, where reported, also differed, indicating tradeoffs with fecundity. Spatial examples include Choristonewra fumiferana (Harvey 1983) and Malacosoma disstria (Parry et al. 2001). A cohort subgroup example is that of Bupalus piniaria (Gruys 1970). Although not precisely understood, such tradeoffs are probably adaptive. They raise the possibility not only of their wider occurrence among capital breeders but that reproductive bulk, R— the mathematical product of fecundity x mean weight of one egg (Roff 1992)—might constitute a reproductive response as useful as fecundity. The expression R = b(W - R) + aremoves R from W on the right-hand side of the equation so that maternal weight alone remains. This equation would be suitable where fully independent response and explanatory variables are desirable. ACKNOWLEDGMENTS I thank J. L. Stanis for researching Choristoneura fumiferana (Relation 53 in Table 1); R. D. Moon for analytical guidance, stimulating discussions, and useful comments on the manu- script; also M. A. Roberts and an unnamed reader for useful manuscript reviews; J. M. Muggli for laboratory assistance; and J. L. Moe for computer support. For translations, I thank K. Nishida and D. A. Andow (Japanese), S. S. 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Studien zur Okologie, insbesondere zur Bevilkerungslehre der Nonne, Lymantria monocha L. Z. Angew. Entomol. 20: 1-50. Received for publication 23 December 2004, revised and accepted 18 May 2005 VOLUME 59, NUMBER 3 Journal of the Lepidopterists’ Society 59(3), 2005, 161-165 16] PHENOLOGICAL “RACES” OF THE HESPERIA COLORADO COMPLEX (HESPERITDAE) ON THE WEST SLOPE OF THE CALIFORNIA SIERRA NEVADA ARTHUR M. SHAPIRO Center for Population Biology, University of California Davis, California 95616, USA E-mail: amshapiro@ucdavis.edu AND MATTHEW L. FORISTER Dept. of Ecology and Evolution, State University of New York, Stony Brook, N.Y. 11794-5245, USA ABSTRACT. On the west slope of the California Sierra Nevada, the subspecies Hesperia colorado harpalus (formerly generally known as H. comma yosemite) is univoltine, flying in midsummer at mid-elevations. In a number of sites, mostly on serpentine and other unusual soils, a variable but usually slightly darker colorado entity flies at similar elevations in September and October. The two are indistinguishable by male genitalia and thus far by mitochondrial DNA sequences. The autumn “race” is usually parapatric with the summer one and is yuan with it at one known site. The hypothesis that the autumn “race” represents Sierran popula- tions of subspecies tildeni of the orth Coast Range is not consistent with our DNA data. Due to complete temporal isolation, the two “races” of H. comma are functioning as effective biological species. Additional key words: serpentine, allochronic isolation, speciation, phylogeography, mitochondrial DNA. Among the many mechanisms suggested for sympatric speciation, temporal (allochronic) isolation has been posited frequently, but rarely supported on further inquiry (Alexander and Bigelow 1960, Harrison and Bogdanowicz 1995, Huang et al. 2000 but see Feder et al. 1994). Allochronic isolation between close relatives is itself common enough, however, whether or not it is a cause of speciation (Coyne and Orr 2004). Two of the most compelling cases for allochronic speciation - sympatric or not - occur in North American Lycaenid butterflies of the genera Apodemia (Pratt and Ballmer 1991) and Euphilotes (Pratt 1994, Pratt and Emmel 1998). In both genera, allochronic “races” or “biotypes” have arisen within morphospecies, reflecting adaptation to specific hosts in the large Polygonaceous genus Eriogonum. The feeding biology of these insects requires close tracking of host flowering phenology, which varies widely among species. The “races” are frequently sympatric, even over wide areas, but since they are completely isolated they are functioning as effective species. They may have diverged too recently to show significant differentiation at the molecular level (Peterson 1995). Allochrony is also a factor in incipient speciation by “host races” in the Cupressaceous-feeding Lycaenid genus Mitowra (Nice and Shapiro 2001). In this case also, butterfly phenology is most likely a function of host phenology (i.e. the availability of new growth), as host association is a better predictor than altitude of adult flight period in areas where races overlap geographically (though the relationship between host phenology and the preference and performance of butterflies is complex, see Forister 2005). We here report a case of sympatric, allochronic “races” in the Hesperia colorado (Scudder) complex (Hesperiidae) in the Sierra Nevada of California, with no obvious adaptive relation to host phenology. ECOGEOGRAPHY OF THE “RACES” These skippers belong to a circumpolar (Holarctic) complex historically called collectively Hesperia comma (L.). Recent authors have split off the Nearctic members of this complex, except