ISSN 0024-0966 Journal of the Lepidopterists Society Published quarterly by The Lepidopterists’ Society THE LEPIDOPTERISTS’ SOCIETY EXECUTIVE COUNCIL James K. Apams, President Nikias Wauiperc, Vice President Susan J. Wetter, Immediate Past President Ernest H. WittiaMs, Secretary Gary G. ANWEILER, Vice President Ketiy M. Ricuers, Treasurer Marc Epstein, Vice President Members at large: William E. Conner Akito Kawahara Robert M. Pyle Rebecca Simmons Jane M. Ruffin John A. Shuey Charles V. Covell Jr. Erik B. Runquist Andrew D. Warren EprrortaL Boarp Joun W. Brown (Chairman) Micuaen E. Touiver ( Journal) Lawrence F. Gait (Memoirs) Priv J. SCHAPPERT (News) Joun A. Snyper (Website) Caria M. Penz (at large) Honorary Lire MEMBERS OF THE SOCIETY Cuarces L. Remincton (1966), E. G. Munroe (1973), [An F. B. Common (1987), Lincoin 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, > 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 Museum, 900 Exposition Blvd., Los Angeles, CA 90007-4057. Cover illustration: Idea leuconoe (Large Tree Nymph) mating. Photo taken by Carol Butler at the American Museum of Natural History butterfly conservatory. VOLUME 59, NUMBER 4 18] JoUuRNAL OF Tue Lepstsorrerists’ Socrery Volume 59 JAN 0 2006 20 Number 4 Journal of the Lepidopterists’ Society 59(4), 2005, 181-199 LIBRARIES A REVIEW OF CALLOPHRYS AFFINIS (W. H. EDWARDS), WITH DESCRIPTIONS OF TWO NEW SUBSPECIES FROM NEW MEXICO AND MEXICO GLENN A. GORELICK Dept. of Biological Sciences, Citrus College, Glendora, CA 9174 email: ggorelick@citruscollege.edu ABSTRACT: The subspecies of Callophrys affinis (W.H. Edwards) are discussed, with emphasis on biosystematics, ecology and distribution. C. apama (W.H. Edwards) is treated as a subspecies and two new subspecies are figured and described from New Mex- ico and Mexico. Additional key words: allopatric, sympatric, submesial band (=macular band; postmedian line), ecotype The holarctic genus Callophrys Billberg, thecline lycaenids recognized by the absence of a tail and green scaling on the undersides of the wings, is widely represented i in western North America by the C. affinis (W. H. Edwards) and C. sheridanii (W. H. Edwards) complexes. Species of the C. sheridanii complex are characterized by a complete or nearly complete submesial band of predominantly white-scaled maculations on the hindwing undersides. This complex includes nominate C. sheridanii (W.H. Edwards), C lemberti Tilden, and C. comstocki Henne. Members of the C. affinis complex, on the other hand, are recognized by significant intrapopulation variation in the shape and relative completeness of the submesial pe on the ventral hindwings. This complex includes C. affinis (W.H. Edwards): ! apama (W.H. Edwards), treated here as a subspecies of C. affinis; C. dumetorum (Boisduval), synonymous with C. viridis (H. Edw.) (Emmel et al 1998): and C. perplexa (Barnes & Benjamin), formerly C. dumetorum (Bdy.) (Emmel et al 1998). Gorelick (1971) treated the members of these complexes as a superspecies (as defined by Mayr 1965) although the study described herein suggests otherwise. Populations of Callophrys affinis (W. H. Edwards) occur throughout western North America as shown by Stanford & Opler (1993). Life histories of the described subspecies were published by Scott (1986). They typically occur in the Transition zone, including steppe habitats. The localities of adult specimens used in this study are depicted in Figure 1. These butterflies are on the wing between April and September, typically in mountain or steppe habitats between 1372—2500m throughout the species range. Like other Callophrys (s str.), some variation may exist in the iridescent color of the green scaling on the ventral surfaces of the hindwings. This variation relies on light interaction with detailed scale architecture to yield the green color. rather than pigment (Ghiradella 1989). The function of this appears to be thermoregulatory (Gorelick 1971). The most readily chosen diagnostic characteristic examined in the C. affinis complex has typically been the number and arrangement of the white maculations composing the submesial band (Clench 1944, Tilden 1963, Ferris 1971a). Owing to substantial adult variation (intrapopulation, ecotypic and clinal), the most revealing taxonomic studies of Callophrys also include life cycle observations, flight period(s), mating and oviposition behavior, as well as habitat and hostplant descriptions (Gorelick 1971, Emmel et al 1998). First instar and mature larvae of Callophrys taxa and other closely related thecline lycaenids are described and figured by Ballmer and Pratt (1988, 1992). Biological and systematic studies of Callophrys species are limited in number, perhaps due to the dearth of exploration in geographically isolated mountain ranges in both the southwestern United States and northern Mexico. Since the 1960s, searches in these areas have resulted in new Callophrys distribution records (Clench 1965, Gorelick 1971, Ferris and Brown, 1981, Mueller 1982, Cary and Holland 1992, Brown et al 1992, Hinchliff 1994, 1996). Bailowitz and Brock (1991) mention the occurrence of . affinis (W.H. Edwards) . apama (W.H. Edwards) . chapmani Gorelick . albipalpus Gorelick . washingtonia Clench echROBPseO qanaaaann ppppp p > p Fic. 1. Distribution of Callophrys affinis (W. H. Edwards) C. affinis apama in the Santa Catalina Mountains of southeastern Arizona with respect to this apparent isolation. Such isolation is also seen in such areas as the La Sal Mountains of eastern Utah, shrub-laden arroyos and prairie breaks of Huerfano, Lincoln/Washington and Crowley counties in eastern Colorado (Stanford 1994), southwestern Nebraska, the Sacramento Mountains of southern New Mexico, and the Sierra Madre Occidental of northern Adult populations of this species were found to be relatively abundant with considerable variation in the northern and central portions of its range, but scarcity and relative loss of variation were observed in the southernmost montane portion of its range in northern Mexico. Mexico. SYSTEMATICS First in the C. affinis species group to be described was Thecla dumetorum, named by Boisduval (1852). He believed C. dumetorum to be a local race of the European species, T. rubi L. (Gorelick 1971). The taxa Thecla affinis and Thecla viridis were described by William Henry Edwards in 1862. In 1882, William Henry Edwards described Thecla apama on the basis of white maculations of the ventral hindwings (VHW) lined with a reddish brown band. F.M. Brown (1970) designated a lectotype for this species and referred to . affinis/homoperplexa blend apama/homoperplexa blend . homoperplexa Barnes and Benjamin JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY ears # sae Rar Sear Ror av ae 7 1 ore Se ry pel SU Hotta psig |, | gee ee Het He five specimens deposited in the AMNH that are probably paratypes. After incorporation into the genus Callophrys, a second subspecies, C. apama homoperplexa, was named by Barnes and Benjamin (1923) from Colorado on the basis of significant "reduction of white lines and their corresponding black and brown bands, the spots being usually quite disconnected." Barnes and Benjamin correctly pointed out that the total absence of these white spots or maculations on the wing undersides in females renders them quite similar to Callophrys dumetorum perplexa from San Diego, described by them in 1923. Unlike other C. affinis complex members, females of C. perplexa are distinct in their dorsally brown to golden brown wing scaling and grass green to golden green- scaled ventral hindwings. They also implied the necessity of examining a_ significant number of specimens from each locality to best insure proper identification of C. apama subspecies due to the existence of "intermediates to typical C. apama." Haskin and Grinnell (1912) treated Thecla dumetorum (Bdv.), T. viridis (W. H. Edw.), & T. affinis (W. H. Edw.) as synonyms. Succeeding publications list these species as separate taxa (Klots 1951, dos Passos 1964, Howe 1975, Miller & Brown 1981). In his study on the genus Callophrys in North America, Clench (1944)) presented a key to the species wherein he separated C. apama from C. perplexa (formerly C. dumetorum) (Emmel et al 1998) on the basis of the VOLUME 59, NUMBER 4 convexity of the outer of the forewing. Subspecies were principally identified by use of these white maculations on the hindwing undersides, and a new taxon, C. affinis washingtonia, was described from Washington. More recently, Pyle (2002) treats C. perplexa and C. affinis as separate taxa, as do Brock & Kaufmann (2003), and Opler & Warren (2003). The variability of Callophrys affinis adults is well described by Ferris and Brown (1981), who treated C. apama as a_ separate species. The diagnostic characteristics and intrapopulation variation discussed sufficiently separate C. apama from closely allied C. affinis. They also described the appearance of the submesial band of white spots on the ventral hindwing (VHW) in some Rocky Mountains specimens as varying from a nearly continuous irregular line to those in which such white spots are caitarellly absent. This description applies to specimens from large populations that were studied from Boulder and Jefferson counties, Colorado, which are treated here as C. affinis homoperplexa. Using ventral hindwing features, _ phenotypic intermediates between C. affinis affinis and C. affinis homoperplexa in Wyoming, Colorado and Nebraska suggest intergradation. Recognizing this intergradation, Scott (1986) was the first to treat both homoperplexa and apama as subspecies of C. affinis. In their treatment of the butterflies of New Mexico, Toliver et al (1994), Cary and Holland (1992), Holland and Cary (1996) list C. apama from the Jemez and Sangre de Cristo ranges of northern New Mexico as C. apama homoperplexa. This was confirmed by eight specimens collected by the author in late May and early June of 1996. Nevertheless, all four specimens reared from four second generation larvae taken from the Jemez Mountains emerged in August 1997 with a nearly complete submesial band more typical of C. affinis apama (Fig. 2), also suggesting intergradation. Two paratypes of “Thecla apama homoperplexa” also have a nearly complete submesial band with abundant brown scaling on the mesial side of the white maculations on the V HW, like that shown in figure 3. margin MATERIALS AND METHODS Ecological and behavioral observations were conducted in the state of Durango (Mexico), and in selected localities in Coconino Co., Arizona, Lincoln Co., New Mexico, Albany Co., Wyoming and Cheyenne Co., Nebraska, between 1981 and 1997 to gather comparative biosystematic information. On these trips, adults, immature stages, larval hostplants, and _ soil samples were collected. Field notes and photos were taken, including habitat descriptions and hostplant selections, as well as mating and ovipositional behaviors. Live immature stages were transported in an ice chest by automobile and live females were shipped by overnight express within twelve hours of their capture to Dr. John Emmel, Hemet, California. In this manner, newly-eclosed adults emerged from ova obtained by oviposition in a lab setting using closely related hostplants. In addition to the rearing of immature stages, 12 C. a. apama ova and first instar larvae were examined using a stereoscopic microscope. No fewer than 12 first instar larvae of C. a. apama were collected and preserved in a larval fixative solution, along with a similar number of mature larvae. These and the field-collected adults are currently housed in the author's personal collection. In most cases, observations were made on flights of a minimum of 12 individual adults at a given roadside or trailside locality, mostly during 2nd generation flight periods. Such observations of C. affinis were conducted on both sides of the Continental Divide. On the west side of the Mogollon Plateau, a portion of the Colorado Plateau characterized by Hubbard (1965), C. affinis populations were studied at four localities: base of San Francisco Peaks, north of Flagstaff in north-central Arizona, 2287m; the vicinity of Rose Peak, Apache National Forest in southeastern Arizona, 2287—2439m: the Zuni Mountains of western New Mexico, 2287-2317m: and the Sierra Madre Occidental west of Durango, Mexico, 2287-2439m. On the east side, studies were conducted on isolated limestone outcrops with silt-laden sandstone layers and in cherty limestone/volcanic ash arroyos that occur in northeastern Colorado and southwestern Nebraska, 1372-1524m, and in the Sacramento Mountains of south-central New Mexico, 2287-2317m. Using both field-collected and reared adult specimens, selected body and wing characteristics were compared. The characteristics chosen for comparison generated the nominal data entered in Table 1. RESULTS Ova are disc-shaped with abundant, variably shaped trabeculae. Green in color, the ova were found to be similar in size and texture to those described for Callophrys perplexa (=dumetorum) and C. dumetorum (=viridis) (Gorelick 1971, Emmel et al 1998) and for C. sheridanii (Ferris 1973). No changes in color occurred up to the day of eclosion. Intrapopulation variation in the submesial band of ventral hindwing maculations was found to conform to the following generalizations along a north to south gradient: 1. Specimens taken in northern Colorado (Larimer, Veld counties), southwestern Nebraska (Banner and 184 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TABLE 1: Characters used in the analysis of the Callophrys affinis complex in the western United States and Mexico CHARACTER Subspecies of Callophrys affinis: apama chapmani apamax — homoperplexa albipalpus affinis x washingtonia affinis homoperplexa homoperplexa 1. VHW vein tips: outer-brown; outer-brown; uniform uniform outer-brown; uniform light uniform pale uniform pale to M3-2A mesial-black; mesial-dark brown brown inner-white brown to light light brown inner-white — brown; inner- (very weak) brown white 2.VHW margin- prominent prominent — intervenous white line white line intervenous intervenous intervenous white females intervenous intervenous white line typically thin; weakly white line white line line absent white line; white line; outside green absent developed — absent absent outside green outside green scaling, Cul scaling, M3 scaling, Cul to 2A to 2A to 2A 3. Wing fringes: basally gray _ basally light basally dark basally brown; basally basally light basally light _ basally brown; VEW brown; brown; brown; distally distally pale — brown; brown; brown; distally pale to mesially light mesially gray pale distally pale distally pale distally white white brown; brown; distally pale- distally pale tipped VHW: M1-Cu2 basally basally basally brown; basally brown; basally basally light basally light _ basally brown; brown; brown; distally pale distally pale to brown; brown; brown; distally pale to mesially dark mesially gray white distally pale distally pale — distally pale white brown; brown; to white or distally pale distally mixed to white mixed pale VHW: 2A-3A — distally dark distally basally brown; basally brown; basally basally light basally light basally light brown; brown; mesially mesially brown; brown; brown; brown; mesially mesially mesially white; distally white; distally mesially mesially white mesially white; distally light white; white; dark brown = dark brown — white (prominent); white; brown distally dark distally dark (reduced); distally dark distally brown; brown; distally dark brown mixed elongated elongated brown white scales white scales at vein tips at vein tips 4. FW apex obtuse- obtuse- obtuse-angled obtuse-angled obtuse- slightly nearly nearly pointed angled angled angled obtuse-angled pointed 5. FW outer slightly asinapama asinapama asinapama as inapama_ as inapama nearly right- nearly right- margin: male crenated angled angled between Cul & M3 female slightly slightly slightly slightly slightly nearly right- nearly right- nearly right- rounded rounded rounded rounded rounded angled angled angled 6. VFW costa pale brown — brown to light brown light brown light brown light brown __ gray grayish to yellowish, yellowish, concolorous concolorous 7. VFW coloration light brown light brown _ light brown to typically grayish to grayish to M2 gray to Cu2; gray from inner inner margin to Cu2; to Cu2; Cu2; brown grayish to Cu2; light mostly green margin to Cu2; brown above brown above above Cu2; light brown above mostly green brown above | 8. Antennal annuli 15-16 16-17 16-17 17-18 16-17 16-17 17 mostly 17 white 9. Facial hair tuft typically bent asin apama asinapama asinapama asin apama_ sparse; bent sparse; bent _ sparse; prostrate forward; forward forward thicker laterally scale color mixed gray as in apama__ light gray light gray light gray to light gray white to gray white to gray and brown white 10. FW stigma- lighter than lighter than _ typically typically lighter than _ typically typically typically male ground color ground color concolorous concolorous ground color concolorous concolorous concolorous VOLUME 59, NUMBER 4 TABLE 1: Continued 185 scaling male CHARACTER Subspecies of Callophrys affinis: apama chapmani apamax homoperplexa albipalpus affinis x washingtonia affinis homoperplexa homoperplexa 11. Labial palp black black black dorsally; black dorsally; black some black; — mixed black thin; mixed black dorsally; dorsally; white laterally white laterally dorsally; mostly white and white and gray white white white laterally laterally laterally ventral setae thick; black; thick: black: thick; black; — thick; black; black; black; mixed black; mixed black: mixed with abundant abundant abundant abundant abundant with white with white white scales scales scales distal segment black; white- typically black; white- _ typically typically mostly black; white; white- mixed black and tipped tricolored: tipped black; white- white; white- white-tipped tipped white black with 4- tipped tipped 5 brown scales ventrally; white-tipped 12. Green scaling- bright typically olive 1st bright green Ist slightly light green _ pale to yellow VHW (Kelly & (emerald) green in generation: generation: — yellowish green Judd, 1976) green males; strong bright green moderate to green (dark) green 2nd bright green infemales generation: 2nd bright to generation: yellow green bright green 13. Maculations complete complete nearly typially Ist variable; both typically one typically absent or VHW band: band & complete; incomplete; generation: — black and spot in cell one spot in cell maculations tricolored as maculations in variable; absent in brown scaling Cu2; brown Cu2; nearly in Cul & in apama; M1-M3 may mesial black females 2nd absent in scaling complete-M2 to Cu2 mesial brown be absent; scaling weak generation: males; brown absent; black 3A in some; black displaced scaling also black scaling _ or absent typically scaling absent scaling scaling reduced to laterally; forms a weak (12 or complete or in females greatly fewer than 12 tricolored- contiguous _ fewer scales) nearly reduced to scales or absent white, black, band complete - fewer than and brown M2 to 3A 12 scales or only absent 2nd generation (males): inner brown scales reduced to 8 or fewer; black scaling absent VFW 4-5 4-5 3-4 weakly 4-5 weakly 4 weakly 4orfewer; 4orfewer; 2 or fewer; greatly maculations maculations developed or developed or developed or greatly greatly reduced or absent prominent weakly black absent; absent; absent; reduced or reduced or white; black mesially contrasting contrasting contrasting —_ absent absent mesially brown scaling brown scaling brown mesially mesially scaling mesially 14. Dorsal features grayish to grayish brown to brown to brown to orange brown grayish grayish brown brown orange brown orange brown orange brown brown female orange asindpama asinapama asinapama asinapama_ as in apama uniform uniform orange brown with orange brown dark brown brown margins 186 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 2— 10: Callophrys affinis subspecies from the U. S. and Mexico. 2. C. affinis apama x homoperplexa blend, 4, C. affinis homoperplexa, 3, Boulder Co., CO. 4: C. affinis homoperplexa, 3, Jefferson Co., CO. 5: C. affinis affinis x homoperplexa blend, Albany Co., WY. 6. C affinis apama, 3, Coconino, Co., AZ. 7. C. affinis affinis, 3, Box Elder Co., UT. 8: C. affinis washingtonia, 6, Goose Lake, British Columbia, Canada 9: C. affinis albipalpus, n. ssp. holotype ¢, Lincoln Co., NM. 10. C. affinis chapmani, n. ssp. allotype °, Durango, Mexico VOLUME 59, NUMBER 4 Cheyenne counties) and southeastern Wyoming (Laramie Range, Albany County) showed wide variation in the submesial band. Most males examined (31 of 38, §1.6%) exhibited up to six maculations with both brown and black scaling, while 7 (18.4 %) females exhibited a complete band that lacked brown scaling. In the Laramie Range, specimens treated as C. a. homoperplexa by Hardesty & Groothuis (1993) had slightly-pointed forewing apices that more resembled those of C. affinis affinis. Similarly, the southwestern Nebraska specimens thus appear to be intergrades between these two taxa, as suggested by Scott & Scott (1980) & Scott (1986). The phenotypic intermediates in Wyoming were studied on the eastern edge of the city of Laramie in a ravine dominated by Cercocarpus shrubs along with scattered Juniperus. Stands of Cercocarpus montanus Raf. were found to be the preferred male perching sites between | and 4 PM. Despite numerous stands of the flowering larval hostplant, Eriogonum flavum (Table 2) in this ravine, adults of these intermediates do not occur in most years (C. Ferris pers. com.). Their relative scarcity was also noted during this study, a fact that rendered a reliable comparison of macular band scaling between this population and other populations in Albany and Laramie counties, Wyoming, and Weld County, Colorado inconclusive. A total of 12 of 12 males (100%) from these localities lacked brown and black scaling mesial to the white maculations. 2. Only 18 of 221 (8%) specimens examined from localities in Colorado (Denver southward) and New Mexico (Albuquerque northward) possessed a nearly complete white band bearing more than 12 black scales in each maculation (Fig. 3). The majority (203, 92%) possessed six or fewer white maculations that included eight or fewer black macular scales (Fig. 4). Intergradation between C. affinis apama (Fig. 6) and C. a. homoperplexa in eastern Wyoming and southwestern Nebraska (Fig. 5) also occurs in the Jemez Mountains of northern New Mexico (Fig. 2). A lack of sympatry exists between C. affinis affinis (Fig. 7) and C. a. apama in Arizona, as shown by Scott (1986). This lack of sympatry probably led to earlier treatments of the two taxa as separate species. Sympatry with C. a. affinis is also absent within the known range of C. affinis washingtonia in the Pacific Northwest (Guppy and Shepard 2001) (Fig. 8) yet C. a. affinis is narrowly sympatric with C. perplexa in Washington (Hinchliff 1996). 3. In southwest Colorado, sharp VHW maculation differences were found to occur within and between populations. These varied from only a few VHW white maculations typical of C. affinis homoperplexa (16 of 23 187 specimens, 70%) to those with more prominent brown- bordered maculations, indicative of C. affinis apama (7 of 23 specimens, 30 %). 4. In New Mexico (southwest of Albuquerque), Utah, and Arizona, most of the specimens examined (142 of 145, 98%) possessed a complete or nearly complete black-and brown-bordered macular band that exhibited little variation, typical of C. affinis apama. 5. Specimens from a population in the Sacramento Mountains near Ruidoso (Lincoln County, New Mexico), lacked hindwing maculations or exhibited an incomplete, curved macular band lacking in brown scaling or containing 12 or fewer brown scales in each. This was seen in 15 of 20 (75%) specimens (Fig. 9). This brown macular scaling was very greatly reduced or absent in both sexes, as were black macular scales in 19 of 20 specimens (95%). One male and one female examined from two localities in the southern portion of the Sacramento Mountains (Otero County, New Mexico) possessed a complete macular band typical of C. a. apama yet its hostplant, C. fendleri A. Gray, is absent from these localities. These two specimens from the first decade of the 20th Century bore incomplete collection data and no further sightings or captures of completely banded specimens ishsee e yee recorded from the Sacramento Mountains. 6. A complete submesial white band was exhibited by all specimens seen from the Sierra Madre Occidental range of Sonora, Chihuahua, and Durango states (21 of 21, 100%). This feature was also accompanied by a darker green scale color on the VHW surfaces (Fig. 10) except on Chihuahua specimens, which exhibited a VHW green color similar to that seen in typical C. affinis apama from Arizona and New Mexico (15 of 15, 100%). The contiguous brown scale border of the submesial band was notably wider, more like that seen in specimens from the state of Durango. These specimens suggest discontinuous distribution in C. a. apama. In summary, populations west of the Continental Divide and south of Utah should now be treated as C. affinis apama with a complete macular band and a nearly complete brown border to this band. These tend to exhibit a relative loss of maculations east of the Continental Divide suggesting possible clinal variation from west to east. Variable specimens from southwestern Nebraska, southeastern Wyoming and northern Colorado suggest local breakdown of this pattern. The populations of C. affinis studied east of the Continental Divide with respect to the VHW submesial band appear to be local adaptations (ecotypes) that most closely resemble C. affinis homoperplexa. Using both reared and_ field-collected specimens, consistent 188 structural differences were found to exist in adult specimens of C. affinis from the Sierra Madre Occidental in the states of Sonora and Durango and the Sacramento Mountains of New Mexico. Such differences were noted in the labial palps as well as on the ventral hindwings. These character differences suggest two rather distinct populations that are now, using both structural and biological features, designated as new subspecies: Callophrys affinis albipalpus ssp. nov. (Figs. 13, 14) Description: holotype male: Head: Antennae black with 16 white annuli; distal three white annuli contiguous ventrally; club annuli black, orange brown distally; basal segment black with a single white maculation ‘laterally: - labial palp scaling black dorsally, white laterally, with distal segment predominantly “white with white tip. Dorsal surface of forewing: FWL = 12.1 mm. Ground color uniform orange brown with contrasting brown venation and light brown stigma; outer margin dark brown with bicolored wing fringe (basally brown, distally w! hite). Dorsal surface of hindwing: Ground color, outer margin and fringe as in forewing; fringe tricolored between vein 2A and 3A (basally brown, mesially white, distally dark brown).Ventral surface of forewing: Ground color grayish from posterior margin to vein Cu2; light brown to vein R5; abundant scattered bright green scaling between vein R5 and costa; incomplete band of five laterally- -displaced submesial white maculations between veins R4 and Cul; two maculations with very weak black scaling mesially; apical region to vein Cul lateral to submesial maculation with bright green scaling; outer margin brown with bicolored fringe (basally gray brown, distally pale). Ventral surface of hindwing: Ground color bright green with an incomplete submesial band of six white maculations displaced laterally between veins Cul and 2A; basal brown scaling in each maculation weakly developed; mesial black scaling significantly reduced to three or fewer scales or absent; outer margin as in forewing: anterior portion of fringe bicolored (basally light | brown, distally pale); tricolored between veins 2A and 3A (basally light brown, mesially white, greatly reduced, & distally dark brown). Allotype female: Head: Antennae as in holotype male with 17 white annuli; labial palp scaling mostly white; distal segment predominantly white with white tip. Dor sal surface of forewing: FWL = 12.5 mm. Ground color orange brown with 1.2 to 1.5 mm wide dark brown outer margin; wing venation and fringe as in holotye male. Dorsal surface of hindwing: Ground color as in forewing, with dark brown outer margin narrower than 0.5 mm; venation and fringe as in forewing; mixed brown and white scales between veins 2A and 3A. Ventral surface of for ewing: Ground color as in holotype male, with brown scaling reaching the costal vein; submesial maculations absent; apical and subapical region very weakly green; fringe light brown basally; pale distally. Ventr: al surface of hindwing: Ground color as in forewing; incomplete submesial band with five white maculations; maculations with reduced brown scaling basally; mesial black scaling greatly reduced to three or fewer scales per maculation; outer margin brow n with white scales forming a line between veins Cul and 2A: fringe as in holotype male. Types. All type specimens were collected in the Sacramento Mountains of Lincoln County, New Mexico. The holotype male and allotype female were reared from a Ist generation female taken at the New Mexico State University's Montgomery Biological Research Laboratory (now privately ow ned property), 10 km north of Ruidoso, 2134m, Sacramento Mountains on ae 28, 1982. They emerged in August 1982. Twelve paratypes (6 ¢ ) bear the same data as the holotype male and _allotype ae ea paratypes were collected at the type locality on VI-4-1981 by the author (2d, 42) and 1d reared ex ovum in 1982. Other field-collected paratypes include 2 males from Cedar Creek Camp, 10 km north of Ruidoso, 2134m, Sacramento Mountains, VI-30-1961, F., P. & J. Rindge, collectors JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY (Camegie Museum). The holotype male and allotype female and seven paratypes will be deposited in the Natural History Museum of Los Angeles County, Los Angeles, California. Two paratypes will be deposited at each of the following institutions: Florida Museum of Natural History, American Museum of Natural History, California Academy of Sciences, Carne -gie Museum of Natural History, National Museum of Natural History, Essig Museum of Entomology (University of California, Be orkeley). One paratype has been placed i in the Entomology Research Museum (University of California, Riverside) and one remains in the collection of the author. Etymology. The name given to this new taxon describes the dominant white scaling on the distal segment of the labial palpi observed in 18 of 20 (90%) specimens of both sexes, both and _field- collected. Diagnosis. Males and females of C. a. albipalpus typically exhibit predominantly white scaling along the labial palps. The distal segment is entirely white. “This trait, along with the absence of black scaling along the submesial band of the ventral hindwing, distinguishes