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Endotricha flammealis (Denis & Schiffermiiller), type species characteristic male genitalia of Endotrichini. Fic. 4. Endotricha flammealis (Denis & Schiffermiiller), aedeagus. Rs anastomosed with Sc+R, (Endotrichini); median vein non-pecti- nated” [italics ours]. Whalley (1961) recognized the problem with the definition of the hindwing character that separated the two groups: “In several cases they have been said to anastomose where, as close examination shows, they merely run very close together (e.g., Rostripalpus Hampson).” The lack of anastomosis of Rs and Sc+R, varies in other groups within the Pyraloidea besides the Pyralini, and it has been documented as highly ho- moplasious at the generic level among the genera of the Pococera complex of the Epipaschiinae (Solis 1993) and at the species level (Shaffer & Solis 1994). Other groups where the majority of the taxa lack the anastomosis of Rs and Sc+R, but where there are ex- amples where the two veins barely anastomose have been observed in representatives of the New World Cacotherapiini (Galleriinae) and some genera in the Crambinae (e.g., Pseudoschoenobius Femald). Based on our observations of the distribution of this hind- wing character in other groups within the Pyraloidea, distribution of the hindwing character within the Pyral- inae, and lack of concurrence with the characters of the male genitalia, we propose that the hindwing venational character is not reliable in the separation or definition of taxa at suprageneric levels in the Pyralinae. Minet (1982, 1985) was the first to maintain that the Pyralinae were paraphyletic because characters used by past workers were plesiomorphic. He stated: “Les Pyralinae semblent paraphylétiques par rapport a des taxa tels que les Endotrichinae, les Chrysauginae ou les Epipaschiinae (dont ils ne different que par un en- VOLUME 53, NUMBER 1 6 Fic. 5. Wing venation (idealized); arrow indicates lack of anas- tomosis. F1c.6. Wing venation (idealized); arrow indicates anas- tomosis. semble de caracteres plesiomorphes: palpes maxil- laires bien développés, ailes antérieures sans écailles hérissées, etc.),” [“The Pyralinae appear to be para- phyletic in comparison with taxa such as the En- dotrichinae, the Chrysauginae or the Epipaschiinae (in that they share a group of plesiomorphic characters: maxillary palpi well developed, forewings without raised scales, etc.).”] but he retained pyralines and en- dotrichines at the subfamily level. Whalley (1963), in his study of Endotricha, found that the retention of Ragonot’s concept of the Endotrichinae as a subfamily was not warranted and proposed the Endotrichinae as a tribe of the Pyralinae. He did not offer a reason or characters to support this conclusion. Solis and Mitter (1992) agreed with Minet that previous characters used to define the two taxa were plesiomorphic states, but they treated the endotrichines as a tribe within the Pyralinae according to Whalley (1961) because Minet (1985), in his study of the tympanal organs, presented no apomorphies for the Pyralinae, Pyralini, or En- dotrichini. Solis and Mitter (1992) proposed one char- acter of the female genitalia as a synapomorphy for the Pyralinae, but proposed none for the Pyralini or En- dotrichini. RESULTS Adult genitalic morphology. Previous authors (Whalley 1961, 1963, Munroe & Shaffer 1980) did not use genitalic characters to define the Pyralinae, Pyralini, or Endotrichini, although they used genitalic morphology at the species level for their studies. Solis and Mitter (1992) proposed a character of the female genitalia (corpus bursae barely extending cephalad be- yond segment 7) to support the monophyly of the Py- ralinae. This study, however, was based on a small sam- ple size, a character that remains untested. Pyralini (Figs. 1, 2, 7): Description: Male: Uncus same width throughout or less nar- row than the base, flat or spatulate, ventrally with spine clusters ab- sent; uncus arms laterally not large and earlike; downcurved gnathos with arms strongly developed, with well-developed medial, narrow spike terminating in a small dorsally curved hook; tegumen strongly sclerotized; vinculum well developed; juxta simple, rarely heavily sclerotized, spiny catena (baso-medial portion of anellus) present or absent or laterally sclerotized, and heavily spined, sometimes anellus reflexed with heavy sclerotization; transtilla absent or, if present, membranous, rarely well developed and heavily sclerotized; valva variable in shape, same width to apex or more narrow distally, basal and costal process absent or present, if present well developed or not, without saccular process, ventral surface of valva bearing hair- like setae not arranged in radiating rows, costal setae absent; vesica of aedeagus with or without clusters of spinelike comuti, vesica sometimes spined, reflexed with heavy sclerotization, or with broad bands of sclerotization its entire length. Female: Segment 8 and as- sociated membranes either short, or long and extensible; apophyses anteriores and posteriores long, stout or slender; ostial end of ductus bursae membranous, with small, well-sclerotized to large, heavily sclerotized compact pouches present or absent; ductus bursae long and narrow with areas of minute spines immediately below antrum or other sclerotized areas; corpus bursae large, signum variable, ab- sent, or if present from scobinate patches, usually within single large area, to long and spinelike. Endotrichini (Figs. 3, 4, 8): Diagnosis: Uncus broadest at apex; uncus arms laterally large, earlike; gnathos medially broad, spatulate, platelike. Description: Male: Uncus broadest at apex, ventrally with spine clusters present or absent (uncus process of Whalley); uncus arms laterally large, earlike [socii of Whalley; socii, according to Klots (1956) are paired processes on either side of the base of the uncus; these structures are not socii, but the most lateral elements of the uncus arms of the Pyralidae (Solis & Mitter 1992)]; downcurved gnathos arms strongly or weakly developed, usually with a well-de- veloped medial, broad, spatulate, and upturned central plate; weakly sclerotized tegumen; vinculum well developed; juxta simple, some- times with spiny manica; transtilla present, usually heavily sclero- tized; valva usually same width to apex, may bear basal process and saccular process; ventral surface bearing hairlike setae in rows point- ing toward base of valva; prominent, reflexed, sometimes spear- shaped costal setae may be present arising from costa near apex: aedeagus with vesica bearing sticklike or clublike cornutus varying in shape and length. Female genitalia: Segment 8 and associated mem- branes long and extensible; apophyses anteriores and posteriores long and slender; ostial end of ductus bursae minutely spined, with- out pouches; antrum sclerotized; ductus bursae short, minutely spined, or very long and membranous; corpus bursae large with signum scobinate. Although we can provide synapomorphies in states of the uncus and gnathos (see diagnosis above) for the Endotrichini in the male JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY ts PO eters at baiceatas UCOM OSHA DSU Cite “ke Fic. 8. Endotricha flammealis (Denis & Schif- Pyralis farinalis Linnaeus, type species; characteristic female genitalia of Pyralini. fermiiller), type species; characteristic female genitalia of Endotrichini. 7 Fic. VOLUME 53, NUMBER 1 genitalia, we were unable to find synapomorphies for the Pyralini where the states are either shared with the Endotrichini or with re- lated subfamilies. The genitalic characters of the Pyralinae are either variable at lower taxonomic levels or plesiomorphic, i.e., found in all related subfamilies. We provide a description of the genitalia for En- dotrichini and Pyralini because a concept based on the morphology of the male genitalia was used to assign taxa in the Endotrichini and Pyralini in the Australian and Neotropical checklists. Larval morphology. Comparison of the caterpillars of En- dotricha flammealis with Pyralis farinalis, Aglossa caprealis, and Herculia psammioxantha (with caterpillars of other subfamilies of the Pyralidae as outgroups) did not result in any apomorphic char- acters to support the monophyly of the Pyralinae, Pyralini, or En- dotrichini. Historically, the Hasenfuss (1960) concept of the Pyrali- nae consisted of present-day galleriines, pyralines, and phycitines (he did not include chrysaugines or epipaschiines in his study); he considered Endotricha as a pyraline. The larvae of Endotricha have a pinaculum ring on SD1 of A9, a synapomorphy for the Pyralidae (the plesiomorphic state, the ab- sence of the pinaculum ring on SD1 of AQ, occurs in the Cram- bidae). In sum, we found that E. flammealis larvae lack the unique characters assigned to other subfamilies and have the same ple- siomorphic setal character states assigned to the larvae of the Pyral- inae. The Epipaschiinae and Pyralinae both lack a pinaculum ring on any other segment other than A9 (in comparison to the presence of a pinaculum ring on T2 of the Phycitinae, T3 of the Chrysauginae, and Al of the Galleriinae; presence in each segment is the derived state, although the pinaculum has been secondarily lost in several genera and/or species of each subfamily). Based on work by Hasen- fuss (1960) and Allyson (1977) the Epipaschiinae and Pyralinae are separated from each other by the distance between the ventral setae on A7 and AQ. In the Epipaschiinae the two ventral setae are closer together on A7 than those on AQ and in the Pyralinae the two ven- tral setae on A7 and AQ are equidistant (the plesiomorphic condition shared by other subfamilies of the Pyralidae). Taxonomic placement of genera. Recently, gen- era from southeast Asia and Australia previously placed in the Endotrichinae with anastomosed Rs and Sc+R, in the hindwing, but with genitalic characteristics of the Pyralini were transferred to the Pyralini in the Australian checklist (Shaffer et al. 1996) based on the genitalia morphology. Those genera transferred from the Endotrichinae to the Pyralinae were based on the genitalia morphology: Gawna Walker, Curena Walker, Arescoptera Walker, Scenedra Meyrick, Tanyethira Turmer, Scenidiopsis Turner, Perisseretma Warren, and Perula Mabille. According to the definition based on genitalic mor- phology given above, there are no known species of Endotrichini in the Western Hemisphere, but four genera, Perforadix Sein, Micronix Amsel, Micromastra Schaus, and Taboga Dyar