those of the far Northwest, as a separate species, H. colorado (Scudder). The complex was studied phylogeographically by Forister, Fordyce and Shapiro (2004), whose findings broadly support this division. Within the Nearctic range (excluding the far Northwest), these authors found the maximum geographic structure for the mitochondrial gene (COI) they studied to be in California, with two major genetic discontinuities centered around the Sierra Nevada and Transverse Ranges. The subspecific nomenclature of the populations at issue is very confused. Historically, east-slope Sierra Nevada populations were placed in subspecies harpalus (W.H. Edwards) and west-slope ones, which are phenotypically quite different, in subspecies yosemite Leussler. Scott (1998) found that the conventional usages were incorrect and that these two names are in fact synonyms and harpalus is the correct name for the west-slope entity, while the correct name of the east- slope one is H .c. idaho (W.H. Edwards). We review this taxonomy because in using the subspecies as redefined by Scott in this paper, we risk causing confusion in the context of virtually all prior publications on the group. The name applied to the Inner North Coast Range subspecies (tildeni H.A. Freeman) is unaffected. Both west-slope harpalus and east-slope idaho are univoltine (as is the entire complex) and fly in early-to- midsummer. Coast Range tildeni fly later, typically from August through October. In the early 1970s one of us (AMS) discovered a population near 1500m in Nevada County, CA on the Sierran west slope that flew only in September and October. AMS continued to monitor this population annually. In 1988 it was included in a permanent monitoring site on his altitudinal transect across California and has thus been visited approximately biweekly except in winter since then. Its autumnal flight period has remained constant throughout. It is limited to a serpentine barren with sparse vegetation, much bare rock and only one nectar source, a distinctive dwarfed ecotype of rabbitbrush, Chrysothamnus nauseosus (Pallas) Britton (Asteraceae), during its flight season. Populations of summer-flying H .c. harpalus occur in the same canyon at 850m and 1525m_ on metasedimentary substrates (phyllite and complex schists) at distances of 4 and 11 km respectively, as well as at 2100m on an andesitic mudflow and granodiorite, 40.5 km away. The phenology of these three Washington 8 "| serpentine i non-serpentine 4 number of years observed - T July 1st August 1st September 1st Fic. 1. Histogram showing the phenology of the two H. colorado races at Washington (Nevada Co.) since 1988. Vertical bars correspond to five day increments (the beginning of July, August, and September are shown for reference). For example, there were five years in which non-serpentine H. colorado individuals were observed at this site during the first five days of August. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY populations (identified as Washington, Lang Crossing, and Donner Pass) is compared with the autumnal one (identified as Washington Serpentine) in Figs. 1 and 2. The serpentine population (Washington serpentine) is later-flying than the 850m Washington population (a) Lang Crossing number of years observed July 1st August 1st September 1st Donner Pass number of years observed July 1st August 1st September 1st Fic. 2. (a) Histogram showing the number of times that H. colorado individuals have been observed throughout the year at Lang Crossing (Nevada Co.) since 1974, and (b) at Donner Pass (Nevada Co.) since 1973. As in Fig. 1, time on the x-axis is expressed in vertical bars indicating observations made within five day intervals. (there is no overlap), and only overlaps with the 1525m Lang Crossing population by a few days. Individuals at the 2100m Donner Pass population have been observed as late into the fall as at the serpentine population, but this is clearly a phenological effect of high-elevation conditions (there is only one brood at Donner Pass). Once alerted to the presence of an autumnal race on unusual soils, AMS and his associates began looking for additional autumn-flying populations, primarily on serpentine. Gervais and Shapiro (1999) reviewed the VOLUME 59, NUMBER 3 distributions of edaphic-endemic butterflies in the Sierra Nevada, including the autumn “race” of H. colorado (there called comma). They reported populations on serpentine and gabbro soils in Nevada, Placer and El Dorado Counties on the west slope, as well as one on limestone in Calaveras County and one on an undetermined substrate in Mariposa County (reported to us by Oakley Shields). There are undoubtedly more populations to be found, and the association with unusual soils may be more apparent than real since we mainly looked for them on such substrates. The populations we identified all appear to be parapatric with summer harpalus on “normal” substrates nearby, except one (Drum Powerhouse Road, Placer