this taxon from its closest allies, C. a. apama and C. a. homoperplexa. Range and habits. C. a. albipalpus is endemic to the Sacramento Mountains of south central New Mexico. It prefers the sun-exposed canyons and disturbed areas that exist in the vicinity of New Mexico State highways 37 and 48 at or above 2100m. First generation males and females often obtain nectar from the flowers of Melilotus indicus (L.), while second generation individuals prefer those of Eriogonum reared jamesii. Bivoltine in most years, it appears to be strongly influenced by ecological succession and thus relocates for both nectaring and oviposition. E. alatum appears to be the preferred larval hostplant (Table 2). Male hilltopping has not been observed. Callophrys affinis chapmani ssp. nov. (Figs. 11, 12) Description: Head: Antennae black with 17 white annuli basal segment bicolored (black mesially, white laterally); club annuli black, terminally orange brown; labial palp scaling Blac white laterally; distal segment black with white tip. Dor sal surface of forewing: FWL = 13.2 mm. Ground color uniform gray brown with concolorous venation; stigma gray; outer margin salle dark brown; fringe brown basally, pale distally. Dorsal surface of hindwing: Ground color and outer margin as in forewing; fringe tricolored (brown basally, pale mesially, orange brown distally): fringe at tips of veins Cu2, 2A, and 3A with elongated white scales. Ventral surface of forewing: Ground color grayish brown from posterior margin to vein Cu2; brown to costa; incomplete band of five submesial white maculations between vein R4 and Cul, displaced laterally between vein M3 and Cul; white maculations very weakly black- scaled mesially; basal, apical and subapical regions lateral to submesial maculations with olive green scaling; outer margin brown with tricolored fringe (basally “light Broun mesially gray brown, distally pale along anterior margin, wah distal portion becoming brown near tornus). Ventral surface of hindwing: Ground color olive green, with complete submesial band of white maculations displaced ‘laterally between veins Cul and 2A; basal brown scaling of maculations form a contiguous band 0.8 to 1.0 mm wide; prominent black scaling mesially; outer mar gin brown with white scaling that forms a line between veins 3A and Cul: fringe VOLUME 59, NUMBER 4 189 Fics. 11-14: New subspecies of Callophrys affinis 11. C. affinis chapmani, holotype male, 38 km east of El Salto, Hwy. 28, 2439 m, collected ex ovum on July 13, 1985 -emerged: August 28, 1985, G.A. Gorelick, collector 12. C. affinis chapmani, allotype fe >male, 38 km east of E] Salto, Hwy. 28, 2439 m, Durango, Mexico, collected ex ovum on July 13, 1985 -emerged: August 27, 1985, G.A. Gorelick, collector 13: C affinis albipalpus, allotype female, 10 km north of Ruidoso, 2134 m, Lincoln Co., NM, ciollected ex female on June 28, 1982 - Emerged: August 15, 1982, G.A. Gorelick, collector 14: C. affinis albipalpus, holotype male, August 15, 1982, G.A. Gorelick, collector female on June 28, 1982 - Emerged: tricolored (basally brown, mesially gray brown, distally mixed pale and gray brown): fringe between veins 2A and 3A distally dark brown; elongate d white fringe scaling at tips of veins Cu2, 2A and 3A. Allotype female: Head: Antennae with 17 white annuli, the distal three contiguous ventrally; basal and distal segments as in holotype male; labial palps white -scaled late rally; distal segment brown, white- tipped. Dorsal surface of forewing: FWL= 13.8 mm. Ground color uniform orange brown with 1.0 mm wide dark brown outer margin; fringe as in holotype male. Dorsal surface of hindwing: G round color as in forewing, with 0.8 mm wide outer margin; fringe as in holotype male; fringe between veins Cu2 and 3A light brown distally; fringe at tips of veins 2A and 3A with elongated white scales. Ventral surface of forewing: Ground color as in holotype male; incomplete band of four nondisplaced white submesial maculations between vein R5 and Cul with prominent black scaling mesially; basal, apical and subapical area lateral to submesial maculations dark green; outer margin and fringe as in holotype male. Ventral surface of hindwing: Ground color dark green; submesial band of white maculations as in holotype male; contiguous brown scaling 1.5 to 1.8 mm wide; outer margin, tips of veins, and fringe as in holotype. Types. The holotype male and allotype female were both collected and reared ex ova by the author, 38 km east of El Salto, 2439m in the Sierra Madre Occidental range of Durango, Mexico on July 13, 1985. The adults emerged on August 28, 1985. These will be deposited in the Natural BESSY Museum of Los Angeles County along with four paratypes (2d, ) from the same locality. The remaining 14 paratypes both reared a field collected) were taken between the city of Durango and the village of El Salto on Mexico Hwy. 28 ( 1964-1986). Two of these paratypes will be distributed to each of the same institutions listed for C. affinis albipalpus. Etymology. This patronym honors Dr. Thomas Algernon Chapman (1842-1921), Scottish physician and 10 km north of Ruidoso, 2134 m, Lincoln Co., NM, collected e lepidopterist. Dr. Chapman named and described the related species, Callophrys avis from sage scrub plant communities in SE France, the Iberian Peninsula and North Africa (Chapman, T.A., 1909). Additionally, he published a life history complete with a description of each _ larval along with photographs and comparisons to closely related C. rubi in detail atypical for his time (Chapman, T.A., 1910). Diagnosis. Unlike the closest ally, C. a. apama, the ventral hindwings of both sexes of C. chapmani exhibit a distinct mesial row of dark brown scales between veins M3 and 2A near the tips. The mesial brown scaling along the maculations forms a single continuous band one mm. in width. The green scaling of the ventral hindwing is typically olive green in males and dark green in females. The labial palp scaling is tricolored (black and brown, with white tips). Range and habits. C. a. chapmani is multivoltine and ony occurs in the Sierra Madre Occidental range above 2200m in the states of Sonora and Durango in Mexico. It frequents disturbed areas that include sun- exposed fields and arroyos along roadsides. Both sexes prefer to perch on and obtain nectar from their larval hostplants, Ceanothus huichagorare and Eriogonum atrorubens. stage, 190 ECOLOGY Adult C. affinis are univoltine in the northern part of their range, appearing between late March and June in Colorado and Wyoming and into July for higher elevation populations (Ferris and Brown, 1981). Bionomics of C. affinis from the Pacific Northwest are presented by Christensen (1981). Colorado prairie populations are double-brooded in some _ years, emerging in May with a second brood appearing in late July arelion August (A.D. Warren, pers. comm.). In western Nebraska, flight occurs only in June and early July. In northern Arizona, relatively large populations were observed above 2287m elevation between late May and early July. These may be bivoltine since adults are also on the wing in August and Septembers. Unlike most populations of (G affinis where bivoltinism is unproven, the Sacramento Mountains adults were collected between early June and late August, suggesting bivoltinism in most years. In the Sierra Madre Occidental of Durango, Mexico, specimens obtained between April and September reveal a multivoltine existence. Unlike related California Callophrys that exhibit a preference for open areas (Ferris 1971b) and hilltopping/territorial behavior (Gorelick, 1971), C. affinis apama adults preferred to visit flowering vegetation along open roads and canyon bottoms and stayed relativ ely close to stands of their larval hostplant. This tendency was also mentioned by Ferris (1971b) pertaining to Wyoming populations. Behavioral similarities to C. affinis affinis such as hilltopping have been reported in several Colorado and New Mexico localities (Stanford 1977, S.J. Cary pers. com.). No hilltopping behavior was observ ed in the C.-a. affinis/homoperplexa blends of Albany and Laramie counties in Wyoming or in the populations studied in Banner and Chey. enne counties in Nebraska. Male C. affiis specimens from several localities in the vicinity of Aspen, Colorado (Pitkin County), exhibited variation in VHW maculations. These males exhibited hilltopping behavior (R.E. Stanford pers. com.). The sparse, non- contiguous VHW maculation pattern of these Pitkin County specimens is reminiscent of such patterns observed in many homoperplexa specimens from other Colorado localities. These maculation patterns indicate the existence of a probable blend zone between C. a. affinis and C. a. homoperplexa in this area. In that no significant ecological or behavioral isolation is evident, these taxa are undoubtedly conspecific as per the discussion by Kohn & Orians (1962). Adults of C. a. apama encountered in montane localities west of the Continental Divide in Arizona, JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY New Mexico, and Utah frequently visited the flowering shrub, Ceanothus fendleri A. Gray (Rhamnaceae). This plant typically blooms between May and July and is used both as a larval hostplant and as a nectar source. Utah populations also used C. fendleri along with Eriogonum racemosum Nuttall (Polygonaceae), even when these occur together in the same locality. In addition, C. affinis affinis in UT uses many species a Eriogonum: EE. alatum, E. heracleoides Nuttall. umbellatum Torrey, and E. racemosum (Jack L. Fae pers. com.) (Table 2) Mexican populations typically occur in association with Eriogonum atrorubens Engelmann and Ceanothus huichagorare Loesner, a species closely allied to C. fe OU On these plants, nectaring was observed in the morning and late in the day, with oviposition usually occurring during the early afternoon hours on both plants In most localities in the Front Range of the Rocky Mountains and east of the Continental Divide in the U.S., Ceanothus fendleri was the preferred hostplant whenever present. In its absence, species of Eriogonum were adopted: E. flavum Nuttall (Wyoming), E. umbellatum (Colorado), and E. alatum (New Mexico). While C. affinis apama appear to be on the wing virtually everywhere one finds abundant Ceanothus fendleri above 2287m, this association in regards to potential Eriogonum hosts was not observed. In Mexico, Eriogonum atrorubens occurs in oak- -juniper grasslands of Chihuahua and Zacatecas, and in the Sierra Madre Oriental of southern Coahuila (Reveal 1967), areas providing no records or evidence of any Callophrys. In this regard, Gilbert and Singer (1975) pointed out that many butterfly species are much more restricted in their distributions than are their potential hostplants. Males employed shrubs such as Cercocarpus species (New Mexico, Colorado, Wyoming), Rhus aromatica Ait. (Nebraska, Colorado, New “Masico): Quercus gambellii Martin and Drew, and Q. undulata Torrey (Colorado, New Mexico) as perching sites used to initiate courtship flights. Such behavior was observed between 9 AM and 4 PM. They may also perch on tall grass, as reported by Scott (1975). Courtship and mating displays were frequently observed in sun-exposed terrain either on low vegetation, larval hostplants, or on male perching sites between 9 AM and 12 PM. C. affinis apama oviposition behavior was similar to that seen in other Callophrys (Gorelick, 1971). Females preferred to remain in close association with the larval hostplant throughout their flight period, employing it also as a nectar source between 9 AM and 12 PM. After some abdominal probing, females placed a single egg on each floral tip VOLUME 59, NUMBER 4 TABLE 2: Host plant associations for Callophyrs affinis subspecies Locality® Arizona 1. Strayhorse Cmpgd.,White Mountains, Greenlee Co. 2. Schultz Pass, Coconino Co. New Mexico 3. Zuni Mountains, Valencia Co. 4. Jemez Mountains, Los Alamos Co. 5. Bandelier National Monument, Sandoval Co. 6. Sacramento Mountains, Lincoln Co. Utah 7. Singletree Cmpgd. Wayne Co. 8. 7.4 mi. south of Torrey, Wayne Co. 9. 3.2 mi. north of Long Valley jet., Kane Co. 10. Hwy 24, Piute Co. 11. 28 mi. SE Moab 12. Rich Co. 13. 4 mi. NE of Accord Lakes, Sevier Co. 14. Wilkerson Pass, Park Co. 15. 2.6 mi. SW of Lynn, Box Elder Co. WYOMING 16. Woodruff, Lincoln Co. 17. nr. Evanston, Uintah Co. 18. east of Laramie, Albany Co. 19] Ceanothus Eriogonum Eriogonum Eriogonum Eriogonum Eriogonum Eriogonum Ceanothus fendleri alatum racemosum umbellatum heracleoides flavum atrorubens —_ huichagorare C. affinis apama C. affinis apama C. affinis apama C. a. apama x homoperplexa C. a. apama x homoperplexa C. a. albipalpus C. affinis C. affinis apama apama C. affinis C. affinis apama apama C. affinis apama C. a. affinis C. affinis apama C. a. affinis C. a. affinis C. a. affinis C. a. affinis C. a. affinis C. a. affinis C. a affinis x homoperplexa TABLE 2: continued JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Ceanothus fendleri Locality® Eriogonum alatum Eriogonum racemosum Eriogonum umbellatum Ceanothus huichagorare Eriogonum heracleoides Eriogonum flacum Eriogonum atrorubens Nebraska 19. Sidney rest area Interstate Hwy 80, Cheyenne Co. 20. Bull Canyon, Banner Co. Mexico 21. Sierra Madre Occidental, Durango °Locality details: C. a affinis x homoperplexa C. a affinis x homoperplexa C. a. chapmani Gig chapmani Strayhorse Cmpgd., Hwy 666, 8000', White Mountains, Greenlee Co. [GAG] .5 mi. N. of Flagstaff Schultz Pass, 7800', Coconino Co. [GAG] . Jemez Mountains, nr. Los Alamos, Burnt Mesa, Hwy 4, 7000', Los Alamos Co. [GAG] Js 2 3. Zuni Mountains, 8000', Valencia Co. [GAG] 4 5 . Bandelier National Monument, 7000! Sandoval Co. [SJC] 6. Sacramento Mountains, 5 mi. north of Ruidoso, 7000', Lincoln Co. 7. Singletree Cmpgd., Hwy 12 11 mi. south of Torrey, 8300', Wayne Co. |JLH] 8. 7.4 mi. south of Torrey, North Slope Rd, 8000', Wayne Co. (JLH] 9, 3.2 mi. north of Long Valley jet., Hwy 89, 7200' Kane Co. [JLH] 10. Hwy 24, 1 mi. south of jet., UT Hwy 24/25, 8300', Piute Co. [JLH] 11. 4.5 mi. SSW Buckeye Reservoir, 7700', 28 mi. SE Moab [JLH] 12. 1.4 mi. north of SW comer of Lincoln Co., WY, 6600', Rich Co. [JLH] 13. 4 mi, NE of Accord Lakes, 8400', Sevier Co. [JLH] 14, 2 mi. E. of Wilkerson Pass, 9100', U.S. Hwy 24, Park Co. [JLH] 15. 2.6 mi. SW of Lynn, 6900', Box Elder Co. vee 16. 8.2 mi. NE of Woodruff, 6800', Lincoln Co. [JLH] 17. 10 mi. N. of Evanston, 6700', Uintah Co. |JLH] 18. east of Laramie, 4450' Albany Co. (young stages unobserved; adults occur in association with Eriogonum flavum) 19. Sidney rest area Interstate Hwy 80, 4200', Chey venne Co. (young stages unobserved; adults occur in association with Er riogonum flavum) 20. Bull Canyon, 17 mi. SW of Harrisburg, 4500! Banner Co. (young stages unobserved; adults occur in association with Eriogonum flavum) 21. Sierra Madre Occidental, 13 mi. E. of El Salto, 8000", state of Durango [GAG} between 10 AM and 12 PM. First generation females oviposited at the tip of the shoot, preferring the floral calyx (Figs. 15-16). First instars fed on new leaves at the tip, or on unopened sepals and petals when available. Later instars moved to the relatively immature leaves a few centimeters from the tip of the stem where they fed after consuming the flower bud. Mature larvae consumed the young leaves. Females in later broods oviposited on sepals or terminal leaves and emerging larvae fed on the flowers, moving from flower to flower on the stem. Mature larvae generally assumed the color of these flowers (Figs. 17-21). While eclosions were noted in most field-collected eggs during the study, each of eight eggs collected on the flowers of C. fendleri growing below San Francisco Peaks (8 km north of Flagstaff, Arizona) was sucked dry by a hemipteran (J. Emmel pers. com.). Field- collected larvae below San Francisco Peaks, and Rose Peak along U.S. Hwy 191 in eastern Arizona were subjected to occasional parasitism by Apanteles sp., a braconid wasp. Along a 1-kilometer length of road at both localities, it was not uncommon to find up to 60 first and second instar larvae in a single day. These larvae exhibited mortality from disease and parasitism of up to 45%. In captivity, mature larvae crawled from the leaves and flowers onto the substrate and attached themselves to the surface or underside of the dead leaves where they pupated. In canyons along New Mexico state highways 37 and 48 amongst stands of Pinus ponderosa Douglas ex Lawson, Quercus undulata, Artemisia dracunculoides Pursh., and Eriogonum jamesii Benth., where Ceanothus fendleri was not found, attempts to collect eggs and larvae of C. affinis on Eriogonum jamesii were unsuccessful despite its use by both females and males as a nectar source. In the Sierra Madre Occidental, west of Durango above 2439m, oviposition and mating behavior was found to be the same as observed in C. a. apama. A VOLUME 59, NUMBER 4 193 Fics. 15-21: Immature stages of Callophrys affinis. 15-16. Egg of C. affinis chapmani on sepal of Eriogonum atrorubens flower, Durango, Mexico. 17. Mature larva of C. affinis apama on Ceanothus fendleri, Greenlee Co., AZ. 18. Mature larval of C. affinis affinis on Eriogonum umbellatum, Lincoln Co., WY. 19. Mature larva of C. affinis albipalpus on lab host, Eriogonum grande rubescens, Lincoln Co., NM. 20. Mature larva of C. affinis chapmani on Eriogonum atrorubens, Durango, Mexico. Durango, Mexico search of a grassy meadow adjacent to farmlands along the highway 21 km east of El Salto (vicinity of Llano Grande) yielded 143 Callophrys eggs (116 living, 11 dead, 16 eclosed) collected in four days (July) on abundant stands of Eriogonum atrorubens. In addition, 77 first instar larvae were collected, with up to 6 larvae 21. Mature larva of C. affinis chapmani on Eriogonum atrorubens, on the same plant. Parasitism, infertility, and larval mortality reduced this number to eight specimens successfully reared to adulthood. Approximately 25 first instar larvae were preserved for later study. Along the roadside 45 kin east of El Salto, an area characterized by Pinus engelmanni Cama durangensis Martinez, 194 Arbutus sp., Quercus sp., and Arctostaphylos sp. between 2439-2500m, a total of four variable instar, green-colored larvae were taken on the same day from Ceanothus huichagorare. No red-colored larvae were found on the abundant E. atrorubens growing nearby yet two variably red and green larval morphs were collected on E. atrorubens bloom stalks growing along the roadside 38 kn east of El Salto. Field-collected adults were rare. No more than three adults of either population were typically seen on any given July day during the study. Interestingly enough, all modern records of C. affinis from the Sacramento Mountains are from areas burned by wildfires in the 1970s. The known larval hosts for C. affinis in New Mexico (C. fendleri, E. alatum) are early successional plants. Outside of occasional wildfires and other disturbances, a decline of hostplants thus appears to be due to ecological succession (Pratt 2001). This suggests that the occurrence of C. affinis is determined primarily by the ecological structure of its habitat or its successional stage rather than the larval host, and it may therefore use any suitable hostplant(s) that are present at a given site. In addition, erosion brought on by excessive summer rainfall and the potential loss of Eriogonum atrorubens due to grazing and local human consumption add to selective pressures operating on the population of C. affinis along Hwy 28 in the Sierra Madre Occidental of the state of Durango i in Mexico. CONCLUSIONS Macular band variation, phenotypic intergrades, synchrony and similar hostplant choices and behaviors warrant the inclusion of C. apama as a subspecies of C. affinis. Additionally, both behavioral and geographic isolating factors appear to have arisen in the southernmost distribution of C. affinis. This scenario conforms well to that described by Cox et al (1977), Holland (1988) and Shields (1996) in regards to geographic isolation in southwestern North American butterflies, ACKNOWLEDGEMENTS I would like to thank the following for assistance given in plant identification, library searches, the collection or loan of adult specimens, and/or the obtaining of records pertinent to this study: George T. Austin, Jim Brock, Steve Cary, Felix Delgado Castaneda, Neil Dankert, Julian Donahue, Scott Ellis, Dave Faulkner, Clifford Ferris, Greg Forbes, Jack Harry, Julie Harvey, Darlene Judd, Gloria Harjes, Joel M. Johnson, Peter Jump, Steve Kohler, Boris Kondratieff, Tim Lowrey, Noel McFarland, Larry Muller, Doug and Mary Mullins, Jane Mygatt, Paul Opler, Jose Pinedo Nesta , John Rawlins, Jim Reiser, James Reveal, Kilian Roever, Rick Rogers, Barbara Rugely, Andy Sanders, James Scott, Virginia Scott, Oakley Shields, Steve Spomer, Ray Stanford, Greg Thorn, Dieter Wilken and Nina Zitani. A collecting permit was obtained from the Direccion JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY General de la Fauna Silvestre, Mexico City, (under the auspices of the Museum of Natural History of Los Angeles County) for collecting the specimens used in the type series described herein. Young stages of C. a. chapmani and C. a. albipalpus were reared in my absence by Greg Ballmer and John Emmel, respectively. John Emmel, Don Frack, and Jack Harry provided photos of the young stages. At Citrus College, Barbara Rugeley and Tina Gutierrez provided library resources. I also received technical assistance in photographing the adults from Darrell Carr, and graphic illustration in Figure 1 by Christopher Guerra and Dr. Richard Fernandes. I examined adult specimens in private collections, California Academy of Sciences, Museum of Natural History-Los Angeles County, California Academy of Sciences (San Francisco), Essig Museum of Entomology (Univ ersity of California, Berkeley), Entomology Research Museum (Univ ersity of California, Riverside), Bean Museum (Brigham Young University), Univ ersity of Colorado (CU Museum- Entomology section), C.P. Gillette Museum of Entomology (Colorado State University), American Museum of Natural History (New York) and the Camegie Museum (Pittsburgh). Plant records were accessed from Rancho Santa Ana iBoveaite Gardens, Museum of Natural History-Los Angeles County, University of California, Riverside, University of New Mexico, New Mexico State University and the University of Wyoming. This paper was reviewed by Steve Cary, Paul Opler and Gordon Pratt. Grateful thanks go especially to Andrew D. Warren, whose comments, suggestions, and contributions of distribution records from the Pacific Northwest greatly enhanced this study. To Richard Holland, I offer my sincere thanks for the years (and miles) of assistance and patience necessary to successfully complete this review. LITERATURE CITED BarLowitz, R. AND J. Brock. 1991.Butterflies of southeastern Ari- zona. Sonora Arthropod Studies, Inc. Tucson. ix + 342 pp. BALLMER, GREGORY R., & G.F. Pratt. 1988. A survey of the last instar larvae of the Lyceanidae (Lepidoptera) of California. Journal of Research on the Lepidoptera. 27(1):1-81. —— 1992. Loranthomitoura, a new genus of Eumaeini (Lepidoptera: Lycaenidae: Theclinae). ). Tropical Lepidoptera. 3(1):37-46, BARNES, WILLIAM & F. H. BENJAMIN. 1923. iC aleps apama race homoperplexa" . Contributions on the Natural History of the Lepidoptera of North America. 5:64-69. 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A biosystematic study of two species of Cal- lophrys (Callophrys) in California (Lycaenidae). Journal of the Lepidopterists' Society. 25 (Suppl. 2):1- 41. Guppy, CrisPIN S., & J.H.SHEPHARD. 2001. Butterflies of British Co- lumbia. University of British Columbia Press, Vancouver. 414 pp. Harpesty, R.L., & D.R. Grooruuts. 1996. Butterflies of the Laramie Mountains, Wyoming (Lepidoptera: Rhopalocera). Journal of. Re- search on the Lepidoptera. 32:107-123. HasKIN, J.P. & F. GRINNELL. 1912. Thecla dumetorum and T. affinis, a study (Lepid.). Entomological News. 23:3-8. HINCHLIFF, J. 1994. An Atlas of Oregon Butterflies. Oregon State University Bookstore, Corvallis, Oregon. v + 176pp. . 1996. An Atlas of Washington Butterflies. Oregon State Univer- sity Bookstore, Corvallis, Oregon. vi + 162pp. HOLLAND, R. 1988. A new subspecies of Speyeria atlantis (Nymphal- idae) from south-central New Mexico. Bulletin of the Allyn Mu- seum. No. 115. pp. 1-9. —— & STEVEN Cary. 1996. Butterflies of the Jemez Mountains of northern New Mexico. Journal of the Lepidopterists' Society. 50(1): 61-79. Howe, W.H., Editor. 1975. The Butterflies of North America. Dou- bleday and Co., Garden City, New York. xiii + 633 pp., 97 pls. HupparbD, J.P. 1965. The summer birds of the forests of the Mogollon Mountains, New Mexico. The Condor. 67:404-5. KELLY, KENNETH L., AND DEANE B. JuDD. 1976. Color —Universal Language and Dictionary of Names. U.S. Department of Com- merce, Washington, D.C. 184 pp. Kxots, A.B. 1951. A Field Guide to Eastern Butterflies. Houghton Mifflin Co., Boston, Massachusetts. xvi + 349 pp. KOHN, ALAN J., & GORDON Orians. 1962. Ecological data in the clas- sification of closely-related species. Systematic Zoology. 11:119- 127. Mayr, E. 1965. Animal species and evolution. Harvard University Press. Cambridge, MA. 797 pp. MILLER, L.D., & FM. Brown. 1981. A catalogue/checklist of the but- terflies of America north of Mexico. Memoirs of the Lepidopter- ists' Society 2:vii + 1-280. MUELLER, STEPHEN J. 1982. Butterflies of the Dixie National Forest, Utah (including Bryce Canyon National Park & Cedar Breaks National Monument). Utahensis. 2(3):25-31. Opcer, P.A. 1999. A Field Guide to Western Butterflies. 2nd ed. Houghton Mifflin Co., Boston. 540pp. — & A.D. WARREN. 2003. Butterflies of North America. 2. Scien- tific names list for butterfly species of North America, north of Mexico. Contributions of the C.P. Gillette Museum of Arthropod Diversity, Colorado State University, Fort Collins. 80 pp. (up- 195 dated periodically at: http: www.biology.ualberta.ca/old_site/ uasm//Opler&Warren.pdf) Pratt, GORDON F., 2001. Survey for the Sacramento Mountains Green Hairstreak (Callophrys apama=affinis). (Unpublished). Filed with the New Mexico branch, U.S. Fish & Wildlife Service. PYLE, ROBERT MICHAEL. 1981. The Audubon Society Field Guide to North American Butterflies. Chanticleer Press, New York. 916 pp. bo, Butterflies of Cascadia. Seattle Audubon Society. Seattle. WA. 420 pp. REVEAL, J.L. 1967. Notes on Eriogonum - II. Variation in Eriogonum atrorubens. SIDA. 3(2): 82-86. Scorr, J.A. 1975. Mate-locating behavior of westemm North American butterflies. Journal of Research on the Lepidoptera. 14(1):1-40. ——1986. The Butterflies of North America. Natural History and Field Guide. Stanford University Press. Stanford, CA. 583 pp.. 64 Is. mae G.R. Scorr. 1978(80). Ecology and distribution of the but- terflies of southem central Colorado. Journal of Research on the Lepidoptera. 17(2):73-128. SHIELDS, O. 1996. Geographic isolation in southwestern North Ameri- can butterflies (Lepidoptera, Rhopalocera). Nachrichten des en- tomologischen Vereins Apollo. 17(1):71-92. STANFORD, R.E. 1977. Zone 3: Rocky Mountains: p. 8 in: Field Sum- mary 1976. News of the Lepidopterists' Society 1977 (2):3-15, 1S- PAI, —— 1994. Zone 4: Rocky Mountains: p. 31 in: Season Summary 1993. News of the Lepidopterists' Society 1994 (2):25-33. —— & PA. OpLer. 1993. Atlas of westem USA butterflies including adjacent parts of Canada and Mexico. Denver and Fort Collins, CO. x + 275 pp. TILDEN, J.W. 1963. An analysis of the North American species of the genus Callophrys. Journal of Research on the Lepidoptera. 1(4):281-300. ToLiver, M.E., R.HOLLAND, AND S.J.Cary.1994. Distribution of but- terflies in New Mexico (Lepidoptera: Hesperioidea and Papil- ionoidea). 2nd ed. Published by the authors., Albuquerque, New Mexico. 393 pp. Received for publication 2003; revised and accepted 19 September 2005 APPENDIX: MATERIAL STUDIED Callophrys affinis affinis (W.H. Edwards) no locality, W.H. Edwards collection, designated lectotype, F.M. Brown (1967), 16 no locality, W.H. Edwards collection, designated lectotype . FM. Brown (1967), 19 no locality, W.H. Edwards collection, designated paratype. FM. Brown. (1967), 16 NEVADA: Elko Co: Spruce Mtn., Pequop summit, Steptoe Valley., TS1N R64E Sec. 33, VI-8 (no year or collector given) Lander Co: Kingston Canyon, VI-18-1970, 1s. C.D. Ferris. coll., Austin summit, VI-13-2002, 2m, P.A. Opler, coll. Lincoln Co: Highland Peak, Highland Range, VI-17-1990. 1c, 12. GT. Austin, coll. Nye Co: Antone Creek, Toquima Mtns.; VI-13 & VI-15-1933, 29. FW. Morand, coll.;Jett Cyn., V-25/26-1974, 1c, 12, J. DeBenedictis & W.E. Knoshaug, colls. White Pine Co: Mt. Wheeler, VI-2 to 6-1929, 2m, FW. Morand, coll.: Wilson Creek Range, base of Mt. Wheeler, 8700', VI -7 to 8-1985, 1, 12. G.A. Gorelick & G.T. Austin, colls., Indian Creek, Schell Creek Range, V-10-1984. 