have been historically placed within the Endotrichinae due to the anastomosing of Rs and Sc+R, in the hindwing. Perforadix, Micronix, and Micromastra were inadvertently excluded from the Neotropical Pyraloidea checklist (Shaffer & Solis 1995). Taboga was included in the Neotropical check- list, but needed to have its position in the Pyralinae verified. We found that Perforadix belongs in the Pyraustinae and Micronix belongs in the Crambinae, both hereby transferred, and, of the four, only Micro- mastra and Taboga remain in the Pyralinae. Table 2 is a complete list of the Pyralinae (Pyralini) of the West- erm Hemisphere (Munroe 1983, Shaffer & Solis 1995). Sein (1930) placed Perforadix sacchari Sein, com- monly known as the Sugarcane root caterpillar, in the Endotrichinae. This species is a major pest of sugar cane in Puerto Rico and nearby islands. Sein (1930) il- lustrated the morphology of all life stages in great de- tail and described its biology and methods of control. The author failed to designate types or even list type specimens, but we found seven specimens each with a small label “P.R./Sein” and a red label “Cotype/Comell U. No. 6087” at Cornell University. According to Sein (1930), W. T. M. Forbes, who was at Cornell Univer- sity at the time, identified the material and presumably he also labelled the material as cotypes. We designate one specimen (male) as the lectotype and the other 6 specimens as paralectotypes (material in poor condi- tion, abdomens are missing), and they are labelled as such in the collection at Cornell University. We stud- ied additional material collected by Sein, identified by H. G. Dyar, as stated by Sein (1930), and dissections by Carl Heinrich at the USNM and found that Per- foradix is a synonym of Sufetula Walker, new syn- onymy, in the Pyraustinae. We discovered that based on the morphology of the tympanal organs (i.e., cram- bid “open” tympanal organs with a praecinctorium) it belongs in the Crambidae. Based on the external and genitalic morphology after comparison with other spe- cies in the genus, including the type species, it belongs in the genus Sufetela Walker. It is interesting to note that P. sacchari was originally identified for Sein by H. G. Dyar as Sufetula grumalis Schaus, a species presently placed in Sufetula (Munroe, 1995:76). We also examined another species, Sufetula pygmaea Hampson, and found it does not belong in the Pyra- loidea, but in the Noctuidae (Nolinae): Nola pygmaea (Hampson), new combination. Amsel (1956) described Micronix nivalis in the En- dotrichinae. Nothing is known about the biology of this Venezuelan species. We were unable to locate the type, a male, but Amsel provided a photograph of the adult and poor line drawings of the male genitalia and wings. The hooded uncus and the costal process of the male genitalia characteristic of crambines are very ev- ident in the illustration; therefore, we transfer this monotypic genus to the Crambinae. Although we can- not determine its placement within the Crambinae, we suggest that it belongs in the tribe Crambini. Schaus (1940) placed Micromastra isoldalis in the Endotrichinae. Nothing is known about the biology of this Puerto Rican species. Dyar (1914) described Taboga inis in the Endotrichinae. The type series is from 8 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TABLE 2. Pyralini (Pyralidae: Pyralinae) of the Western Hemisphere Aglossa Latreille, [1796] Hypsopygia Hiibner, [1825] Euclita Hiibner, [1825] Agriope Ragonot, 1894 acallalis Dyar, 1908 baba Dyar, 1914 cacamica (Dyar, 1913) (Pyralis) caprealis (Hiibner, [1800—09])(Pyralis) capreolatus Haworth, 1809 cuprealis Hiibner, [1825], missp. aenalis (Costa, 1836)(Pyralis) domalis Guenée, 1854 incultella (Walker, [1866])(Acrobasis) enthealis (Hulst, 1886)(Tetralopha) cuprialis Heinrich, 1931, missp. costiferalis (Walker, 1886) (Pyralis) costigeralis (Walker, [1865] (Pyralis), preoce. (Walker, 1862) cuprina (Zeller, 1872) (Pyralis) disciferalis (Dyar, 1908) (Pyralis) electalis Hulst, 1866 furva Heinrich, 1931 gigantalis Barnes & Benjamin, 1925 oculalis Hampson, 1906 pinguinalis (Linnaeus, 1758)(Pyralis) marmorella (Geoffroy, 1785)(Tinea) marmoratella (Villers, 1789)(Tinea) pinguiculatus (Haworth, 1809)(Crambus) guicciardti Constantinio, 1922 Arispe Ragonot, 1891 Uscodys Dyar, 1909 cestalis (Hulst, 1886)(Anerastia) atalis (Dyar, 1908)(Uscodys) concretalis Ragonot, 1891 ovalis Ragonot, 1891 Catocrocis Ragonot, 1891 Catacrocis Ragonot, 1892, missp. lithosialis Ragonot, 1891 Dolichomia Ragonot, 1891 amoenalis (Méschler, 1882) (Asopia) isidora (Meyrick, 1936)(Pyralis) binodulalis (Zeller, 1872)(Asopia) craspedalis (Hampson, 1906) (Tegulifera) datames (Druce, 1900)(Pyralis) decetialis (Druce, 1900)(Pyralis) graafialis (Snellen, 1875)(Asopia) impurpuratalis (Dognin, 1910)(Pyralis) nigrapuncta (Kaye, 1901)(Pyralis) olinalis (Guenée, 1854)(Pyralis) trentonalis (Lederer, 1863)(Asopia) himonialis (Zeller, 1872)(Asopia) infimbrialis (Dyar, 1908)(Herculia) phanerostola (Hampson, 1917)(Paractenia) planalis (Grote, 1880)(Asopia) enniculalis (Hulst, 1886)(Asopia) occidentalis (Hulst, 1886)(Asopia) plumbeoprunalis (Hampson, 1917)(Herculia) resectalis (Lederer, 1863)(Asopia) thymetusalis (Walker, 1859)(Botys) New combination devialis (Grote, 1875)(Asopia) vernaculalis (Berg, 1874) (Asopia) Herculia Walker, 1859 Buzala Walker, 1863 Cisse Walker, 1863 Bejuda Walker, [1866] Bleone Ragonot, 1890 Herculea Amsel, 1956 [index], missp. tabidalis (Warren, 1891)(Pyralis) costalis (Fabricius, 1775)(Phalaena) fumbrialis ([Denis & Schiffermiiller], 1775)(Pyralis) purpurana (Thunberg, 1784)(Tortrix) hyllalis (Walker, 1859)(Pyralis) Mapeta Walker, 1863 Homalochroa Lederer, 1863 cynosura Druce, 1895 omphephora Dyar, 1914 schausi Druce, 1895 xanthomelas Walker, 1863 aestivalis (Lederer, 1863)(Homalochroa) divisa (Boisduval, 1870)(Pyralopsis) Micromastra Schaus, 1940 isoldalis Schaus, 1940 Neodavisia Barnes & McDunnough, 1914 Davisia Barnes & McDunnough, 1913, preoce (Del guercio, 1909 [Hemiptera]) melusina Ferguson, Blanchard, & Knudson, 1984 singularis (Barnes & McDunnough, 1913)(Davisia) Ocrasa Walker, [1866] Parasopia Méschler, 1890 nostralis (Guenée, 1854)(Pyralis) helenensis (Wollaston, 1879)(Pyralis) tenuis (Butler, 1880)(Pyralis) dissimilalis (Méschler, 1890)(Parasopia) sordidalis (Barnes & McDunnough, 1913)(Herculia) psammioxantha (Dyar, 1917)(Herculia) venezuelensis (Amsel, 1956)(Herculia) tripartitalis (Herrich-Schiaffer, 1871)(Asopia) Pseudasopia Grote, 1873 cohortalis (Grote, 1878)(Asopia) florencealis (Blackmore, 1920)(Herculia) intermedialis (Walker, 1862)(Pyralis) sodalis (Walker, 1869) (Pyralis) squamealis Grote, 1873 phoezalis (Dyar, 1908)(Herculia) Pyralis Linnaeus, 1758 Aletes Rafinesque, 1815, nom. nud. Ceropsina Rafineques, 1815, nom. nud. Spyrella Rafinesque, 1815, repl. name Asopia Treitschke, 1828 Sacatia Walker, 1863 Eutrichodes Warren, 1891 farinalis Linnaeus, 1758 domesticalis (Zeller, 1847)(Asopia) fraterna Butler, 1879 manihotalis.- Matsumura, 1900 (not Guenée, 1854) meridionalis Schmidt, 1934 orientalis Amsel, 1961 manihotalis Guenée, 1854 vetusalis Walker, [1859] gerontesalis Walker, [1859] laudatella (Walker, 1863)(Sacatia) despectalis Walker, [1866] miseralis Walker, [1866] achatina Butler, 1877 haematinalis (Saalmiiller, 1880)(Asopia) gerontialis (Meyrick, 1888)(Asopia), emend. centripunctalis (Gaede, 1916)(Endotricha) pupalis Strand, 1919 compsobathra Meyrick, 1932 Taboga Dyar, 1914 inis Dyar, 1914 VOLUME 53, NUMBER 1 Panama. Study of the genitalia of the type series at the USNM of both of these species confirm their placement within the Pyralini, and not in the Endotrichini. In the Australian checklist (Shaffer et al. 1996:173) the headings of the Endotrichini and Pyralini were mislabeled and difficult to change at proof stage. The Endotrichinae should have been titled the Endotrich- ini and placed under the heading of the Pyralinae. In addition, the genus Macna Walker was inadvertently included in the Pyralinae (Pyralini), but it should have been placed in the Chrysauginae. In the Neotropical checklist (Shaffer & Solis 1995:80) the Pyralini should have been included as a subheading under Pyralinae to indicate the tribal placement of the genera found in the Western Hemisphere. DISCUSSION The Pyraloidea, one of the larger superfamilies of the Lepidoptera, has over 15,000 described species, yet much remains to be done in taxonomy, and, more so, with the phylogenetic relationships. A taxonomic study usually begins with a checklist or a catalogue of described species as an inventory to document those that have already been described. A checklist may re- fine the placement of taxa and can clearly mark taxon transfers, as well as provide other information, such as misplaced taxa. By definition, a checklist or catalog does not adequately state or discuss the taxonomic problems solved or those that remain to be solved. We have described the morphological reasons for the placement of taxa in the Pyralini or Endotrichini in two checklists (Shaffer et al. 1996, Shaffer & Solis 1995). We have also summarized the taxonomic and phylogenetic status of the two tribes included in the Pyralinae. The genitalia of the Endotrichini are clearly different from those of the Pyralini, but authors have dealt only with a few genera in both taxa and, as we have shown, have used the same plesiomorphic or ho- moplasious characters since Meyrick (1890) to define higher level taxa. We retain the two tribes in the classi- fication system for the sake of stability and retention of character information, but acknowledge that the Pyralini is likely a paraphyletic group. Moreover, a pre- liminary study of an independent character set, the lar- val stage, provides no obvious synapomorphies for the Pyralinae or the Pyralini. Our observations of the genitalia and larvae of the Pyralini and Endotrichini are made with the expecta- tion that they may prove useful in a future phyloge- netic study of the Pyralini genera that includes an en- tire suite of adult and immature characters. Any future study should also include pupal and perhaps behav- ioral characters, although caution is suggested con- cerning the latter due to the convergent nature of be- havioral characters. A phylogenetic analysis of the gen- era of the Pyralini would be the first attempt to test the paraphyly of the Pyralini with respect to the Endotri- chini, with the possibility that results may also invali- date the traditional tribal concept. Such a study may also provide characters to support the monophyly of the Pyralinae. ACKNOWLEDGMENTS We thank for their hospitality: E. R. Hoebeke, Comell University, Ithaca, New York, USA, for help while studying the Sein material and A. Hausmann, Zoologische Staatssammlung, Munich, Germany, while studying the Amsel material. We also thank Linda Lawrence, scientific illustrator, Systematic Entomology Laboratory, U.S. De- partment of Agriculture, for the figures. The following provided re- views or helpful comments: Vitor O. Becker, Marianne Horak, Douglas Ferguson, James Pakaluk, and David Smith. LITERATURE CITED ALLYSON, S. 1977. The study of some North American larvae of the genus Tetralopha Zeller (Lepidoptera: Pyralidae: Epipaschi- inae). 