County, in the Bear River drainage) in which 163 they are actually sympatric, but allochronic, in an area where serpentine and metasedimentary rocks are intimately interdigitated, and some of the serpentine is unusually mesic. We have been unable to find any male genitalic differences between these “races” (C.D. MacNeill, pers. comm.). The phenotypes of the autumn “race” are about as variable as summer harpalus, though on average slightly darker with a more defined ventral hind wing pattern. Fig. 3 illustrates the phenotypes. We have not identified any consistent character in ae sex which can be used to identify flight date correctly. should be noted that some individuals of the sae “race” rather closely resemble the apparent hybrid swarm between east- and west-slope subspecies in the Fic. 3. Photographs of H. colorado specimens (showing left dorsal and ventral oe (a) seven females and four males, serpentine “race” from Washington, (b) three males, non- serpentine “race” ’ from Washington, (c) three males and three females, from Lang Crossing. 164 Feather River Canyon farther north in the Sierra. These animals fly very Molecular-genetic evidence bearing on the identity of early, in May and June.) the autumn “races” is discussed below. DISCUSSION An endemic late-season serpentine subspecies of Hesperia colorado, described from the Siskiyou mountains of far northern California, was named mattoonorum by McGuire (1998). This entity is darker on average than nearby non-serpentine populations, but is also v. aviable: Those near by populations are extremely variable and confusing. Scott (1998) synonymized the usual name applied to them, oregonia (W.H. Edwards), with harpalus as well. In our judgment his fixation of the type locality of oregonia as “Sierra Nevada Mts., west of Carson City?” is unjustified by the historical and biological facts. In any case, Shapiro, Palm and Weislo (1981) recorded rildeni-type foothill H. colorado in the Trinity Alps from 7-29 August; “light” oregonia (from rain-shadow areas) from 16 August - 20 September, and “dark” oregonia (from wetter areas) from 12 July through 25 September. The very long type series of mattoonorum was collected in go ee Shapiro (1991) illustrated a hypervariable population series from a serpentine site in the nearby Trinity Divide, collected 5 September. Because of the very extended flight seasons of far northem California colorado, it is difficult to assess the significance of the late flight season of mattoonorum. No mattoonorum were available for molecular-genetic study, but several specimens from the upper foothills of the Trinity Alps above French Gulch were used. When Gervais and Shapiro (1999) discussed the geography of edaphic-endemic Sierran butterflies, they proposed the hypothesis that the autumn “races” of H. colorado were actually populations of the late-season- flying Inner Coast Range subspecies tildeni, signifying a double invasion of the Sierra. This was suggested by the fact that most of the edaphic-endemic entities they found were much more widespread in the Coast Range (and had been largely or entirely overlooked in the Sierra) and that their ranges in the Sierra, including autumn colorado, were largely concordant, suggesting a common history. ( Forister, Fordy ce and Shapiro (2004) found for a portion of the COI mitochondrial gene, coast Range tildeni possessed a unique haplotype (D) not found in far-northern California and Oregon or in the Sierra Nevada (see Fig. 1 in Forister et al. (2004) for the distribution of “haplotypes in the Western United States). The Sierran east and west slopes shared no haplotypes, except in the thoroughly mixed Feather JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY River hybrid population. They sequenced COI from 5 M ashington serpentine animals (Nevada County), all of which were haplotype A, the most common haplotype on the Sierran west slope and in Oregon. Haplotype A and is two mutational steps removed from the tildeni haplotype D does not occur in Coast Range tildeni, in the statistical parsimony network described by Forister et al. (2004). Three Lang Crossing harpalus were A, two B; one summer harpalus from Drum Powerhouse Road was A (B is one mutational step from A). The serpentine series seems unusually invariant, but cannot be distinguished from near by summer harpalus. On the other hand, the hypothesis that it is really tildeni is not supported by our data since haplotype D was not found. Two individuals from the foothills of the Trinity Alps were haplotype B, one I (B is connected to I by one mutational event, and I is two steps removed from A). Forister et al. (2004) found no geographically patterned variation for the nuclear gene wingless in North America south of British Columbia. We are thus unable at this time to identify any genetic discontinuity between summer harpalus and the autumn “race” in the Sierra Nevada. It is possible that a survey of nuclear variation encompassing a larger