1c, G.T. Austiin, coll.; Snake Creek, 8000', Snake Range, VI-27-1972, 1c. 19. Scott L. Ellis, coll. UTAH: Box Elder Co: 2.6 mi. SW Lynn, 6900', coll. ex larvae on Eriogonum heracleoides (emerged IV-21-1988), 2¢, J.L. Harry, coll. 196 Garfield Co: Panguitch-Cedar Break Rd., Markagunt Plateau, VI-20-1972, 24,12, Scott L. Ellis, coll. Juab Co: Eureka, 7500! elev., no date, 46, R.C. Williams collection; VI-18- 1910, 1d, 19, T,. Spalding, coll. Park Co: 2 mi. E. of Wilkerson Pass, 9100', U.S. Hwy 24, coll. ex larvae on Eriogonum alatum, VII-7-1988 (emerged: VIII-18-1988), 1¢, 19, J.L. Harry, coll. Piute Co: UT Hwy 24, 1 mi. So. of jct., UT Hwy 24/25, $300!, coll. ex larvae on Eriogonum. racemosum (emerged X-19-1986),1¢, J.L. Harry, coll.; 0.7 mi up Hwy 25, Fish Lake Road, VI-21-1971, 19, J.A. Scott, coll. Rich Co: 1.4 mi. north of SW corner of Lincoln Co., WY (0.1mi.W. state line), 6600!, coll. ex larva on E. umbellatum, VI-1983 (emerged VIHI-9-1984), 1d; coll. ex larvae on Eriogonum umbellatum, VI-1985 (emerged II-1986), 4<, J.L. Harry, coll.; 4.5 mi. NE Woodruff, east side of Crawford Mtns., 6800',42, coll. ex larvae on E. umbellatum (emerged II-13-1986), J.L. Harry, coll. Sanpete Co: 18 mi. w. of Levan, VI-22-1979, 1¢, W.H.Whaley, coll. Sevier Co: Fish Lake, VI-24-1927, 1¢, 12, Ashby Boyle collection; Old Woman Plateau, VII-2-1986, 1¢, C.A. Miles, coll.; 0.1 mi. N. Hogan Pass, 8800’, V-5-1987, 26, J.L. Harry, coll.; 4 mi. NE Accord Lakes, Duncan Draw, $400, coll. ex larvae on Eriogonum.alatum (emerged V-1988), 2¢, 4°, J.L. Harry, coll. Summit Co: Snyderville, VII-27-1980, 1°, Ken Tidwell, coll.; North fork, Provo River, Uintah Mtns., 7500!, VI-29-1965, 1d, 2°, John Justice, coll.; 8000', VII-5-1964, 22, Ken Tidwell, coll.; VI-15-1963, 1¢;VH-5-1964, 12, J. Don Eff, coll. Uintah Co: Blue Mountain Plateau, Dinosaur National Monument, VI- 1968, 2¢, Scott L. Ellis, coll. Utah Co: Provo Peak, VII-30-1982, 1¢, C.A. Miles, coll., Silver Lake, VH- 15-1899?, 3¢, 2°, Henry Skinner, coll., VII-17-1949, 1¢, 69, A.J. Snyder, coll. County unknown®: City Creek Canyon, VII-12 to 15-1899, 2¢, Henry Skinner, coll. °cited as the type locality erroneously by Clench (1944); corrected type locality (Fort Bridger, WY) published by F.M. Brown (1970). WYOMING: Carbon Co: Bottle Creek Campground, southwest of encampment, 8400', VI-25-1972, 19; VI-17-1981, 1d, 19; VI-21-1982, 19; VII-2-1982, 1d, C.D. Ferris, coll. Fremont Co: Sinks Canyon Rd. at Slate Creek, VII-1-1979, 1, J. Brock, coll. Natrona Co: Casper Mountain, V-30-1988, 1¢, Karolis Bagdonas, coll. Teton Co: Teton Mtns., VII-8 to VI-17-1937, 4¢, 3°, R.W. Wind, coll. [Nevada State Museum]; Jenny Lake, Teton Mtns., VI-14-1931, 19, no coll.; 7¢, 14°, no date, Wallace-Bauer coll'n; VI-15 to 17-1956, 40, 2°, J. Don Eff, coll.; VI-12-1937, 1d, 12, R.G. Wind, coll.; Teton National Park along Snake River, VI-1/15-1979, 3¢, 22, T. McGann and U. of Wyoming (Zoology Dept.) Albany Co: Pole Mountain, $500', VIT-21-1977, 1° , C.D. Ferris, coll.; VI- 1- 1996,1¢, G.A. Gorelick, coll. Lincoln Co: Cokeville, VI-16-1979, 13, W.H.Whaley, coll.; 8.2 mi. NE Woodruff, Wildhorse Spring, 6800!, coll. ex larvae on E. umbellatum (emerged II-12-1986), 5¢, J.L. Harry, coll.; 4.5 mi. E. of Rex Peak, 0.4 mi. SE rd jet., 6900', coll. ex larvae on E. racemosum, VI-1985, (emg'd IH-24-1985), 19, J.L. Harry, coll.; coll. ex larvae on E. racemosum, VI-1986, (emerged IH-11-1986), 3d, 42, J. L. Harry, coll. Uintah Co: Evanston, VI-4-1885, 1¢, Evanston, 0.5 mi. E. of county road, 6700', coll. ex larvae on E. umbellatum (emerged IHI-12-1986), 3¢, J.L. Harry, coll. Sublette Co: Trail's End Campground., Elkhart Park, 9350', VII-22-1991, 2°, Larry D. Beutler, coll. Sweetwater Co: Pine Buttes, VII-1942, 1¢, 12, L. Bauer, coll.; 25 mi. south of Bitter Creek, VI-1942, 1, L. Bauer, coll. IDAHO: Franklin Co: Birch Creek, 5700', VI-6-1990, 12, C.D. Ferris, coll. Oneida Co: 3rd Creek Trailhead, 3 mi. east of SR 36, 6400', VI-23-2002, 2, R.E. Stanford, coll. MONTANA: Beaverhead Co: Polaris, VI-2-1943, 12, H.A. Howland, coll.; Lemhi Pass, 7340-7400', VIT-15-1978, 19, J.F.G. Clarke, coll.; VII-21-1995, 1¢, J. Verhulst, coll. ; Big Horn Co: Bighorn Canyon, 4610', V-28-1995, 16, C. Harp, coll. Carbon Co: East Rosebud Canyon, 1750m, VII-5-1966, 1d, C. Durden, coll.; VI-7-1992, 13, B. Vogel, coll.; East Rosebud Lake, 1850m, VI-27-1966, 1¢, C. Durden, coll. ex Skinner coll'n; 10 mi. No. of JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fergus Co; Little Snowy Mountains, VII-13-1976, 1°, S. Kohler, coll. Gallatin Co: Hebgen Lake, VI-21-1962, 1¢, J.A. Scott, coll. Granite Co: Black Pine Road, VI-26-1990, 1¢, S. Kohler, coll. Hill Co: Beaver Creek, Bear Paw Mountains, VI-20/28-1982, 10¢, 29, N. G. Kondla, coll. Judith Basin Co: 15 mi. SW of Utica, 4800', VI-28-1998, 1¢, R. Stanford, coll. Madison Co: Camp Creek, east of Melrose, 5485', VI-7-2001, 1¢, S Kohler, coll.; off Camp Creek Road, east of Melrose, VI-18-2001, 1°, S. Kohler, coll. Missoula Co: Ninemile Prairie, VI-9/18-1979, 9¢, 1°, S. Kohler, coll.; Ninemile Prairie, up ridge to north near Hwy. 200, 3900', VI-8-2001, 1d, S. Kohler, coll.; Shoofly Meadows, VII-4-2003, 1¢, D. Thompson, coll. Powell Co: S. Helmville , VI-5-1989, 2¢, S. Kohler, coll. Silver Bow Co: Rocky ridge trailhead, 2.5 mi. W. of Feely Divide Creek, VI- 19-2003, 1¢, S. Kohler, coll. Sweet Grass Co: East side, Crazy Mountains, VI-27-1966, 2c, J. Scott, coll.; VII-17-1966, 19, J. Scott, coll.; near Big Timber Creek, 4400', VI-28-1966, 1¢, J.A. Scott, coll.; Swamp Creek Road, Crazy Mountaiins, VII-12-1978, 3¢, 2°, S. Kohler, coll Wheatland Co: breaks west of U.S. Hwy. 191, 4950', 13 mi. south of Harlowton, VI-28-1998, 1d, R. E. Stanford, coll. COLORADO: ° Garfield Co: near. Rifle, 2¢, 2° (no date or collector given); Roan Plateau, 9200', VI-18-1996, 2¢, C.P. Slater, coll. Grand Co: Beaver Creek, VI-8-1977, 1¢, J. Donald Eff, coll.; Cottonwood Pt., 9800-10,000', 4 mi. south of Hot Sulphur Sprs., VI-3-1992, 49,; VI-30-1993, 5d, 22; VII-1-1993, 2d, 19, all A.D. Warren, coll.; County Road 50, 4 mi. east of jet., U.S. Hwy 40, 8 mi. west of Hot Sulphur Springs, VI-29-1991, 1°,; VII-14- 1991, 3¢, A.D. Warren, coll. Pitkin Co: nr. .Aspen Airport, 8100', VI-16-1973, 1¢; VI-13-1979, 3¢, R.E. Stanford, coll.; Aspen, V-25-1992, 1¢, R.E. Stanford, coll; 0-3 mi. west of. Aspen, VI-12 to 15-1969, 3¢, 19 ; VI-13-15-1979, 4, R.E. Stanford, coll. [GM- CSU] Routt Co: Steamboat Springs, 7200', VI-20-1973, 2d, R.E. Stanford, coll. [GM-CSU]; Little Snake River, 8000', VI-20-1973, 26, R.E. Stanford, coll. [GM- CSU] Summit Co: | mi. W. of Keystone, VI-30-1990, 12, A.D. Warren, coll. ° Additionally, both male and female adult specimens identified as C.a. affinis are housed in the collection of A.D. Warren [ADW] and were taken from localities in Rio Blanco and Moffat counties. C. affinis washingtonia Clench CANADA: BRITISH COLUMBIA: Oyama Lake, IV-22-1983, 1¢ ; Vernon, V-5-1983, 4d, 12 ; Goose Lake, V-7- 1984, 26 , all C.S. Guppy, coll.; Kalamalka Lake, Cosen Bay, nr. Vernon, V-25 1976, 6¢, J. Shepard, coll., Stubb Creek, Hwy 3, 8mi. E. Grand Forks, V-5-1973 1d, J. Shepard, coll.; Douglas Lake Rd., 30 mi. E. Merritt, V-22-1976, 1d, J. Shepard, coll, WASHINTON: Columbia Co: Blue Mtns., 4800! eley., VI-17-1961, E.J. Newcomer, coll.; along Tucannon River, Blue Mountains, VI-8-1963, 1°, R.E. Woodley, coll.; south of Gilbirth Spring, T9N R40E sec. 35NE Y4, 4550', BMTR.15, VII-3- 1964, 1d, J. & S. Shepard, coll. Douglas Co: Pine Creek, 700m, V-18-2000, 33, J. & S. Shepard, coll. Kittitas Co: Manastash Ridge, view point, Hwy. I-82, 2800', V-1-1983, 5d, 2°, J. Hinchliff, coll.; Manastash Ridge, 2672', V-18-1984, 1¢, J. & S. Shepard, coll Lincoln Co: Hawk Creek, 2300', V-17-1985, 2¢, J. & S. Shepard, coll.; head of Hawk Creek, V-28-1984, 1d, J. & S. Shepard, coll.; Hawk Creek Cmpgd., no. of Creston, V-19-1979, 2, 19, J. Shepard, coll. Okanogan Co: Brewster, V-9-1939, 1¢, J.C. Hopfinger, coll. (paratype); V- 14-1956, 12, J.C. Hopfinger, coll. (CMNH]; Barker Mtn., 3800!, VI-3-2000, 1d, J. & S. Shepard, coll.; Strawberry Mtn. Lookout, 4200', VI-21-1961, 1d, Jon Shepard, coll. Stevens Co: Pascal Cemetery, nr. Fort Spokane, V-28-1984, 2¢, J. & S. Shepard, coll. Whitman Co: Steptoe Butte, VI-15-1964, 1¢, 12, Jon Shepard, coll. OREGON: Baker Co: FR 11 to Bald Mtn., VI-19-01, 3°, Vern Covlin, coll.; Cave Creek Rd., 4000', Burnt River, w. of Durkee, V-27-1973, 1d (very worn), J. Hopfinger, VOLUME 59, NUMBER 4 coll. Crook Co: Ochoco Mtns., Hwy. 126, ca. 1 mi. NE Nat. For. boundary, VI- 10-2003, 26, A.D Warren, coll. Gilliam Co: Lonerock, 2000', VI-7-1961, 2°, D.L. Bauer, coll. Grant Co: Aldrich Mtn., Malhaur Nat. For., VI-2-1992, 6991', 5d, 19, J. Hinchliff, coll.; V-31-2001, 43, 12, A.W. Warren, coll.. Harney Co: Pueblo Mtns., 6500', Arizona Creek, VI-23-1979, 1d (worn), Mark Smith, coll.; summit of King Mtn., ca. 6400', VI-14-99, 9¢, 42, Vern Covlin, coll. Malheur Co: Trout Creek Mtns., 7000', 10 mi. WNW McDermott, VI-8- 1980. 1c, 12, Mark Smith, coll.; 5-6 mi. N. of Beulah Reservoir, Bendire Mtn. Rd., 4700', V-17-2001 276, 3°, A.D. Warren, E. Runquist, D. McCorkle, collectors Malheur/Baker Co. line: south of Rd. 16, 2-3 mi. SW Hwy. 26, VI-19-02, 45,12. A.D. Warrren, coll. Umatilla Co: summit of Tower Mtn., ca. 6700', VII-5-99; VI-7-00; VI-14-01, 183, 82, Vern Covlin, coll. Wallowa Co: ca. 10 mi. N. of Wallowa on rd. to Troy, VI-23-02, 19, A.D. Warren, coll. IDAHO: Adams Co: near mouth of Wildhorse River, V-12-1959, 19, S.G. Jewett, coll. Boise Co: 5 mi. west of. Idaho City, Cold Spring Creek, VI-8-1976, 19, JEG. Clarke, coll. Camas Co: Willow Creek Cyn., V-27-00, 1¢, A.D. Warrren, coll. Callophrys affinis affinis/ homoperplexa blend WYOMING: Albany Co: east of Laramie, TI5N R73W S1, 4450', VII-17-1971, 26; VI-30- 1972, 1d; VII-1-1972, 19; VII-6-1972, 16; VII-8-1972, 12; VII-10-1972, 29; VII- 18-1982, 12, VI-18 to VII-1-1996, 12d, 4°, C.D. Ferris, coll., VI-26/30-1996, 6d, G.A. Gorelick, coll. Laramie Co: 20 mi. W. Cheyenne, Hwy 210, VI-22-1985, 1d, P.A. Opler, coll.; 24 mi. W. Cheyenne, Hwy 210, VI-15-1985, 1¢, P.A. Opler, coll.; Curt Gowdy St. Pk., 24 mi. W. Cheyenne, Hwy 210, VII-6-1997, 56 , G.A. Gorelick, coll; § mi. S. Cheyenne, VI-18-1985, 1¢, 29; VII-22-1985, 1° , P.A. Opler, coll. Carbon Co: near Battle Mountain, 7500', VI-20-1973, 1d, R.E. Stanford, coll. NEBRASKA: Morrill Co: 6 mi. S. Redington, VI-27-1974, 1¢, W.T. Morgan, coll. Cheyenne Co: Hwy IS0, Sidney rest stop, 4200', VI-29-1996, 203, 49; VII- 10-1997, 1d, 12. G.A. Gorelick, coll.; VI- 1 to VI-29-1996, 8< , Jim Reiser, coll. Banner Co: Bull Canyon, 17 mi. SW Harrisburg, 4500', V-23-1994, 4¢ , Jim Reiser, coll.; VI-30-1996, 46, 22 , G.A. Gorelick, coll. COLORADO: Jackson Co: Sand Dunes (North Sand Hills SRMA), northeast of Walden, VI-18-1978, 1d, Peter Eades, coll. (probable blend) Larimer Co: County road 80C, 7.8 mi. E. Jct., county road 89, VII-6-1996, 1d, A.D. Warren, coll. (very worn specimen - probable blend) Weld Co: 7 mi. NW Carr, VI-13-1985, 1¢, P.A. Opler, coll.; § mi. NW Carr, VI-18-1985, 11, 42 , P.A. Opler, coll.; 9 mi. NE Grover, 5455', VI-8-1991, 19 , R_E. Stanford, coll.; Pawnee Buttes, Pawnee National Grasslands, 13 mi. south of Wyoming Border (north of Colorado state Hwy. 14), VI-21-1978, 1d, Peter Eades, coll. Callophrys affinis apama (W.H. Edwards) “ARIZ”, W.H. Edwards collection, designated lectotype by F.M. Brown, 1967, 1d “ARIZ”, W.H. Edwards collection, designated paratype by H.R. Clench, 1973, 19 ARIZONA: Apache Co: Greer, VI-12-1936, 1¢, G.H. & J.L. Sperry, coll.; Greer, White Mtns., VII-11-1995, 1¢, Ken Davenport, coll.; Wheatfields Creek, 8000', Chuska Mtns., VII-8-1978, 1¢, R.W. Holland, coll., Lukachukai Creek Campground, 7000', Chuska Mtns., VI-14-1971, 1¢, R.W. Holland, coll.;. McNary, VI-20-1981, 12, G.A. Gorelick, coll.; Imi. E. McNary, VI-21-1981, 93, 32, G.A. Gorelick, coll.; 2 mi. S. McNary, VI-21-1981, 12 , G.A. Gorelick, coll. Gila Co: Tonto Creek Campground, nr. Kohls Ranch, VI-26-1956, 19, no coll.; “Gila Co.”, VI-1902, 26, O.C. Poling, coll.; Sierra Ancha Mtns., VII-7-1933, 3d, 32, Harold M. Bower, coll; VII-4 & VII-10-1934, 26, D.K. Duncan, coll.; Globe, VI-4-1934, 1°, no coll.; VIT-11-1933, 24, no coll. Yavapai Co: School House Gulch, Prescott, VII-9-1993, 4¢, Ken Davenport, coll.; Prescott, VII-7-1917,1¢ , J. Gunder coll'n; VII-9-1993, 3¢ , 197 Ken Davenport, coll.; E. of Prescott, VIH-11-1952, 12, Lloyd Martin. coll; no locality, VII- 1928, 1d , O. Buchholz coll'n.; Wolf Creek, S. of Prescott. VII-20- 1959, 63, 3° , R.F. Sternitzky, coll. Greenlee Co: Strayhorse Campground, White Mountains, VII-28-30-1937. 25d, 492; VII-1-3-1937, 29d, 119, all Don Meadows, coll.; VII-7-1958. 12. J.W. Tilden, coll.; 8 mi. S. Strayhorse Cmpgd., 8000', Hwy 666, VI-25-1981, 152. 32, G.A. Gorelick, coll.; Rose Peak, VII-7-1958 12 , J.W. Tilden, coll.: 4 mi. S. Rose Peak, 7600', Hwy 666, VI-24-1981, 56 29, G.A. Gorelick. coll; White Mountains, VI-16-1947, 1d, 19 , E.P. Mellon II , coll., VI-28-1937, 12. M-E. Smith, coll., VI-26-1936, 1° ; VI-20 to 26-1936, 5d, 12; VII-30-1937. 12. all L.P. Grey, coll.; VI-22-1936, 26 ; VI-28-1937, 1d ; VII-1-1937, 3d, 19 , all T.-M. Dunkle, coll.; VI-26-1937, 4m, 1°; VI-1-1937, 3d, 12, T. Dunkle, coll.; VI-22- 1942, 33, 49 , H.A. Freeman, E.R. Hulbirt, colls.; VII-3& 4-1951, 3d, 12: VI-16- 1948, 1d, E.R. Hulbirt, coll.; White Mountains, VI-8-1929, 1d ,L.I. Hewes. coll: VII-18-1961, 26 , Keith Brown, coll.; Hannagan Mdws., White Mtns., VI-20- 1930, 36 , J.C. Hopfinger, coll.; VI-20-1937, 26, 2°, J. Baker, coll.; Blue Mtns.. VI-30-1933, 1c , E.R. Hulbirt, coll. Graham Co: Mt. Graham, 1882, 2¢ , Herbert K. Morrison, coll. (Skinner collection) (paratype); Graham Mtns., VI-4-1932, 16 , R.G. Wind, coll. Coconino Co: Lockett Lake Road, 7400', VI-5-1989, 1¢ , Ken Davenport. coll.; Flagstaff, VII-27-1916, 2¢, J.A. Comstock coll'n.; Schultz Pass, 7800', San Francisco Peaks, V-27-1960, 16, Lee D. Miller, coll.; 5 mi. NW Flagstaff, VI-4- 1967, 19; V-19-1968, 5d, 19: VI-11-1969, 1d, R. Funk, coll.; Schultz Pass Rd.. 4 mi. N. Flagstaff, V-29-1985, 5d, 49, G.T. Austin, coll.; VII-21-1982. coll. ex larvae on Ceanothus fendleri (emerged IV-5-1983), 43, 22; VI-30-1986, coll. ex larvae (emerged II to V-1987), 5d 19, Dave Daniels, coll.; Alpine Garden Club tract, 5 mi. NW Flagstaff, VI-14-1970, 8d, 3°, R.Wielgus, coll. Cochise Co: vic. Fort Grant, SW of Mt. Graham, 1d , W.H. Edwards collection (lectotype), Huachuca Mtns., VI-23 to 30, no year, 126, Barnes coll'n.; VI-6-1910, 49 ; VI-6-1916, 26 J.A. Comstock coll'n; VI-22-1910, 2¢, 22 . E.I. Huntington coll'n; IX-8-1915, 19 ; V-24 to 30-1919, 1d .W.G. Wright coll'n: V-16-1923, 1d, V-24-1930, 1d, R.C. Williams coll'n.; Carr Peak, Huachuca Mtns., VI-19-1940, 7d ; VI-23-?- , 2d, all A.C. Twomey, coll., Chiricahua Mins.. VI-20-30-1908, 64, 12 , Victor L. Clemence, coll.; VI-30-1916, 1¢ , VW. Owen. coll.; VII-10-1958, 2¢, 12 , J.P. & G.C. Hubbard, coll. Navajo Co: Santa Rita Mtns., 5000-8000', July, 1¢, F.H. Snow, coll.; VI-20- 1903, 12, Stephens, coll.; Pinal Mtns., VII-5 & 14-1900, 2d; V-30-1925, 1d, no coll.; Navajo Mtn., VI-14-1936, 1d, no coll., VI-19-1935, 22. R.G. Wind, coll: VII-12 & 13-1933, 4d, 12 , H.N. Hultgren, coll., VI-21-1935, 16, R.G. Wind. coll.; Williams Creek, North fork White River nr. McNary, V-29-1932, 1c: VI-7- 1932, 2d, 12, no coll.; White Mtns. Pima Co: Baboquivari Mtns., VII-15 to 30-1903, 2¢, 12 , O.C. Poling, coll.: VIII-1 to 15, 1924, 15d, 32, J. Gunder, coll.; Catalina Mtns., 7500', VI-13-1937. 46, 82, O. Buchholz coll'n.; 8500', VI-21-1936, 3¢, 12 , C.F. dos Passos coll'n.: Mud Springs, Santa Catalina Mtns., 6500', VII-17 to 20 -1916, 1¢, 2° . no coll.: Catalina Mtns., 8500', VI-22 to 26-1936, 2d, 2° , L.P. Grey, coll. UTAH: Garfield Co: Blue Spruce Camp, 18 mi. north of Escalante, 8000', VII-5- 1963, 1d, F. Rindge, coll. [AMNH] Iron Co: Burnt Peak Road, summit, 2412m, VI-21-1971, 1c, 22. J.A. Scott. coll. San Juan Co: Monticello, VI-25-1930, 1° , Ashby D. Boyle coll'n; 4.5 mi, SSW Buckeye Reservoir, 7700',coll. ex larvae on E. alatum (emerged VI-9 to 12- 1989), 9d , 32, J.L. Harry, coll.; Navajo Mtn., 6000', VI-16 to 20-1936, 21° . Allyn Museum collection. Kane Co: U.S. Hwy 89, 3.2 mi. N. Long Valley Jct., 7200', coll. ex larvae on E. racemosum (emerged V-21-1987), 6¢, 2°, J.L. Harry, coll. Wayne Co: Singletree Campground, 8300', 11 mi. So. Torrey, collected ex larvae on C. fendleri and E. racemosum (emerged IV-25-1987), 53, J-L. Harry. coll.; 7.4 mi. so. Torrey, 8800', N. Slope Rd., coll. ex larvae on C. fendleri and E. racemosum (emerged IV-20-1987), 1d, 19, J.L. Harry, coll. Grand Co: 4.4 mi. no. Mt. Waas, 18 mi. E. Moab, 8500', VI-26-1987, 1c . J.L. Harry, coll. NEW MEXICO: Catron Co: 2-3 mi. E. of Mogollon, 7200-7600', VII-11-1961, 12 ,. AMNH collection. McKinley Co: Grasshopper Canyon, 6800', 10 mi. S. Ft. Wingate, VI-26- 1981, 43, 29, G.A. Gorelick, coll.; Grasshopper Spring, 7500', Zuni Mtns., V-7- 1977, 29: V-23-1977, 3d; V-16-1976, 3d , all R.W. Holland, coll.; Tohatchi Peak, 8300', Chuska Mtns., VI-18-1978, 24, R.W. Holland, coll.; 1 mi. N. Tohatchi 198 Outlook, 8400', Chuska Mtns., VI-18-1978, VI-18-1978, 19, all R.W. Holland, coll.; Ft. Wingate, V-25 to VII-1-1909, 12¢, 8°, AMNH collection. Prop Cyn., 8000', Zuni Mtns., V-14 & V- 29-1976, 3c ; V- 30-1977, 2¢: VIII-15-1976, 1d , R.W. Holland, coll. Valencia Co: Pole Cyn., §000-8500', Zuni Mtns., V-6-1977, 1¢; V-30-1977, 2d ; VI-12 to 24-1977, 3d, 12; VII-29-1977, 16 , all R.W. Holland, coll.; Ojo Redondo, Zuni Mtns., V-16-1970, 12, R.Bailowitz., coll. Grant Co: McMillan Campground, 13 mi. N. Silver City, 6000-7000', VII- 16 to VII-19-1964, 250, 189 ;Cherry Creek Campground, 6900', 13 mi. N. Silver City, VII-18-1964, 3¢, all F., P., & M. Rindge, colls.; Cherry Creek Cyn. & McMillen Cyn., 6700-7300', Pinos Altos Mtns., June-July 1976, 286, 292, C.D. Ferris, coll.; Pinos Altos Mtns., Pinos Altos, VH-4 to 10 -1958, 3d, 12 , J.P. Hubbard, coll.; Pinos Altos Mtns., Cherry Creek, VII-6-1958, 3 II-19-1959, 1d , J.H., coll.; VII-10-1958,1¢, J.P. & G.C. Hubbard, colls.; Cherry Creek, 7100! Gila Nat'l Forest, VII-1-1977, 1¢ , C.D. Ferris, coll., vic. McMillan Cyn., 7400', Gila Nat'l Forest, VII-1-1977, 2° , C.D. Ferris, coll.; Signal Peak Rd., Gila Nat. For., VI-29-1978, 1d, C.D. Ferris, coll.; 12-14 mi. N. Silver City, 7000', VIT- 6-1985, 8° , C.D. Ferris, coll.; VII-3-1986, 4° ,7¢, G.A. Gorelick, coll. *Otero Co: Cloudcroft, VI-19-1902, 19, no coll.; High Rolls, V-22-?, 1¢, both in Carnegie Museum collection [CMNH] (°Received from the Engel collection, these very early captures bore no collector labels and are probably IV-29-1972, 1d; VI-4-1978, 1d ; mislabeled specimens) MEXICO: Chihuahua: Mun. Casas Grandes, head of Rio Piedras Verdes, 7200', 30 15'N, 108 15'W, 1899, 12, Townsend, coll. [CMNH]; Madera, 7200', VII-6- 1947, 2°, W. Gertsch & M. Cazier, colls. [AMNH]; VIII-1968, 2¢, T. Escalante, coll., Arroyo Mesteno, Sierra del Nido, 7600', VII-21-1959, 12 , W.C. Russell, coll; 10 mi. E. Namiquipa, VII-3-1947, 8¢ , M. Cazier & W. Gertsch, colls.[AMNH]; near Babicora, 5.7 mi. E. jct., road to Madera, VII-29-1984, 1° , D.Daniels, coll. Callophrys affinis apama/ C. a. homoperplexa blend NEW MEXICO: Los Alamos Co: Burnt Mesa, Hwy 4 west of Los Alamos, Bandelier National Monument, 1¢, VI- 20 -1997; coll. ex larvae on C. fendleri, VI-22-1997 (Giersed VII to IX-1997), 62, 22. G.A. Gorelick, coll. Sandoval Co: Jemez Springs, 7000', V-25 to VI-13-1913, 96, 132 ; V-21- 1921, 12 ; VI-2-1921, 32 , all J. Woodgate, coll., Bandelier National Monument, 7000', VII-17-1999, 2d, 22, S.J. Cary, coll. Callophrys affinis homoperplexa Barnes & Benjamin NEW MEXICO: Colfax Co: Dale Mtn., 8400', Johnson Mesa, VI-21-1997, 1¢, S.J. Cary, coll.; Tolby Campground, Hwy 64, 7000', June 2, 1996, 16, 12, G.A. Gorelick, coll. Santa Fe Co: Hyde State Park, § mi.NE. Santa Fe, 8700', Sangre de Cristo Mtns., VII-29 to 31-1964, 19 , F, P., & M. Rindge, colls.; Chupidero Cyn., 8450', Sangre de Cristo Mtns., VII-23-1934, 1d , M. Hebard, coll. Rio Arriba Co: San Antonio Mtn., 9000', VI-21-1978, 3¢, J.A. Scott, coll. San Miguel Co: near Hot Springs, 7000', Las Vegas, VII-1882, 1d, 29, F.H. Snow, coll. Union Co: Capulin Mountain National Monument, VI-28-1968, 19, no coll. COLORADO: Larimer Co: Horsetooth Mtn. Park., 5 mi. SW. Ft. Collins, V-6-1977, 1¢ , J. Buchholz, coll.; VI-23-1985, 2d ; V-5-1985, 1d , R.E. Stanford, coll.; V-30 to VI- 8-1987, 5d, P.A. Opler, coll.; VI-29-1997, 29, P.A. Opler, coll., VI-29-1997, 2¢ , G.A. Gorelick, coll.; Lory State Park, 5 mi. W. Ft. Collins, VI-8-1985, 2d , P.A. Opler, coll.; VI-29-1997, 26, Andrew D. Warren, coll.; “A” Mountain on Horsetooth Res. Rd., V 1-25-1995, 19, A.D. Warren, coll.; 4 mi. W. of Hwy. 1-25 on Woodman Rd., VI-2-1992, 1¢, A.D. Warren, coll. Jefferson Co: Apex Trail at Heritage Square, V-13-1990, 1d; V-11-1991, VI-6-1990, 43, A.D. Warren, coll.; Indian Hills, nr. Denver, VI-21-1936, 1d, W.D. Field, coll.; Mother Cabrini Shrine, Hwy. 40 W. of Denver, VI-10-1961, 2¢ DN 14-1966, 19, RJ. Jae, coll.; VII-5-1965, 12 , John A. Justice, coll.; V-26-1990, 2d, A.D. Warren, coll., Chimney Gulch, V-24 to30-1907, 3¢ ; VI-8-1915, 2d , E. Gane coll. (from Barnes collection, incl. 1 paratype); V1-6-1937, 1d ; V-13- 1938, 1¢ , Bob Potts, coll.; Chimney Gulch, 1/4 mi south, V-6-1966, 1d , J.A. Scott, coll.; V-25-1927, 1d , E.I. Huntington coll'n; VI-6-1937; Rooney Ranch, VI-17-1956, 1°, R.J. Jae, coll.; Lookout Mtn., nr. Golden, 6500-7200', VII-8- 1967, 5 VI-11 to 18-1968, 7, 29 , all Mike Fisher, coll.; Castle Rock, near Golden, V-21-1966, 1d , J.A. Scott, coll.; Golden, V-24 to 30 (no year), 2d ; VI- 8 to 15 (no year), 2¢ ; V-16 to 23 (no year) , 1d , V-24-1930 , 1d, all Bares and JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Benjamin (paratypes); Golden, VI-1910, 2¢, F. Skinner, coll.; V-28-1939, 2°, C.C. Albright, C.D. Schryver, colls.; VI-11-1939, 1d, C.C. Albright, coll.; V-2- 1943, 1d, C.D. Schryver, coll.; Clear Creek Canyon, V-8 to 15-1907, 1¢ , Ernest J. Oslar, coll.; Clear Creek, 6000-7000', VI-20-1922, 1d , G.P. Engelhardt, coll.; VII-3-1984, 1¢ , P.A. Opler, coll.; Red Rocks Park, 6000', V-11 & 25-1956, 26 , R. J. Jae, coll.; V-24-1964, 1d, 19 , VI-4-1966, 2<, Ns Scott, coll; Red Rocks Park, VI-10-1971, 1d , R.J. Jae, coll.; Red Rocks Park, 2 mi. NW Morrison, VI- 5-1968, 1d, 12, Mike Fisher, coll.; Ralston Butte, V-23 & 24-1994, 2. 1988, 4¢ ; VI-10-1994, 26 ; VII-10-1985, 29, all J.A. Scott, coll.; sea, V-26- 1984, -15 to VI-25 (1984-94), 17d, 3; 72 , all J.A. Scott, coll.; Tinytown (trail to Mt. Lindo), V-31-1993, 1d, 19; VI-7-1992, 19; V-27-1991, 192; V-29-1989, 1d; VI-9-1998, 1¢, all A.D. Warren, coll.; Lookout Mtn., V-29-1988, 2¢ ; VI-15-1995, 5d , J.A. Scott, coll.; Indian Peak, V-14-1994, 2¢ , J.A. Scott, coll.; E] Dorado Mtn., VI-11- 1994, 1d, J.A. Scott, coll.; Zion, 2100m, V-28-1991, 1d , C.P. Slater, coll.; Mt. Zion, V-30-1981, 1¢; V-14-1994, 2d;V-26-1984, 22, ].A. Scott, coll.; VI- 1-1990, 1d, 19; V-28-1989, 2d; V-30-1991, 1d; V-28-1998, 26, A.D. Warren, coll.; Guy Hill, VI-18-1992, 19 , J.A. Scott, coll.; Falcon County Park, V-30-1984, 1d ; VI-15-1989, 2d; VI-6-1994, 2°, J.A. Scott, coll.; Mt. Falcon, V-27-1994, 29, J.A. Scott, coll.; Tucker Gulch, VI-12-1982, 1¢ , C.P. Slater, coll.; VI-15-1995, 1d , J.A. Scott, coll.; Apex Gulch, VI-5-1990, 2¢ ; VITI-20-1990, 1¢ ; VI-20-1995, 4¢ ; V-19-1994, 1d V-29-1991, 2¢ , all J.A. Scott, coll.; Coal Creek, VII-17-1991, 1d, J.A. Scott, coll.; Green Mtn., V-10 and VI-3-1986, 2¢, J.A. Scott, coll.; Crawford Hill, VI-5-1980, 5d, 12, J.A. Scott, coll.; N. Crawford Gulch/Canyon, V-30-1981, 1d; V-24-1988, 1¢ , J.A. Scott, coll.; Golden Gate Cyn., VI-12-1956, 1d , Lincoln F. Brower, coll.; ° Type locality of C. a. homoperplexa. Boulder and Denver Boulder Co: Eldora, VI-15-1941, 12, P.S. Remington, coll.; Boulder, VI-20- 949, 32 , J.C. Hopfinger, coll.; “July” , 1¢ , (paratype), no coll.; Boulder Canyon, VI-18 to 20-1933, 19, 1d, P.S. & C.I. Remington, colls.; Mtns. No, of Boulder Cyn., VI-2-1965, 2° , John Justice, coll.; Baird Park, Gregory Cyn., V- 23-1965, 1d , J.A. Scott, coll.; nr. Baird Park, Gregory Cyn., VI-7-1964, 12, VI- 14-1965, 1¢ , John Justice, coll.; V-4-1965, 2d, V-12 to 19-1965, 6d , J.A. Scott, coll.; Gregory Cyn., 5900', V-29-1954, 16; V-20-1966, 1d, J. Don Eff, coll., V-23- 962, 1d; VI-3-1962, 19; V-19-1966, 1d, J.A. Scott, coll.; V-1 to 6-1966, 8¢, 12 , S.A. Johnson, coll.; V-20-1970, 1¢ , R.J. Jae, coll.; V-4 to 23-1965, 1¢ , J.A. Scott, coll.; IV-28 to V-30-1966, 21d, 22 , J.A. Scott coll.; Cardinal Hill, VI- 1-1957, 6, J. Don Eff, coll.; Sunshine Cyn., 7200', VI-1-1947, 3d , V-28-1953, 1d, all J. Don Eff, coll.; V-30-1962, 22 , J.A. Scott, coll.; 1 mi. NW Nederland VII-2-1989, 3 , J.A. Scott, coll.; nr. summit, Flagstaff Mtn., VIH-15-1975, 1d , J. Vernon, coll.; Flagstaff Mtn., V-23-1948, 2d; VI-19 to 21-1953, 2d , all J. Don Eff, coll.; V-30-1961, 3d , J.A. Scott, coll.; V-29-1951, 26 , O. Buchholz coll'n ; Four Mile Cyn., VI-17 to 19-1953, 42, 3d; V-16 to 19-1958, 19, 26; VI-25-1965, 19; V-15- 966, 2¢, 12, all J. Don Eff, coll.;V-9-1959, 2¢ , J. Don Eff, coll.; Four Mile Creek Cyn., nr. Crismon, V-16 to 24-1958, 6d, 12; VI-5-1959, 2d; V-2 to 17-1962, , 22, all J. Don Eff, coll.; Packer's Gulch, nr. Crismon, V-15-1966, 1d, J. Don Eff, coll.; Lefthand Cyn., V-4-1932, 1d ; V-30-1951, 1d, V-24-1953, 53; V-29- 954, 2d, all J. Don Eff, coll.; VI-28-1952, 16, O. Buchholz collection; V-23- 962, 1d; VI-9-1968, 1d; VI-8-1994, 1d, J.A. Scott, coll.; Magnolia Road, VI-13- 972, 1d , J. Don Eff, coll.; 2 mi. SW Boulder, 6500', V-18-1963, 1d, U. Lanham, coll.; Sugarloaf, V-30-1955, 24, J. Don Eff, coll.; V-20-1958, 22, E.R. Hulbirt, coll; Iron Mtn., Sugarloaf Rd., V-13-1962, 1d, J. Don Eff, coll. Denver Co: Ruby Hill, 5390', V-25-1994, 19, R.E. Stanford, coll [GM-CSU] Gilpin Co: No. Clear Creek, 7200', VI-2 to7 (1977-8), 4d, 19, J.A. Scott, coll.; VII-3-1982, 1d, B.E. Stanford, coll.; East Portal, Moffat Tunnel, VI-19- 1977, 1d , J.A.Scott, coll.; Aspen Springs, 8900', VI-8-1985, 16, R.E. Stanford, coll., Blackman Meadows Trail, 8400-8700', Golden Gate State Park., V-24- 1975, 1d, R.E. Stanford, coll. Gunnison Co: Almont, VI-20 to 30-1925, 2° , J.D. Gunder, coll.; One Mile Camp, § mi. NE Almont, 8200', VII-8-1957, 2d, 19, F. & P. Rindge, colls.; 5 mi. W. Gunnison, 7000', VI-17-1966, 1¢, 12 ; Iola, 7600', VI-19-1967, 12 ; Curecanti Creek, 8000', VI-2-1962, 2d, all Scott L. Ellis, coll.; Blue Mesa Reservoir, jct. Hwys 50 &149, VI-4-1996, 3d, G.A. Gorelick, coll. Logan Co: County Road 9, 6.5 mi. N. of Ft. Morgan, chalk bluffs, V-28- 2000, 1¢, 12, R E. Stanford, coll. Morgan Co: S/N scarps, Wildcat Creek, SR 52, 7-11 mi. N. of Ft. Morgan, V-28-2000, 4d, R.E. Stanford, coll. Adams Co: | mi. W. fairgrounds, 5100', V-25-1994, 1d, R.E. Stanford, coll. Custer Co; Davenport Camp, 36 mi. So. Florence, 8500', VI-30-1967, 3d, 12, F., P., & M. Rindge, colls.; Silver Cliff, VI-29-1968, 1d , J.A. Scott, coll.; Paratypes exist also from VOLUME 59, NUMBER 4 Washout Creek, VII-4-1967, 1d , J.A. Scott, coll.; Smith Creek Campground., VI-13-1971, 1°, J.A. Scott, coll.; _ mi. E. Smith Creek Campground., Hwy 82, VII-28-1970, 1d , J.A. Scott, coll.; VI-3-1996, 1d , G.A. Gorelick, coll.; Bull Domingo Mine, VI-18-1973, 1¢, J.A. Scott, coll.; Ben West Hill, VI-12-1970, 1¢ . J.A. Scott, coll.; 3.2 mi. W. of Wetmore, Lewis Creek Trail, VI-11-1991, 12, A.D. Warren, coll. Fremont Co: | mi. So. Cotopaxi, VII-1-1969, 1¢ , J.A. Scott, coll. Park Co: nr. Antero Jct., 9300-9500', VI-23-1973, 1d, R.E. Stanford, coll. Arapahoe Co: Piney Creek, 6000', VI-2-1974, 1¢; VI-10-1979, 2d, R.E. Stanford, coll.; Piney Creek, Smoky Hill Rd., 6700', VII-4-1965, 1¢, J.-A. Scott, coll.; V-20-1993, 5d; V-28-1993, 3d; VI-5-1992, 1d; VI-6-1992, 2d; V-29-1994, 3d, A.D. Warren, coll. Pueblo Co: 2 mi. up Greenhor Trail, Greenhorn Campground, VI-22- 1967. 1¢, J.A. Scott, coll.; Greenhorn Trail, w. of Rye, VI-4-1971, 2¢, J.A. Scott, coll.; Beulah, VII-7-1899, 1¢, W.D. Kearfott, coll.; VI-12-1970, 4¢, J.A. Scott, coll.; 4 mi. NW Beulah, VI-29-1970, 12, J.A. Scott, coll.; Los Animas Co: near Weston, VII-13 to 19-1975, 2¢, 12, J. Vernon, coll. Douglas Co: Indian Creek Campground., VII-6-1995, 1d, J.A. Scott, coll.; Mitchell Gulch @ Founders Village, 6400', VII-31-2000, 33; Surrey Ridge, VII- 17-1989, 1¢; Hidden Pointe area, ca. 8 mi. NNW Castle Rock, VI-13-1998, 1¢; VII-4-1998, 19: VI-11-1998, 1d; VI-6-1998, 12; VI-18-1998, 26; VI-7-1998, 12; VII-16-1998, 16; VI-24-1999, 3d, 22: Newlin Gulch, ca. 8 mi. NNE Castle Rock, VI-28-1999, 26, 12.; VIH-17-1999, 4¢; canyon N. of Palmer Lake, VII-29-2000, 1d; Daniels Park, VII-12-1998, 1d, 12; V-31-1998, 1d; McMurdo Gulch, 3 mi. NE of Castle Rock, VI-17-1999, 26; Hunt Mountain, ca. 6 mi. S. of Castle Rock, VII-24-1999, 1¢; VII-11-1999, 2d; VI-26-1999, 33, 12; Wolfensburger Road, ca. 2 mi. W. of Castle Rock, VI-19-1999, 32; mouth of Jarre Canyon, VI-4-1998, 1d, all A.D. Warren, coll. Elbert Co: Running Creek Field Sta., 6950', T9S, R65W Sec 26, SE 1/4, VI- 25-1976, 12, S. Condie, coll.; Pohl Ranch, 6850', T6S, R65W, Sec 26, SW 1/4, 1d, no coll.; near Elbert, 7000-7400', VI-26-1976, 1d , R.E. Stanford, coll.; south side of County Line Rd., 6.2 mi. E. of Gun Club Rd., VI-10-1993, 3¢, A.D. Warren, coll. Teller Co: 4 mi. SW. of Florissant, VI-26-1989, 1d, 12, A.D. Warren, coll.: Florissant Fossil Beds, 8550', R70W, T13S, Sec 30, Maytag, VI-22-1976, 1d, S. Condie, coll. Ouray Co: 4 mi. N. Ouray, 7400', VI-5-1966, 1¢, F. M. Brown, coll. El Paso Co: Mt. Hermon Mon. Area, 7500', VII-8-1956, 4¢, no coll.; Star Ranch, VI-26-1931, 1d, F.M. Brown, coll.; VI-27-1965, 1d, no coll.; 4 mi. W. of Hwy. I-25 on Woodman Rd., VI-2-1992, 1d, A.D. Warren, coll. Chaffee Co: nr. Mt. Princeton Hot Sprs., Hwy 162, 7500', VI-20-1966, 1¢, 12, Scott L. Ellis, coll.; 1 mi. WSW of Buena Vista, VI-13-1992, 2, A.D. Warren, coll. Dolores Co: West Fork Camp, 20 mi. NW Dolores, 7800', VI-30-1957, 3¢, F. & P. Rindge, colls. Clear Creek Co: Fall River Rd., 2 mi. W. Idaho Sprs., 7500', VI-4-1967, 1d; VI-3 to 8-1968, 2¢ , Mike Fisher, coll.; Clear Creek, 7000-8000', V-27-1979, 1¢ . RE. Stanford, coll. Montrose Co: Alpine Trail Rd., VI-22-1973, 1¢, no coll.; north .rim, Black Canyon National. Monument, 7809', VI-30-1967, 1d, 29, Scott L. Ellis, coll. Lincoln Co: N. microwave tower, 6.5 mi. W. CO Hwy 71, 5850', VI-30- 1993, 12, R.E. Stanford, coll; CO Hwy 71, 5400', 21.6 rd.mi. N. Limon, Li/Wa Co. line, VI-30-1993, 1d, R.E. Stanford, coll. Archuleta Co: NE of Pagosa Sprs., June 30, 1965, 1¢, sight record, JA. Scott Saguache Co: County Road 31 above Vulcan, west slope portion of county, VI-16-2001, 2c, Michael S. Fisher, coll. Callophrys affinis albipalpus ssp. nov. NEW MEXICO: Lincoln Co: Cedar Creek Camp, 5 mi. N. of Ruidoso, 7000', Sacramento Mtns., VI-30-1961, 2d, F., P., & J. Rindge, colls.; New Mexico State University's Montgomery Biological Research Laboratory, 5 mi. n. of Ruidoso, 7000’, Sacramento Mtns., VII-26 to 28-1978, 22 , VII-21 & 22-1980, 2° , all Greg Forbes, coll. (R.W. Holland collection.); VII-4-1981, 2 G.A. Gorelick, coll.; VI-28-1982, 1d, ex ovum (Emgd: IV-27-1983); VII-1982, 72, 72, ex ova reared on lab host, Eriogonum grande rubescens (Emgd: VIII-1982) Callophrys affinis chapmani ssp. nov. MEXICO: Sonora: Mun. Nacori Chico, 7300',11.6 mi. N.of Mesa Tres Rios and 44.9 mi. so. of Huachinera, 108 49'W, VII-2-1979, 12, R.W. Holland, coll., 13.7 mi. 199 N. of Mesa Tres Rios and 42.8 mi. S. of Huachinera, 7600'. 108 49'W, VIJ-2- 1979, 1d , R.W. Holland, coll. Durango: E] Salto, 27.7 mi. E., 8000', VII-18-1964, 12, J-A. Powell. coll.; VII-11-1981, 12, R.W. Holland, coll., VII-15-1981, 1¢,12, G.A. Gorelick, coll: VIII-5-1981, 12, G.A. Gorelick, coll., VII-10/11-1983, 2¢, 42, G.A. Gorelick, coll., VII-15-1983, 29, G.A. Gorelick, coll.; 13 mi. E. El Salto, Hwy. 28, 8000". VII-10-1983, 192, G.A. Gorelick, coll.; Mun. Llano Grande, 16-17 mi. E. El Salto, Hwy. 28, VII-13-1985, 2d, 32, coll. ex ova on Eriogonum atrorubens (Emgd: VII-28-1985) G.A. Gorelick, coll; 16.3 mi. E. El Salto, Hwy. 28, [V-14- 1986, 1°, Greg Ballmer, coll. ex larva on Eriogonum atrorubens. 