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Review and phylogenetic analysis of the subfamilies of the Pyralidae (sensw stricto)(Lepidoptera: Pyraloidea). Syst. Entomol. 17:79—90. WHALLEY, P. E. S. 1961. A change of status and a redefinition of the subfamily Endotrichinae (Lep. Pyralidae) with the description of a new genus. Ann. Mag. Nat. Hist. 3(13):733-736. . 1963. A revision of the world species of the genus En- dotricha (Zeller) (Lepidoptera: Pyralidae). Bull. Brit. Mus. (Nat. Hist.) Entomol. 13(2):395—454. Received for publication 15 February 1998; revised and accepted 29 September 1998. MANUSCRIPT REVIEWERS, 1998 The merit of a scientific journal depends on the quality of its reviewers as well as its authors, but the former are usually unknown to the readers of the published articles. The Journal relied on the expertise of 60 reviewers last year to provide 66 evaluations of manuscripts. It is with much gratitude that the Journal acknowledges the services of the people listed below, from whom manuscript reviews were received in 1998. Those who reviewed two or more manuscripts are denoted by asterisks. Adamski, David, Washington, DC Aiello, Annette, Balboa, Panama Arnold, Richard, Pleasant Hill, CA Austin, George, Las Vegas, NV Ballmer, Greg, Riverside, CA Boggs, Carol, Stanford, CA Borkin, Susan, Milwaukee, WI Bowers, Deane, Boulder, CO Brower, Andrew, Corvallis, OR Brown, John, Washington, DC Brown, Richard, Mississippi State, MI Burns, John, Washington, DC Calvert, William, Austin, TX Carde, Ring, Riverside, CA Cho, Soowon, Berkeley, CA Collins, Michael, Nevada City, CA Covell, Charles, Louisville, KY Drummond, Boyce, Florissant, CO Dussourd, David, Conway, AR Eichlin, Tom, Sacramento, CA Emmel, Tom, Gainesville, FL Epstein, Marc, Washington, DC Ferguson, Douglas, Washington, DC Fink, Linda, Sweet Briar, VA Friedlander, Timothy, North Potomac, MD Gall, Lawrence, New Haven, CT Goldstein, Paul, New York, NY Hawkins, Bradford, Irvine, CA Jenkins, Dale, Sarasota, FL Kelley, Scott, Boulder, CO Landry, Bernard, Aylmer, QB Landry, Jean Francois, Ottawa, ON Lederhouse, Robert, Lockport, NY MacNeill, Donald, San Francisco, CA McCabe, Timothy, Albany, NY *Metzler, Eric, Columbus, OH Miller, Lee, Sarasota, FL Miller, William, Saint Paul, MN Peigler, Richard, San Antonio, TX Poole, Robert, Rockville, MD *Powell, Jerry, Berkeley, CA Prowell, Dorothy, Baton Rouge, LA Pyle, Robert, Gray’s River, WA Rindge, Frederick, New York, NY Robbins, Robert, Washington, DC Scholtens, Brian, Charleston, SC Scriber, Mark, East Lansing, MI *Shapiro, Arthur, Davis, CA Shaw, Scott, Laramie, WY Shepard, Jon, Nelson, BC *Shuey, John, Indianapolis, IN Sims, Steve, Maryland Heights, MS Singer, Michael, Austin, TX Strand, Michael, Madison, WI Swengel, Ann, Baraboo, WI *Tuck, Kevin, London, England Webster, Reginald, Fredericton, NB Weller, Susan, Saint Paul, MN West, David, Blacksburg, VA Williams, Ernest, Clinton, NY Journal of the Lepidopterists’ Society 53(1), 1999, 11-21 POPULATION BIOLOGY AND WING COLOR VARIATION IN HELICONIUS ERATO PHYLLIS (NYMPHALIDAE) RENATO ROGNER RAMOS Av. Francisco Glicério 615/33, 11065-405, Santos, Sao Paulo, Brazil AND ANDRE VICTOR LUCCI FREITAS Museu de Historia Natural, Instituto de Biologia, Universidade Estadual de Campinas, CP 6109, 13083-970, Campinas, Sao Paulo, Brazil* ABSTRACT. Twenty-six months of mark-recapture study of three populations of Heliconius erato phyllis in SE Brazil showed these popu- lations to be relatively stable through time. The adults showed high longevity, similar to that of other tropical Heliconius. Sex ratio was male bi- ased, and males showed longer residence times than females. The number of red raylets on the ventral hindwing showed seasonal variation, con- sidered to be numeric polyphenism. Of all the species of Heliconius, H. erato is proposed to be the most tolerant of variation in habitat, larval and adult resources (“ecological plasticity”). These traits are proposed as reasons for the wide distribution of this species in tropical America. Additional key words: mark-recapture, Heliconiini, Passifloraceae, red raylets, polyphenism. Even if Heliconiine butterflies are among the best- studied tropical insects, with much known about their systematics, population ecology, behavior, immature biology, host plant relationships, coevolution, mimicry, chemistry, genetics, and conservation (Ehrlich and Gilbert 1973, Brown & Benson 1974, Gilbert 1975, Araujo 1980, Brown 1981, Tumer 1981, Sheppard et al. 1985, Mallet 1986a, 1986b), data are not available for most species and geographic populations of Helico- nius; and generalizations based on other well-known species and regions may fail to explain local patterns and processes. For example, the current idea that Heli- conius maintain relatively stable population numbers through time (based mainly on Tumer 1971 and Ehrlich & Gilbert 1973) is not true for Heliconius erato phyllis (F.) near the limits of its range in temperate southern Brazil (Saalfeld & Araujo 1981). Heliconius erato is the most widespread species of the genus, present in several different habitats and for- est types from Mexico to north Argentina (Emsley 1964, 1965), and its subspecies phyllis has the widest geographic distribution (Brown 1979, Sheppard et al. 1985) and environmental tolerances (Araujo 1980). In the southern populations of H. erato (30°S), periodic variation in several features of wing color pattern, es- pecially the hindwing red-raylets, has also been noted (Pansera & Araujo 1983, Oliveira & Araujo 1992). The present study describes features of a population of H. erato phyllis in a subtropical rainforest in southeastern Brazil (24°S), 6° (660 km) farther north of the southern limits of the species distribution, and reports cyclical annual variation in two wing color-pattern elements. *To whom all correspondence should be addressed. STuDy SITES AND METHODS A mark-release-recapture (MRR) study was carried out in three areas in SAo Paulo state, southeastern Brazil. The main study area was “Morro do Voturua” (MV) (46°22’W, 23°57’S), in the city of Sado Vicente (Fig. 1). The site was originally covered with lowland subtropical rainforest (Ururahy et al. 1987). The an- nual rainfall reaches 2500 mm and the average annual temperature is 21°C (Setzer 1949, Prodesan 1969, Nimer 1972), with the mean of the coldest month 18.2°C and of the warmest month 25.3°C (Santos 1965) (Fig. 2, methods following Santos 1965 and Wal- ter 1985). Most of the area is covered by secondary forest on low hills (100—200 m elevation). Similar work was done in two nearby sites: the “Vale do Rio Qui- lombo” (VQ, Fig. 1), a road along a river valley with much secondary vegetation and flowers, 12 field days over three months; and the “Morro do Japui’ (MJ, Fig. 1), a large hill facing the ocean SW of MV with similar vegetation, 47 field days over seven months (see details on these areas in Freitas 1993). Mark-release-recapture studies were made in Morro do Voturud during 26 months, from 13 August 1994 to 30 September 1996, 1—4 times per week, to- taling 153 field days (about 4 hours/day). Butterflies were captured with an insect net, individually num- bered on the underside of both forewings (in the red central macula) with a black permanent felt-tipped pen, and released. Age (based on wing wear), forewing length (in mm), point of capture, sex and food sources were recorded (as in Freitas 1993, 1996). The age of individual butterflies was estimated based on wing wear, initially using the six categories described by 12 —— SAO VIC MORRO DO VOTURUA (MV) JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY ATLANTIC OCEAN =* TRAILS —<— STREAMS GBB BUILDINGS Fic. 1. (A) study area (Sao Vicente Region) in Southeastern Brazil. In regional map (B), VQ = Vale do Rio Quilombo, MJ = Morro do Japuf, MV = Morro do Voturua (details in C). Ehrlich and co-workers (Ehrlich & Davidson 1960, Brussard & Ehrlich 1970, Ehrlich & Gilbert 1973). These six categories (freshly emerged, new, intermedi- ate, old, very old, tattered) were regrouped into three categories: new = freshly emerged and new individu- als, intermediate = intermediate individuals, and old = old, very old, tattered (as in Freitas 1993, 1996). Age structure was calculated as the daily proportion of each category, and grouped into monthly means. Three wing-color-pattern elements were recorded (Fig. 3): the number of red basal spots, the number and shape of the red raylets on the ventral hindwing, and the color and size of the spot in the inner angle of space Cul—Cu2 (“cubital spot”) on the dorsal forewing (nomenclature following Miller 1970). Data from Au- gust and September 1994 at MV were regarded as the “Winter 1994” sample; each of the following seasons represents observations for 3 consecutive months. The MRR data from MV were analyzed by the Jolly- Seber method (Southwood 1971) for estimating popu- lation parameters (software developed by R. B. Francini, UNISANTOS). In most cases, only males were analyzed because of the low number of females. Daily results were tabulated as “number of individuals captured per day” (NICD), and “number of individu- als present per day” (NIPD). To estimate the NIPD, recaptured individuals were considered to be present in the population on all previous days since the day of first capture (=marked animals at risk). RESULTS Population Dynamics In Morro do Voturua, the NICD for males varied from one to 13 (mean = 5.9, SD = 2.4; n = 153 days), with 19 days with n = 3 and 13 days with n = 10 males (Fig. 4). The number of newly marked individuals captured in each month (=monthly recruitment) also varied, more in the first year than in the second (Fig. 5). In nearby Morro do Japui, the male NICD ranged from 1 to 5 (mean = 2.3, SD = 1.1, n = 47 days). The area covered in these two sites is similar (Fig. 1, see