portion of the genome but we have been unwilling to sample these mostly very small populations to the extent needed for statistical rigor. Unlike the Eriogonum-feeding Lycaenids with phenological “races,” there is no obvious connection between the seasonality of these Hesperia and their presumed host plants, perennial bunchgrasses. We do not know the hosts used by them at any of the sites mentioned except Lang Crossing, where summer harpalus routinely uses the naturalized European ¢g grass Anthoxanthum odoratum L. (Poaceae). This grass does not occur at any of our other sites in the region. Hesperia colorado overwinters as an egg, and larvae feed on young growth in the spring. The flight season of the adults thus appears decoupled from any need to match host phenology, although it could be related to the ability of the eggs to tolerate late-summer desiccation prior to the onset of winter. Until appropriate experiments are done, we will be unable to state whether the autumn “races” are genetically programmed to emerge at that season or are somehow induced physiologically to do so as a result of the nutritional properties of their hosts — which in turn could be affected by occurrence on serpentine vs. non- serpentine substrates. We do not know if the northern entity mattoonorum is in any way connected with the Sierran serpentine populations, or whether autumn in the Sierra are all derived from a single would reveal such differences, “races” VOLUME 59, NUMBER 3 ancestor or have arisen repeatedly in various locations from local summer harpalus, in the manner of many plant ecotypes which are generated over and over again. Our nearest sample to mattoonorum, from French Gulch (Trinity County), is not that entity and is thus not informative. CONCLUSIONS 1. Local autumn-flying “races” of Hesperia colorado occur within the range of summer-flying subspecies harpalus on the west slope of the Sierra Nevada. Most of these known to date are on serpentine or other unusual soils. 2. These autumn “races” average a little darker than nearby harpalus but have no definitive wing, genitalic, or molecular characters to allow them to be identified without collection dates. 3. Despite similar seasonality and biogeographic arguments, molecular phylogeography has not supported the hypothesis that the Sierran autumn “races” are actually the Inner Coast Range subspecies tildeni. 4. Further study is needed to distinguish between genetic and environmental/physiological factors as determinants of the aberrant phenology of these “races.” ACKNOWLEDGEMENTS We thank C.D. MacNeill for reviewing specimens for taxo- nomic purposes, H.B. Shaffer for the use of his laboratory, and J-A. Fordyce for help in the phylogeographic study. The phylo- geography was conducted under National Science Foundation Fellowship DGE-0202740 to MLF as well as grants from the Graduate Group in Ecology, Center for Biosystematics and Cen- ter for Population Biology at U.C. Davis. LITERATURE CITED ALEXANDER, R.D. & R. S. BIGELOW. 1960. Allochronic speciation in field crickets, and a new species Acheta veletis. Evolution, 14: 334-346. Coyne, J.A. & H. A. ORR. 2004. Speciation. Sinauer Associates, Sun- derland, Mass. FEDER, J.L., S.B. Opp, B. WLAZIO, K. REYNOLDS, W. Go & S. SPIsAK. 1994. Host fidelity is an effective premating barrier between sympatric races of the apple maggot fly. Proc. Natl. Acad. Sci. USA, 91: 7990-7994. ForisTerR, M. L. 2005. The influence of host-plant phenology on Mi- toura nelsoni Boisduval (Lepidoptera: Lycaenidae). Ann. Ento- mol. Soc. Am., 98: 295-301 ForisTeR, M.L., J.A. ForpycE & A.M. SHAPIRO. 2004. Geological barriers and restricted gene flow in the Holarctic skipper Hespe- ria comma (Hesperiidae). 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Population genetic evidence of re- stricted gene flow between host races in the butterfly genus Mi- toura (Lycaenidae). Ann. Entomol. Soc. Amer., 94: 257-267. PETERSON, M.A. 1995. Phenological isolation, gene flow and develop- mental differences among low- and high-elevation populations of Euphilotes enoptes (Lepidoptera: Lycaenidae). Evolution, 49: 446-455, Pratt, G.F. 1994. Evolution of Euphilotes (Lepidoptera: Lycaenidae) by seasonal and host shifts. Biol. J. Linn. Soc., 51: 387-416. Pratt, G.F. & G.R. BALMER. 1991. Three biotypes of Apodemia mormo (Riodinidae) in the Mojave Desert. J. Lepid. Soc., 45: 46-57. Pratt, G.F. & J.F. EMMEL. 1998. Revision of the Euphilotes enoptes and E. battoides complexes (Lycaenidae). pp. 207-270 in T.C. Emmel, ed., Systematics of Western North American Butterflies. Mariposa Press, Gainesville, Fla. Scott, J.A. 1998. New western North American butterflies. Papilio (N.S.), 11: 1-12. SHapirO, A.M. 1991. Significant additions to the butterflies of the Trinity Alps and Mount Eddy, Northern California. J. Res. Lepid., 29: 33-36. SHapiro, A.M., C.A. PALM & K.L. WcIsLo. 1981. The ecology and biogeography of the butterflies of the Trinity Alps and Mount Eddy, Northern California. J. Res. Lepid., 18: 69-152. 