200, Journal of the Lepidopterists’ Society 59(4), 2005, 200-211 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY CLOUD FOREST BUTTERFLY FAUNA OF THE PANTEPUI— POOR OR POORLY KNOWN? DESCRIPTION OF NEW SPECIES AND RECORDS OF NEW GENERA OF PRONOPHILINA: ERETRIS AGATA AND OXEOSCHISTUS ROMEO (NYMPHALIDAE: SATYRINAE) TOMASZ W. PYRCZ Zoological Museum of the Jagiellonian University, Ingardena 6, 30-060 Krakow, Poland email: pyreztomasz@hotmail.com AND STEVEN FRATELLO 11 First Street, W. Islip, New York, USA email: sfratell@suffolk.lib.ny.us ABSTRACT. Two new species of Pronophilina (Nymphalidae, Satyrinae) - Eretris agata and Oxeoschistus romeo - are described from the Guyana shield, also known as the Pantepui, a region of table mountains situated in southeastern Venezuela, northwestern Guyana and adjoin- ing Brazil. They are only the third and fourth genera of the diverse neotropical montane subtribe Pronophilina (Nymphalidae, Satyrinae) re- ported for this vast biogeographical region. Their affinities are evaluated indic: ating close relationship with the Pronophilina of the Venezuelan Cordillera de La Costa. The current state of knowledge of the Pronophilina in the Pantepui is discussed. Low figures of diversity are interpreted as a result of isolation from the center of origins and diversity in the central Andes, and partly as a consequence of heavy undersampling. Additional key words: Andes, affinities, bamboo, endemic species, Guyana shield, island biogeography, Protopedaliodes, species richness. INTRODUCTION Pantepui is a term originally proposed by Mayr and Phelps (1955) to designate the region of table mountains situated in southern Venezuela and adjacent areas of Brazil and Guyana. Even though throughout the years there were many contradictions and various definitions of Pantepui (Huber. - 1987: Brown, 1979), all the authors concurred that this vast area is worth identifying as a separate biogeographical entity identified by a number of biotic (i.e. endemic taxa) and abiotic features. Nearly a century ago, Strand (1912) described from the Pantepui the first species of butterfly belonging to the predominantly Andean subtribe Pronophilina (Miller, 1968; Adams, 1985; Pyrez & Wojtusiak, 2002)— Pedaliodes roraimae (see fig. 5). His discovery passed completely unnoticed (\ filoria & Pyrez, 1995) ) and only recently, during the last decade, was there an increase of interest in the butterfly fauna of the Pantepui in general (Neild, 1996) and the Pronophilina in particular, resulting in the description of several new taxa. Four of them, Pedaliodes demarmelsi Viloria (1995), P. chaconi Viloria (1998), P. terramaris Viloria & Pyrez (1999) and P. yutajeana Viloria & Pyrez (1999) are closely related to each other and to P roraimae. Protopedaliodes kukenani, described by Viloria & Pyrez (1994) from the top of Kukenan and Roraima, represents a well differentiated endemic genus. Viloria & Pyrez (1999) described a further two species belonging to this genus: one from Roraima — Pr. ridouti, and one from the Auyan Tepui — Pr. profauna. All of the species described hitherto from the Pantepui belong to the Pedaliodes complex (Viloria, unpublished Ph.D. thesis). The two new species described herein belong to other genera and sections of the subtribe Pronophilina previously not reported in South America outside the Andes and their peripheral ranges (Sierra Nevada de Santa Marta, Cordillera de La Costa) — Eretris Thieme (1905) and Oxeoschistus Butler (1867). Their discovery is very significant as it emphasizes how little the Pronophilina fauna of the Pantepui is known, how much research is still to be done, and also raises interesting zoogeographical issues. Subtribe Pronophilina The genera Eretris and Oxeoschistus belong, according to Miller (1968), to the neotropical subtribe Pronophilina (considered by Harvey (1991)and Lamas et al. (2004) as a sub-tribe within the tribe Satyrini), an entirely neotropical section of the worldwide subfamily Satyrinae (Nymphalidae). The Pronophilina can he divided into three gr oups based on ecological and morphological criteria: Chileno-Patagonian (considered by Lamas & Viloria 2004 as belonging to the predominantly Australian tribe Hypocy stini), Caribbean (only the genus Calisto Lathy) and tropical montane. The latter dominant section comprises approximately 460 species (Lamas et al., 2004) distributed in the cloud forests and paramos of Central and South America. In continental Central America 18 species are restricted to the highest mountains of Panama and Costa Rica, and 10 to Guatemala and the neighboring states of Mexico. In South America, the vast majority, more than 95%, VOLUME 59, NUMBER 4 occur in the Andes and its peripheral ranges: the Sierra Nevada de Santa Marta and the Cordillera de La Costa. Seven species are found in the highlands of southern Brazil. The second region outside the Andean Cordilleras where tropical montane pronophilines occur is the Pantepui. The species belonging to the subtribe Pronophilina occur in well-defined and sometimes very narrow bands of altitude. Their altitudinal zonation has been attracting the attention of naturalists since the early twentieth century (Fassl, 1911, 1915 and 1918; Kriiger, 1924 and 1925), and recently was more extensively discussed by Adams & Bernard (1977, 1979 and 1981), Adams (1985 and 1986) and Pyrez & Wojtusiak (1999, 2002). Endemism is also a notable feature of the Pronophilina. In each north Andean range, between 10% and 30% of taxa are endemic at the specific level (Adams, 1985). The endemism ratio is higher in most isolated units, such as the Sierra Nevada de Santa Marta, the Cordillera de Mérida and also the Pantepui. Adams (1985) concludes that the average altitude range of the endemic species in all three Colombian Cordilleras is set higher, close to the upper forest limit, than that of the non-endemic species. However, Pyrez & Wojtusiak (1999) show, based on sampling in western Colombia, that this is not always the case. The highest diversity of the subtribe is reported close to the upper forest limit, varying from range to range, but generally falling at 2600-3000m (Adams, 1985; Pyrez & Wojtusiak, 1999: 2002). Several species occur in the paramo grassland above the timberline as high as 4500m. Very few Pronophilina were reported below 1000m. Adult Pronophilina, like other satyrines, are mostly sedentary (Adams, 1986; Pyrez & Wojtusiak, 2002) and even inside the cloud forest they are generally restricted to particular habitats (Adams, op. cit.). Most species show restricted vagility and even though this aspect of their behavior was not studied rigorously. field observations indicate that adults move little vertically or horizontally and keep close to their host plants or roosting places (DeVries, 1987). The biology of the Pronophilina remains largely unexplored but it appears that their larvae feed mostly on montane bamboo, in the Andes chiefly belonging to the genus Chusquea (Poaceae) (Schultze, 1929: Adams & Bernard, 1981; DeVries, 1987; Pyrez & Greeney, in prep.). However, Pelz (1997) reared an Ecuadorian species, Parapedaliodes parepa (Hewitson), on a substitute secondary grass, Poa festuca (Poaceae), whereas Pyrcz et al. (1999) report that several species also use Guadua bamboo as their host plants. There are indications that the Pronophilina, or at least some of them, are oligophagous. In Ecuador, Pyrcz (unpubl.) reared the 201 first to fourth instars of Junea doraete (Hewitson) on several different species of Chusquea. Bamboos are most abundant in clearings, roadsides, at the forest edge and along paths. Adults of all species of Pronophilina are attracted to decomposing organic matter, particularly to feces, carrion and rotten fruits. Therefore, the use of baited traps provides a very good method of sampling Consulted collections: BMNH: The Natural History Museum. London, United Kingdom. MBLUZ: Museo de Biologia de la Universidad del Zulia, Maracaibo, Venezuela. MCC: Mauro and Clara Costa, Caracas, Venezuela. MIZA: Museo de Entomologia. Universidad Central, Maracay, Venezuela. MZUJ: Muzeum Zoologiczne Uniwersytetu Jagielloriskiego, Krakow, Poland. SIW: Smithsonian Institution, Washington, USA. TWP: Tomasz Wilhelm Pyrez, Warsaw, Poland. Acronyms used: FW: forewing. HW: hindwing. D: dorsal surface. V: ventral surface. Eretris agata Pyrcz, new species (Figs. 1, 2, 6, 14) Diagnosis: Compared to allied species: E. encycla (figs. 7, 11-13, 15), E. oculata (fig. 16) and E. calisto (fig. 17)(all C. & R. Felder), the male of E. agata has no red markings along the VHW postmedian band’s basal edge. The upperside is darker —blackish—not dark choc olate brown (which is noticeable in freshly emerged individuals). It is larger than nominotypical E. encycla from the Cordillera ‘de la Costa, being approximately the size of the Cordillera de Mérida population. But E. agata has bigger ocelli than both and a different HW shape, somewhat ovaloid rather than squarish. Description: Male (Fig. 1): Head: Eyes chocolate brown, lustrous, covered with dense and long setae; labial palpi covered with dark brown hair; antennae 8-9mm, orange brown, basal half covered ventrally with white scales, club only slightly thickened compared to shaft, two terminal segments black. Thorax: dorsally blackish brown, legs beige, tibiae cov ore ventrally with gray hair. Abdomen: dorsally and laterally blackish. brown, ‘ventrally gray. Wings: FW length: 22-24mm, mean: 23.2mm; n=5; apex blunt, distal margin straight. HW outer margin slightly undulated, inner margin incised below tornus. DFW ground color glossy blackish brown in freshly emerged individuals, gradually fading and becoming lighter as the wings are exposed to sunshine: distal one third with a distinctively lighter shade from costa to tornus; a nearly straight darker submar. ginal line running parallel to distal margin; fringes dark brown. DHW ground color same as on the FW, a very slightly lighter shade submarginally at apex. occasionally extending along submarginal area, especially in older individuals; a thin parallel undulated blackish brown submarginal line, darker than the ground color; an even thinner marginal line parallel to outer margin; fringes dark brown. VFW ground color glossy brown, lighter than on the upperside; four oblique dark brown lines, a median one, slightly arched across discal cell, a postmedian one from costa towards anal margin, fading away before touching it, nearly straight except for two curves between vein Cul and anal margin; a submarginal one, thinner than two basals, parallel to distal margin, slightly Srecular from costa to vein M2, then nearly straight: a marginal line, very thin, parallel to the distal margin. VHW ground eailee from base to postmedian line dark brown, slightly suffused with tiny golden scales along inner margin and postdisc: al area, which is noticeable only in freshly emerged individuals: a shallowly curved dark brown median line, darker than the ground color; a thicker chocolate brown postmedian line from costa to inner margin near tornus, roughly parallel to distal margin, slightly irregular and undulated but 202 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 1-5, Some Pronophlina from the Pantepui. without any deeper curves; a series of six submarginal rounded ocelli ringed with light orange and with small white triangular pupils, one in each cell from Rs—M1 to Cu2-1A, the biggest in Cul-Cu2, the second biggest in Rss—M1, the remaining approximately the same size, half the size of the b iggest, not touching the lines basad or distad of them, except the one in Cu2-1A, which crosses the submarginal line; a chocolate brown submarginal line, parallel to distal margin, thinner than the postmedian one; a thin dark brown marginal line parallel to outer margin. Genitalia (Fig. 6): Uncus stout, ended with a short hook; gnathos short, less than one-fourth the length of uncus, slat-like; valvae long and thin with a characteristic shallow curve in the middle; aedeagus tubular, simple; saccus shallow. ; Female (Fig. 2): Head, thorax and abdomen as in male. Wings: 0 1. Eretris agata female male dorsum/venter; dorsum/venter; 3. Oxeoschitus romeo, n. sp., male dorsum/venter; 4. Oxeoschistus romeo, n. sp., female dorsum/venter; 5. Pedaliodes roraimae male dorsum/venter n. sp., 2. Eretris agata, n. sp., Wing shape as in male. FW length 23.5mm. DFW ground color dull dark brown, slightly lighter than in the male; color light brown distally from a postmedian oblique darker line running from costa to inner margin near tornus; a dark brown submarginal iin, parallel to distal margin; a thin dark brown marginal line. parallel to distal margin. DHW ground colour dark brown, same as in basal two-thirds of ‘the FW; a aia red postmedian line with a dark brown distal edge roughly parallel to distal margin from mid costa towards inner margin fading away at vein Cu2: a double dark brown and light brick- red submarginal scalloped line parallel to outer margin, fading away between vein Cul and tornus; a thin yellow marginal band edged basally with dark brown; a prominent oval submarginal ocellus in cell Cul-Cu2, ringed with yellow and with a white pupil. VEW ground color medium brown, light brown distally from the postmedian line; VOLUME 59, NUMBER 4 203 _ ‘. Fics. 6-9. Male genitalia of Pronophilini. 6. Eretris agata, n. sp. (aedeagus extracted in lateral and dorsal view). 7. Eretris encycla encycla (aedeagus extracted in lateral and dorsal view). 8. Oreeschicne romeo, n. sp. (aedeagus extracted in lateral view, valva extracted in dorsal view) 9. Oxoeschistus puerta ssp. (aedeagus extracted in lateral view). the pattern of dark brown lines as on the upperside but edged with crimson red, the median and submarginal ones distally, the postmedian one basally; a thin yellow marginal band, edged basally with dark brown; a small rounded subapical ocellus in cell MI1-M2, ringed with yellow and with a minute white pupil. VHW ground color medium brown; the pattern of dark brown lines as in the male, except that median and submarginal ones are edged distally with a thin crimson red line, whereas the postmedian is edged basally with a band of the same colour which overshadows most of the brown; submarginal ocelli of the same size and color as in the male; a thin yellow marginal basally dusted with dark brown. Type material: Holotype ¢: Venezuela, Estado Bolivar, Sierra de Lema, road El] Dorado - Santa Elena de Uairén km 132, 1983, 13 50-1400m, M.C. Costa leg., MIZA Maracay; Paratypes:9 6 and 1 : same locality as the holotype, 22 93.11.2004, M.C. Costa & T. Pyrez leg. (2d in MCC, 1 ¢ in BMNH, 1d in MIZA,5¢and12 in TWP). Etymology: This species is dedicated to Agata, a daughter of its discoverers Mauro and Clara Costa, Italo-Venezuelan lepidopterists from Caracas. Behavior: Eretris agata was discovered along the E] Dorado — Santa Elena de Uairén asphalt road between kilometer 130 and 135 (figs. 19-23). All the individuals were collected along the cloud forest edge when moving from inside the forest to the roadside, apparently looking for sunning spots. Several stands of an unidentified bamboo, its most likely host plant, were found some 20-30 meters inside the forest and more individuals were observed there. They were keeping to sunny gaps, occasionally patrolling and perching on large leaves at 3-5 meters above the ground. In the same habitat another species of Pronophilina was observed, Pedaliodes roraimae showing similar behavior but venturing more often into open areas, as well as another cloud forest Euptychiini satyrine, Forsterinaria sp. Remarks: The genus Eretris (Thieme) comprises approximately fifteen species distributed in South and Mesoamerica (Lamas, et al., 2004). Three species are 204 found in Guatemala/Mexico and Panama/Costa Rica, and the remainder in the Andes. There has been no cladistic analysis of Eretris, but the genus is morphologically highly homogenous and_ presents several plausible synapomorphies. It can be recognized from other Pronophilina by several characters emphasized in the original generic description (Thieme, 1905), such as short antennae reaching roughly one third the length of costa and the lack of a FW upperside male androconial patch. The most outstanding feature of the color pattern are fully developed submarginal ocelli on the VHW and a rarely present vestigial M1-M2 ocellus on the VFW. The wing venation with the HW cross-vein ml—m2 bent inside discal cell is typical of the Pronophilina (Miller, 1968; Viloria, Ph.D.). Male genitalia of Eretris are very particular and can be recognized by a short, flattened, singularly sculptured aedeagus, short subunci and _ thin slat-like sculptured valvae. The systematics of Eretris is very demanding because the differences between most species are very subtle. Additionally there is considerable infraspecific (Adams, 1986) and individual variation. The most obvious and significant taxonomic characters in the color pattern are all on the VHW: the shape and color of postmedian, non- submarginal and marginal lines, the size and number of submarginal ocelli and the color between the submedial line and the anal margin. Oxeoschistus romeo Pyrcz & Fratello, new species (Figs. 3, 4, 8) Diagnosis: This species is immediately recognized from other congeners by the highly irregular shape of the FW orange band char acterized by deep basal and distal intrusions compared to regular, nearly linear, edges of other related species, especially O. simplex Butler and O. puerta (Westwood) (fig. 9). Description: Male (Fig. 3): Head: eyes ‘Blaclash brown, setose; palpi white covered with black hair; antennae two fifths the length of costa, dark brown, club formed gradually, slightly thicker than ‘shaft. Wings: FW length 31mm; apex blunt; distal margin slightly convex; HW outer margin scalloped. FW and HW fringes medium brown. DFW basal and postbasal area medium brown; median area crimson red progressively darkening distally; a wide, brick orange band extending from postmedian to subm: irginal area with an irregular basal e dge and deep incursions of dark brown along costal margin in the subapical area, along vein M3 and in cell Cu2—-1A, and a dentate distal edge; dark brown rounded submarginal spots M1-M2, M3-Cul and the largest of all in Cul—Cu2; submarginal area dark brown becoming gradually lighter towards distal margin. DHW basal and median area medium brown; a wide brick orange band extending from postmedian to submarginal area with a nearly ‘straight i inner edge except for a shallow postdiscal incision and a dentate distal ec dge; a series of dark brown rounded spots situated in the middle of the band smaller than on the FW in Rs—M1, M1—-M2, M3—Cul, Cul—Cu2 and two in Cu2-1A, all of them surrounded by a crimson red halo; submarginal area dark brown reaching distal margin from costa to vein M2; an orange marginal band from vein M2 to torus where merging JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY with the postmedian orange band. VFW color pattern similar as on the upperside; median orange band slightly lighter, whitish along basal edge towards costa, subapical area along costa and inside cell R5—M1 suffused with chestnut; subapical black spot in MI—M2 with a white pupil; subapical pale yellow spot in cell R5-M1; apical and submarginal areas suffused with beige scales. VHW ground color rufous oun median line nearly straight, from costa to vein 1A basally edged with white; a band extending from postmedian to submz arginal area shaped as on the uppe srside, white along basal and distal end, rufous or: ange in the middle, and a series of six black dots with white pupils situated in the middle of the band from Rs—M1 to Cu2-1A, two in the latter, except in cell M2-M3 where instead a lighter pale yellow patch. Genitalia (Fig. 8): Uncus twice the length of tegumen, hooked; gnathos two-thirds the length of uncus, curved upwards; saccus deep, Se like; valvae approximately the same width in basal and distal part with slightly serrate distal extremity and curved inwards; aedeagus slightly longer than valvae, smooth and straight. ea (Fig. 4): Head, thorax and abdoten similar to the male. Wings: FW length 33 mm; DFW and DHW similar to the male; orange band a shade lighter, pale orange. VFW similar to the male; orange band a shade lighter, pale orange; VHW similar to the male but lighter, suffused with white scales along distal edge of the submarginal line. Type material: Holotype 6: Guyana, N. slope Mt. Roraima, 2"4 Camp, 1300m, 5°16N, 60° 44W, 12.1IL.—16.1V.2001, Romeo Williams, Wiltshire Hinds leg., STW. Paratype 9: same data. Etymology: This beautiful montane satyrine is gratefully named after Romeo Williams, a Guyanese national. A friend of the junior author, Romeo has been an indispensable team member for the recent lepidoptera expeditions to a number of Guyana's remote montane regions. Bey ond these expeditions, Romeo's bush skills, work ethic and dedication have added immeasurably to numerous other biological field endeavors in Guyana's hinterlands. Remarks: The genus Oxeoschistus Butler comprises thirteen species, including one highly polytypic species — Oxeoschistus simplex Westwood with five recognized subspecies. The genus Dioriste Thieme is considered herein as a subjective junior synonym of Oxeoschistus following Lamas et al. (in press). Accordingly, seven species of Oxeoschistus are found in the Andes and six in Central America. One species, O. puerta, is represented by separate subspecies in Costa Rica, Panama, Colombia and northern Venezuela. The genus is therefore particularly well represented in Central America, as compared to other genera of Pronophilina (except for the genus Drucina Butler, endemic to Central America), and the fact that there are as many species outside and within the Andes is unique for the subtribe. A few genera are represented in Central America by one or two species, rarely more (Pedaliodes Butler, Pseudomaniola Rober), and many have no representative at all, including Corades Doubleday, Junea Hemming, Daedalma Hewitson and Mygona Thieme among others. Adult Oxeoschistus are recognized by large size for the standards of the subtribe, with an average FW VOLUME 59, NUMBER 4 12 Fics. 10-18. Erestris taxa related to E. agata. 10. Eretris encycla ssp. (I S-W Venezuela) 12. Eretris encycla ssp. (Tandapi, W Ecuador) sp. (Sierra de Lema, Venezuela) 15. Eretris encycla ssp. ( calisto (Zamora, S Ecuador) 18. Eretris sp. (San Andres, S Ecuador) length 3-3.5cm, blunt FW apex and undulate HW margins. Most species have conspicuous orange or yellow upperside markings usually shaped as a median band. VHW is characterized by usually fully developed large postmedian ocelli present in all cells (except M2-M3) in most species. Antennae are slender, approximately 2/5 of the FW length. Eyes are setose as in other Pronophilina (Viloria, Ph. D.). Male genitalia can be recognized by a long uncus, and particularly long subunci, in some species approaching the length of the uncus, elongate valvae without any secondary process and a serrate dorsal surface, deep saccus and long, straight aedeagus. Although, there was no cladistic analy sis of the Subtribes these morphological characters indicate Pseudomaniola Réber or Pronophila Doubleday as possible sister genera (Pyrcz, in prep.). Affinities Currently, there are 10 species of Pronophilina known to occur in the Pantepui. Three of them belong to an endemic genus Protopedaliodes (presenting ), whose affinities within the speciose Pedaliodes complex are unclear (Viloria, Ph.D.). Five species belonging to Pedaliodes—P. roraimae, P. demarmelsi, P. chaconi, P. terramaris and P. yutajeana— are closely several specialized morphological structures related 205 Las Golondrinas, N-W Ecuador) 11. Eretris encycla ssp. (Mérida, 13. Eretris encycla encycla (Colonia Tovar, N Venezuela) 14. Eretris agata n Santa Lucia, S-W Ecuador) 16. Eretris oculata (Jorge Chavez, N Peru) 17. Eretris allopatric taxa. They could eventually be considered, due to the slight morphological differences, as subspecies of P. roraimae. Viloria & Pyrez (1995) discuss the affinities of P roraimae and indicate that its closest Salient morphological similarities can be observed in the wing shape and colour pattern and more importantly in the shape of the male androconial patch and male genitalic structure (Viloria et al., 2001). P. pisonia occurs in the Venezuelan Cordillera de La Costa and the parallel Cordillera del Interior in low elevation cloud forests, at 1200—1700m. E. agata presents a series of wing color pattern similarities to: E. oculata (C. & R. Felder) distributed along the easter slopes of the Andes between calisto (C. & R. Felder) found in central and eastern Colombia, eastern Ecuador and northern Peru and E. encycla (C. & R. Felder) occurring in the Venezuelan Cordillera de La Costa and Mérida, and in western Colombia and Ecuador (figs. 10-18). The VHW ocelli of E. agata are intermediate in size as compared to other congeners: ally is P. pisonia (Hewitson). Colombia and northern Peru, smaller than in E. oculata, about the same size as in some populations of E. encycla (an undescribed subspecies) from the Western Cordillera in Colombia and Ecuador, and larger than in E. encycla. The shape 206 of the VHW postmedian band is most similar to E. oculata, less sinuate than E. calisto and E. encycla, both similar in this respect. E. agata has no orange or yellowish scales in the VHW anal area or basad to the postmedian line, similar to E. encycla (although some individuals of the nominate subspecies do have some yellow scaling) and E. oculata, and contrary to E. calisto, which always presents some yellow or orange pattern. The exclusive character of E. agata is that all three VHW lines are all brown, whereas in all other compared species they are crimson red. Additionally, the typical incision of the anal mar gin of the HW is shallower in E. agata than in other species, and the HW is ovaloid rather than squarish, in this respect it resembles an undescribed species of the “calisto” stock from southern Ecuador and northern Peru (Pyrcz, in prep.). Male genitalia of E. agata show some similarities to E. oculata, egasctllly characteristic valvae with a shallow curve in the middle and a tubular, simple aedeagus. In E. encycla the aedeagus is somewhat inflated in the middle and shorter. However other sclerites, tegumen, uncus, vinculum and transtilla, are nearly identical to E. encycla and E. calisto. These two species have rather long gnathi, roughly one third to half the length of uncus. The gnathi of E. agata are slightly shorter. On the other hand, the gnathi of E. oculata are rudimentary short tips, and in this respect are unique compared to other congeners. Oxeoschistus romeo, as already pointed out in the new species diagnosis, basically presents all typical generic characters of the color pattern. However its DFW median orange band pattern is unique because it is modified as compared to allied species. Instead of a regular band with roughly parallel outer and inner edges, it presents an extremely irregular, erratic shape with several deep intrusions. Diagnostic features can be pointed out in the VHW pattern, especially the fact that the white-edged postmedian band is unbroken, smooth and runs roughly parallel to the outer margin. This character is found in O. simplex distributed throughout Colombia, O. puerta found in the Venezuelan Cordillera de La Costa, Mérida, Perija and northem Central and Eastern Cordillera in Colombia, and O. duplex. The latter is found in central (Junin, Pasco) and southern (Cuzco, Puno) Peru and Bolivia (Yungas). The male genitalia of O. romeo provide a series of very useful diagnostic characters. They clearly indicate O. puerta and O. simplex as the closest allies. In all these species, the gnathi are thinner than the uncus, about two-thirds its length and hooked upwards, the saccus is very long, and the aedeagus is nearly straight. The valvae of O. simplex are singularly serrate in the distal-dorsal one-third and occasionally present small aK JURNAL OF THE LEPIDOPTERISTS’ SOCIETY processes. The valvae of O. puerta and O. romeo are shorter and more compact, dorsal teeth are not as prominent and do not have any secondary process (figs. 8-9). The Protopedaliodes above considerations indicate, the genus taken apart, close faunal affinities between the Pronophilina of the Pantepui and the Pronophilina occurring in the Venezuelan Cordillera de La Costa: P. pisonia, E. encycla and O. puerta. It is therefore sound to consider that this range was the primary source of cloud forest Pronophilina for the Pantepui. The colonization of the Pantepui might have taken place through ecological corridors across the Orinoco plains during cold glacial phases in the Pleistocene, when temperatures were 5°-9° cooler than at the present (Bush, 2002, 2004). These ecological conditions might have favored the growth of premontane forest vegetation (including Chusquea) at much lower elevation than currently (Cowling et al., 2001); thus allowing the dispersal of pronophilines, especially the species inhabiting lower elevations (below 1500m). This scenario is pepe considering that at the southern limit of the latitudinal range occupied by the subtribe, where climate is cooler with low winter temperatures, several species of Pronophilina live in the lowlands. For example, in southern Brazil and northern Argentina Praepedaliodes phanias (Hewitson) occurs commonly at 200m and feeds on local Aulonemia sp. bamboo (Pyrez, unpubl). In south-central Bolivia, near Santa Cruz de la Sierra, a local population of Physcopedaliodes physcoa (Hewitson) occurs in a relictual forest at 400m. Interestingly enough, the same species is found in central Peru (Chanchamayo) at higher elevations—1200—1600m (Pyrez, unpubl.). Diversity Eretris agata and Oxeoschistus romeo represent the ninth and tenth species and only the third and fourth known genera of the subtribe Pronophilina in the Pantepui. A faunal comparison (Appendix) showing data from ten Andean localities gathered along altitudinal transects (only sympatric or parapatric species occurring along the same slope are considered) yields the immediate conclusion that the Pronophilina fauna of the Pantepui is extremely species poor as compared to the Andes. The most straightforward factors responsible for this disproportion would be the lower ecological diversity and the smaller area of montane habitats in the Pantepui as compared to the Andes. Whereas in the Andes suitable habitats for the Pronophilina are found within the wide elevational belt roughly between 1000-4000m corresponding with cloud forests and paramo, in the Pantepui they are restricted by the VOLUME 59, NUMBER 4 highest elevations of the table mountains, which is generally below 2500m, except for a few higher tepuis slightly exceeding this elevation (La Neblina, Roraima, Kukenan). In the Andes, cloud forests form a nearly continuous belt extending over 2500km along the eastern slopes from Bolivia to Venezuela, and over 1000km along the western slopes from northern Peru to northern Colombia. In the Pantepui, on the other hand, cloud forests are scattered and restricted to the slopes of isolated table mountains (Huber, 1995). However, there are also significant differences in the observed species richness between various localities within the Andes. The index falls steadily from central towards northern Andes, from more than 100 species in southern Ecuador (Zamora) and northern Peru (Molinopampa) to 70-80 in central Colombia, and 55 in western Venezuela (El Tama), roughly 50% of the figure in Zamora. These areas do not differ in ecological diversity and suitable habitat area (Vuilleumier & Monasterio, 1986), therefore, other factors are to be involved. The decrease in species richness is more radical in the peripheral ranges of the northern part of the continent, the Venezuelan Cordilleras de Mérida, Perija and La Costa, and the Colombian Santa Marta range. Sharp diversity break-downs correspond with several topographical and ecological barriers. A very important one is the so-called TAchira depression, a deep valley separating the Colombian Cordillera Oriental from the Cordillera de Mérida (Vuilleumier & Ewert, 1978; Pyrez & Viloria, in press). Other barriers include: the Valle del César separating Perija from the Sierra Nevada de Santa Marta (Adams, 1985) and the Lara plateau separating the Cordillera de Mérida from the Cordillera de la Costa (and the parallel, low Cordillera del Interior). The fauna of the Cordillera de la Costa (awaiting monographic treatment, Pyrcz et al., in prep.) accounts for 24 species of Pronophilina, less than half of that of El Tama, and merely one fourth of the species- rich southern Ecuador. This yields the conclusion that these barriers act as a ‘filter’ for the dispersal of pronophilines from the center of diversity in the main Andes (corresponding with the putative center of radiation of the subtribe, situated in the central Andes between Ecuador and Bolivia, Pyrez, in prep.) into peripheral ranges. The tepuis are separated from the already impoverished Cordillera de La Costa by vast lowlands, the Orinoco plains (300-400km) and from the Eastern Cordillera by the upper Llanos (>500km). If a simple area/distance from source calculation is carried out according to the classical island geography model (MacArthur & Wilson, 2001), a dramatic decrease in the 207 Pantepui is to be expected. Low diversity figures for the Pantepui may also be partly a consequence of the severe undersampling of this region. Admittedly, the sampling for pronophiline butterflies in the Pantepui has so far been negligible. The few specimens collected throughout the years are an outcome of random or even eal collecting, either by scientists other than entomologists (usually geologists, botanists or ornithologists who most frequently visited the region), or tourists. The reasons for this poor knowledge of the Pronophilina fauna in the Pantepui are multiple. First of all, Venezuelan authors (DeMarmels et al., 2003) point out that the region is very remote and difficult and costly to reach. It certainly is true in the case of the summits of the most remote tepuis such as La Neblina, Tapirapecd, Duida or Marahuaca. These tepuis can be accessed easily only by helicopter. Otherwise, a long arduous expedition is necessary usually entailing canoeing for several days and many days of jungle trekking. Both. ways are very costly, which is a very strong barrier for any biological research. However, the a few tepuis situated in the eastern part of the Guayana highlands, especially Roraima, Kukenan, Auyan and a handful of other smaller table mountains, are relatively easy and inexpensive to reach. As a matter of fact, one of the new species described herein was collected in the area called La Escalera, along the asphalted main road leading from Km. 88 to Santa Elena de Uairén, which is crossed daily by dozens of vehicles! La Escalera can be considered as the only relatively well sampled area within the entire Pantepui. In the local cloud forest at 1300-1400m only two species of Pronophilina were documented—Pedaliodes roraimae and Eretris agata, and a third—Oxeoschistus romeo—is likely to occur. The question arises: is the reported diversity low in relative terms as compared to the Andes? Not necessarily, considered that we are discussing diversity at 1300-1600m. As already stated, the highest diversity of the subtribe in the Andes falls at elevations between 2500 and 3000m (Adams, 1985: Pyrez & Wojtusiak, 1999, 2002). Below 1600m diversity is at its lowest. For comparison, in the Andean Western Cordillera in Colombia—Tambito (Pyrez & Wojtusiak, 1999) and Ecuador—Las Golondrinas (Pyrez & Wojtusiak, in prep.), where total species richness along sampled elevational gradients (1600—2500/2600m) exceeds 60, only three species were reported below 1600m. Interestingly enough, they are representatives of three genera reported from La Escalera: Pedaliodes phrasiclea Grose-Smith, Eretris depresissima Pyrcz and Oxeoschistus simplex (Butler), which underlines a structural similarity to the Pantepui fauna at a similar 208 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 19-23. Type locality of Eretris agata, new species. 