also Freitas 1993). In Vale do Rio Quilombo, the NICD for males was greater than in Morro do Voturua (mean = 14.5, SD = 4.1, n = 12 days) (Fig. 6A). The area covered in this site was about six times greater than in the previous two, corresponding closely to the NICD ratio for Morro do Japui (6.3) and suggesting an equivalent density, 40% of that in Morro do Voturua. Jolly-Seber analysis for males in Morro do Voturua (Fig. 7) gave estimated numbers from 3 to 88 individ- VOLUME 53, NUMBER 1 (mm ) RAINFALL wn i JASONDJFMAMJJVASONDJFMAMJJAS 1994 1995 (MONTHS ) 1996 Fic. 2. Climatic diagram of the Sao Vicente Region during the study period (format following Santos 1965 and Walter 1985). Hatched = humid periods, black = superhumid periods. uals (mean = 13.5, SD = 9.9, n = 152), with most days (n = 90) between 10 and 30. The NIPD for males in the same site ranged from 1 to 20 (mean = 9.8, SD = 3.8, n = 154). The male population was stable through- out the year, with small fluctuations in number of indi- viduals apparently not related to season, but with lower numbers of individuals observed at the end of winter in the three years (Figs. 4, 5). The results show that NIPD is better than NICD as an index of population size. In species with stable pop- ulations composed of long-lived resident individuals, efficient marking and recapture can give nearly equal population size estimates by the Jolly-Seber method and simple NIPD. Sex Ratio The sex ratio of individuals captured and marked was male biased in all sites (Table 1, Figs. 6B and 8). In MV males dominated in all but one month of the study; all the captured individuals were male in four months, including April of both years (Fig. 8). In all, 263 males and 74 females were captured and marked. Males were recaptured from one to 16 times and fe- males from one to five times; 154 males and 30 fe- males were recaptured at least once. The proportion of recaptured males (58.5%) was statistically the same as that of females (40.1%) (y? = 2.6, df = 1, P > 0.20). In VQ the proportions of recaptures of males (30.8%) and females (30.5%) were equal. Age Structure Comparing the new individuals vs. intermediate+old, most of the first captures in MV were of “new” indi- viduals from both males (71%), and females (54% 13 new), even if this number was considered greater in males (x? = 7.6, df = 1, P < 0.005). Most of the butter- flies captured on each day (after the first) were previ- ously marked individuals (mean = 71.4%, SD = 23.4, n = 152 days). The age structure during the 26 months in MV was not stable, with decreases in proportion of “new” individuals in the winter (Fig. 9). In VQ, the proportion of recaptures/day was high (Fig. 6C), and the age structure quite stable (Fig. 6D), but these results should be taken with care due to the short period of study. Residence Time In MV, males had a longer residence time (mean = 37.6 days, SD = 25.8, n = 154) than females (mean = 22.6 days, SD = 23.0, n = 30) (t = 2.96, df = 182, p = 0.004). Estimated residence time of males (“life ex- pectancy” of Cook et al. 1967) was 28.1 days. The max- imum individual permanence (survival) was at least 127 days for a male and 89 days for a female (Table 2). In MJ, residence time was calculated only for males (mean = 18 days, SD = 14.6, n = 22), since the recap- ture rate of females was low (only 3 recaptures). In VO maximum permanence could not be calculated (due to the short period of study and low number of recap- tures), but the residence time for males (mean = 20.1 days, SD = 12.8, n = 32) and females (mean = 15.0 days, SD = 10.6, n = 11) were not significantly differ- ent (t = 1.17, df = 41, p = 0.248). Vagility In MV, most adults showed home range behavior, being observed in the same site for several days, often using the same flower resources. The total distance RR Fic. 3. Wing pattern variation in Heliconius erato phyllis (back- ground black, solid = red, dotted = yellow). Colored spot in the in- ner angle of space Cul—Cu2 (cubital spot, CS) present (left, red macula almost fused with this), or absent (right). Five basal red spots (RBS) and six red raylets (RR) shaped like small lines (left); four basal red spots and four red raylets shaped like dots (right). 14 oN A TOTAL MALES CAPTURED/DAY yu fH @ JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY SOE OED My OD ES ONO ya A I od D8 @ 1994 1995 (MONTHS) 1996 Fic. 4. Daily captures for males of H. erato phyllis in Morro do Voturud, August 1994 to Sep- tember 1996. flown by adults varied among individuals. In general, two classes of individuals could be identified: those al- ways observed in the same site (residents) and those seen in two or more separate points in the study area (vagile). Most of the males (71%) and females (85%) recaptured were vagile, while among the residents, some individuals appeared to have tightly restricted home ranges. One male was captured 17 times (twice on the same day) at the same place, and at least half of the resident individuals were captured more than 4 times. The distance traveled by vagile individuals var- ied from 50 to 660 m (mean = 270.6, SD = 122.9m,n = 97) for males, and 80 to 600 m (mean = 236.3, SD = 140.9, n = 23) for females (Table 3). Some individuals showed great mobility in a single day. In one case, one male captured at 0900 was observed 500 m away at 1000 h, returning at 1030 h to within 50 m of the first site of capture. Males were commonly seen near nec- tar sources and females were mostly observed near the larval food plants. Wing Size and Color Pattern in Morro do Voturua Four or five red basal spots are present on the ven- tral hindwing of Heliconius erato butterflies. This trait was very stable, as only three out of 337 individuals an- alyzed (males and females) showed the fifth basal spot (Fig. 3). TOTAL N2 MALES MARKED/MONTH ASONOD,JFMAMJ peice ta altar Clt 1994 1995 ( MONTHS ) 1996 Fic. 5. Monthly recruitment of males of H. erato phyllis in Morro do Voturud, August 1994 to September 1996, as total number of individuals marked for the first time in each month. The number of red raylets in butterflies of both sexes showed a bimodal distribution, ranging from one to seven, with peaks at three and five (Table 4). The average number increases in summer and autumn and decreases in winter and spring (Table 4). The shape of the red raylets varied from a rounded dot to a small red line. Individuals with red raylets shaped like dots (65.8%) were more frequent than those with lines (34.2%). The proportions of each type also varied throughout the year (Fig. 10A). The cubital spot on the dorsal forewing could be ab- sent, present, or fused with the transverse subapical red bar. If present and not fused, the spot could be red or yellow. This character was very variable, but there was a Clear majority of “spot to be present and yellow’, with no regular pattern of variation throughout the year (Fig. 10B). The forewing length of males ranged from 27 to 42 mm, the average varying along the year, with greatest values being observed during the summer (Fig. 11). The average forewing length of males (mean = 36.25 mm, SD = 2.99, n = 250) was greater than that of females (mean = 35.21 cm, SD = 2.96, n = 71; t = 2.58, df = 319, p = 0.01). This difference was not found when only recaptured individuals were considered (t = 1.31, df = 181, p = 0.191). When comparing the recap- tured vs. not recaptured within sexes, recaptured indi- viduals were smaller. This difference was almost sig- nificant for males (t = 1.933, df = 248, p = 0.054), but not for females (¢ = —0.172, df = 2.83, p = 0.864). Natural History of Adults Adults began activity around 0800 h in summer and 0930 h in winter, but this could vary within a season according to the weather. On some cold days during winter, activity began only after 1030 h. Beak marks (from handling and release by, or es- cape from, bird predators) were found on the wings of individuals from all populations, but a different pro- portion of individuals with damaged wings was ob- VOLUME 53, NUMBER 1 TABLE 1. Sex ratio of marked individuals of H. erato phyllis in the three study sites. Study site Males Females Sex ratio 72 values Morro do Voturua 263 74 3.5:1 105.9 Morro do Japui 50 6 8.3:1 34.6 Vale do Rio Quilombo 104 36 3:1 33 served in each site. Beak marks were seen in MV on 18 males (6.8%; 14 later recaptured) and 1 female (1.3%); in MJ, on 10 males (20%) and no females; in VQ, on 18 males (17%) and 8 females (22%). Adults were almost always observed feeding on flow- ers, but a few individuals were observed feeding on damaged fruits of Rubus rosaefolius Smith (Rosaceae), and in one case a captured individual was observed bearing a seed of this plant on its proboscis. The most visited flower in the MV site was Lantana camara L. (Verbenaceae) (more than 150 records). Two varieties of L. camara occur in the study area, one with red-yel- low flowers and the other (much more common) with white flowers. Some individuals were also observed visiting flowers of Rubus rosaefolius (30 records), As- clepias curassavica L. (Asclepiadaceae) (13 records), Epidendrum fulgens Brongn. (Orchidaceae) (3 records), Gurania sp. (Cucurbitaceae) (3 records), and Impatiens walleriana Hook. f. (Balsaminaceae) (2 records). Pollinia of Epidendrum fulgens and Asclepias curassavica were observed attached to the proboscis of some captured individuals. A new male was once observed visiting the inflorescence of Heliconia sp. (Musaceae). Individuals in MV were rarely observed feeding on flowers of Asteraceae, although several species of Mikania, Eu- patorium, and Vernonia are common there. The few Asteraceae that were observed being visited include Mikania lundiana D.C. (3 records), Trixis antimenor- rhoea Mart. ex. Baker (3 records), Eupatorium laevi- gatum D.C. (3 records), and Emilia sonchifolia D.C. (2 records). In the VQ site (where L. camara is not so common), the Asteraceae Bidens pilosa L. and Titho- nia speciosa Hook ex Gris. were the most visited spe- cies with 90 and 45 feeding records respectively. Also in this site Stachytarpheta polyura (L.) (Verbenaceae) was commonly used (15 records). Some adults in MV appeared to show “trap-line” (learned sequence) be- havior for gathering pollen and nectar (see Ehrlich & Gilbert 1973 and Ehrlich 1984). Over a two-month period, one male was observed visiting a dense Lantana patch at about 1000 h, and then in another patch 150 m away after 30-40 min- utes. Several individuals showed similar patterns throughout the two years of study. Occasional