166 Journal of the Lepidopterists’ Society 59(3), 2005, 166-169 HESPERIA LINDSEYI MCCORKLEI ( HESPERIDAE JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY ): A NEW SUBSPECIES OF SKIPPER FROM SOUTHWESTERN OREGON, USA PAUL M. SEVERNS Department of Botany and Plant Pathology, 2082 Cordley Hall, Oregon State University, Corvallis, OR 97331. Email: severnsp@onid.orst.e du AND DONALD G. SEVERNS 2526 N. 21st, Springfield, OR 97477 ABSTRACT. A new subspecies of Hesperia lindseyi Holland is described from southwestern Oregon, United States. This sub- species, Hesperia lindseyi mecorklei, is the northernmost known taxon of the Hesperia lindseyi group, which is mostly distributed throughout Californian savanna and chaparral plant communities. H. /. mecorklei presently appears to be isolated to the Jackson Creek and South Umpqua River drainages in Douglas County, Oregon. Additional key words: skippers, oak savanna, chaparral Hesperia lindseyi (Holland, 1930) Hesperiidae is a taxon that occupies various grasslands, chaparral, and savanna habitats of California, with outlying populations in southern Oregon and northwestern Nevada (MacNeill 1964, Dornfeld 1980, Hinchliff 1994, Pyle 2002). Recently, Emmel et al. (1998) designated three new subspecies of Hesperia lindseyi geographically distinct phenotypes that were originally delimited by MacNeill (1964). Hesperia lindseyi macneilli Emmel, Emmel and Mattoon, 1998 occurs from coastal southern Marin Co., California north to southern Humbolt County and was considered the darkest of the H. lindseyi subspecies. H. 1. eldorado Emmel, Emmel, and Mattoon, 1998 inhabits the western foothills of the Sierra Nevada from its most based on southern distribution in E] Dorado Co., California north to Butte County. The third subspecific taxon described, H. 1. septentrionalis Emmel, Emmel and Mattoon, 1998 occupies the northernmost latitudes of the three H. lindseyi subspecies. It occurs from low to middle the southwestern Oregon, elevations in Siskiyou Mountains in northern California and and extends Basin of south-central Oregon into northwestern Nevada. eastward across the Klamath The immature stages of Hesperia lindseyi consume grasses, primarily Festuca idahoensis and Danthonia californica Boland Poaceae (MacNeill 1964, 1975). These grasses are generally shade intolerant and are commonly encountered in drier, mesic grasslands, scrub, savanna and chaparral plant communities (Hitchcock 1970). In Oregon, Hesperia lindseyi septentrionalis, is confined to the southern portion of the state. It is commonly taken by collectors in oak (Quercus kellogii Newb. and Q. Fagaceae) savanna remnants near the Oregon/California border, in garryana Dougl. ex Hook. and near the town of Klamath Falls, and occasionally in the Warner Mountains of Lake County (Warren 2005). On 6 June 1996 the northernmost known population of H. lindseyi was discovered by Donald G. Severns in southern Douglas County, Oregon, approximately 12 km east of the town of Tiller along Jackson Creek in the South Umpqua River drainage. This population of H. lindseyi is approximately 70 km north of the nearest known populations of H. lindseyi septentrionalis, is phenotypically distinct, and exists in an isolated remnant of oak savanna surrounded by Douglas fir (Psewdotsuga menziesii Mirbel Franco Pinaceae) forests. In the following pages we describe a new subspecies of Hesperia lindseyi, comment on its life history, and discuss the geographic bounds for the new taxon. Hesperia lindseyi mccorklei Severns and Severns, new subspecies Description. Male. Mean forewing width 11.8 mm, ranging from 10.5 to 13.0 mm (n= 38). Dorsal surface (Fig. 1): Forewing bright orange and slightly iridescent; outer black margin extends into inner two thirds of forewing, often surrounding and defining the apical and subterminal spots; post-stigmal patch appears larger and darker when compared to all other Hesperia lindsey. Fringe is brownish-buff with the vein terminals marked by the presence of iin scales. Hindwing ground color same as forewing; black hindwing margins extend to distal end of the macular arm band. Lighter orange areas correspond with macular band spotting, which te sal to contrast with the ground color. Fringe unbroken throughout the hindwing margin, ranging from cream to buff. Ventral (Fig. 1): Pale orange-brown ground color becoming inwardly buff, abruptly transitioning to black near thorax; apical and subterminal spots surrounded by a field of golden-green scales with intermittent melanic scales cov ering approximately one third of distal forewing. Apical and subterminal spots cream with a slight greenish tinge, area below stigma is black. A black terminal line runs the length of the forewing separating the fringe from the rest of the wing; vein terminals are marked with enlarged areas of black scales extending from terminal line to fringe edge. Hindwing ground color same as area surrounding the