19. Chusquea? sp. bamboo in a forest clearing. 20. Bamboo node detail. 21. Chusquea? sp. bamboo, likely host plant of E. agata. 22. Young Miconia sp. (Melastomataceae). 23. Swampy forest with bamboo clumps VOLUME 59, NUMBER 4 altitude. There are sight reports of further species and genera of Pronophilina in the Pantepui by experienced lepidopterists. Mauro Costa (pers. comm.) observed for a while an individual butterfly, which he identified as Lasiophila or Mygona, as he was able to notice its diagnostic pattern of the HW _ underside. This observation took place on the above-mentioned road at Km 130 (1400m). Andrés Orellana (pers. comm.) observed for some time a butterfly on the summit of Roraima, brown with some golden sheen, which he believes could have been a Lymanopoda Westwood (a species rich Andean genus of the — subtribe Pronophilina). The junior author has participated in two large-scale expeditions to major Guyana tepuis in the Pakaraima Mts: Mt. Wokomong (ca. 1675m) in November 1993 and Mt. Ayanganna (ca. 2050m) in April 1999. During the Mt. Wokomong expedition, approximately one week was spent at an elevation between ca. 1425m—1575m with plenty of bright sunshine. Even though a medium- sized bamboo was very common along the summit ridge (ca. 1575m), no pronophilines were captured or definitively seen. On the Mt. Ayanganna expedition there was the same result, with a little over a week spent at elevations (ca. 1375m—1675m) where at least some pronophilines were expected. Again there was plenty of brilliant sunshine, but on Mt. Ayanganna, bamboo did not seem common on the small part of this huge mountain massif that we explored. It was disappointing that we spent no time collecting on the summit plateau (ca. 2050m). On a previous botanical expedition, botanist Terry Henkel reported seeing dark medium- sized butterflies common on the summit plateau when the sun was shining. Were these pronophilines, and if so, could they differ from species on the not too distant Mt. Roraima? All this emphasizes an urgent need for well-organized sampling and thorough research of the cloud forest butterfly fauna of this extremely interesting region of South America. There is little interest among local entomologists in promoting and carrying out research, and especially field work, in this part of Venezuela. Unfortunately, when it comes to foreigners, the restrictions imposed by Venezuelan laws (these laws having as their primary goal the protection of the environment and native Amerindian culture) in recent years, have practically created such bureaucratic obstacles that permits are nearly impossible to obtain rendering any investigation very difficult (see Viloria & Pyrez, 1999 comments on this issue). ACKNOWLEDGEMENTS 209 The senior author would like to thank Mauro and Clara Costa for their outstanding help in Venezuela and Dr. Janusz Wojtusiak (Krakow) for kindly supporting throughout the years his research on the Pronophilina. The 2004 trip to Venezuela was made possible thanks to an intemal grant of the Zoological Museum of the Jagiellonian University in Krakéw, Poland (BW/2003). The Mount Roraima ornithology expedition was largely funded through a National Geographic Research and Exploration Grant. The Smithsonian Institution Division of Birds graciously allowed lepidoptera collecting on their expedition. Christopher Milensky, Museum Specialist in the Division of Birds, facilitated this effort and made sure the specimens and data returned to the Smithsonian. Chris, Wiltshire Hinds, a University of Guyana Biology Student at the time, and Romeo Williams all did some lepidoptera collecting, with Romeo and Wiltshire doing the bulk of this. Dr. Scott Miller, then Chairman of the Smithsonian Entomology Department, made Smithsonian personnel and equipment available for the photos. Dr. Patricia Gentili-Poole, Museum Specialist, took the excellent digital photos. The authors would like to thank Dr. Keith Willmott (BMNH) for performing genitalic dissections and drawing the male genitalia of O. romeo. LITERATURE CITED Apams, M. J., 1985. Speciation in the Pronophiline Butterflies (Satyri- dae) of the Northem Andes. J. Res. Lepid. 1985, Supplement No.1: 33-49. —. 1986. Pronophiline butterflies (Satyridae) of the three Andean Cordilleras of Colombia. Zool. J. Linn. Soc. 87: 235-320. ApaMs, M. J. & BERNARD, G. IL, 1977. Pronophiline butterflies (Satyridae) of the Sierra Nevada de Santa Marta, Colombia. Syst. Ent. 2: 263-281. —— 1979. Pronophiline butterflies (Satyridae) of the Serrania de Valledupar, Colombia-Venezuela border. Syst. Ent., 4: 95-118. ——. 1981. Pronophiline butterflies (Satyridae) of the Cordillera de Mérida, Venezuela. Zool. J. Linn. Soc. 71: 343-372. Busu, M. B., SILMAN, M. R. & UrrEGO, D. H., 2004. 48.000 years of climate and forest change in a biodiversity hot spot. Science, 303: 827-829. CowLInG, S. A., Mastin, M. A. & Syxes, M. 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New York: Oxford University Press, xii + 649 pp. Received for publication 16 October 2004; revised and accepted VOLUME 59, NUMBER 4 211 Appendix: Pronophilini species richness in 11 localities along altitudinal transects Genera San Colonia Manaure LaCulata Betania ChoaChi Puracé Runtin Zamora Molino- Roraima Lorenzo ‘Tovar pampa Arhuaco 1 0 0 0 0 0 0 0 0 0 0 Cheimas 0 0 0 i} 0 0 0 0 0 0 0 Corades 4 3 4 6 t rf 7 8 10 8 0 Daedalma 0 0 0 0 1 2 2 3 3 3 0 Drucina 0 0 0 0 0 0 0 0 0 0 0 Junea 0 0 0 0 1 2 2 2 2 2 0 Apexacuta 0 0 0 0 0 0 1 1 iL 2 0 Lasiophila 1 1 1 1 2 3 3 3 4 4 1? Mygona 0 1 IL 1 1 2 2 2 2 2 0 Oxeoschistus 0 1 1 1 1 2 2 2 2 3 1 Pseudomaniola 1 1 1 1 2 1 1 1 4 4 0 Pronophila 2 2 3 2 3 3 3 4 6 5 0 Thiemeia 0 1 0 0 0 0 0 1 1 1 0 Pedaliodes° tf § 12 15 19 29 34 32 42 42 1 Paramo 1 0 0 0 0 0 0 0 0 0 0 Dangond 0 0 il 0 0 0 0 0 0 0 0 Redononda 0 0 0 1 0 0 0 0 0 0 0 Protopedaliodes 0 0 0 0 0 0 0 0 0 0 2 Eretris 1 1 3 2 3 6 5 5 5 5 1 Lymanopoda 2 2 6 5 6 10 9 10 10 10 1? Manerebia 1 1 2 2 4 4 3 4 rh 6 0 Steroma 1 1 1 1 1 1 1 1 2 3 0 Steremnia 0 0 0 0 1 pH 3 3 5 5 0 Diaphanos 0 0 0 1 0 0 0 0 0 0 0 Idioneurula 0 0 0 0 1 1 0 0 0 0 0 Tanusiussa 0 0 0 0 1 0 1 1 0 0 0 Tamania 0 0 0 0 1 0 0 0 0 0 0 Total species 22 24 35 37 55 75 79 83 107 105 5 (7) Total genera 11 12 12 14 17 15 16 17 16 16 4 (6) ° Pedaliodes sensu lato Localities co-ordinates and faunal data source: - San Lorenzo: Colombia, Sierra Nevada de Santa Marta, north slopes, 11°08'N 74°03'W (Adams & Bemard, 1977) - Manaure: Colombia, Serrania de Perija, west slopes, 10°23N 72°58'W (Adams & Bemard, 1979; Viloria, Msc. thesis, unpubl.) - La Culata: Venezuela, Cordillera de Mérida, Sierra de la Culata, southeast slopes, 8°41'N 71°08'W (Adams & Bernard, 1981; Pyrez & Wojtusiak, 2002) - Colonia Tovar: Venezuela, Cordillera de La Costa, north slopes, 10°26'N 67°15'W (Raymond, 1982; Pyrcz, in prep.) - Betania: Venezuela, Cordillera Oriental, Sierra del Tama, north slopes, 7°27'N 72°26'W (Pyrez & Viloria, in press) - Choachi: Colombia, Cordillera Oriental, Cundinamarca, east slopes, 4°33'N 73°57'W (Adams, 1986; Pyrcz, 1999) - Puracé: Colombia, Cordillera Central, Cauca, east slopes, 2°22'N 76°16'W (Adams, 1986; Pyrez, 1999) - Runtiin: Ecuador, Cordillera Oriental, Tungurahua, north slopes, 1°25'S 78°25'W (Pyrecz et al. 1999; Pyrez & Viloria, 1999, Pyrez, 2000, unpubl.) - Zamora: Ecuador, Cordillera Oriental, Nudo de Sabanillas, east slopes, 3°58'S 79°03'W (Pyrez et al. 1999; Pyrez & Viloria, 1999, Pyrez, 2000, in press) - Molinopampa: Peru, Cordillera Oriental, Chachapoyas, southeast slopes, 6°10'S 77°34'W (Pyrcz, 2004) JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Journal of the Lepidopterists’ Society 59(4), 2005, 212-218 THE HIGHLY SEASONAL HAWKMOTH FAUNA (LEPIDOPTERA: SPHINGIDAE) OF THE CAATINGA OF NORTHEAST BRAZIL: A CASE STUDY IN THE STATE OF RIO GRANDE DO NORTE José ARAUJO DUARTE JUNIOR Programa de Pés-Graduagao em Ciéncias Biolégicas, Departamento de Sistematica e Ecologia, Universidade Federal da Paraiba, 58059-900, Joao Pessoa, Paraiba, Brazil. E-mail: josejunior_lep@yahoo.com.br AND CLEMENS SCHLINDWEIN Departamento de Botanica, Universidade Federal de Pernambuco, Av. Prof. Moraes Rego, s/n, Cidade Universitaria, 50670-901, Recife, Pernambuco, Brazil. E-mail:schlindw@ufpe.br ABSTRACT: The caatinga, a thorn-shrub succulent savannah, is located in Northeastern Brazil and characterized by a short and irregular rainy season and a severe dry season. Insects are only abundant during the rainy months, displaying a strong seasonal pat- tern. Here we present data from a yearlong Sphingidae survey undertaken in the reserve Estagdo Ecolégica do Se rid6, located in the state of Rio Grande do Norte. Hawkmoths were collected once a month during two subsequent new moon nights, between 18.00h and 05.00h, attracted with a 160-watt mercury vapor light. A total of 593 specimens belonging to 20 species and 14 genera were col- lected. Neogene dynaeus, Callionima grisescens, and Hyles euphorbiarum were the most abundant species, together comprising up to 82.2% of the total number of specimens collected. These frequent species are residents of the caatinga of Rio Grande do Norte. The rare Sphingidae in this study, Psewdosphinx tetrio, Isognathus australis, and Cocytius antaeus, are migratory species for the caatinga. Ninety percent of specimens and 75% of species were collected during the rainy season. With the arrival of the dry sea- son, when bushes and trees lose their leaves and the herbaceous layer disappears, Sphingidae abundance decreases rapidly. Rich- ness and abundance of species through the year is highly seasonal and correlated with rainfall. Erinnyis alope alope, Eumorpha fas- ciatus, E. vitis vitis, and Manduca brasilensis were recorded for the first time in NE-Brazil. Additional keywords: Callionima grisescens, dry savannah, Hyles euphorbiarum, Neogene dynaeus, Northeastern Brazil, Sphin- gidae. INTRODUCTION sporadic visits to the field counts 14 species (Duarte & Motta 2001). Both surveys add up to a total of 32 species for NE - Brazil. The caatinga, a 834,666km? thorn-shrub succulent savannah (Andrade-Lima 1981), is the predominant vegetation in Northeastern Brazil (Fig. 1). The region is characterized by seasonal rainfall that defines the phenophases of the vegetation. During the dry season, which lasts approximately eight months (May—December), the vegetation loses its leaves. Sporadically there are years with almost no rain and drought can be severe (Andrade-Lima 1981, Sampaio et al. 1991, Machado et al. 1997). Therefore, the seasonal characteristics of the Bienes climate and vegetation must influence the seasonality of Sphingidae and the development from an immature stage to adult (Janzen 1983, Haber & Frankie 1989, Darrault & Schlindwein 2002). The annual development cycle of the Sphingidae is not known for the caatinga region. Generally, the relationships between larvae of Sphingidae are widely distributed and comprise approximately 1200 species, which occur predominantly in tropical regions (Lemaire & Minet 1999). Sphingidae are generally among the first Lepidoptera to be surveyed in a region. Despite this, studies related to the taxonomy, geographical distribution, relationship with host plants, and adult food sources are scarce for Sphingidae (Kitching & Cadiou 2000). In Brazil to date, 180 species of Sphingidae have been recorded in regional surveys (Brown 1986), more than half in the Amazon basin: 61 species in the state of Amazonas (Motta 1998) and 90 in the state of Para (Moss 1920). Seventeen species were recorded in the state of Sao Paulo (Coelho et al. 1979), 55 in the state of Paranda (Laroca & Mielke 1975), and 65 in the state of Rio Grande do Sul (Biezanko 1982; Corseuil et al 2001). In Northeastern Brazil the Sphingidae fauna is still poorly studied. The only published data concern a community of Sphingidae and associated plants from an area of tabuleiro nordestino (a kind of cerrado, isolated from the Central Brazilian Cerrado) on the coast of the state of Paraiba (Darrault & Schlindwein 2002): this study recorded 24 species of Sphingidae for the area. A preliminary list for the caatinga region, based on Sphingidae and their host plants are highly specific. In many cases, the larvae accumulate toxic compounds from the plants that protect the larvae against predators. Such host plants include species of Apocynaceae, Euphorbiaceae, Solanaceae, and Rubiaceae (Biezanko VOLUME 59, NUMBER 4 to w Seat a aw, ata {ae Bi OS Esec-Serido ic Ses zs : yi Maranhéo Te ~ \ | / Ceara Baws eae iE ~ = @ 4 P Norta x a ak 4 ee “Paatal ee me Los z= BRASIL ce : A : eeceatae) a) x9 3 ~/ Algaoas “A ; go uo Si Bahia a N NE-Brazil Caatinga Fic.1. Map of Brazil indicating Northeastern Brazil, the distribution of the caatinga and the location of the study site “Estagao Ecologica do Seridé (ESEC - Serid6)”, 1982, Janzen 1983, Moss 1920). The relationships between adult Sphingidae and sphingophilous flowers, however, does not seem to be specific (Darrault & Schlindwein 2002, Haber & Frankie 1989). Pittaway (1993) affirms that a species may be submitted to seasonal cycles of abundance and scarcity, or its occurrence throughout the year may be constant for an area. According to this author, species abundance may also decline or increase in response to environmental conditions and to the degree of migration. Here we present data for the composition, species richness and abundance of Sphingidae through the year in the caatinga in the nature reserve Estagao) Ecol6gica do Serid6 in the state of Rio Grande do Norte, NE- Brazil. MATERIALS AND METHODS Study area The study was undertaken in the nature reserve Estacgdo Ecolégica do Seridé (ESEC-Serid6) in the municipality of Serra Negra do Norte, state of Rio Grande do Norte, NE - Brazil (Fig.1). The reserve covers an area of 1,166.38 ha located between 06°35! and 06°40'S and 37°20! and 37°39'W, at an altitude of approximately 170 m. The climate is hot and dry (mean annual temperature of 28°C), with a rainy season that varies from January/February to April/June (mean annual rainfall of 497 mm). The rest of the year is characterized by a severe drought (IBAMA 1989) (Fig. state of Rio Grande do Norte. Source: FREIRE, 2003. 2). The temperature and rainfall data were obtained from the climate station in the reserve. The vegetation is an arbustive to arboreous caatinga, with an herbaceous stratum that is only well-dev eloped during the rainy period (Prado 2004). Sampling Sphingidae were collected between March 2002 and May 2003 (with the exception of May 2002). The moths were attracted by a 160-watt mercury light source positioned against a white wall. The light dispersed over an extensive area. Specimens were collected during two consecutive new moon nights per month from 18.00h to 05.00h on the following day. The moths were killed with an injection of ethyl acetate between the thorax and abdomen. The specimens were then placed in entomological envelopes and prepared in the laboratory. The enc aention of the material collected was based on D’Abrera (1986) and Kitching & Cadiou (2000). Specimens were deposited in the entomological collection of the Federal University of Pernambuco (UFPE, Recife), the entomological collection of the Department of Systematics and Ecology of the Federal University of Paraiba (UFPB, Joao Pessoa), and in the Museum of Natural History of Serid6 (located in the ESEC Serid6 reserve). Three abundance criteria were established using Rabinowitz et al. (1986), based on the number of specimens collected per species: rare (1 to 2 specimens), common (3 to 19 specimens), and abundant (20 to 50 specimens). 250 lalla JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 7 300,0 + 20,0 + 100,0 Precipitation mm/month mar apr may jun ju A ® 200 + 3 a Y 150 + Oo 2 g 3 al @d 100 + S =] Ky = -\ \ 50 + ‘\ ZEN jul aug sep oct nov dec op jan feb mar apr may Months MM Species richness Precipitation during study period Abundance Prec. Pluriannual 1992-2002 Fic. 2. Annual and pluriannual precipitation and abundance and richness of species of Sphingidae of the Estagao Ecolégica do Seridé between March 2002 and May 2003 (May 2002 was not sampled). The following indices were calculated: Shannon- Wiener, Spearman's correlation coefficient and Pearson's correlation coefficient. The indices were calculated using the BioStat version 2.0 (Ayres et al 2000) and Krebs version 5.1. (Krebs 1999). RESULTS Sphingidae species richness and abundance In the nature reserve Esec-Serid6, 593 specimens of 20 species were recorded (Table 1). From this total, 90% of specimens and 75% of species were collected during the rainy months (February to June) (Fig.2). From October to December, only three specimens of two species were recorded, Erinnyis ello ello and Xylophanes tersa tersa (Fig. 2). In October, no hawkmoths appeared at the light traps. Neogene dynaeus (38%), Callionima grisescens elegans (28%), and Erinnyis ello ello (8.8%) were the predominant species, accounting for over two-thirds of all Sphingidae collected (Fig. 3). The six most abundant species comprised 91.7% of the total of specimens, while the six least abundant summed to only 1.7% Fig.3). Neogene dynaeus occurred during the rainy season, from February to June, and Callionima grisescens elegans from February to September (Fig.4). The rare species Cocytius antaeus, Isognathus australis, and Pseudosphinx tetrio, were recorded only in the dry months (Table 1). Spearman's correlation index was 0.6196 for species abundance versus rainfall and 0.5913 for species richness versus rainfall, showing significant correlations. In contrast, correlations for both rainfall and species richness versus temperature, calculated using Pearson's coefficient, were negative. The accumulated species curve was representative, showing no increase during the last months (Fig.5). DISCUSSION Species richness, abundance and seasonality Our results show that species abundance and richness of Sphingidae are highly seasonal in the caatinga region. g y ga reg by This seasonality is influenced the — distinct 225 / 38,0% Net di ogene dynaeus Pr Callionima grisescens 52/.8,8% Erinnyis e ello Hyles euphorbiarum 4417,.4% /5,0% Agrius cingulatus ZOE 27 14,6% Xylophanes ttersa Eumorpha fasciatus 8/1,3% Mauduca r.rustica 8 Ginnyis lassauxii 5/0,8% Erinnyis aalope A > Protambulyx strigilis S Enyo Llugubris p>! O° Erinnyis 0 obscura Eumorpha vitis vitis Manduca brasilensis Eumorpha llabruscae fP2/03 Perigonia pittieri Cocytius antaeus lsognathus australis 1 /.0,2% Pseudosphinx tetrio Fic.3. Number of specimens and relative abundance of species of Sphingidae recorded between March 2002 and May 2003 at the Estagao Ecolégica do Serido. VOLUME 59, NUMBER 4 TABLE 1. Species of Sphingidae recorded in the Estagao Ecolégica do Serid6, municipality of Serra Negra do Norte, state of Rio Grande do Norte, Northeast Brazil, between March 2002 and May 2003. Abundant? Species Month Rare’® Common’ Macroglossinae Macroglossini Hyles euphorbiarum (Guérin & Percheron, 1835) Feb - Mar, Aug Xylophanes tersa tersa (Linnaeus, 1771) Apr, Jun-Sep, Dec Dilophonotini Callionima grisescens elegans (Rothschild, 1894) Feb-Sep Erinnyis lassauxii (Boisduval, 1859) Mar xX Erinnyis obscura obscura (Fabricius, 1775) Jan, Apr-May XS Erinnyis alope alope (Drury,1773) Feb, May, Jul-Aug X Erinnyis ello ello (Linnaeus, 1758) Jan-Mar, May-Aug, Dec Enyo lugubris lugubris (Linnaeus, 1771) Mar-Apr X Isognathus australis Clark, 1917 Sep X Perigonia pittieri Lichy, 1962 Feb xX Pseudosphinx tetrio (Linnaeus, 1771) Sep xX Philampelini Eumorpha fasciatus (Sulzer,1776) Feb, Apr xX Eumorpha labruscae labruscae (Linnaeus, 1758) Mar, Aug x Eumorpha vitis vitis (Linnaeus, 1758) Feb-Mar x Smerinthinae Ambulycini Protambulyx strigilis (Linnaeus, 1771) Jun-Jul xX Sphinginae Acherontiini Agrius cingulatus (Fabricius, 1775) Feb-Ago Sphingini Cocytius antaeus (Drury,1773) Aug xX Manduca brasilensis Jordan, 1911 Feb-Mar, Jun xX Manduca rustica rustica (Fabricius,1775) Mar-Apr, Sep xX Neogene dynaeus (Hiibner,[1827]-[1831]) Feb-Jun *The abundance categories follow Rabinowitz et al (1986). Ot phenophases of the vegetation as well as by the rainy period. Most Sphingidae occurred only during two months (March and April) of the rainy season; 65% of species occurred in these months. Other insect groups are probably also influenced by biotic and abiotic factors in the caatinga to the same degree. In the rainy season, leaves of host plants are available to larvae. The species Erinnyis ello ello showed an even distribution of abundance throughout the year. This was probably related to the existence of leaves of the host plants of native and cultivated species of Euphorbiaceae like Manihot for the larvae and sphingophilous flowers for the adults. No species occurred continuously throughout the entire year. The high seasonality of the Sphingidae was expected as, during the leafless dry season, the caatinga offers no food for the larvae and no nectar sources for the adults. Almost all perennial plants that flower during the dry the ornithophilous or chiropterophilous (Machado & Lopes 2004). season in caatinga are melithophilous, Number of specimens JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY other states of NE-Brazil. The species may show sporadic occurrence in the contact zone of caatinga with neighboring ecosystems. Hyles euphorbiarum and Erinnyis ello ello, like most of the other species recorded, are widely distributed in the Neotropical region (D'Abrera 1986, Biezanko 1982, Hodges 1971, Kitching & Cadiou 2000). Nevertheless, rare species like Pseudosphinx tetrio, Isognathus australis, and Cocytius antaeus were found exelusiv ely during the dry period, when there were no host plants for the larvae in the leafless environment. Therefore, we suppose that these Sphingidae migrated from adjacent regions, such as the non-seasonal Atlantic forest with its These hawkmoths probably do not reproduce in the caatinga. P. tetrio and C. y distributed. Isognathus. australis was considered endemic to NE- Brazil (Schreiber 1978). Our survey the occurrence of four species not previously recorded for NE-Brazil (Erinnyis alope alope, Eumorpha fasciatus, E. vitis vitis, and Manduca brasilensis), increasing to 36 rain evergreen vegetation. antaeus are widely shows mar apr jun jul aug sep O Neogene dynaeus EH yl es euphorbiarum oct mov dec jan feb mar Callionima gris B Agrius cingulatus apr may cens GWeErinnyis ello @ xylophanes tersa Fic.4. Seasonality of the six most frequent species of Sphingidae at the Estacao Ecologica do Seridé6 between March 2002 and May 2003. Caatinga- resident and migratory Sphingidae The abundant Sphingidae in the caatinga occur exclusively during the rainy season. Therefore, Neogene dynaeus (Fig. 7a), Callionima grisescens elegans (Fig.7b), Hyles euphorbiarum, and Erinnyis ello ello are caatinga-resident species closing the whole ontogenetic cycle during the short period of rainfall. The host plants for their larv ae, as well as sphingophilous flowers for the adults, thus should be plants of the caatinga vegetation. Schreiber (1978) considered Neogene dynaeus and Callionima grisescens elegans to be endemic to the state of Pernambuco, one of the neighbor states of Rio Grande do Norte. Yet the author did not specify with which vegetation types these species are associated. Based on our data, they are endemic to the caatinga region, and they probably also occur in caatinga of the ACCUMULATED NUMBER OF SPECIES 0 A A A A a A A ' as Sey mar apr jun jul aug sep oct nov dec jan feb mar apr may Months Fic.5. Accumulated number of species of Sphingidae recorded between March 2002 and May 2003 at the Estagao Ecolégica do Serido. VOLUME 59, NUMBER 4 217 Venezuela: Clavijo & Chacin, 1992 (Tropical rain forest) Para/Brazil: Moss, 1920 (Tropical rain forest) Costa Rica: Haber & Frankie 1983 (Tropical dry forest) Rio Grande do Sul: Corseuil & Specht, 2001(various vegetation types, subtropical) Roraima/Brazil: Motta, 1998 (Tropical rain forest) Parana/Brazil: Laroca & Mielke, 1975 (Atlantic rain forest) French Guyana: Beebe & Fleming, 1945 (Tropical rain forest) Number of Species —_ o fo) 20 0 [4 < 4 a a Ste aad 20 é 3 a A 62 = a ‘ 2 7) Fs wi < fe) =o {e) > oa oO Ya [ad Peru: Lamas, 1985 (Andine forests) Uruguay: Biezanko et al., 1962 (grasslands, subtropical) Paraiba/Brazil: Darrault & Schlindwein 2002 (cerrado- “Tabuleiro Nordestino”) Rio Grande do Norte/Brazil: This study (caatinga) Galapagos Islands/ Ecuador: Kernbach, 1962 po Norte, Br |] Parana, Br GuayANa Peru Parasa, BR Rio GRANDE GaLaPpacos Fic. 6. Number of species of Sphingidae recorded in surveys undertaken in the Neotropical region. Fic. 7. The most common hawkmoths of the caatinga. a) Neogene dynaeus b) Callionima grisescens elegans the number of Sphingidae in this region. When compared to Northern, Southern, and Southeastern Brazil and to other countries of the Neotropical region, the species richness recorded here was low (Fig.6). The abundance, on the other hand, was high. The accumulated species curve shows that the number of species in this locality is representative and probably may not or only slightly be increased in a prolonged survey. ACKNOWLEDGEMENTS We thank the staff of the nature reserve Estagdo Ecolégica do Seridé: Adson Borges de Macedo, Arménio Aratijo, Ridalvo Medeiros, George Aratijo, and Manuel Irmao for their support and IBAMA for the license to work at the reserve. Dr. Olaf H. H. Mielke (Federal University of Parana, Curitiba), Catarina Motta (National Research Institute of Amazonas, INPA, Man- aus) and Dr. LJ. Kitching (The Natural History Museum, Lon- don), kindly confirmed the identification of some specimens. Furthermore, we thank and André Mauricio Melo Santos for support in statistics, Arthur C.D. Maia for his help with the pho- tos, Dr. Celso Feitosa Martins (Federal University of Paraiba, Joao Pessoa), Reisla Oliveira Darrault for valuable suggestion on the manuscript, and Scott Heald (Cornell University) for revis- ing the English. The study was supported by grants of CAPES to J.A.D. and CNPq to CS. LITERATURE CITED ANDRADE-LIMA, D. 1981. The caatinga's dominium. Revista Brasileira Botanica, Recife, 4:149-155. Ayres, M., JR. AYRES, M., D.L, Ayres & A.S. Santos. 2000. BioEstat 2.0: aplicagées estatisticas nas areas de ciéncias biolégicas e médi- cas. Sociedade Civil Mamiraua e CNPq, Brasilia. BEEBE, W. & H. FLEMING, 1945. The Sphingidae (Moths) of Kartabo, British Guiana and Caripito, Venezuela. Zoologica, New York, 30:1-6. BIEZANKO, C. M,. 1982 Sphingidae da Zona Sueste do Rio Grande do Sul. Revista Centro Ciéncias Rurais, Santa Maria, 12(1): 59 75. BIEZANKO, C, M., A. RUFFINELLI & C. S. CARBONELL.1962. Lepi- 218 doptera del Uruguay. Notas Complementarias. II. Revista Facul- dade Agronomica. (Univ. de la Republica), 50:47-117. Brown, K. S. JR. 1986. Diversity of Brazilian Lepidoptera: History of study, methods for measurement, and use as indicator for genetic, specific and system richness, p. 221-253. In Bicudo, C.E & Menezes, N.A. (Ed.). Biodiversity in Brazil: a first approach. Sao Paulo, 326p. Cravyo, A. J. & E. M. CHacin. 