distur- bance on forest edges (removal of some flower patches) 15 TABLE 2. Permanence of marked H. erato phyllis in the “Morro do Voturua”. Days elapsed between marking and last recapture rep- resent the minimum permanence (MP) for each individual. MP Males P(%) Females P(%) Total 1-20 51 33.1 18 60.0 69 21-40 38 24.7 5 16.7 43 41-60 34 22.0 5) 16.7 39 61-80 21 13.6 ] 3.3 22 81-100 9 6.0 1 3.3 10 >100 1 0.6 0 0.0 1 Total 154 100.0 30 100.0 184 resulted in disappearance of some individuals from the study area for up to two weeks. Four Passiflora species were recorded as larval food- plants in MV: Passiflora alata Dryand, P. edulis Sims (both used rarely), P._ jileki Wawra, and P. capsularis L. Other species may be used in the same area and in the neighboring sites. The behaviors observed in oviposit- ing females and larvae were the same as reported in the literature (see Brown 1981). DISCUSSION Populational Ecology and Adult Biology The population parameters of H. erato phyllis seen in this study agree with those reported for other tropi- cal Heliconius (Turner 1971, Ehrlich & Gilbert 1973, Araujo 1980, Mallet & Jackson 1980, Ehrlich 1984, Mallet et al. 1987, Quintero 1988). The MV popula- tion was reasonably stable in numbers during the two years of study if compared with species of other sub- families such as Ithomiinae and Troidini, both showing pronounced fluctuations in numbers through time (Vasconcellos-Neto 1980, Brown et al. 1981, Francini 1989, Freitas 1993, 1996, Tyler et al. 1994, Pinto & Motta 1997). Other features observed for H. erato in the present study were the continual recruitment of new individuals, low density of adults, absence of sud- den increases and decreases in numbers of individuals, and non-cyclical age structure. The population seems to be less affected by climate than those observed by Benson (1978) for populations in a drier site in Rio de Janeiro or Saalfeld and Araujo (1981) for populations TABLE 3. Maximum distance (m) traveled by adults of H. erato phyllis in MV. Residents Vagile up to 50 51-150 150—500 >500 Males 39 17 77 3 Females 4 8 13 2 Total 43 25 90 5 16 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TABLE 4. Frequency distributions, sample size (N), mean (X), and standard deviation (SD) of the number of “red raylets” for month of first capture in butterflies of both sexes captured in the “Morro do Voturua’. Number of Red raylets t to > Month/year Aug/1994 Sep/1994 Winter/1994 Oct/1994 Nov/1994 Dec/1994 Spring/1994 Jan/1995 Feb/1995 Mar/1995 Summer/1995 Apr/1995 May/1995 Jun/1995 Autumn/1995 Jul/1995 Aug/1995 Sep/1995 Winter/1995 Oct/1995 Nov/1995 Dec/1995 Spring/1995 Jan/1996 Feb/1996 Mar/1996 Summer/1996 Apr/1996 May/1996 Jun/1996 Autumn/1996 Jul/1996 Aug/1996 Sep/1996 Winter/1996 Total — —_ —_ — FPOOrRrFOWFOORrRNAN ODE ORR HRANKHRWOOKEUONDYHNK OO NWRWNrFRrFOWFRONTUNHFNODNOKKNNOHOOH KANN OW OW — AWOonWAnNnrnNn®@bPRPRNRP WWF ONWWrrR WOANAHADBHLADL WONOrFOGOCGCOCOOCOFORrFOONNNFORFONFFrOOCOCONOCOLhD — Ol — ie) ee} i OL =] much farther south. In the latter populations, three distinct phases were reported: a period of rapid growth in early spring, a period of maximum density in sum- mer through autumn, and an abrupt decline in winter (Saalfeld & Araujo 1981, Romanowsky et al. 1985). In the periods of maximum density, the number of indi- viduals captured in these populations could be very high (more than 150 individuals reported in one roost- ing site) in relation to the maximum values obtained in the present study (from 20 to 30 individuals) (Saalfeld & Araujo 1981, Romanowsky et al. 1985). The average time of residence and the life expec- tancy reported in the present study are both high, like those of other Heliconius (Turner 1971, Benson 1972, Ehrlich & Gilbert 1973, Araujo 1980, Quintero 1988), including the populations from southern Brazil (Ro- manowsky et al. 1985). The values are higher than those of many Ithomiinae, Pieridae, and Papilionidae 1 Zz X+ SD ur 2 1 0 20 3.3 + 14 2 0 0 15 3.3 + 1.0 4 1 0 35 3.3 + 1.2 3 0 0 8 3.9 + 1.0 4 3 0 21 4.0 + 1.2 5 1 0 17 3.7 + 1.2 12 4 0 46 3.9 + 1.2 10 6 0 33 4241.3 2 5 0 9 5.0 + 1.6 8 3 1 20 4.5 +14 20 14 1 62 44+1.4 5 3 ] 9 5.3 + Ld 8 0 0 17 3.9 + 1.3 5 2 0 16 4141.3 18 5 1 43 4.3+1.3 5 1 0 25 3.4 + 1.3 1 1 0 6 3.3 + 2.1 1 0 0 11 Qu & 12; if 2 0 42 3.2 + 1.4 0 2 0 8 3.6 + 1.6 1 1 0 13 3.6 + 1.6 1 ] 0 5 3.6 + 1.8 2 4 0 26 3.6 + 1.4 5 2 0 11 44+14 4 5 0 10 5.3 + 1.0 5 5 0 15 4.6+1.4 14 12 0 36 4.7+1.3 4 6 1 16 5.1 + 1.2 1 1 0 4 40+ 1.2 2 ] 0 i 40+1.4 7 8 1 27 4.6+1.4 2 0 0 9 3:0 + 1.5 0 2 0 3 47+ 2.3 0 0 0 8 2.2 + 0.8 2 2 0 20 2.9 + 1.5 86 52 3 337 3.9 + 1.4 in the same or nearby sites (Table 5; see also Young & Moffett 1979, Vasconcellos-Neto 1980, Brown et al. 1981, Freitas 1993, 1996, Tyler et al. 1994, Pinto & Motta 1997). The high values for residence are not re- lated to geographic location of the populations, since they are higher in Heliconius erato than in butterflies of other groups in the same and nearby sites (Table 5). In part the high values of residence reported in He- liconius could be related to the movements and dis- persal of the adults. Some Heliconius and Eueides but- terflies are known to have restricted home ranges throughout their lives, and even if adults can fly dis- tances up to 5 km within the forest (Brown 1981, Pansera & Araujo 1983, Romanowsky et al. 1985, Mal- let et al. 1987), small areas of open field could act as efficient barriers to dispersal (Romanowsky et al. 1985). This pattern contrasts with that of “nomadic” butterflies such as Ithomiinae and Troidini (see discus- VOLUME 53, NUMBER 1 > Pa) A 6 J Ww a 20 > Ee = 1© oO Q CJ 2 B fe} = q x 7) ° Zz ie) > eS > a ° Z a K= > = er > 7) 1994 1995 1996 Fic. 11. Mean forewing length of males of H. erato phyllis in Morro do Voturud, August 1994 to September 1996 (based on monthly recruitment). Histogram bars = monthly means, line exten- sions = standard deviation. VOLUME 53, NUMBER 1 19 TABLE 5. Residence values (in days) for some butterfly species in Southeasterm Brazil. Mean mr = mean minimum residence, mo = maximum for one individual. Males Females mean mr mo mean mr mo Sao Vicente Region Nymphalidae Heliconius erato phyllis \ Morro do Voturua Heliconius erato phyllis ' Morro do Japui Melinaea ludovica paraiya Reakirt * Melinaea ethra Godart ? Placidula euryanassa ** Mechanitis lysimnia lysimnia Fabricius” Dircenna dero celtina Burmeister 2 Heterosais edessa Hewitson ” Hypothyris ninonia daeta (Boisduval) * Pseudoscada erruca (Hewitson) * Ithomia drymo Hiibner * Actinote pellenea pellenea Hiibner ° Actinote brylla Oberthiir ° Papilionidae Parides anchises nephalion Godart ° Other regions Nymphalidae Heliconius erato phyllis * Aeria olena olena Weymer * Actinote zikani D’ Almeida ® Pierella lamia Sulzer '° Pieridae Eurema elathea (Cramer)! dry season Eurema elathea™ wet season 37.6 + 25.8 127 22.6 + 23.0 89 18.04 + 14.6 47 -= 22 14.1+ 141 55 -- 13:7 + 12:9 AT — — 8.4 + 8.3 45 7.2 +5.7 23 15.5 + 15.4 67 18.3 + 16.2 72 9.4 + 6.9 23 10.9 + 10.5 49 12.9 + 15.1 65 13.5 + 10.3 39 16.3 + 11.1 40 14.2 + 16.2 52 9.9 + 9.6 25 — 33 7.0 + 7.3 17 12.3 + 12.1 26 3.2 + 2.6 12 — 6 — 16 — 16 14.1 + 8.2 30 9.0 + 3.6 12 30.7 + 29.0 112 — — 8.7 +5.9 24 11.7+2.5 14 3.9 + 1.3 7 — — as 60 ae _ 10.9 + 9.1 52 9.7+9.8 54 8.8 + 5.6 28 6.9 + 4.2 21 Superscript numbers : 1, this study; 2, Freitas 1996; 3, Freitas 1993; 4, Freitas, unpubl. data from Morro do Japuf; 5, Francini 1989; 6, Freitas, unpubl. data from Morro do Voturué; 7, Francini, unpubl. data from Lavras, MG, Brazil; 8, Vanini and Freitas, unpubl. data from Campinas, SP; 9, Francini, unpubl. data from Paranapiacaba, SP; 10, Freitas, unpubl. data from Sete Barras, SP; 11 Vanini, Bonato and Freitas, unpubl. results from Campinas, SP. Ecological Plasticity in Heliconius? Several features of H. erato vary in different parts of the neotropics. Although in the Sao Vicente region and Trinidad the populations are stable, maintaining con- stant low numbers throughout time (Turner 1971, R. B. Francini, pers. obs.; this study), in southern Brazil they show strong fluctuations over the year, with some extinction in colder years (Saalfeld & Araujo 1981, Ro- manowsky et al. 1985). Heliconius erato is reported as feeding on more than 37 species and 6 genera/subgen- era of Passifloraceae (Benson et al. 1976, Brown 1981, Spencer 1988), even using Passiflora edulis and P. alata, host species normally rejected in most popula- tions, on forest edges and urban areas (this study, A. V. L. Freitas, pers. obs., and unpublished results by L. S. Otero and K. S. Brown Jr.). The differences in use of flower resources among areas in this study shows that H. erato phyllis rapidly responds to variations in re- source availability in different sites and seasons. Heliconius erato, especially the subspecies phyllis, seems to be able to persist in many kinds of climate and vegetation. Colonies of this subspecies are present in virtually any kind of vegetation in southeastern and southern Brazil, including primary and secondary for- est edges, urban parks, plantations of Pinus and Euca- lyptus, riparian forests, savannas and sandy soil forests, and in tropical, subtropical, and temperate environ- ments (Araujo 1980 and pers. obs. of the authors). This is probably related to the ability of this species (espe- cially the subspecies phyllis) to use a great range of lar- val and adult resources, and change behavior and pref- erences according to the environment (=ecological plasticity). Such statements could apply not only to H. erato, but also to H. sara and H. ethilla populations in south- eastern and southern Brazil, both with similar general- ist habits, and could help to explain patterns of distri- bution of the species of this genus. In contrast to the three species cited above, four other Heliconius of southeastern Brazil have much more restricted habits. Heliconius melpomene nanna Stichel, H. numata robi- gus and H. nattereri Felder & Felder, are restricted to limited sectors of forest habitats, the last one with a 20 very small range, a poor competitor with H. sara and other lowland species (Brown 1972, W. W. Benson, pers. comm.); H. besckei Ménétriés (a co-mimic of H. erato) is typical of Atlantic forest mountains, in pri- mary and secondary forests at medium high altitudes, descending to sea level only in winter (Brown & Mielke 1972 and unpublished results by the authors). Even though environmental tolerance may help ex- plain distributions of the species of Heliconius, other factors need to be investigated. Benson (1978) argues that the availability of new host plant shoots in the dry winter results in high competition and is responsible for the absence of H. melpomene in a seasonal site in South Brazil. In the Sao Vicente region, however, this seasonality is much less pronounced, and periods of severe drought are virtually absent (Fig. 2); new shoots of Passiflora species seem to be abundant throughout the year, and though a detailed study needs to be carried out, this could not explain the absence of H. melpomene south of Rio de Janeiro State. Although these ideas and many others were discussed by Gilbert (1984), there still seem to be more questions than answers. Population studies need to be undertaken with other species of Heliconiini, especially the more widspread species of Heliconius, such as H. ethilla and H. sara. The presence of seasonal polyphenism may indicate species with broad tolerances to different environ- ments, as has been suggested for other polyphenic but- terflies (Shapiro 1976, Kingsolver & Wiernasz 1991, Van Dyck et al. 1997), although in Heliconius this could be apparently without adaptive consequences. These studies could guide future research in population biol- ogy of butterflies, and help in the understanding of the ecology of tropical insects. ACKNOWLEDGMENTS We thank R. B. Francini, K. S. Brown Jr., W. W. Benson, B. A. Drummond, D. Bowers, A. M. Araujo, A. Caldas, J. R. Trigo, L. E. Gilbert, J. Mallet, A. Gomes-Filho, G. Machado, M. R. Pie, G. W. Chaves and C. E. G. Patto for discussions and comments on the manuscript. R. B. Francini helped in field work and in population analysis. Climatological data were kindly provided by the Brazilian Air Force. A. V. L. Freitas thanks the CNPq for a fellowship. LITERATURE CITED Araujo, A.M. 1980. 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REITSMA & J. G. M. WILSON. 1994. Seasonal polyphenism in the wild: survey of wing pattern in five species of Bicyclus butterflies in Malawi. Ecol. Entomol. 19:285-298. Younc. A. M. & M. W. Morretr. 1979. Studies on the population biology of the tropical butterfly Mechanitis isthmia in Costa Rica. Amer. Midl. Nat. 101:309-319. Received for publication 15 February 1998; revised and accepted 2 December 1998. Journal of the Lepidopterists’ Society 53(1), 1999, 22-28 A HOSTPLANT-INDUCED LARVAL POLYPHENISM IN HYALOPHORA EURYALUS (SATURNIIDAE) MICHAEL M. CoL.Lins! Research Associate, Invertebrate Zoology, Camegie Museum of Natural History ABSTRACT. A hostplant-induced larval polyphenism is described in Hyalophora euryalus. Larvae reared on madrone (Arbutus menziesii) and manzanita (Arctostaphylos patula) have greatly reduced or no lateral nor abdominal scoli; sibs reared on the conifer Douglas-fir (Pseudot- suga menziesii) possess fully expressed scoli. Other native hosts (Ceanothus integerrimus and Prunus emarginata) do not induce the polyphenism. The third instar appears to be the critical stage during which the polyphenism is determined. Size and fecundity of adults reared on madrone and Douglas-fir are comparable. The evolutionary basis of this polyphenism is discussed in terms of increased crypsis on the ap- propriate host. Madrone may be an ancestral host, a member of the Madro-Tertiary flora with which H. ewryalus is closely associated. Douglas- fir may have been important during the re-invasion of northern and boreal regions following the Pleistocene. Mature larvae of the allied Cal- losamia also have a similar “nude” larval phenotype, suggesting a possible ancestral genetic potential to evolve the polyphenism. Additional key words: caterpillar, crypsis, developmental plasticity, life cycle, phenotypic plasticity, polymorphism. Although known as the “Ceanothus silk moth’, Hyalophora euryalus (Boisduval) is polyphagous and occupies an unusual range of West Coast plant com- munities including the deserts of Baja California, Coast Range and Sierran chaparral, Central Valley ri- parian habitat, Great Basin scrub, and conifer forests in the Sierra Nevada and the Cascades (Tuskes et al. 1996). In the central Sierra Nevada the larvae feed on at least eight genera of shrubs and trees, representing six plant families, including the conifer Douglas-fir (Pseudotsuga menziesii Mirb. |Franco]). By dispersing their populations over a range of plant communities, hostplant generalists may benefit from reduced search time for ovipositing females, and partially escape from predators and parasitoids associated with specific plant species or plant communities (Janzen 1984a). Never- theless, a large, palatable larva like H. euryalus must also depend on crypsis to survive. Its larval phenotype inevitably represents a compromise in camouflage value among such a wide variety of foliage shapes, col- ors, and lighting regimes. The host-induced larval polyphenism reported here appears to represent an evolutionary response to this dilemma. The larvae of H. euryalus differ from congeners in the last two instars in the tendency of scoli to be smaller relative to total larval size, and in being armed with fewer and smaller spines on their scoli (Fig. 1) (Collins 1997, Tuskes et al. 1996). This trend reaches its extreme expression in a “nude” fifth instar larval phe- notype in which all but the two pairs of dorsal thoracic scoli, the first dorsal abdominal pair, and the caudal scolus are very reduced or absent. I have observed this phenotype in approximately a third of wild collected larvae in the central Sierra Nevada, and have recorded it in populations from as far north as Victoria, British 'Mailing address: 11091 Miwok Path, Nevada City California 95959 Columbia and as far south as Baja California. No other Hyalophora taxon expresses this reduction in scoli. The environmental control of scoli expression was discovered fortuitously when I divided a batch of H. euryalus ova laid by a single female into two lots, one of which I reared on Arbutus menziesii Pursh (madrone; Ericaceae) and the other on Pseudotsuga menziesii (Douglas-fir; Pinaceae). Both are common hosts for H. euryalus throughout the Sierra Nevada and Cascade Range. All larvae on madrone developed into the nude morph (Fig. 2), while their sibs on Douglas-fir all expressed fully developed scoli (Fig. 3). In this paper I report the results of a controlled breed- ing program to verify these findings and to further in- vestigate the genetic basis for this host-induced poly- phenism. In the discussion I offer the interpretation that the nude larval morph is especially cryptic on madrone and the full expression of scoli is cryptic on Douglas-fir. MATERIALS AND METHODS Stock of Hyalophora euryalus from northern Cali- fornia was derived from a female collected as a cocoon near Donner Lake, Placer Co., and mated to a wild male from Nevada Co. Subsequent generations were produced by mating reared females to wild Nevada Co. males. One other brood resulted from mating a reared female from Victoria, British Columbia, Canada, stock to a wild male from Nevada Co., Cali- fornia. Ova were obtained by confining females in pa- per bags and cutting out sections with clusters of ova attached. Ova were incubated during May in a screened-in insectary at 1000 meters in Nevada Co., California. Newly eclosed larvae were confined with twigs of hostplant in 15 cm x 4 cm plastic petri dishes. After 2 to 3 days larvae were transferred to nylon mesh sleeves placed on Arbutus menziesii, Pseudotsuga menziesii, and other indigenous hosts. Larval growth VOLUME 53, NUMBER 1 23 Fic. 1. Hyalophora euryalus fifth instar reared on Ceanothus integerrimus; most common phenotype in mid-latitude California Sierra Nevada, with reduced, but entire, scoli. Fic. 2. Fifth instar H. ewryalus of the “nude” phenotype reared on Arbutus menziesii (madrone). Dorsal and lateral abdominal scoli reduced to near absence. FIG. 3. with fully developed scoli, induced by feeding on this host. Fic. 4. which induces the “nude” phenotype. Siblings of larva in Fig. 3. and phenotypes expressed were monitored periodi- cally under these natural conditions. Fifth instar larval phenotypes were scored as a presence or absence of abdominal scoli. For larvae reared on Arbutus or Arc- tostaphylos (manzanita) compared to Pseudotsuga this was virtually a qualitative trait, although some larvae expressed much reduced button-like scoli. Larval morph scores were analyzed using a Chi-square test, Fifth instar H. euryalus reared on Pseudotsuga menziesii (Douglas-fir) Fifth instar H. euryalus reared on Arctostaphylos patula (manzanita), assuming a null hypothesis that a simple polymor- phism existed based on a single major genetic locus, independent of hostplant. The possible effect of the maternal hostplant was tested by mating a female reared on Pseudotsuga and another on Arbutus each to wild males, and rearing larvae from both matings as split broods on both hosts as before. 24 TABLE 1. Effect of hostplant on expression of fifth instar larval scoli in Hyalophora euryalus. Brood Pseudotsuga Arbutus oxd scoli nude scoli nude Victoria, B.C. x Nevada Co., Calif. 1994 26 0 0 25 Donner L., Placer Co. x Nevada Co., Calif. 