1992. Sphingidae (Insecta: Lepi- doptera) Reportados como plagas en cultivos Venezolanos: Clave para las espécies. Boletim Entomologico Venezolano. N.S. 7(2): 119-125. CoELHO, I. P., S. SILverRA NETO, J. F. S. Dias, L. C. Forti & F. M. Lara. 1979. Fenologia e andlise faunistica da familia Sphingidae (Lepidoptera); através de levantamentos com armadilha luminosa em Piracicaba SP. 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W. FRANKIE. 1983. Checklist of insects. Checklist of Sphingidae: p. 645-650 In D.H. Janzen, (Ed.). Costa Rica Nat- ural History, Chicago, §16p. HaBeEr, W. A & G. W. FRANKIE. 1989. A tropical hawk-moth commu- nity: Costa Rican dry forest Sphingidae. Biotropica. 21(2): 155 - Qe Hopces, R. W. 1971. The Moths of America North of Mexico. R.B. D. Publication & E. W. Classey. fasciculo 21, London, 158 p. IBAMA 1989. Caracterizagao da Estagao Ecolégica do Seridé RN. Natal- RN - Abril. PROJETO PNUD/FAO/BRA. JANZEN, D. H. 1983. Costa Rica Natural History. Univ.Chicago Press. Chicago, 816p. KERNBACH, K. 1962. Die Schwirmer einiger Galapagos Inseln (Lep. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY SPHINGIDAE). Opuse. Zool. 63:1-19. KiTCcHING, L. J. & J. M. Capiou 2000. Hawkmoths of the world: an an- notated and illustrated revisionary checklist (Lepidoptera: Sphin- gidae). Cornell University Press, Ithaca, 227 Kress, C. J.1999. Ecological methodology. Benjamin/Cummings, Menlo Park, Calif. 624. Laroca, S. & O. H. H. MIELKE. 1975. Ensaios sobre ecologia de co- munidades em Sphingidae na Serra do Mar, Paranda, Brasil.(Lep- idoptera). Revista Brasileira Biologia, Curitiba,. 35 (1): 1-19. Lamas, G, 1985. The Castniidae and Sphingidae (Lepidoptera) of the Tambopata Reserved Zone, Madre de Dios, Peru: a preliminary list. Revista Peruana Entomologia. 27:55-58. LemalrE, C. & J. MINET. 1999. The Bombycoidea and their relatives, 321-353. In De Gruyter (Ed.). Lepidoptera: Moths and Butter- flies. 1. Evolution, Systematics, and Biogeography. Part 35. Hand- book of Zoology. IV, Berlin and New York. Macnapo, I. C. S. & A. V. Lopes. 2004. Recursos Florais e Sistemas de Polinizag&éo e Sexuais em Caatinga. In Editora Universitaria UFPE. Ecologia e Conservagao da Caatinga. Recife 12:515-563. ——., L. M. Barros & E. V. S. B, Samparo. 1997. Phenology of Caatinga Species at Serra Talhada, Pe, Northeast Brazil. Biotrop- ica. 29 (1): 57 - 68. Moss, A. M. 1920. Sphingidae of Para. Brasil. Early stages, food plants, habits, etc. Novitates. Zoologicae, 27:334-357. Moma, C.S. et al. 1998. Aspectos da esfingofauna (Lepidoptera, Sphingidae), em area de terra-firme, no Estado do Amazonas, Brasil. Acta AmazOnica, 28 (1): 75-92. Pirtaway, A. R. 1993. The hawkmoths of the western Palaearctic. London & Colchester, UK, 240p. Prapo, D. E. 2004. As Caatingas da América do Sul. Universitaria, UFPE (Ed.). In: Ecologia e Conservagao da Caatinga. Recife-Pe. 43-73. eae: D., S. Catrns & T. DILLON. 1986. Seven forms of rarity and their frequency in the flora of the British Isles. In M. E. Soulé (Ed.). Conservation biology: The science of scarcity and di- versit. University of Michingan, Sunderland, Massachusetts. 584 pp. SantpntO) E. V.S. B., M. F. A. V. Santos, & M. R. RIBEIRO. 1991. Semelhangas Vegetacionais em Sete Solos da Caatinga. Pesquisa Agropecuaria, Brasilia, 27(2): 305 - 314. SCHREIBER, H. 1978. Dispersal centres of SPHINGIDAE (Lepi- doptera) in the Neotropical Region. Biogeographica 10: 195 p. Received 18 October 2004; revised and accepted 12 May 2005 VOLUME 59, NUMBER 4 Journal of the Lepidopterists’ Society 59(4), 2005, 219-222 BEHAVIORAL OBSERVATIONS ON THE EARLY STAGES OF JAMIDES CELENO (CRAMER) (LYCAENIDAE) AT CAT TIEN NATIONAL PARK, VIETNAM: AN OBLIGATE MYRMECOPHILE? Rop Eastwoop Australian School of Environmental Studies, Griffith University, Nathan, Qld 4111, Australia, email: r.eastwood@griffith.edu.au R.L. KITCHING Australian School of Environmental Studies, Griffith University, Nathan, Qld 4111, Australia, email: rkitching@griffith.edu.au AND Hut Buu MANH Cat Tien National Park, Nam Cat Tien Commune, Tan Phu, Dong Nai Province, Vietnam ABSTRACT. The life history of Jamides celeno has been well documented across its range. The larvae feed on plants from six families and are attended by ants from seven genera. This paper documents a new host plant record and additional attendant ant species from observations made in Cat Tien National Park, Vietnam. We discuss ecological and behavioral traits of the early stages of J. celeno and their associated ants, and suggest that the categorization of J. celeno as a facultative myrmecophile may be incorrect. Additional key words: Ant feeding, Camponotus, facultative, Formicidae, mutualism, Polyommatinae. The lycaenid butterfly, Jamides celeno (Cramer) is a sexually dimorphic species in the — subfamily Polyommatinae. It is widespread in south and east Asia distributed from Sri Lanka, India and Nepal to Taiwan and south China, and throughout the Malaysian Archipelago to New Guinea and the Solomon Islands (Corbet et al. 1992). In Vietnam, J. celeno is one of the most common species of butterflies, occurring in many habitats ranging from primary evergreen forest and forest gaps to cultivated areas (Monastyrskii & Devyatkin 2002). Larvae of J. celeno feed on a variety of plants in the families Fabaceae, Caesalpinaceae, Sterculiaceae, Meliaceae, Zingiberaceae —_ and Marantaceae (Robinson et al. 2001), and the early stages have been recorded in association with ant species across seven genera (Corbet et al. 1992, Ballmer 2003, Cleary & Grill 2004). Larval association with ants, termed myrmecophily, is a well-known phenomenon in the Lycaenidae (Newcomer 1912, Hinton 1951, Pierce et al. 2002). Symbiotic interactions fall into two broad categories: obligate, where the survival of the butterfly species is dependent on the presence of ants; or facultative, where butterfly larvae may or may not have ants in attendance (Fiedler 1991a, Eastwood & Fraser 1999). As a rule, obligate myrmecophiles associate with a single species or a group of closely related species of ants, whereas facultative myrmecophiles typically associate with a variety of ant species, often from different genera or subfamilies (Pierce et al. 2002). Accordingly, recent workers have categorized J. celeno as a facultative myrmecophile (Fiedler 1996, Ballmer 2003). During a survey of arthropod biodiversity in Nam Cat Tien Park, Vietnam, we observed J. celeno adults and early stages on a regular basis. The Nam Cat Tien section of Cat Tien National Park is located in Dong Nai Province 150 km north of Ho Chi Minh City in the monsoonal tropical region of southern Vietnam. It contains the largest remaining lowland tropical forest in southern Vietnam. A semi-evergreen regeneration forest dominated by Lagerstroemia spp. (Lithraceae), it has an average annual rainfall of ~2300 mm and average temperatures ranging from 15 °C to 35 °C. The local Chau Ma and S'Tieng people have practiced shifting cultivation in and around the park for generations. Our visit in July 2002 coincided with the wet season, which lasts from April/May to October/November. This paper documents new ecological and behavioral traits of J. celeno early stages and their associated ants, and discusses the category of myrmecophily to which J. celeno is usually assigned. OBSERVATIONS Adult J. celeno were often encountered in open areas adjoining tracks and roads through Cat Tien National Park, but many adults were also observed within the forest where their presence usually indicated the proximity of a breeding site. Understory plants were typically less than 3 m in height and larvae were encountered on leaf flushes about 1.8 m above ground (1.82 m + 0.63 SD, n=11). Larvae formed loose aggregations (2.36 + 0.92 SD individuals, n=11) and fed exclusively on the soft new growth flush of Euodia meliaefolia Benth. (Rutaceae) growing in the understory. Eggs were laid on the new leaves when there was sufficient foliage to sustain a small cohort of larvae. Leaf flush varied from pale green to pale yellow in color before changing to a darker green as the leaves hardened. Larvae were either green or yellow, but always matched the color of the leaves on which they were resting. Mature larvae became brown before leaving the tree to pupate. Larvae rested under the foliage during the day and began feeding late in the afternoon (ca. 1600 h). Their feeding patterns followed the flush of new growth as it progr essed but larvae did not appear to be food-limited. Nevertheless, the butterfly had a window of only about 2-3 weeks on each tree in which the life cycles of multiple broods could be completed before the leaves hardened. Larval duration was uupgee six days from egg hatching to pre- pupa, and pupal eon for three individuals was seven (n=2) and eight days. Of the eleven larval aggregations observed, five ant two subfamilies were in Camponotus (Tanaemyrmex) sp. (n=3); Camponotus (Myrmosaulos) singularis (F. Smith) (n=2); Polyrhachis species in attendance: (Myrmhopla) rufpes F. Smith (n=1); Anoplolepis gracilipes (F. Smith) (n=2) (all Formicinae); and Crematogaster sp. (n=2) (Myrmicinae). Ant and butterfly voucher specimens are lodged at the Museum of Comparative Zoology, Harvard University (RE-02- A057, 58, 61, 73, 87, 88, 90, 91, 94, 105, 106, 107, 108, 160). Two larvae were found untended on different plants but the following day one (presumably the same larva) was tended by six GC singularis and the other mature larva could not be round on the plant or in the leaf litter. Six to ten ants typically tended larval aggregations, thus providing a significant deterrent to potential enemies of the larva. On one occasion a single Crematogaster was found tending a larva together with Camponotus When Crematogaster came into contact with one of the several singularis. the Camponotus, however, it leapt off the leaf. On another occasion several Crematogaster were observed tending three J. celeno larvae but later the same day the larvae were tended by Camponotus singularis. Ants were observed antennating the entire dorsal surface of the larvae and at times the ant's tongue could be seen extending to touch the larval cuticle as well as the dorsal nectary organ (see Bell 1918 for a very detailed description of the morphology of J. celeno early stages). J. celeno larvae pupated at the base of the host- -plant or in the leaf litter surrounding the base of the tree. Larvae tended by Camponotus (Tanaemyrmex) sp. pupated in a protective chamber or byre constructed by the ants from soil and leaf debris at the base of the tree where ants remained in attendance. Other ant species, including C. singularis, which did not construct a byre, were not found tending pupae. A. gracilipes were found in proximity to, but not tending, ihe e cecal larvae in the leaf litter. Individuals of both species of Camponotus, but JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY especially C. singularis, were observed chewing on leaf edges where J. celeno larvae had been feeding. C. singularis the leaves and continued chewing the leaf edges, after the lycaenid larvae had left remained on to pupate, ev entually leaving a rough sawtooth pattern along the leaf margin. Ev idence of ant chewing was also adjoining those where lycaenid larvae had been feeding. A few small stingless seen on the edges of leaves bees were attracted to the ly caenid feeding sites. DISCUSSION The presence of ants at leaf margins in proximity to feeding lepidopteran _ larvae has been recorded previously (Ford 1945, Owen 1971, Fiedler 1991a); the ants apparently taking phloem exuding from leaves damaged by butterfly larval feeding. It is known that adult leafcutter ants in South America derive most of their nutrition from leaf phloem as they cut or chew the leaves before feeding the pulp to their fungal symbionts (Holldobler & Wilson 1990). We could find no other records of ant chewing on leaves damaged by herbivores or around the leaf edges, and it is not known if ants behave in this manner in the absence of butterfly larvae. The construction of a byre or phytophagous lycaenid larvae, or to ‘farm! honeydew- producing phenomenon among nectar-gathering ants (H6lldobler & Wilson 1990; Anderson & McShea 2001). Several lycaenid species in the Australian genus Ogyris, namely O. genoveva (Hewitson), O. zosine (Hewitson), O. idmo (Hewitson), and O. otanes (C. Felder & R. Felder), have an obligate relationship with Camponotus ants (Eastwood & Fraser 1999). In these associations, butterfly larvae feed nocturnally on mistletoe high up in the trees but hide during the day in ealene constructed by the ants at the base of the trees (Eastwood 1997). Accordingly, it was surprising to find that Camponotus ants in Vietnam constructed a similar gallery to house insects generally, is a well-known structure for J. celeno, suggesting that the behavior is a plesiomorphic trait in Camponotus that evolved before the evolution of mutualistic associations with Ogyris or Jamides. A similar behavior has been recorded with Mexican C. (abdominalis) atriceps F. Smith, tending the riodinid Anatole rossi Clench (Ross 1966), and South American Camponotus tending Polyommatine lycaenids (Benyamini 1995). It seems that the butterfly larvae on all three continents are independently exploiting a trait in Camponotus ants, which are pre-adapted to construct byres around free-living insect mutualists. It is also interesting that J. celeno larvae behaved differently in choosing pupation sites when attended by different ant species. The Camponotus (Tanaemyrmex) sp. may have shepherded penultimate lycaenid instars into the byre. VOLUME 59, NUMBER 4 However, relationships between lycaenid larvae and ants are known to differ depending on the attendant ant species (Axén 2000; Fraser et al. 2001). As is the case with other Jamides species (Fiedler 1996), J. celeno did not display preference for a particular ant species, but the regularity with which we found larvae and ants together suggests that ant attendance is important for the survival of this species in the study area. Furthermore, in Thailand, Ballmer (2003) recorded 48 J. celeno larvae feeding within Pueraria phaseoloides (Roxb.) Benth. (Fabaceae) flowers and all were tended by ants, although the ants were from six genera. Cleary and Grill (2004) recorded 1079 J. celeno larvae in Borneo predominantly attended by A. gracilipes, and found there were significantly more caterpillars on plants with ants present than on plants without ants. This present study recorded five ant species from four genera, and additional attendant-ant species are noted! in the literature. Although many attendant-ant species are recorded, the regularity of attendant-ant records and the high proportion of attendance levels suggest that tending ants may play a significant role in the survival of J. celeno overall. ie celeno larvae have also been found without ants (Parsons 1999, Cleary & Grill 2004); however, many other obligate lycaenid species have likewise been found untended (e.g. Eastwood & Fraser 1999). Furthermore, a recent survey of the obligate myrmecophile Jalmenus evagoras (Donovan), sation was thought to associate preferentially with a single species of ant, showed this butterfly associated wath seven species (from the same genus) across its range (RE unpublished data). Accordingly, it is plausible that for some obligate myrmecophiles, attendant-ant species specificity may be less important. The categorization of lycaenid-ant relationships was based historically on ant attendance in non-tropical Lepidoptera (e.g. Malicky 1969, 1970; Henning 1983; Fiedler 1991a,b). Lycaenids are regarded as facultative if they associate with a variety of ant species, and obligate if they consistently associate with only one or a few dlosdy related ant species. Ants that attend obligate myrmecophiles are typically aggressive, are spatially or temporally dominant and form long-lived colonies (Atsatt 1981: Pierce 1987: Fiedler 1991la, 2001; Eastwood & Fraser 1999). Thus, they are a predictable resource that provides optimal protection for lycaenid larvae. In the rainforest, however, an extremely diverse and heterogeneous could necessitate association with many ant species if lycaenid larvae were to survive attacks from predators and parasites. In particular, the protection conferred by mutualistic ants could be an important and effective defence against ant fauna predacious ants, which are ubiquitous and known to attack lepidopterous larvae in south east Asian lowland rainforests (Floren et al. 2002). While lycaenid survival rates may be variable, depending on the species of tending ant (Eastwood 1997; Fraser et al. 2001), association should significantly improve the survival rate overall. In particular, it seems likely that pupae tended by Camponotus (Tanaemyrmex) sp., would have a higher survival rate since the ants protect them in an rdereround chamber. So, although the relationship that J. celeno has with ants is less specific than generally observed for obligate myrmecophiles, lack of specificity in tending ants may be a function of the heterogeneity of the ant community and not necessarily because J. celeno has a facultative relationship with ants. In fact. we suggest that the dependence that J. celeno has on attendant ants falls somewhere in the upper bounds of the continuum between facultative and obligate. Thus, the categorization of lycaenid-ant relationships, in this instance, into obligate or facultative myrmecophiles based on the number of ant partner species may be overly simplistic or misleading (e.g. Fiedler 1991a, 1996, 2001, Eastwood & Fraser 1999, Ballmer 2003). J. celeno was recorded feeding on a single host plant at Cat Tien Park, but the record is most likely a temporal phenological preference or local host plant preference since many host plants have been recorded for this butterfly, and local preferences, especially on new growth, are recorded elsewhere (Cleary & Grill 2004). Multiple host plant use, often across several families of plants, is a characteristic of lycaenids having obligate relationships with ants (Pierce & Elgar 1985). The larvae of J. celeno are also difficult to detect because of their ability to adopt the color of their substrate and their tendency to hide under foliage during the day; so, in addition to the protection afforded by ants, J. celeno employs multiple strategies for survival. These observations highlight how much we still do not know about the intimacy of lycaenid-ant relationships and how much we can still learn about their joint survival strategies and the evolutionary forces shaping their relationships. ant ACKNOWLEDGEMENTS We thank Tran Van Mui, Park Director and his staff, for their hospi- tality and for allowing us to collect and study insects at Cat Tien National Park. Thanks also to Tran Van Binh (Cat Tien National Park) for identify- ing the plants; and David Lohman (Harvard University), Rudy Kohout (Queensland Museum) and Sitorn Hasin (Thai Ant Museum - Kasetsart University) who identified the ants. We thank Stefan Cover (Harvard Uni- versity) for useful discussion and for providing references: and Greg Ballmer (University of California, Riverside) for advice on the identifica- tion of Jamides celeno. The biodiversity survey in Nam Cat Tien formed part of an ACIAR Project directed by Professor R. A. I. 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Received for publication 23 February 2005; revised and accepted 9 August 2005 VOLUME 59, NUMBER 4 223 Journal of the Lepidopterists’ Society 59(4), 2005, 223-225 POPULATION BIOLOGY OF TWO SPECIES OF HELICONIUS (NYMPHALIDAE: HELICONIINAE) IN A SEMI-DECIDUOUS FOREST IN SOUTHEASTERN BRAZIL RAFAEL BARRETO DE-ANDRADE AND ANDRE VICTOR LUCCI FREITAS Departamento de Zoologia and Museu de Histéria Natural, Instituto de Biologia, Universidade Estadual de Campinas, CP 6109, CEP 13083- 970, Campinas, Sao Paulo, Brazil email: baku@unicamp.br ABSTRACT. Populations of two species of butterflies, Heliconius erato and Heliconius ethilla, were studied during 17 months in SE Brazil. For H. erato, the number of individuals present per day varied from one to 10. Time of residence was 24. 9 +18.97 days, with the maximum 59 days. Males were recorded traveling distances up to 200 m. A study of wing color patterns of H. erato showed results similar to those of previous studies, including the variation in the number of red raylets. For H. ethilla, the number of indi- viduals present per day varied from one to 15. Time of residence was 32.6 + 23.93 days, with the maximum registered of 106 days. Males can travel distances up to 650 m. For both species, the population peaks occurred in March and May of both years. The sex ratio of individuals captured was male biased for most of the months. Age structure was not stable, though intermediate-age indi- viduals dominated in every month. The difference between male and female mean forewing lengths was not significant. These popu- lation features agree with patterns previously observed in southern Brazil populations, though with much lower population num- bers. Additional key words: mark-recapture, Heliconiini, red raylets Although butterflies in the genus Heliconius are among the most studied tropical species, with many publications covering diverse aspects of their biology, such as systematics, ecology, genetics and evolution (Turner 1971, Ehrlich & Gilbert 1973, Cook et al. 1976, Araujo 1980, Mallet & Jackson 1980, Brown 1981, Ehrlich 1984, Mallet et al. 1987, Quintero 1988, Mallet & Gilbert 1995, Ramos & Freitas 1999, Jiggins et al. 2001, Gilbert 2003), data from population biology of most species are still unavailable or incomplete (Ramos & Freitas 1999). The work by Ehrlich & Gilbert (1973) brought important information on the biology of Heliconius ethilla and is, so far, the only published population report of this species. In contrast, Heliconius erato has been studied in many different places in the Neotropics (Turner 1971, Benson, 1972, Araujo 1980, Saalfeld & Araujo 1981, Romanowsky et al. 1985, Ramos & Freitas 1999), including studies of genetics and variation on wing color patterns (Saalfeld & Araujo 1981, Pansera & Aerie 1983, Sheppard et al. 1985, Oliveira & Araujo 1992, Ramos & Freitas 1999). Most of the published population studies cover only three different regions in the Neotropics (Trinidad, Rio Grande do Sul and coastal Sao Paulo state) and none in most seasonal semi-deciduous forests or cerrados. In a recent paper, Ramos & Freitas (1999) suggested that H. erato is an ecologically plastic species, in use of resources (larval and adults), behavior and population parameters. In this context, the descriptions of different populations of H. erato are important to support a broader view about the natural history of this species. The objective of the present study is to describe and compare population parameters of H. erato phyyllis (Fabricius) and H. ethilla narcaea (Godart) in a small forest fragment of semi-deciduous forest — in Southeastern Brazil, reporting also the cyclical annual variation in four color pattern elements in H. erato. STUDY SITES AND METHIDS The study was carried out in the 250-ha semi- deciduous forest Reserva da Mata da Santa Genebra (22° 49'S, 47° O7'W), a municipal forest reserve in Campinas, Sio Paulo State, SE Brazil. The annual rainfall is near 1400mm and the average annual temperature is 20.6°C (data from the Campinas Agronomical Institute). The climate of the region is aren acterized by a dry, cold season (May to August) and a wet, warm season (September to April) (more details and a climatic diagram can be found in Vanini et al. 2000). Detailed descriptions and maps of the study area are available in Morellato & Leitao-Filho (1995). Butterflies were marked, released and recaptured (MRR) along a trail (1150m long divided into 50m sectors) during 17 months, from December 24, 2001 to May 29, 2003 (1-3 times a week), for a total of 91 d. ays (about 3 hours/day). Butterflies were net- captured, individually numbered on the underside of both forewings with a black permanent felt-tipped pen, and released. Individual characteristics of each individual 294 (age, forewing length, point of capture, sex, food sources and color patterns) were recorded for later analysis (as in Ramos & Freitas 1999). based on six categories (freshly emerged, intermediate, old, very old, Phe was used as an additional measure of age of individual butterflies (following Ehrlich & Davidson 1960, Brussard & Ehrlich 1970, Ehrlich & Gilbert 1973). These six categories were grouped into three (new, intermediate and old) for analy sis (as in Freitas 1993, 1996). Age structure was calculated f through the monthly means of daily proportions of ‘each category. Wing wear, new, ‘or males only, Individual vagility was measured as the maximum less distance from capture to recapture points. For Heliconius erato, four color-patterm variations were recorded: number and shape of the red raylets on the ventral hindwing; presence and color of the cubital spot on the dorsal forewing (more details and pictures in Ramos & Freitas 1999); number of “light yellow squares” on the apical ventral hindwing; and ie color of the prolegs (entirely yellow or with tarsal portion red). The MRR data were analyzed by the Jolly-Seber method for estimating population parameters ( (software developed by R. B. Francini, UNISANTOS) for obtaining the “estimated numbers”. Only males were Ronvidered for this analysis due to the low number of females captured. “number of individuals captured per day” including recaptures, and “number of individuals present per day” (NIPD). To estimate NIPD, recaptured individuals were considered to be present on all days between the first and last capture. For H. erato, population parameters and residence time were calculated from January to May 2002 recorded as (NICD), Daily results were and the color patterns data analyzed from December 2001 to 2002, December due to low numbers of captured 150- 120 + 90 4 604 Number of Individuals Months Fic. 1 - Number of males H. erato from December 2001 to May 2003 in Santa Genebra, Campinas, SP. Solid circles = NIPD, open circles = estimated number based on Jolly-Seber (bars = 1 standard error). JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY individuals on other periods. For H. ethilla analysis were made form December 2001 to May 2003. The sex ratio was calculated through the monthly means of daily proportions in NIPD. RESULTS Heliconius erato phyllis Population biology. From December 2001 to May 2003, the NICD for males varied from one to five (mean = 1.6; SD = 0.91; n = 52 days). The calculated NIPD varied from zero to 12 (mean = 3.4; SD = 2.03; n = 52 days). Estimated numbers based on Jolly-Seber (Fig. 1) suggests that population numbers are not very much higher than numbers obtained by the NIPD. The population peaks occurred in March and May of both years, although in 2003 the increase in butterfly numbers was about half of that in the same period of 2002. In other months of both years the butterfly numbers remained low and stable (Fig. 1). Sex ratio. During this study, 47 males and 12 females were captured, giving a male biased sex ratio (x? = 19.59; p < 0.001; DF = 1). Males dominated in every month considered (Fig. 2). Males were recaptured from one to six times and females from one to four times; 23 males and three females were recaptured at least once. Age structure. Even though the proportion of “intermediate” individuals remained greater in every month except for May 2002, the proportions of “new” and “old” individuals were unstable during the months 100 “ cy w E 5 50 cs Cc a 2 } a 0 ULE) -pMiee AM, J F M A M 2002 Months 2003 Fic. 2. Sex ratio of H. erato from January 2002 to May 2003 in Santa Genebra, Campinas, SP. considered. Figure 3 shows the proportion of the three age classes from January to May 2002. Residence time. The residence time varied from three to 59 days (mean = 24.9 days; SD = 18.97). Life expectancy (following Cook et al. 1967) was 11.21 days. Vagility. Only males were analyzed due to the low number of females. The greatest distance recorded along the trail flown by an individual was 200 meters (mean = 65.2 m; SD = 61.12 m; n = 23). Most of the individuals were recaptured at least once in a different VOLUME 59, NUMBER 4 bo bo Ol TABLE 1 - Frequency distributions and sample size (N), of the three color patterns recorded in H. erato butterflies of both sexes captured in the Santa Genebra from December 2001 to December 2002 (2003 not included due to the low number of captures). For number of red raylets and number of light yellow squares, data are presented as mean + standard deviation. Month/year proleg color red raylets Light yellow squares N yellow red tip Dec/01-Jan/02 2 2 5+0 4.25 +05 4 Feb/02 6 1 4.14 + 1.07 3.5 + 0.55 fl Mar/02 3) 2 4.3+1.16 3.4 + 1.35 10 Apr/02 9 3 3.58 + 1.16 3.17 + 1.59 12 May/02 6 1 3.71 + 1.5 3.29 + 0.49 i Oct/02—Dec/02 2 0 3.5 + 2.12 4+1].4] 2 100 Percentage among age classes 2002 Months Fic. 3 - Age structure of males H. erato from January 2002 to May 2003 in Santa Genebra, Campinas, SP. Black = new individuals, gray = intermediate, white = old. site from that of first capture (18 out of 23 recaptured individuals). Wing size. The forewing length of males varied from 30 mm to 39 mm (mean = 35.2 mm; SD = 2.69 mm; n = 47) and of females from 32 mm to 40 mm (mean = 36.1 mm; SD = 2.57 mm; n = 12). This difference was not significant (t = 1,03; DF = 17,7; p = 0,314). Resource utilization. For 39 visits recorded, 69% were on Lantana camara L. (Verbenaceae), 18% on Chromolaena odorata (L.) R. King & H. Robinson (Asteraceae) and 13% on flowers of other genera, including Lippia sp. (Verbenaceae) and Manettia sp. (Rubiaceae). In both years, L. camara flowers decreased after May, and the visits to other nectar sources increased. Color patterns. The number of red raylets varied from 1 to 6 in both sexes (mean = 3.88; SD = 1.20: n = 58). Monthly analysis for this trait was done only for 2002, and some months were grouped due to the small number of marked individuals. The average number decreases in colder months (Table 1). The total number of individuals with dot shaped red raylets (41) was greater than the number of individuals with line shaped red raylets (17) (¥? = 9.12; p < 0.005; DF = 1) and fits in 3:1 distribution (x? =0.09; p = 0.75; DF=1). Four categories of cubital spot on the dorsal forewing were recorded with uneven distribution. The predominant category was “yellow” (22 individuals), followed by “absent” (15), “red” (5) and “fusing with the subapical red bar” (5). The number of “light yellow squares” varied from 0 to 5 in males (mean = 3.0: SD = 1.33: n = 45) and 4 or 5 in females (mean = 4.2; SD = 0.38; n = 12). Monthly analysis for this trait was done only from December 2001 to December 2002, and some months were grouped due to the small number of marked individuals. The average number decreases in colder months (Table Il)), Two color patterns of the prothoracic legs were recorded. The number of individuals with “yellow” legs (33) was greater than the number of individuals with “yellow with red tips” (9) (y? = 13.09; p < 0.001; DF = 1) and also fits in a 3:1 distribution (y? = 0; p=0.99; DF = 1). Monthly analysis for this trait was done only for 2002, and some months were grouped due to the small number of marked individuals (Table 1). Heliconius ethilla narcaea Population biology. The NICD for males varied from one to six (mean = 2.0; SD = 1.18; n = 90 days). The NIPD varied from one to 15 (mean = 6.4; SD = 4.43; n = 90 days). As in H. erato, estimated numbers based on Jolly-Seber (Fig. 1) are not very much higher than actual numbers obtained by the NIPD. The population peaks (based on NIPD) occurred in March and May of both years, and in the remaining months the population numbers remained low and stable (Fig. 4). Sex ratio. During this study, 123 males and 35 females were captured giving a male biased sex ratio (7 = 47.9; p < 0.001; DF = 1). Males dominated in every month considered, except for December 2002 (Fig. 5). 