1996 23 0 ] 22 Sib above x Nevada Co. 1997a 7 0 0 y As above 1997b 9 0 0 13 Total: 65 0 1 63 Chi? for pooled data = 128.0 .0005 >> p. To assess the possible effect of hostplant species on reproductive fitness, fecundity of females was recorded for Pseudotsuga vs. Arbutus and compared to pub- lished data for other hosts. Since saturniid females eclose with a full complement of mature ova and oviposit virtually all their ova, the total number of eggs laid yields a direct index of fecundity when divided by forewing length to standardize for variation in adult size (Collins 1997). Unmated females were dissected to determine fecundity. In an attempt to determine the critical instar in which induction of larval morph occurs, I initially switched one batch of 20 third instar larvae from Ar- butus to Pseudotsuga. All larvae died within a few days of transfer, apparently from either refusing to feed on or an inability to metabolize this new host. This test was repeated in 1998 with lots of ten sibling larvae each reared in cages on either Pseudotsuga or Arbutus, then switched to the other hostplant of the pair during the third instar. A control lot was reared continuously on the common foothill host, Ceanothus integerrimus. In addition, a fourth lot was reared on Ceanothus until the third instar, and then switched to Pseudotsuga. In- dividuals from a different brood were reared on Pru- aus emarginata (Rosaceae). TABLE 2. Effect of maternal parental hostplant on induction of larval morph in Hyalophora euryalus. Siblings of 1997 broods in Table 1. Female parental host Pseudotsuga Arbutus Host scoli nude scoli nude Arbutus 0 13 0 2g Arctostaphylos 0 14 0 9 Pseudotsuga JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TABLE 3. Phenotype and survival of Hyalophora euryalus sibling larvae switched to new host as third instars. Fifth instar phenotype No. switched Number ———————_ as 3rd instar survived fullscoli intermediate* nude 1998a: ? progeny of 1997a x wild Nevada Co. d Host switch Arbutus - Pseudotsuga 4 4 4 0 0 Ceanothus - Pseudotsuga 5 4 2 il 1 Pseudotsuga - Arbutus 8 3 0 a 2 Ceanothus integerrimus (Control, not switched) 7 ~l (oe) S nS 1998b: 2 progeny 1997a x sib. Sleeved as ova on live hostplant, not switched. Prunus emarginata 2 9 13 Arctostaphylos patula 0 0 13 Pseudotsuga menzeisii 6 0 0 * Larvae with the intermediate phenotype possessed entire but reduced lat- eral scoli, and no or only remnant dorsal abdominal scoli. RESULTS Larval phenotypes for broods reared on Pseudotsuga vs. Arbutus are shown in Table 1. A nearly complete di- chotomy in phenotypes for both broods was seen in re- lation to hostplant, disproving the null hypothesis that the expression of scoli reduction is due to a simple ge- netic polymorphism independent of hostplant. During the first two rearing seasons, survival on both hostplants was nearly 100%. In 1997 larvae in some sleeves suffered heavy parasitism by the bra- conid Cotesia. Although exact data were not collected, I observed that larval growth rates on Arbutus were consistently faster in all broods than those for larvae reared on Pseudotsuga. Notes on approximate average time to cocoon spinning show that larvae on madrone matured about one week to 10 days faster than on Douglas-fir. No effect of maternal hostplant on larval phenotype was seen (Table 2), as larval morphs showed the ex- pected dichotomy regardless of the hostplant of the fe- male parent. In addition, larvae reared on Arc- tostaphylos completely expressed the nude phenotype in the last instar (Table 2, Fig. 4). Observation of larvae showed that the reduction of scoli was nearly as pro- nounced in the fourth instar on Arbutus and Arc- tostaphylos, although the majority possessed scoli as small knobs, especially the lateral rows. In all broods, third instars possessed fully developed scoli. However, of six third instar larvae reared in 1997 on Pseudo- tsuga, three had all scoli heavily pigmented with black, VOLUME 53, NUMBER 1 TaBLE 4. Effect of hostplant on fecundity. Index = no. ova/forewing length. No. ova Forewing length Index Arbutus California: Placer Co. x Nevada Co. 228 55 4.15 130 57 2.28 151 57 2.65 Canada, Victoria B.C. x California, Nevada Co. 143 57 Dell Avg. 163.0 56.5 2.90 Pseudotsuga California: Placer Co. x Nevada Co. 2QA47 60 4.12 192 62 3.10 154 5D 2.80 Canada, Victoria B.C. x California, Nevada Co. 178 57 Onl, 209 59 3.54 Avg. 196.0 58.6 3.34 Prunus emarginata California, Nevada Co.; avg. 12 pairings (Collins 1997) 175 58.3 2.99 Ceanothus integerrimus California, Nevada Co. 179 57 3.14 as is seen in Hyalophora columbia columbia and Hyalophora columbia gloveri, while the remaining three possessed yellow dorsal and light blue lateral scoli as is typical of H. euryalus (Tuskes et al. 1996). Of 15 third instars examined in madrone broods, none possessed black scoli, nor were such dark forms seen among madrone broods in previous seasons, although careful notes were not made previously on third instar coloration. In 1998 among third instar larvae on Pseudotsuga two possessed black lateral scoli with yel- ' low dorsal scoli, three had blue tipped with black lat- eral scoli and yellow dorsal scoli, none were all black, and two had the normal blue lateral and yellow dorsal scoli coloration. All ten third instar larvae on madrone possessed the yellow and blue pattern. The effects of host-switching on survival and larval phenotype are shown in Table 3. Initial losses from the 25 lots of 10 neonates were highest in the lot begun on Arbutus, due apparently to wandering off the host- plant. Only 2 larvae were lost before the third instar for those begun on Pseudotsuga. None of the controls on Ceanothus was lost. After host switching, the lot switched from Pseudotsuga to Arbutus suffered the greatest loss with only 3 of 8 surviving. Larvae switched from Arbutus to Pseudotsuga expressed fully developed scoli. However, larvae switched from Cean- othus to Pseudotsuga showed different phenotypes: one larva was intermediate with small but distinct lat- eral scoli and very reduced or absent dorsal scoli; two other fifth instar larvae had large scoli and one was of the nude phenotype. Of the larvae switched from Pseudotsuga to Arbutus two expressed the nude phe- notype but one other was intermediate. Control larvae reared continuously on Ceanothus displayed both morphs; some had fully developed scoli, while others had the nude morph. The brood reared on Prunus emarginata (Table 3) also expressed all three larval morphs in the final in- star, while siblings reared on Arctostaphylos and Pseudotsuga expressed the expected nude vs. scoli phenotypes. These results suggest that Prunus emar- ginata and Ceanothus integerrimus are neutral with respect to the host-induced polyphenism. The host switching experiments suggest that the third instar is the stage that is responsive to host cues, thus controlling final instar phenotype, because exposure during the first two instars can be counteracted by sub- sequent exposure to other hostplant taxa. Although I did not attempt to subject fourth instars to host switching, casual observations showed that the fourth instar phe- notype was usually an accurate predictor of final instar phenotype. Fourth instars with very reduced scoli often produced nude fifth instars. However, two fourth instars reared on Prunus and possessing fully developed scoli changed to the nude phenotype in the fifth instar. Sibling females reared on Pseudotsuga or Arbutus did not differ in fecundity (Table 4). Average number of ova laid, forewing length, and fecundity index were larger for those reared on Pseudotsuga, although the sample size was too small to justify calculating statisti- cal significance. Collins (1997) reported an average and SD for these parameters, respectively, of 175.5 + 36.8, 58.3 + 4.6, 2.99 + 0.48 for a sample of 12 H. eu- ryalus reared on Prunus emarginata. DISCUSSION In discussing the evolutionary significance of larval phenotypic plasticity in H. euryalus, it is important to distinguish between the terms polymorphism and poly- phenism. In a polymorphism, genetic differences among 26 individuals produce discrete phenotypes. A polymor- phism is a population phenomenon; the frequency of al- termate morphs in the population reflects the frequency of those genes controlling the expression of each morph, and the phenotype of a given individual is de- pendent on its genotype. A polyphenism is the expres- sion of a specific phenotype in response to environ- mental cues, which regulate gene action through a neural-molecular pathway. (“Phenotypic plasticity” is also a widely used term (Stearns 1989), although Williams (1992) objects to its non-genetic connotation). Every individual in a polyphenic population theoreti- cally could be genetically identical for the loci in ques- tion, and phenotypic variation within the population then would be a consequence of individual exposure to variable environmental cues. In a seasonal polyphenism, immatures in the population respond to a reliable sea- sonal cue, such as photoperiod, to produce “spring” and “summer” adult phenotypes in pierids (Shapiro 1989) or in the saturniid genus Actias (Miyata 1974, 1986); or the “wet” and “dry” seasonal forms of the neo- tropical saturniid Rothschildia lebeau (Janzen 1984b), and in the African butterfly Bicyclus (Windig et al. 1994). Fewer cases of larval polyphenism in Lepidoptera have been carefully documented. Greene (1989) showed that strikingly different cryptic phenotypes are produced in the geometrid Nemoris when larvae feed on oak flower buds in the spring versus leaves during the summer rainy season in southeast Arizona. Fink (1995) demonstrated that hostplant partly controls a color polyphenism in Eumorpha (Sphingidae) larvae, but was unable to do controlled breeding experiments due to the difficulty of pairing these moths in the lab. The number and spacing of the large, silvery, lancet- shaped scoli of certain Southwestern Sphingicampa (Saturniidae) appear to be influenced by hostplant leaflet size and number (Tuskes et al. 1996; P. Tuskes, pers. comm.). Plant secondary compounds may act as cues in controlling these polyphenisms, although a pu- pal color polyphenism in certain Papilio is controlled by light level and other environmental cues (West 1995, Sims & Shapiro 1983). My interpretation of the nude larval morph in H. euryalus is that this phenotype is more cryptic on madrone than the morph with fully developed scoli. The leaves of madrone are large, typically 10-14 cm in length, with entire margins, and light grey-green be- low. Especially when viewed from below, even the ma- ture larva of H. euryalus is inconspicuous in the nude morph as it rests or feeds underneath the large, simi- larly colored madrone leaf. The same argument can be applied to the association with Arctostaphylos; the leaves of most species are also glaucus with smooth, JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY entire margins, but smaller than the foliage of Arbu- tus. By contrast, the larval morph with fully developed scoli appears more cryptic against the foliage of Doug- las-fir because the numerous scoli break up the solid mass and match the visual effect of spots of light shin- ing through a matrix of small needles. In both cases, the cryptic appearance is lost when each morph is viewed against the foliage of the “inappropriate” host- plant. The trend in early instars of darkening of scoli seen in broods reared on Douglas-fir would seem to camouflage these larvae against the dark twigs and fo- liage of Douglas-fir. The early instars of the Larix feeder, H. c. columbia, are always black. The larvae of the related Callosamia also have very reduced scoli in later instars (Tuskes et al. 1996). This condition is most pronounced in the Magnoliaceae specialists, C. angulifera and C. securifera, whose lar- vae are very cryptic hidden under the large leaves of their hostplants. The foliage of Pseudotsuga, in common with other conifers, is rich in terpenes and