226 250- 200 150 - 100 3 60-4 Number of Individuals Fic. 4. Number of males H. ethilla from December 2001 to June 2003 in Santa Genebra, Campinas, SP. Solid circles = NIPD, open circles = estimated number based on Jolly Seber (bars = 1 standard error). Males were recaptured from one to 13 times and females from one to three times; 53 males and 11 females were recaptured at least once. Age structure. The proportion of “intermediate” individuals was most of the months The proportions of individuals varied during the months considered. Figure greatest in considered. “new and “old” 6 shows the proportion of the three age classes from January 2002 to May 2003. Residence time. The time of residence two to 106 days (mean = 32.6 days; SD = eee (following Cook et al. 1967) was 44.61 days. Vagility. Only males were analyzed due to the low varied from 23.93). Life number of females. The greatest recorded distance along the trail flown by an individual was 650 meters (mean = 83.96 m; SD = 94.45 m:n = 53). Most of the individuals were recaptured at least once in a different site from that of first capture (44 out of 53 recaptured individuals). Wing size. The forewing length of males varied from 35 mm to 46 mm (mean = 41.5: SD = 2.14: n = 121) and of females from 38 mm to 45 mm (mean = 41.5; SD = 1.77; n = 35). This difference was not significant (t = 039; DF = 65.4; p = 0.969). Resource utilization. For 128 visits recorded, 52% 26% on C. odorata and 23 including Lippia sp. Manettia sp. (Rubiaceae) and Passiflora suberosa L. (Passifloraceae). As described for H. erato, were on L. camara, % on other (Verbenaceae), flowers of genera, visitation to other nectar sources increased after May when flower production decreased for L. camara. DISCUSSION Population biology. The populations showed three distinct phases: growth in January, February and March, JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Wee aa HH a 7 E ‘5 50 es c a 2 a a 0 JFMAMJJASOND JSFMAM 2002 Months 2003 Fic. 5. Sex ratio of H. ethilla from January 2002 to May 2003 in Santa Genebra, Campinas, SP. peak in April and decline in May and June. The instability in numbers differs from the patterns found in other tropical Heliconius studies (Turner 1971, Ehrlich & Gilbert 1973; Ramos & Freitas 1999) that reported stable populations throughout the year, and was to some extent similar to those studied in southern Brazil (Araujo, 1980; Romanowsky et al., 1985), with marked the number of individuals. However, in southern Brazil, population decline seems seasonal variation in to be correlated with a decrease in ambient temperature (Saalfeld & Araujo, 1981; Romanowsky et al., 1985), whereas at Santa Genebra the decline is probably due to The number of individuals in the studied populations was relatively low (up to 60 decrease in rainfall. individuals based on Jolly-Seber estimates in both species) compared with studies in south Brazil, where captures could be as high as 60 individuals in a single day (Romanowsky et al. 1985). In the periods of population peak, the number of captured individuals in both species was equivalent to those reported in other studies in tropical sites (Turner 1971, Ehrlich & Gilbert 1973; Ramos & Freitas 1999). Rainfall is considered as an important factor limiting population numbers for Heliconius, since it is closely 100 a o e a os 7) o i=) os E 50 oc — o o i=\] a is o 5 o 0 JFMAMJJASOND JSIFMAM 2002 Months 2003 Fic. 6 - Age structure of males H. ethilla from January 2002 to May 2003 in Santa Genebra, Campinas, SP. Black = new individuals, gray = intermediate, white = old. VOLUME 59, NUMBER 4 linked with adult resource availability (see Ehrlich & Gilbert, 1973 and Gilbert, 1984). The seasonal variation in rainfall probably explains the population fluctuation patterns reported in the present paper, which is quite different from stable populations reported in tropical or coastal areas where rainfall is constant throughout the year. The accentuated decline and low population numbers in the Santa Genebra populations can also be a consequence of migration of individuals to surrounding subpopulations in more — suitable environments (as proposed in Saalfeld & Araujo 1981) outside our sampling area. High values of residence time agree with other studies of Heliconius (Turner 1971, Benson 1972 Ehrlich & Gilbert 1973, Araujo 1980, Quintero 1988S, Ramos & Freitas 1999). These numbers may be related to restricted dispersion of adults and to several visits to resources patches. For both species studied here, sex ratio was male biased, agreeing with field results from several butterfly studies (Ehrlich & Gilbert 1973, Mallet & Jackson 1980, Ehrlich 1984, Ehrlich et al. 1984, Freitas 1993, Ramos & Freitas 1999). Differential behavior in the sexes may cause this deviation. The data on resource utilization by both species suggests that they are using all available flowers in the study site. Flowers of L. camara were the most visited by both Heliconius in Santa Genebra. Previous studies showed that Lantana is a major pollen source for some species of Heliconius in several tropical sites (Boggs et al. 1981, Ramos & Freitas 1999). We suggest that intense use of Lantana and other flowers with small pollen grains could be related with more generalist edge species of Heliconius. Comparing both species in the study site, H. ethilla is more abundant and persistent than H. erato, but both species show the same seasonal fluctuation pattern of population. Both species are suggested as plastic species (Ramos & Freitas, 1999 and this paper), but there are more data available for H. erato than for H. ethilla at the moment. Additional studies of H. ethilla, including host plant use, larval performance and population dynamics in different habitats are needed. Color patterns in Heliconius erato. The decline in the mean number of red raylets in colder months agrees with Pansera & Araujo (1983), Oliveira & Araujo (1992) and Ramos & Freitas (1999). The mean number of “light yellow squares” also seems to decline in colder natetit and the population becomes virtually absent during winter and spring. For this reason it is difficult to perform any analysis to help to clarify the direction and origin of the variation of this and other traits. The three-to-one proportion between the two colors of prothoracic legs may be a hint about the heritability of this trait. However, due to the low number of H. erato captured, it is difficult to infer much about the biology of the color traits considered in this study. Further studies could investigate the influence of seasonal climate pattern on the av vailability of resources for long-life butterflies such as Heliconius, and how these factors restrain population numbers. Also, the understanding of temporal and spatial fluctuation patterns of color patterns can elucidate ecological processes related to natural selection and genetic variability. ACKNOWLEDGEMENTS Claudemir R. Dias, Alice Moraes, and Humberto P. Dutra helped in fieldwork. Keith Brown helped in various phases of this work with suggestions in the final version of the manuscript. Carla Penz and an anonymous referee gave valuable suggestions in the last version of the manuscript. Rafael Andrade thanks Tais Mazzola for helping in various phases of this work. This study was supported by Fapesp, PIBIC/SAE and the National Science Foundation (Fapesp grants 00/01484-1 and 04/05269-9: NSF DEB-0316505), and is part of the BIOTA-FAPESP pro- gram (98/05101-S). LITERATURE CITED ArAUJO, A. M. 1980. Estudos genéticos e ecolégicos em Heliconius erato (Lepidoptera, Nymphalidae. Actas TV Congr. Latinoam. Genética 2: 199-206. BENSON, W. W. 1972. Natural selection for Miillerian mimicry in He- liconius erato in Costa Rica. Science 176: 936-939. Boccs, C. L., J. T. SMILEY & L. E. GILBert. 1981. Patterns of pollen exploitation by Heliconius butterflies. Oecologia 48: 284-289. Brown, K. S. JR. 1981. The biology of Heliconius and related genera. Ann. Rey. Ent. 26: 427-456. BrussarD, P. F. & P. R. EHRLICH. 1970. The population structure of Erebia epipsodea (Lepidoptera: Satyrinae). Ecology 51: 119-129. Cook, L. M., E. W. THoMason & A. M. YouNG. 1976. Population structure, dynamics and dispersal of the tropical butterfly Helico- nius charitonius. Journal of Animal Ecology 45: 851-863. EuRticu, P. R. 1984. The structure and dynamics of butterfly popu- lations, pp. 25-40. In: R. I. Vane -Wright & P. R. Ackery (eds.), The biology of butterflies. Academic Press, London. — &S. E. Davipson. 1960. Techniques for capture-recapture studies on Lepidoptera populations. J. Lepid. Soc. 14: 227-229. —— &L.E. GILBERT. 1973. Population structure and dynamics of the tropic butterfly Heliconius ethilla. Biotropica 5: 69-82. ——, A. E. LAuNER & D. D. Murpuy. 1984. Can sex ratio be defined or determined? The case of a population of checkerspot butter- flies. Amer. Nat. 124: 527-539. Freitas, A. V. L. 1993. Biology and population dynamics of Placidula nee a relict ithomiine butterfly (Nymphalidae: Ithomi- inae). 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. GILBERT, L. E. 1984. Biology of butterfly communities, pp. 41-46. In R. I. Vane -Wright & P. R. Ackery (eds.), The biology of butter- flies. Academic Press, London. . 2003. Adaptive novelty through introgression in Heliconius wing patterns: evidence for a shared genetic "tool box" from synthetic hybrid zones, and a theory of diversification, pp. 281-318. In C. L. Boggs, W. B. Watt & P. R. Ehrlich (eds.), Butterflies: Ecology and Evolution Taking Flight. Univ. Chicago Press, Chicago. Jiccins, C. D., M. Linares, R. E. Naissrt, C. SaLazar, Z. H. Yanc & J. MALLET. 2001. Sex-linked hybrid sterility in a butterfly. Evo- lution 55: 1631-1638. MALLET, J. B. & D. A. JACKSON. 1980. The ecology and social behav- iour of the neotropical butterfly Heliconius xanthocles Bates in Colombia. Zool. J. Soc. 70: 1-13. MaLLet, J. L. B., J. T. Loncino, D. Murawski & S. DE GAMBOA. 1987. Handling effects in Heliconius: where do all butterflies go?. J. of Anim. Ecol. 56; 377-386. MALLET, J. & L. E. GitBert. 1995. Why are there so many mimicry rings? Correlations between habitat, behaviour and mimicry in Heliconius butterflies. Biological Journal of the Linnean Society, 55: 159-180. MILLER, L. D. 1970. Nomenclature of wing veins and cells; J. Res. Lepid. 8: 37-48. MorELLATO, R.C. & H. F. Lerrao FILHO, 1995. Ecologia e preser- vacgao de uma floresta tropical urbana; reserva Santa Genebra. Campinas: Ed. da UNICAMP. Ouiverra, D. L. & A. M. Araujo. 1992. Studies on the genetics and ecology of Heliconius erato (Lepidoptera; Nymphalidade). IV. Ef- fective size and variability of the red raylets in natural population. Rey. Brasil. Genet. 15: 789-799. PaNseRA, M. C. G. & A. M. ARAUJO. 1983. Distribution and heri- tability of the red raylets in Heliconius erato phyllis (Lepid.; Nymph.). Heredity 51: 643-652. QuinTERO, H. E. 1988. Population dynamics of the butterfly Helico- nius charitonius L. in Puerto Rico. Caribb J. Sci, 24: 155-160. Ramos, R. R. & A. V. L. FreIvAs. 1999. Population biology and wing color variation in Heliconius erato phyllis (Nymphalidae). J. Lepid. Soc, 53: 11-21. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY RoMANowsky, H. P., R. Gus & A. M. Araujo. 1985. Studies on the genetics and ecology of Heliconius erato (Lepid. Nymph). UI. Population size, preadult mortality, adult resources and polymor- phism in natural populations. Rev. Brasil. Biol. 45: 563-569. SAALFELD, K. & A. M. ARAUJO. 1981. Studies on the genetics and ecology of Heliconius erato (Lepidoptera, Nymphalidae). I: De- mography of a natural population; Rey. Brasil. Biol. 41: 855-860. SHEPPARD, P. M., J. R. G. TURNER, K. S. BROWN JR., W. W. BENSON & M. C. SINGER. 1985. Genetics and the evolution of Muellerian mimicry in Heliconius butterflies. Phil. Trans. Roy. Soc. London 308: 433-613. TURNER, J. R. G. 1971. Experiments on the demography of tropical butterflies, If. Longevity and home range behavior in Heliconius erato. Biotropica 3: 21-31. VANINI, F., V. Bonato & A. V. L. Freitas. 2000. Polyphenism and population biology of Eurema elathea (Pieridae) in a disturbed environment in tropical Brazil. J. Lepid. Soc. 53: 159-168. Received for publication 20 Janurary 2005; revised and accepted VOLUME 59, NUMBER 4 Journal of the Lepidopterists’ Society 59(4), 2005, 229-233 REPRODUCTIVE BULK IN CAPITAL-BREEDING LEPIDOPTERA WILLIAM E.. MILLER Department of Entomology, University of Minnesota, St. Paul, Minnesota 55108 USA, e-mail: mille014@umn.edu. ABSTRACT. Reproductive bulk is the weight of the individual egg load, also the mathematical product of individual fecundity x. mean egg weight. Somatic bulk is its coulerea Interrelations among nonindependent maternal weight, reproductive bulk, somatic bulk, fecundity, egg Ww eight and related variables were explored with eight sets of data; three original, and five previously published, all representing four capital- breeder species in three families. In traditional fecundity. —maternal weight regressions, response and explanatory variables are not statistically independent because maternal weight includes reproductive bulk. Comparative regressions where nonindependent maternal weight was the explanatory variable yielded median coefficients of determination of 0.62 for fecundity and 0.76, or 0.14, for reproductive bulk among the data sets. When maternal weight was rendered independent by subtracting reproductive bulk, the median coefficient of determination for repro- ductive bulk plummeted from 0.76 to 0.12, which exposes the extent to which nonindependence inflates regression test statistics. Reproductive bulk nevertheless increased with both independent and nonindependent maternal weight. Nonindependence does not necessarily invalidate the practical use of traditional fecundity-maternal weight regression models. Reproductive bulk is potentially useful as a response variable in re- gression models of reproductive potential where trade- offs between fecundity and egg weight occur, and where independent maternal weight is otherwise desired as an explanatory variable. Additional key words: fecundity, egg weight, reproductive potential, reproductive effort, Malacosoma disstria, Dendrolimus pini, Chilo partellus, Choristoneura fumiferana 'Capital-breeding' refers to Lepidoptera that depend for reproduction entirely or chiefly on metabolic resources assembled during the larval rather than the adult stage (Boggs 1992, Miller 1996, Tammaru and Haukioja 1996). The great majority of Lepidoptera are capital breeders, as are the great majority of outbreak Lepidoptera (Miller 1996, 2005; Tammaru and Haukioja 1996). Capital breeders that assemble resources entirely during larval growth are termed ‘perfect’, and those that assemble most resources during larval growth but some during posteclosion adult feeding are termed 'imperfect' (Miller 2005). Because reproductive potential in capital breeders is solely or heavily dependent on larvae, initial maternal weight has long been considered an indicator of fecundity (Honek 1993, Miller 2005). Indeed, retrospective analysis of capital-breeder fecundity-maternal weight regressions confirms a consistent relation between fecundity and maternal weight (Miller 2005). Moreover, extrinsic factors such as temperature and diet quality during larval development influence fecundity indirectly through their influence on maternal weight. Although lineal maternal size variables, including forewing length and pupal diameter, are also predictive, weight has traditionally predominated. Trade-offs between fecundity and egg weight for a given maternal weight also can alter fecundity—maternal weight regressions. Trade-offs may vary temporally, spatially, and among subgroups of a cohort Trade-offs are probably adaptive, and two notable examples are the geographic trade-off clines in Choristoneura fumiferana (Clem.) (Tortricidae) and Malacosoma disstria (Hbn.) (Lasiocampidae) (Harvey 1983, Parry et al. 2001). Trade-offs highlight the potential utility of reproductive bulk as a response variable in regression models of reproductive potential (Miller 2005). Reproductive bulk, R, is the weight of the individual egg load, or some meaningful fraction of it such as laid eggs or realized fecundity (Miller 2005). R also is ae iathemmatienl product of fecundity, F, x mean eg weight, E. Somatic bulk, or nonR, is maternal w fe fe R, or W - R. The expressions nonR and W - R are interchangeable. Reproductive effort, RE, is the proportion of maternal weight consisting of egg load, or R/(W + R). These definitions are approximations in that they do not account for energy used in maintenance and in the development of reproductive structures (Roff 1992). Traditional fecundity-maternal weight regressions for capital breeders are usually formulated as F=bW+a (Eq. 1) where W is weight of first-day pupae or of adults at eclosion, b is the slope parameter, and a is the intercept or scaling parameter (Honek 1993, Miller 2005). Reproductive bulk—maternal weight regressions may be analogously formulated as R=bW+a (Eq. 2) W in Eqs. 1 and 2 almost always includes R such that W = nonR + R. Thus R is represented on both sides of these equations, thereby creating __ statistical nonindependence between response and explanatory variables (Roff 1992, Honek 1993, Miller 2005). Nonindependence masks data variability, inflates regression test statistics, and exaggerates the strength of relations. If measures of R are available, statistical ws ier) S independence can be attained with the formulation R=b(W-R)+a (Eq. 3) Probably because fecundity is so obviously important in reproduction, and has so often been regressed on nonindependent maternal weight, pure R in capital breeders has been little investigated. The objectives of this report are (i) to illuminate reproductive bulk, R, by examining its interrelations with nonindependent maternal weight, W +R, ornonR + R, independent maternal weight or somatic bulk, nonR or W - R, reproductive effort, RE, fecundity, F, mean egg weight, E, and (ii) to assess the impact of nonindependence between explanatory and response variables on the reliability of regression models for estimating reproductive potential. MATERIALS AND METHODS I sought capital-breeder data sets where R, nonR, F, and E were given or could reasonably be derived. Before analysis, all weights were standardized to fresh weights at adult eclosion expressed in milligrams. Dry weight in data sources was converted to fresh WwW eight using the factor 3.3 (Honek 1993), and pupal eadn weight to adult fresh weight using the factor 0.54, the reciprocal of 1.85 for the opposite conversion (Miller 2005). Data sets were accepted if they included more than eight mothers and did not require more than two conversions to obtain fresh weights of any variable. Some sample sizes are slightly less than in sources because individual observations were not complete enough for use in this study. I compared test statistics derived from each data set for regressions of F relative to W + R (Eq. 1), R relative to W + R (Eq.2), and R relative to W - R or nonR (Eq. 3). Most statistics were computed with SYSTAT (1992) software. Correlations refer to Pearson r-values, and coefficients of determination to 7. Mean REs were based on individual mother REs. Lack of statistical significance at the 0.05 level is indicated by 'ns'. More specific methodological information is given as the analysis of each data set is discussed below. always RESULTS Eight data sets, marginally to eminently suitable, were found for four capital breeders in three families. Three sets are considered original because they were used only partially in previous publications. Three or more comparative regressions are presented for each data set as well as mean RE and the correlation between F and E. Regression test statistics where W + R rather than W - R is the explanatory variable should be interpreted cautiously because of inherent nonindependence. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY No. 1. In this and the following two analyses, pupal weights, fecundities bulk — for Malacosoma disstria (Hbn.) (Lasiocampidae), a perfect capital breeder, were transcribed from enlarged photocopies of scatterplots in Fig. 2 of Parry et al. (2001). Regression slopes computed { from transcribed values were compared with slopes given in the source, and reproductiv e and F-tests showed that departures were small and nonsignificant. In each of these three data sets two conversions were necessary: fresh first-day female pupal weight to fresh maternal weight at eclosion, and dry reproductive bulk to fresh reproductive bulk. Louisiana results are presented first. Regressions (all n = as F = 0.98(W + R) + 23.4,7° = 0.81, P < 0.01. R = 0.84(W + R) - eS. r= 0.85, P < 0.01. R = 0.05(W - R) + 181.0, 7° = 0.00, ns. Mean RE = 0.76. r,,,. = 0.24, ns. No. 2. Michigan M. disstria. eee (all n = 29): = 0.77(W + R) + 37.4, ° = 0.77, P < 0.01. = 0.70(W + R) - 16.4, 7° = 0.86, P < 0.01. = 0.76(W - R) + 85.7, 7° = 0.17, P = 0.03. jie RE = 0.63. r |... = 0.29, ns. No. 3. Manitoba M. disstria. Regressions (all n = 24): F = 0.76(W + R) - 30.9, 7° = 0.89, P < 0.01. = 0.64(W + R) - 38.8, °° = 0.90, P < 0.01 R = 1.04(W - R) - 2.06, 7° = 0.40, P < 0.01. Mean RE = 0.50. r ,, , = 0.28, ns. No. 4. The Minnesota M. disstria data analyzed below originated in simultaneous laboratory rearings of three subgroups of larvae from a cohort of eggs collected near International Falls on Populus tremuloides Michx. (Salicaceae). The subgroups were reared at constant temperatures of 20°, 25°, and 30°C on foliage of the source foodplant, and the data used here are subsamples. Experimental procedure is detailed further in Miller (2005). TWwo conversions were necessary: fresh first-day pupal weight to fresh maternal weight at eclosion, and dry egg weight to fresh egg weight. Regressions (all n = 24): F = 0.71(W + R) + 6.4, ° = 0.63, P < 0.01. R = 0.56(W + R) - 23.7, 7 = 0.68, P < 0.01. R = 0.27(W - R) + 60.2, 7° = 0.05, ns. Mean RE = 0.44. r REX 0.45, P < 0.05. No. 5. In the following analysis of data for Dendrolimus pini (L.) (Lasiocampidae), a perfect VOLUME 59, NUMBER 4 capital breeder, Eckstein (1911) may have provided the earliest published evidence for relations among the variables of capital-breeder reproductive potential. These are the only data here in which individual maternal weight was reported both before oviposition (gravid condition) and after eggs were laid or removed from ovaries (spent condition). Neither transcription nor conversions were necessary, making this the most experimentally satisfactory data. In the third ce below, W - R was computed indirectly as Wee - (W - We a) and in the fourth, directly as W, spent” gravid Regressions (n=55-5S) F = 0.10(W + R) + 33.8, °° = 0.60, P < 0.01. R = 0.82(W + R) - 220.1, 7° = 0.93, P < 0.01. R = 1.20(W - R) + 491.6, 7° = 0.16, P = 0.01. R= 129(W__) + 477.7, 2 = 0.17, P= 0.01. Mean RE = 0.67. RE = 0. 000093(W + R) + 0.525, r 0:59, P< 0.01. r,, = -0.12, ns. The third and fade regressions above differ only negligibly. No. 6. The source data treated below are from Berger (1989) who reared Chilo partellus (Swinhoe) (Crambidae), an imperfect capital breeder, on artificial diet. After adults eclosed and were weighed and individually mated, Berger recorded numbers and weights of eggs laid on the first three nights of oviposition during which 80% of eggs typically are deposited. It appears that mothers were provided with water for imbibing. The data were transcribed from enlarged photocopies of Berger's Figs. 3 and 4. No conversions were necessary. Regressions (all n = 9): F = 1.80(W + R) + 247.7, °° = 0.20, ns. R = 0.19(W + R) + 6.33, 7° = 0.55, P < 0.01. R = 0.18(W - R) + 12.1, ° = 0.35, ns. Mean RE = 0.25. r,, = -0.10, ns. No. 7. In the two following analyses, data sets for Choristoneura fumiferana (Clem.) (Tortricidae), an imperfect capital breeder, originated with pupae collected near Duluth, Minnesota, from severely infested Abies balsamea (L.) Mill. and Picea glauca (Moench) Voss. (both Pinaceae). Eclosing aaults were divided into two groups, individually paired, and the group discussed first provided with sponges soaked in sweetened water for imbibing. Numbers of eggs laid by mothers of both groups were recorded daily. Eggs were weighed at least twice, the first time on the! iest or neal day of oviposition, and the second and sometimes third on the fourth day of oviposition or later. Forewing length of mothers was measured after they bo Go expired. Experimental procedure is detailed further in Miller (1987). Two conversions, the first involving two steps, were necessary in both sets: forewing length, FL, to dry weight, DW, by the species-specific formula DW = 0). 0057[F L333] (7 = 0.97, df = 211, P < 0.01), from dry to fresh weights. Regressions (all n = 33): F = 2.47(W + R) + 91.9, 7° = 0.31, P < 0.01. R = 0.52(W + R) + 9.7, 7° = 0.34, P < 0.01. R = -0.15(W - R) + 45.9, 7° = 0.09, ns. Mean RE = 0.68. r ,. , = -0.02, ns. and No. 8. These C. fumiferana results are from the second group mentioned above, which was provided with unsweetened water for imbibing. Regressions (all n = 26): F = 0.83(W + R) + 97.8, 7° = 0.06, ns. R = 0.21(W + R) + 12.4, 7° = 0.08, ns. R = -0.18(W - R) + 30.6, 7° = 0.09, ns. Mean RE = 0.44. r REZ 0.11, ns. DISCUSSION Comparison of regressions where fecundity, F, and reproductive bulk, R, were response variables to nonindependent W, that is to W + R or nonR + R (Eqs. 1 and 2), shows coefficients of determination to be higher for R than for F among the eight data sets. This result reveals less variability in R than in F relative to the explanatory variable. Median coefficients of determination were 0.76 for R and 0.62, or 0.14 less, for F, and their intraspecific differences ranged 0.01—0.35, with a median difference of 0.08. This result was expected because R integrates fecundity, F, and egg weight, E, and, unlike F alone, takes account of trade- Offelbenweentl ancliey By contrast, when independent maternal weight, W - R or nonR, was the explanatory variable (Eq. 3), the median coefficient of determination for reproductive bulk among the data sets was only 0.12. This plummeting of the coefficient reflects the extent to which nonindependence artificially elevates _ test statistics. The fact that coefficients of determination for Eqs. 1 and 2 are inflated is not surprising; what is surprising is the extent of inflation. Even if barely satisfactory data sets 7 and 8 for Choristonewra fumiferana are omitted, the median coefficient of determination for reproductive bulk does not change appreciably (nor does that for fecundity). It is also telling that the corresponding coefficient of determination in an eminently satisfactory data set— No. 5 for Dendrolimus pini—dropped from 0.93 to 0.16, and thus reflects the overall trend among the eight data sets. Low coefficients of determination where bo ioe) bo response and explanatory variables were independent— all < 0.40—expose _ the variability in nonindependent data. Clearly, nonindependence in traditional regressions using W + R as the explanatory true variable has led to gross overstatements of the strength of maternal weight in models of reproductive potential. Slopes in regressions of reproductive bulk relative to independent maternal weight or somatic bulk ranged 0.05-1.2 and were positive if not significant in six of the eight data sets. Positive slopes confirm Honek's (1993) reasoning that larger-bodied mothers are necessary— probably sufficient—for greater R as well as greater F. In the two exceptions, Nos. 7 and 8 for Choristoneura fumiferana, negative slopes may be anomalous as suggested by some but not all previous work (Miller 2005), and may be due to conversion errors, although variability in the life system of this species should not be ruled out. Other data sets with slopes not statistically different from zero—Nos. 1, 4, and 6—likewise may be due to transcription and conversion error. As indirectly measured variables become smaller, errors of conversion are magnified. The slope values show that reproductive bulk changes disproportionately relative to independent and nonindependent maternal weight, just as found for fecundity (Miller 2005). Disproportionate change between these variables may have implications for rates of population buildup and decline. As already mentioned, trade-offs between fecundity, EF, and mean egg weight, E, may occur temporally or spatially, and are doubtless adaptive. An example of statistically significant increase in E accompanying increasing F is data set No. 4 for Malacosoma disstria. An example of decrease is evident in Bupalus piniaria (L.) (Geometridae), where egg diameter was a surrogate for E (Klomp 1966) (data omitted here). Klomp noted the counterintuitive surrogate egg weight of individual mothers increased serially, it still decreased overall with increasing fecundity. An example of constant E with increasing F is data set No. 5 for Dendrolimus pini. A remarkable further example is Bombyx mori (Bombycidae) (Fig. 1), illustrated here because it may represent the most situation where even though extensive data in existence concerning egg weight relative to fecundity in a capital breeder. Unfortunately, the corresponding maternal weights were unavailable. Reproductive effort, RE, ranged from 0.20s—0.70s among the eight data sets. RE is expected to vary adaptively in different species and environments (Roff 1992). It appeared static in all but one data set-—No. 5 for Dendrolimus pini—but RE is derived and its test statistics relative to other reproductive variables are also subject to inflation. In Choristoneura fumiferana data sets 7 and 8, mothers provided with sweetened water JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Mean weight of 1 egg (mg) (E) 0.65 0.45 400 500 600 Fecundity (F) 700 FIGURE 1. Relation of mean egg weight, E, to fecundity, F, in 293 races of Bombyx mori. Each point represents one or more females per race. The regression line is described by E = 0.0000109 + 0.56F, r = 0.00, ns. Data were plotted from tables in Hiratsuka (1999). had an RE of 0.68 compared with only 0.44 for those provided with unsweetened water; the 0.24 difference further demonstrates the enhancing role of adult nutrition in reproductive performance (Miller 1987, 1989). Lack of independence between response and traditional explanatory variables does not necessarily invalidate practical use of fecundity-maternal weight regression models for estimating reproductive potential (Roff 1992). Maternal weight, whether independent or not, remains indicative of fecundity and reproductive bulk. In most basic research, however, response and explanatory variables should be statistically independent for reliable results. Further, all weights ideally should be measured directly and in dry form. The use of dry weights would warrant adoption of the stricter terminology of 'reproductive mass! and 'maternal mass'. ACKNOWLEDGEMENTS I thank J. M. Muggli, K. V. Bui, and J. G. Yang for laboratory assistance, R. D. Moon, D. A. Andow and an unnamed reader for highly useful manuscript reviews, and J. L. Moe for computer and software support. LITERATURE CITED Boccs, C. L. 1992. Resource allocation: exploring connections be- tween foraging and life history. Funct. Ecol. 6: 508-518. ECKSTEIN, K. 1911. Beitrage zur Kenntnis des Kiefernspinners Lasio- campa (Gastropacha, Dendi rolimus) pini L. Zool. Jahrb. Abt. Syst. Geogr. Biol. Tiere 31: 59-164. Harvey, G. T. 1983. A geographic cline in egg weights in Choris- toneura fumiferana (Lepidoptera: Tortricidae) and its significance in population dynamics. Can. Entomol. 115: 1103-1108. VOLUME 59, NUMBER 4 Hiratsuka, E. 1999. Silkworm breeding. Balkema, Rotterdam. 500 pp. [Translation of Nihon Sanhinshu Jitsuyokeifu, Tokyo, 1969] Honek, A. 1993. Intraspecific variation in body size and fecundity in insects: a general relationship. Oikos 66: 483-492. Krome, H. 1966. The dynamics of a field population of the pine looper, Bupalus piniarius L. (Lep., Geom.). Adv. Ecol. Res. 3: 207-305. MILLER, W. E. 1987. Spruce budworm (Lepidoptera: Tortricidae): role of adult imbibing in reproduction. Environ. Entomol. 16: 1291-1295. MILLER, W. E. 1989. Reproductive enhancement by adult feeding: ef- fects of honeydew in imbibed water on spruce budworm. J. Lepid. Soc. 43: 167-177. MILLER, W. E. 1996. Population behavior and adult feeding capability in Lepidoptera. Environ. Entomol. 15: 213-226. MILLER, W. E. 2005. Extrinsic effects on fecundity-maternal weight relations in capital-breeding Lepidoptera. J. Lepid. Soc. 59:.143-160, Parry, D., R. A. Goyer and G. J. Lenhard. 2001. Macrogeographic clines in fecundity, reproductive alloction, and offspring size of the forest tent caterpillar Malacosoma disstria. Ecol. Entomol. 26: 281-291. Rorr, D. A. 1992. The evolution of life histories. Chapman and Hall. New York. 535 pp. SYSTAT: statistics, version 5.2 ed. 1992. SYSTAT Inc., Evanston, IIli- nois. TamMaru, T. & E. Hauxroja. 1996. Capital breeders and income breeders among Lepidoptera-consequences to population dy- namics. Oikos 77: 561-564. Received for publication 14 February 2005, revised and accepted 21 September 2005 GENERAL NOTES Journal of the Lepidopterists’ Society 59(4), 2005, 234 PIGGYBACKING NORTHWARD: MOVEMENT OF LEPTOTES CASSIUS (LYCAENIDAE: LYCAENINAE) THOUGHOUT THE SOUTHEAST Additional key words: butterfly gardening, dispersal, Florida, Georgia, Plumbago, South Carolina The widespread distribution of Leptotes cassius Cramer across the many island groups of the West Indies demonstrates its success as a highly effective colonist (Smith et al. 1994). Within the Southeast, L. cassius is primarily restricted to south Florida and the Florida Keys where it is generally the most abundant blue of open disturbed sites or semi-open, scrubby habitats. may jocastoncilly expand its range northward through During favorable years with mild winters, it the peninsula into the central and northeastern counties. Records beyond the state's border are rare but isolated vagrants have been reported from Alabama, Mississippi, Ar kansas, Louisiana, Georgia and South Carolina (Opler et al. 1995, Opler and Malikul 1998, Scott 1986,). The exact origin of such strays is unknown and may represent isolated dispersal events, hurricane or tropical storm-assisted movements, or temporary A fourth alternate seasonal expansion of breeding populations. explanation, though, may provide an explanation for many of the confirmed records. In August 1999, while shopping at a small specialty nursery on Hilton Head Island, Beaufort County, South Carolina, I noticed a small blue flying erratically around a patch of colorful flowers. Upon closer inspection, I was surprised to see that it was a male L. cassius. Immediately following the discovery, I searched the remainder of the nursery property and adjacent open areas, but did not observe any additional adults. The nursery did, however, have several one to three gallon planters of Plumbago auriculata Lam. (Plumbaginaceae) and P. awriculata ‘alba' that between them supported 21 larvae ranging from second to fifth intar. I inquired about the origin ‘of the plants and was informed by the nursery staff that they had arrived earlier in the week from a lar ge grower located in south- Sealy lorida. tween September 1999 to August 2002, subsequent visits were made to the same Hilton Head nursery and to two other similar specialty garden centers located in Bluffton, Beaufort County, South Chatham County, Georgia. During this period, adults, immatures, or in some cases small breeding colonies of L. cassius were found at each nursery in every year except 2002, when adult individuals were found only at the Hilton Head Island location. In each instance, all three nurseries received shipments of Plumbago from the same south-central Florida grower. It is not known if purchased plants livestock led to the establishment of additional local temporary breeding colonies. With the ever-growing popularity of butterfly gardening and native landscaping, it is likely that the frequency of such human-assisted piggybacking of livestock on nursery plant material will continue to increase and thus serve to blur the legitimacy of some county or state records. Carolina and Savannah, harboring LITERATURE CITED Oper, P. A., H. PAVULAAN & R. E. STANFORD (coordinators). 1995. Butterflies of North America. Jamestown, ND: Northern Prairie Wildlife Research Center Home Page. http:/Avww.npwre.usgs.gov/resource/distr/lepid/bflyusa/bflyusa.ht m (Version 30DEC2002). Oper, P. A. & V. MALIKUL. 1998. A field guide to eastern butterflies. Houghton-Mifflin Co., Boston. 486pp. Scort, J. A. 1986. The butterflies of North America. Stanford Univer- sity Press, Stanford, Calif. 583pp. SmitH, D. S., L. D. MILLER, & J. Y. MILLER. 1994. The butterflies of the West Indies and South Florida. Oxford Univsity Press. Ox- ford. 264 pp. JARET C. DANIELS, McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, P.O. Box 112710, University of Florida, Gainesville, Florida 32611-2710. Email: jdaniels@flmnh.ufl.edu. Received for publication 5 December 2004; revised and accepted 13 July 2005 VOLUME 59, NUMBER 4 Journal of the Lepidopterists’ Society 59(4), 2005, 235-236 bo (es) Ol SYNANTHEDON BOLTERI (SESIIDAE) IN MICHIGAN Additional key words: pheromones, Baskerville-Emin, Synanthedon sigmoidea The clearwing moth, Synanthedon bolteri (Hy, Edwards), has a wide-ranging distribution in mostly sub-arctic to arctic areas of North America. This statement is based on scattered records, "... from southern Quebec and Rhode Island west to Colorado and Washington and north to the Northwest Territories and Alaska (Eichlin and Duckworth 1988, p. 86)" Large numbers of sesiids have been collected since the advent of synthetic sex pheromone use for monitoring clearwing moths (USDA 1979). However, to date, no collectors have reported capturing this species with the use of synthetic sex pheromones, even though these chemicals are commonly used for monitoring clearwing moths across the United States for integrated pest management purposes. Historically, $. bolteri specimens collected in Michigan were rare. Two specimens of S. bolteri collected by R.R. Driesbach in southwestern Michigan (Mecosta Co., 31 July, 1941 and Ottawa Co., 17 Aug., 1945) are located in the Michigan State University A. J. Cook Arthropod Research Collection and University of Michigan Museum of Zoology, respectively. In over 30 years of collecting clearwing moths I have encountered only 1 specimen of S. bolteri (Figure 1). The male moth was caught on 23 July 1984 in Dickinson County of Michigan's Upper Peninsula in a malaise trap located within a flowering stand of fireweed (Epilobium angustifolium). Another clearwing moth, Albuna pyramidalis (Walker), was commonly found sitting on fireweed foliage during this same time period. This situation changed in August 2004, when 30 specimens of S. bolteri were collected at two locations over a 5-day period with the use of a Multipher 1® moth trap baited with a grape root borer pheromone lure made by Suterra. This lure composition is 99% E,Z 2.13 -ODDA and 1% Z,.Z 3,13 -ODDA. This lure was chosen to capture Synanthedon sigmoidea (Beut.), which is reportedly attracted to this lure formulation (Taft et al. 1991). The trap location was selected due to the occurrence of large stands of pussy willow (Salix discolor) and several other Salix species (Figure 2). Engelhardt (1946) reported that both of these moth species were associated with low growing Salix species that have been injured by coleopteran larvae. Fourteen specimens of S. bolteri were captured over a 3-day period (1-3 August 2004) in a willow thicket along the Rock River at the US-2 stream crossing (T43N, RIIW, Section 24) in Newton Township, Mackinac County. Sixteen additional specimens were collected over a 3-day period (3-5 August 2004) approximately 12 miles southeast of Sault Ste. Marie (T46N, RIE, Section 34) in Bruce Township, Chippewa County. The habitat at both collection sites was similar. The cumulative degree-day value (Base 10°C) using the Baskerville-Emin method (Baskerville and Emin 1969) was calculated with the assistance of staff from the agricultural weather office at Michigan State University (MSU) for Manistique and Barbeau, Michigan. These were the closest Upper Peninsula weather stations to the capture locations. The cumulative degree-day calculated values for Manistique and Barbeau were 394 and 385, respectively. S. sigmoidea was later collected in southern Ingham County starting on 26 August 2004, when the cumulative degree-day calculated value (Base 10° C) for East Lansing was 1061. This important pheromone and degree-day information should help collectors target S. bolteri and S. sigmoidea in the future. I would like to thank Jeff Andresen at the MSU agricultural weather office for providing the temperature data used to calcu- late the degree-day accumulation information. Also, many thanks to Stewart Gage of MSU for calculating the actual de- gree-day numbers and to Thomas D. Eichlin, Sarah Wolf and Mogens C. Nielsen for helpful suggestions and reviewing this paper. LITERATURE CITED BASKERVILLE, G.L, & P. EMIN, 1969. Rapid estimation of heat accu- mulation from maximum and minimum temperatures. Ecology 50: 514-517. EICHLIN,T. D., & W.D.DuckwortH, 1988. The Moths of America North of Mexico. Fasc. 5.1: Sesioidea, Sesiidae. Wedge Entomol. Res.Foundation, Washington, DC pp.85-86. ENGELHARDT, G.P., 1946. The North American Clear-wing Moths of the Family Aegeriidae. US Nat. Mus. Bull. 190: 84-85. Tart, W. H., D. SmITLey, & J.W. SNow, 1991. A guide to the Clear- wing Borers (Sesiidae) of the North Central United States. North Central Regional Publication. No.394. pp. 30. USDA, 1979. Pheromones of the Sesiidae (formerly Aegeriidae). Sci. and Edu. Admin. ARR-NE-6 December 1979. pp. 81. WILLIAM H. Tart, 1430 Locher Road, Dewitt. Michigan 48820 E-mail: taftw@michigan.gov Received for publication 3 January 2005; revised and accepted 21 October 2005 236 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Ficure 1. S. bolteri collected southeast of Ralph on 23 July 1984, Dickinson County, Michigan. (Photo Credit: P. Carrington) River, Mackinac County, Michigan. (Photo Credit: Wim. Taft) VOLUME 59, NUMBER 4 Soliciting Color Photos for the Cover of the Journal We are soliciting submissions of high quality color photographs for publication on the cover. Submissions of electronic files on CD are preferred, but we will accept high quality printed photographs suitable for scanning. Electronic submissions should be in °.tif or *.psd format, with dimensions of AT LEAST 1640 pixels by 2030 pixels at 300 pixels per inch (120 pixels per cm). Submissions should be mailed to: Dr. Michael E. Toliver Divsion of Science and Mathematics Eureka College, 300 E. College Eureka, IL 61530 Photographers whose work is used will be acknowledged on the inside front cover. Journal of the Lepidopterists’ Society 59(4), 2005, 238-240 Abbot, John, 172-173 Abies balsamea, 152, 157, 231 Acentria ephemerella, 79-80 Aceraceae, 72, 73 Achlyodes, 33, 52 Achroia grisella, 157 Acleris, 72 Acrobasis betulella, 78 Acrolepiidae, 67 Acrolepiopsis, 67 Acronictinae, 118 Actinote, 88, 134-142 zikani, 134-142 Aechmea, 37-38 Aglais, 40-43 Agonoxeninae, 68 Alberta, 61-82 Allochronic (temporal) isolation, 161-165 Alnus, 71, 73, 74 Amelanchier alnifolia, 73 Anacampsis, 69 Anacardiaceae, 37 Ananas comosus, 37 Ancyloxypha numitor, 111 Annonaceae, 115 Anoplolepis gracilipes, 220, 221 Anrenholz, David H., 178-180 Antheraea assama, 156 Anthoxanthum odoratum, 164 Anweiler, Gary, 61-82 Apanteles, 192 Apiosporina morbosa, 70, 71 Apodemia, 85, 92, 161 Apotomis paludicolana, 74 Araliaceae, 107 Arceuthobium, 78 Arctotheca calendula, 155 Aremfoxia, 35 Argyrotaenia, 73 Asclepiadaceae, 111, 118 Asteraceae (“Compositae”), 7, 10, 35, 70, 107, 128, 134, 135, 137, 155, 162, 168, 225 Astragalus, 116, 175-176 Astraptes, 19, 23, 50, 52, 56 Aulonemia, 206 Austin, George, 89-95 Baccharis neglecta, 107 Balanocarpus heimii, 112, 115 Barreto De-Andrade, Rafael, 223-228 Batesian mimicry, 44 Battus philenor, 118 Beilschmiedia, 19-34 Belize, 83-88 Betulaceae, 62, 71, 73, 75, 78 Billbergia, 37 Biodiversity, 89-95, 96, 170, 200-211, 212-218, 219 Biogeography, 61-82, 83-88, 89-95, 110-112, 200- 211, 212-218 Bird, Charles, 61-82 Blastodacna, 68 Boherhavia, 107 Bombyx mori, 152, 157, 232 Borreria verticillata, 137 Bothrops, 39 Brachymeria, 118-119 INDEX FOR VOLUME 59 (New names in boldface) Brassica 10-12 Brassicaceae, 10, 13, 170 Brassolinae, 37-39, 88, 97-101 Brazil, 35-37, 96-106, 134-142, 178, 200, 206, 212- 218, 223-225 Brodiaea elegans, 168 Bromelia, 37-39 Bromeliaceae, 37-39 Brower, Andrew V. Z., 44 Brown, Keith S., 35-37, 59, 96-106, 134-142, 178, 227 Bucculatrix pyrivorella, 143 Bupalus piniaria, 143, 154, 157, 158, 232 Burns, John M., 19-34, 110, 112 Cadra cautella, 155 Caesalpinia bonduc, 174 Callionima grisescens elegans, 212, 214-217 Callophrys, 181-199 affinis affinis, 181-199 affinis albipalpus, 188-189, 191, 193, 194, 199 affinis apama, 181-199 affinis chapmani, 184-186, 158-189, 192-194, 199 affinis homoperplexa, 182-199 affinis washingtonia, 183-187 dumetorum, 181-183 perplexa, 181-199 sheridanii comstocki, 181 sheridanii lemberti, 181 sheridanii sheridanii, 181, 183 viridis, 181-183 Camponotus, 174, 219-222 Canada, 1, 61-82, 186, 196 Capital-breeding, 143-160, 229-233 Caprifoliaceae, 66, 71 Carex, 110-112 Carya glabra, 154 Caryocolum, 70 Caryophyllaceae, 70 Casinaria, 33 Catoptria maculalis, 79 Ceanothus, 187, 189-194, 197, 198 Celastraceae, 115 Cephise, 19, 54, 57 Charaxinae, 87, 97, 98, 100-103, 105 Chilo partellus, 229, 231 Chlorohystricia, 33 Chlosyne, 92, 172-173 gorgone, 172-173 Choristoneura fumiferana, 71, 145, 152, 157, 158, 229, 231, 232 Chromolaena odorata, 225 Chrysothamnus, 107, 128, 162 Chusquea, 201, 206, 208 Cinnamomum, 156 circadian rhythm, 16 Cirsium, 10 Cnephasia jactatana, 144, 155 Cocytius antaeus, 212, 214-216 Coleophora rosaefoliella, 69 Coleophoridae, 69 Coleophorinae, 69 Coleotechnites laricis, 69 Colias, 13, 92 Comandra umbellate, 77 Conservation, 89-95, 96-106, 134-142 Conyza canadensis, 107, 109 Cooper, David, 110-112 Corades, 204, 211 Corcyra cephalonica, 155 Coronilla varia, 175 Costa Rica, 19-34, 40-41, 100, 180, 200, 204, 217 Covell, Jr., Charles V. 178-180 Crambidae. 79-80 Crataeagus, 68, 73 Crematogaster, 112-115, 220 Creonpyge creon, 33 Crocidosema plebejana, 154, 157 Cucullia, 107-109 dorsalis, 107-109 speyeri, 107-109 Cunoniaceae, 22-23 Cyclargus thomasi bethunebakeri, 174 Cyperaceae, 110 Daedalma, 204, 211 Dahlica triquetrella, 65 Dana, Robert P., 175-177 Danainae, 44, 118-119 Danaus, 1, 17, 92, 94, 118, 143 gilippus, 92, 94, 118 plexippus, 1, 17, 92, 118, 143 Daniels, Jaret C., 234 Danthonia californica, 166, 168 Dasypyga alternosquamella, 78 Davis, Andew K., 1-5 Dendrolimus, 156, 230-232 Depressaria atrostrigella, 68 Depressariinae, 67-68 Diatraea saccharalis, 155, 156 Dichomeris, 70 Dipterocarpaceae, 112, 115 Disporum trachycarpum, 67 Ditrysia, 143 DNA, 19, 23, 111, 112, 161, Drucina, 204, 211 Duarte Jiinior, José Araujo, 212-218 Dynastor darius darius, 37-39 Dyscophellus, 23, 55 Eastwood, Rod, 219-222 Ecuador, 6, 37, 178, 180, 201, 205-207, 217 Elachista maritimella, 68 Elachistidae, 67-68 Elachistinae, 68 Endangered ecosystems, 96-106 species, 116-117, 134-142, 175-177 Ennomos subsignarius, 152, 154 Epermenia, 75-77 Epermeniidae, 75-77 Epinotia albicapitana, 75 Epiphyas postvittana, 144, 155, 157 Epityches, 35, 37 Eretris, 200-211 agata, 200-211 calisto, 201, 205, 206 depresissima, 207 encycla, 201, 203, 205, 206 oculata, 201, 205, 206 Ericaceae, 73, 75 Erigeron asperugineus, 107 Erinnyis, 212, 214-216 alope alope, 212, 215, 216 ello ello, 214-216 Eriocrania semipurpurella pacifica, 62 VOLUME 59, NUMBER 4 Eriocraniidae, 62 Eriogonum, 161, 164, 187-196, 199 alatum, 188, 190-192, 194, 196, 197 atrorubens, 189-194, 199 flavum, 187, 190-192 heracleoides, 190-192, 195 racemosum, 190-192, 196, 197 umbellatum, 190-193, 196 Erynnis baptisiae, 111, 177 Eucalyptus, 10, 97, 140 Eucosma, 121-133 emaciatana, 121-133 larana, 121-126, 133 nordini, 121-133 piperata, 121-133 taosana, 121-133 totana, 121-133 Eueides vibilia unifasciatus, 42 Eumorpha, fasciatus, 212, 215, 216 vitis vitis, 212, 215, 216 Euodia meliaefolia, Eupatorium gaudichaudianum, 137 Euphilotes, 92, 161 Euphydryas, 40-43 Euproctis chrysorrhoea, 154 Fabaceae (“Leguminosae”), 23, 73, 115, 174, 175- 177, 219, 221 Fecundity, 42, 143-160, 229-233 Festuca idahoensis, 166 Fissistigma wallichii, 115 Fleishman, Erica, 89-95 Forister, Matthew L., 161-165 Formica, 176 Francillon, John, 172-173 Francini, Ronaldo Bastos, 134-142 Fratello, Steven, 200-211 Fraxinus, 73 Freitas, André, 35-37, 59, 96-106, 134-142, 223-228 Frugivory, 96-106 Gabbro soils, 163 Galleria mellonella, 157 Garsauritis, 35, 37 Gelechiidae, 69-70 Geum triflorum, 64 Gilbert, N., 10-18 Giles, Valerie, 83-88 Glaucopsyche lygdamus palosverdesensis, 116-117 Glyphidocera, 69 Glyphidoceridae, 69 Glyphipterigidae, 67 Glyphipterix montisella, 67 Gonzalez, Jorge, 37-39 Gorelick, Glenn A., 181-199 Gracillariidae, 65-66 Greeney, Harold F., 6-9, 201 Gretchena semialba, 74 Guadua, 201 Habitat fragmentation, 16, 89-95, 96-106 Haloragaceae, 80 Hansen, Dean, 175-177 Hedera, 107 Helianthus, 107 Heliconiinae, 44, 88 (as Acreiinae), 134-142, 223- 228 Heliconius, 40-43, 51, 96-106, 141, 223-228 ethilla narcaea, 223-228 erato phyllis, 223-228 hewitsoni, 40-43 Hesperia, 16, 45-58, 92, 161-165, 166-169 Hesperia (cont.) colorado harpalus, 161-165 colorado idaho, 161, 162 colorado mattoonorum, 164-165 colorado oregonia, 164 colorado tildeni, 161-165 comma, 16, 92, 161-165 “comma” yosemite, 161-165 lindseyi eldorado, 166 lindseyi macneilli, 166-167 lindseyi mccorklei, 166-169 lindseyi septentrionalis, 166-168 Hesperiidae, 6, 16, 19, 45-56, 97, 110, 161, 166, 179-180 Hofmannophila pseudospretella, 156, 158 Holodiscus discolor, 62 Howard, Elizabeth, 1-5 Human-assisted dispersal, 234 Hyalyris, 35, 37 Hyles euphorbiarum, 212, 215, 216 Hyphantria cunea, 156 Hypothyris, 35, 37 Hypsopygia costalis, 77-78 Ichneumonidae, 33 Income-breeding, 143 India, 170-171, 219 Isognathus australis, 212, 214-216 Ithomia, 37 Ithomiinae, 35-37, 97 Jalmenus evagoras, 221 Jamides celeno, 219-222 Janzen, Daniel H., 19-34 Jemadia pseudognetus, 23 Jonaspyge, 23, 57 Journey North, 1-5 Juncaceae, 67 Juncus, 67, 90 Junea, 210, 204, 211 Keckiella antirrhinoides, 41 Kitching, R. L., 219-222 Kovarik, Peter, 83-88 Kurrimia paniculata, 115 Labus, Paul, 83-88 Lampronia, 62-64 Landry, Jean-Frangois, 61-82 Lane, Cynthia, 175-177 Lantana camara, 225 Larix, 69, 78 Lasiocampidae, 118, 144, 229, 230 Lasius, 115 Lauraceae, 19-34, 156 Lepidium ruderale, 170 Leptotes cassius, 234 Lespesia archippivora, 118 Leucanthemum vulgare, 168 Leucoptera spartifoliella, 143 Licaria, 19-34 Lichen, 65 Life history, 6-9, 19-34, 37-39, 42, 54, 66, 69, 73, 110-112, 112-115, 166-169, 170-171, 181-199, 219-222 Liliaceae, 64, 67, 168 Lilium washingtonianum, 67 Limenitidinae, 87, 97 Lipographis, 79 Litsaea, 156 Lobesia botrana, 155, 157, 158 Lomatium, 77 Longcore, Travis, 116-117 Lonicera, 66 239 Loranthaceae, 78 Lotus scoparius, 116 Lozotaenia hesperia, 73-74 Lupinus, 175-176 Lycaeides melissa melissa, 175-177 Lycaenidae, 49, 55, 85-87, 97, 112-115, 118, 174- 175, 178-180, 181-199, 219-222, 234 Lycaeninae, 86-87, 234 Lycianthes, 35 Lymantria, 143, 145, 151, 154, 156 Machaeranthera canescens, 107-109 Machilus bombycina, 156 Malacosoma, 71, 118, 144, 145, 151, 158, 229, 230, 232 Malus, 73, 155 Manduca brasilensis, 212, 215, 216 Mangifera indica, 37 Mango, 37-38 Manh, Hui Buu, 219-222 Mark-recapture, 10-18, 96-106, 134-142, Mattoni, Adriano, 116-117 Mattoni, Rudy, 116-117 Medicago sativa, 175 Meerman, Jan, 53-88 Melastomataceae, 135, 137, 208 Melilotus, 175, 188 Melitaea ismeria, 172-173 Microlepidoptera, 61-82 Micrurapteryx salicifoliella, 65-66 Migration, 1-5, 10, 11, 13-17, 117, 213, 227 Mikania, 134, 135, 137, 140, 141 obsoleta, 134, 137, 141 Miller, William E., 143-160, 229-233 Mimicry, 42, 44, mitochondrial DNA, 23, 161 Mitoura, 92, 161 Mitracarpus hirtus, 137 Mojave desert, 89-95 Monarch butterfly, 1 Miillerian mimicry, 42 Munroessa icciusalis, 80. Murphy, Dennis, 89-95 Mygona, 204, 209, 211 Myrcinia, 137 Myriophyllum spicatum, 80 Myrmecophily, 112-115, 174-175, 176, 219-222 Nakamura, Ichiro, 110-112 Napeogenes sulphurina, 35-37 Nectandra, 19-34 Nemapogon acapnopennella, 64-65 Neogene dynaeus, 212, 214-217 Neomyrina nivea periculosa, 112-115 Nephila clavipes, 138 Noctuana haematospila, 6-9 Noctuidae, 45, 107-109, 118 Nymphalidae, 1, 16-18, 35-43, 83, 85, 87-SS, 96- 103, 118-119, 134-142, 143, 172-173, 180, 200- 211 Obituaries, 45-58 Freeman, Hugh Avery, 45-58 Nicolay, Stan (Colonel), 178-180 Ochromolopis ramapoella, 77 Ocotea, 19-34 Ogyris, 220 Olethreutinae, 74-75, 121-133 Olson, Carl, 118-119 Opuntia, 68 Ortgiesia, 37 Oryza sativa, 156 Oviposition, 1, 3, 4, 35, 40-43, 116, 137-138, 140, Oviposition (cont.) 170-171, 174, 181, 183, 188, 190, 192, 229-233 Oxeoschistus, 200-211 duplex, 206 puerta, 203, 204, 206 romeo, 200-211 simplex, 204, 206, 207 Oxytropis lambertii, 175 Palos Verdes blue, 116-117 Pantepui, 200-211 Paranthrene robiniae, 70-71 Parapedaliodes parepa, 201 Parasitoid, 19, 20, 33, 42, 43, 115, 118-119 Passiflora, 40-43, 226 Patelloa xanthura, 33 Pedaliodes, 200, 202-205, 207, 211 chaconi, 200, 205 demarmelsi, 200, 205 phrasiclea, 207 pisonia, 205, 206 roraimae, 200, 202, 203, 207 terramaris, 200, 205 yutajeana, 200, 205 Pediasia abnaki, 79 Pelochrista, 121, 130-132 emaciatana, 121-133 (new combination) perpropinqua, 121-133 (new synonym) popana, 121-133 powelli, 121-133 Persea, 19-34 Petit, J. C., 107-109 Petit, M. C. 107-109 Phaeostrymon alcestis, 118 Phaneta lapidana, 74 Pheidole, 138 Phenology, 155, 161-165, 212-218 Philosamia ricini, 144 Phocides, 33, 52 Phragmites, 110-112 Physcopedaliodes physcoa, 206 Picea glauca, 73, 152, 231 Pieridae, 10-18, 85, 97, 118, 170-171 Pieris, 10-18, 42, 92 Pinaceae, 69, 73, 74, 152, 156, 166, 231 Pineapple, 37 Pinus, 73, 78, 156, 168 Plantago lanceolata, 155 Platytes vobisne, 79 Plumbago auriculata, 234 Plutella vanella, 66-67 Plutellidae, 66-67 Poanes viator, 110-112 Pohl, Gregory, 61-82 Polyrhachis (Myrmhopla) rufipes, 220 Pontederia cordata, 110 Pontia, 92, 94, 118, 179-171 daplidice moorei, 170-171 Populus, 71, 73, 75, 151, 230 Potamogeton, 80 Potamogetonaceae, 80 Praepedaliodes phanias, 206 Proclossiana eunomia, 16 Prodoxidae, 62-64 Pronophila, 205, 211 Pronophilina, 200-211 Protopedaliodes, 200, 205, 206, 211 kukenani, 200 profauna, 200 ridouti, 200 Prudic, Kathleen, 118-119 Prunella vulgaris, 168 Prunus, 70-73, 75, 79 Pseudomaniola, 204, 205, § Pseudosphinx tetrio, 212, 214-216 esti, TA Pseudotsuga menzie Psychidae, 65 Pupal salvage, 116-117 Pyla aenigmatica, 78-79 Pyralidae, 77-79, 157 Pyrez, Tomasz W., 200-211 Pyrenees, 10-18 Pyrginae, 6-9, 19-34, 45-57, 180, Quadricalcarifera punctatella, 152 Quercus, 73, 154, 166 Quesnelia arvensis, 38 Raworth, D. A., 10-18 Recolonization, 1 Reed, Robert, 40-43 Reproductive bulk, 143-160, 229-233 Rhamnaceae, 23, 75, 154, 190 Rhamnus, 75, 154 Rhodussa, 35, 37 Rhopobota naevana, 74-75 Ridens, 19, 23, 33,54-56 Riodininae. 85-86 Riparian, S9-95 Robbins, Robert K., 178-180 Romero M., Francisco, 37-39 Rosa, 69 Rosaceae, 6, 64, 68-73, 75, 155 Rubiaceae, 137, 212, 225, 226 Rubus, 6, 73, 75, Rudbeckia, 70 Rumex crispus, 155 Saarinen, Emily, 112-115 Saccharum, 155 Salicaceae, 66, 71-73, 75, 151, 230, 236 Salix, 66, 71-73, 75, 236 Santalaceae, 77 Sarcostemma, 118 Satyrinae, 88, 97-103, 105, 200-211 Satyrium, 92, 172 Scada, 35 Schlindwein, Clemens, 212-218 Schreckensteinia festaliella, 75 Schreckensteiniidae, 75 Schutte, Carol, 83-88 Scrophulariaceae, 41 Scythridinae, 68 Scythris mixaula, 68-69 Semioscopis, 67-68 Serpentine soils, 161-165 Sesiidae, 64-65, 70-72, 235-237 Shapiro, Arthur M., 161-165 Sharma, Narender, 170-171 Shuey, John, $3-88 Simyra henrici, 118 Solanaceae, 35, 212 Solidago, 70 Sorghum halapense, 155 Sparganothis unifasciana, 72-73 Sphingidae, 45, 212-218 Spodoptera exigua, 155 Spring Mountains, 89-95 Springs, 89-95 Stellaria nemorum, 70 Streblote panda, 144, 152, Strymon, 87, 92, 174-175, 180 martialis, 174-175 Suriana maritime, 174 Date of Issue (Vol. 59, No.4): 16 December 2005 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Synanthedon, 71-72, 235-237 bolteri, 235-237 Tachinidae, 33, 118 Taleporia walshella, 65 Tapinoma, 112, 114, 115 Taraxacum, 10 Tasmania, 10-18 Tegeticula, 64 Tetragma gei, 64 Theclinae, 112-115, 174-175, 178-180, 181-199 Tibouchina, 135, 137 Tillandsia, 37 Tineidae, 64-65, 143 Tortricidae, 72- Tortricinae, 72-74 Trema micranthum, 174 Trichoplusia ni, 156 Trifolium, 73 Typha, 111 Uehara-Prado, Marcio, 96-106 Ulmaceae, 23, 73, 174 Ulmus, 73 Urodidae, 75 Urtica, 40 Utetheisa ornatrix, 118 Vaccinium, 73, 75 Velaea, 77 Venada, 19-34 advena, 20, 23, 33 cacao, 19-34 daneva, 19-34 naranja, 19-34 nevada, 19-34 Venezuela, 6, 37-39, 178, 200-211, 217 Verbenaceae, 107, 225-226 Viburnum, 71 Vietnam, 219-222 Vitis, 155 Warren, Andrew D., 6-9, 45-58, 59, 166, 168-169, 173, 190, 194 Weinmannia wercklei, 22-23 Wockia asperipunctella, 75 Wright, Donald J., 121-133 Xenolechia velatella, 69-70 Xylophanes tersa tersa, 214, 215 Xyloryctidae,, 68-69 Yanguna cosyra, 33 Ypsolopha dentella, 66 Ypsolophidae, 66 Yucca glauca, 64 Zea mays, 155, 156 Zeiraphera hesperiana, 74 Zera, 23, 53 hosta, 23 Zizania, 110 75, 121-133, 144, 229, 231 EDITORIAL STAFF OF THE JOURNAL Micuaet E. Totiver, Editor Department of Biology Eureka College Eureka, Illinois 61530 USA miketol@eureka.edu Brian ScHottens, Assistant Editor Pec Touiver, Layout Editor Puit DeVries, Book Review Editor Biology Department Natural Imprints Department of Biological Sciences College of Charleston, 66 College Street, 706 Lake Road University of New Orleans, 2000 Lakeshore Dr. Charleston, South Carolina 29424-0011, USA Eureka, Illinois 61530 New Orleans, Lousiana 70148-0001 USA scholtensb@cofc,edu Naturimp@mtco.com pjd@mpm.edu Associate Editors: Gerarpo Lamas (Peru), KeNeLM W. Puitie (USA), Ropert K. Rossins (USA), FeLix Speriinc (Canada), Davin L. Wacner (USA), Curister WikLuND (Sweden), Caria Penz (USA), ANDREW Warren (USA), NOTICE TO CONTRIBUTORS Contributions to the Journal may deal with any aspect of Lepidoptera study. Categories are Articles, Profiles, General Notes, Techni- cal Comments, Book Reviews, Obituaries, Feature Photographs, and Cover Illustrations. 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For authors who are not members of the Society, page charges are $75 per Journal page. Authors unable to pay page charges for any reason should apply to the editor at the time of submission for a reduced rate. Au- thors of Book Reviews and Obituaries are exempt from page charges. Correspondence: Address all matters relating to the Journal to the editor. Address book reviews directly to the book review editor. PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A. CONTENTS A REVIEW OF CALLopHrys AFFINIS (W. H. Epwarps), WITH DESCRIPTIONS OF TWO NEW SUBSPECIES FROM NEw MExIcolAnD Mirxico! (Glenn Ay Gorelick =22=-22-= 2a ae 181 CLOUD FOREST BUTTERFLY FAUNA OF THE PANTEPUI—POOR OR POORLY KNOWN? DESCRIPTION OF NEW SPECIES AND RECORDS OF NEW GENERA OF PRONOPHILINA: ERETRIS AGATA AND OXEOSCHISTUS ROMEO (Nympnauipae: SatyrinaE) Tomasz W. Pyrcz and Steven Fratello ---------------------------- 200 THE HIGHLY SEASONAL HAWKMOTH FAUNA (LEPIDOPTERA SPHINGIDAE:) OF THE CAATINGA OF NORTHEAST BRAZIL: A CASE STUDY IN THE STATE OF Rio GraNnbE DO Norte José Aratijo Duarte Junior and Clemens Schlindwein -=-=======-----— === === = == a DID BEHAVIORAL OBSERVATIONS ON THE EARLY STAGES OF [AMIDES CELENO (Cramer) (LyCcAENIDAE) av Cat Tien Nationa Park, VIETNAM: AN OBLIGATE MYRMECOPHILE? Rod Eastwood, R. L. Kitching and Hui Buu Manh_ ---------------------------------------------------=--- === 5-5-5559 35 nnn 219 PoPULATION BIOLOGY OF TWO SPECIES OF HeLiconius (NyMPHALIDAE: HELICONTINAE) IN A SEMI- DECIDUOUS FOREST IN SOUTHEASTERN BraziL Rafael Barreto de-Andrade and André Victor Lucci Freitas -------------------------------------------------------------------------- 223 REPRODUCTIVE BULK IN CapitaL-BREEDING LepiportTeRA William E. Miller ------------------ 229 GENERAL NOTES PIGGYBACKING NORTHWARD: MoveMENT oF Leprores cassius (LYCAENIDAE: LYCAENINAE) THROUGHTOUT THE SOUTHEAST Jaret C. Daniels. -------------------------------------------------- 234 SYNANTHEDON BOLTERI (SESUDAE) IN Micuican William H. Taft ---------------------------------- 235 INDEX TO VOLUME 59) ---------------------------------------------------------------------------------- 238 | 'ON LIBRARIES UT HOLA wi iN 3 9088 01181 8309 @ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanance of Paper).