so presents a meta- bolic barrier against some insect herbivores (Smith 1989, Gershenzon & Croteau 1991, Harborne 1997). In a study of Lepidoptera diversity associated with Abies and Pseudotsuga, Powell and De Benedictis (1995) list 40 species, of which 73% are conifer spe- cialists. Conifer feeding is not common within the three North American saturniid subfamilies. The few known examples, with the exception of Hyalophora euryalus, are generally feeders on Pinus spp., whose congeners feed on leafy shrubs and trees (Lemaire 1988, Tuskes et al. 1996, Wolfe 1993). Hyalophora eu- ryalus is exceptional because it is primarily a general- ist on shrubs and trees, and also because it was the only species found on Douglas-fir that preferred older needles (Powell & De Benedictis 1995). My work con- firms this preference. First year conifer needles have been shown to contain up to ten times the diterpene acids of older needles (Ohigashi et al. 1981), so feed- ing on older needles may be due to an avoidance of high levels of these diterpene acids. Adaptation to a new host involves many complex fit- ness tradeoffs affecting the evolution of life history traits (Fox & Morrow 1981; Krainacker et al. 1987; Fox & Caldwell 1994; Leclaire & Brandl 1994). Al- though it is difficult to measure the metabolic and other “costs” of conifer feeding in H. euryalus, larvae on Pseudotsuga consistently matured more slowly than sibs reared on Arbutus and one larval brood begun on Arbutus could not switch to Pseudotsuga in the third instar. It is not known if this result was due to a refusal to initiate feeding, or perhaps due to the failure of in- duction of the synthesis of a critical enzyme (cf VOLUME 53, NUMBER 1 Brattsten et al. 1977, Brattsten 1983; Moldenke et al. 1983). No reduction in adult size nor female fecundity was found in broods reared on Pseudotsuga. The foliage of Arbutus menziesii may also contain secondary compounds exacting a metabolic cost from insect herbivores. Ezcurra et al. (1987) list cardiac gly- cosides, quinones, and tannins among others in leaves of the Mexican madrone species, Arbutus xalapensis. Larvae of H. euryalus switched from Douglas-fir to madrone were reluctant to accept madrone (Table 3). Larvae feeding in the Douglas-fir canopy may par- tially escape those predators and parasites normally as- sociated with the smaller trees and shrubs that serve as Hyalophora hosts. The majority of the Lepidoptera fauna feeding on Douglas-fir are microlepidoptera (Powell & De Benedictis 1995), which could attract a different set of predators and parasites than those at- tacking Hyalophora. Danks (1994) asserts that genetic polymorphisms tend to evolve in predictable environments, while polyphenisms are associated with unpredictable envi- ronments. An example of a polymorphism in a pre- dictable environment would be the various color forms of Saturnia mendocino larvae, which match the living green foliage and persistent yellow and mauve dead leaves all reliably present on manzanita (Tuskes et al. 1996). The array of hosts utilized by populations of H. euryalus represent an unpredictable environment. One can theorize that if a H. euryalus population were genetically polymorphic for larval scoli size, females ovipositing on both madrone and Douglas-fir would produce many larvae of the inappropriate phenotype, stranded on large trees against a non-cryptic back- ground. Collins (1997) proposes that conifer feeding in Hyalophora arose as an adaptation to post-Pleistocene environments, in which pioneer populations of H. eu- ryalus fed on Pseudotsuga as they reinvaded northern and montane portions of the moth’s current range, and H. columbia columbia similarly adapted to Larix (tamarack) as it spread north and east where it cur- rently occurs in tamarack bogs. Even coastal popula- tions of H. euryalus in southern California may accept and mature on Pseudotsuga, but the Rocky Mountain subspecies H. c. gloveri lacks this adaptation (Tuskes et al. 1996). Madrone and manzanita are members of the Madro-Tertiary flora, and probably represent ancestral Hyalophora hosts, based on comparative biogeograph- ical evidence and the close association of H. euryalus with modern derivatives of this ancient flora (Collins 1997). Fully developed fifth instar scoli appears to be the primitive condition in the Saturniidae (Ferguson 1971, 2 Minet 1994), and fifth instar scoli are prominently ex- pressed in all Hyalophora taxa except H. euryalus. The neutral condition in H. euryalus is one of variable but reduced scoli in later instars. The evolution of the lar- val polyphenism in H. euryalus may best be described as the host-induced shift in this neutral developmental state toward either suppression of scoli when larvae feed on certain Ericaceae, or a shift in the opposite di- rection toward full scoli expression in larvae feeding on Pseudotsuga. Certain other hosts, such as Ceanothus integerrimus and Prunus emarginata, appear neutral in inducing the scoli polyphenism and a range of de- velopmental variation in scoli expression occurs among siblings reared on these hosts. It is difficult to deter- mine to what extent this represents genetic variation in loci controlling scoli development, given the environ- mental influence seen in this study, but I have ob- served larvae with fully developed scoli on Arctosta- phylos (fig. 1d in Collins 1997). ACKNOWLEDGMENTS Valerie Passoa, Richard Peigler, David West, and an anonymous reviewer read the entire manuscript and offered valuable com- ments. Deane Bowers, Francie Chew, and Art Shapiro kindly sug- gested specific phytochemistry references. LITERATURE CITED BRATTSTEN, L. B. 1983. Cytochrome P-450 involvement in the in- teractions between plant terpenes and insect herbivores, pp. 174-195. In P. A. Hedin (ed.), Plant resistance to insects. Amer. Chem. Soc. Symposium ser. 208. BRATTSTEN, L. B., C. F. WILKINSON & T. EISNER. 1977. Herbivore- plant interactions: mixed function oxidases and secondary plant substances. Science 196:1349-1352. CoLLins, M.M. 1997. Hybridization and speciation in Hyalophora (Insecta: Lepidoptera: Saturniidae): a reappraisal of W. R. Sweadner’s classic study of a hybrid zone. Ann. Carnegie Mus. 66:411—455. Danks, H. V. 1994. Insect life-cycle polymorphism: current ideas and future prospects, pp. 349-365. In H. V. Danks (ed.), Insect life cycle polymorphism. Kluwer Academic Publ. Ezcurra, E., J. C. GOMEEZ & J. BECERRA. 1987. Diverging pat- terns of host use by phytophagous insects in relation to leaf pu- bescence in Arbutus xalapensis (Ericaceae). Oecologia 72:479—480. FERGUSON, D. C. 1971. Bombycoidea (in part), Fasc. 20.2A, pp. 101-153. In R. B. Dominick et al. (eds.), The moths of North America north of Mexico. Classey, London. Fink, L. S. 1995. 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Two ways to be a big tropical moth: Santa Rosa saturmiids and sphingids. Oxford Surv. Evol. Biol. 1:85-140. . 1984b. Weather-related color polymorphism of Roth- schildia lebeau (Saturniidae). Bull. Entomol. Soc. Am. 30:16—20. KRAINACKER, D. A., J. R. CAREY & R. I. VARGAS. 1987. Effect of lar- val host on life history traits of the Mediterranean fruit fly, Cer- atitis capitata. Oecologia 73:583—590. LECLAIRE, M. & R. BRANDL. 1994. Phenotypic plasticity and nutri- tion in a phytophagous insect: consequences of colonizing a new host. Oecologia 100:379—385. LEMAIRE, C. 1988. Les Saturniidae américains. Ceratocampinae. Museo Nacional de Costa Rica, San José. Miner, J. 1994. The Bombycoidea: phylogeny and higher classifi- cation (Lepidoptera: Glossata). Ent. Scand. 25:63—88. Miyata, T. 1974. Studies on diapause in Actias moths (Lepi- doptera:Saturniidae). 1. Photoperiod induction and termina- tion. Kontyti 42:51-63. . 1986. Studies on diapause on Actias moths (Lepidoptera: Saturniidae). V. 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Experimental studies on the evolution of sea- sonal polyphenism, pp. 297-307. In R. I. Vane-Wright & P. R. Ackery (eds.), The biology of butterflies. Princeton University Press. Sims, S. R. & A. M. SHapiro. 1983. Pupal colour dimorphism in California Battus philenor: pupation sites, environmental con- trol, and diapause linkage. Ecol. Entomol. 8:95-104. SmitH, C, M. 1989. Plant resistance to insects: a fundamental ap- proach. Wiley & Sons, New York. STEARNS, S. C. 1989. The evolutionary significance of phenotypic plasticity. Bioscience 39:436—445. TusKEs, P. M., J. P. TurrLe & M. M. Couns. 1996. The wild silk moths of North America. Cornell University Press, Ithaca. West, D. A. 1995. Comparative pupation behavior in the Papilio glaucus group: Studies of Papilio glaucus, Papilio ewrymedon and their hybrids (Lepidoptera: Papilionidae), pp. 93-99. In J. M. Scriber, Y. Tsubaki & R. C. Lederhouse (eds.), Swallowtail butterflies: their ecology and evolutionary biology. Scientific Publishers, Gainesville. WiLuiAMs, G.C. 1992. Natural selection: domains, levels, and chal- lenges. Oxford University Press, New York. WINDIG, J. J., P. M. BRAKEFIELD, N. REITSMA & J. G. M. WILSON. 1994. Seasonal polyphenism in the wild: survey of wing pat- terns in five species of Bicyclus butterflies in Malawi. Ecol. En- tomol. 19:285—298. Wo LF, K L. 1993. The Copaxa of Mexico and their immature stages (Lepidoptera:Satumniidae). Trop. Lepid. 4:Suppl. 1. Received for publication 8 April 1998; revised and accepted 3 De- cember, 1998. Journal of the Lepidopterists’ Society 53(1), 1999, 29-31 A NEW HYPATOPA FROM COSTA RICA (GELECHIOIDEA: COLEOPHORIDAE: BLASTOBASINAE) Davip ADAMSKI Department of Entomology, NHB-127, Smithsonian Institution, Washington, D.C. 20560 ABSTRACT. Hypatopa tapadulcea is described from northwestern Costa Rica. A photograph of the imago and illustrations of wing vena- tion and male and female genitalia are provided. Additional key words: Lepidoptera, Blastobasini, Guanacaste, Puntarenas. About ten years ago a survey of all Costa Rican fauna and flora was undertaken by Instituto Nacional de Biodiversidad (INBio). Among the many organisms collected were microlepidoptera, and through the ef- forts of Costa Rican “parataxonomists”, a large number of specimens of Blastobasinae have accumulated. Through the combined work of many Costa Ricans and invited scientists, an inventory for this neotropical region is possible. This paper describes Hypatopa tapadulcea, and represents one small work among sev- eral major studies planned by the author to make known the rich diversity of this group of moths. Members of the Blastobasinae are generally small to medium-sized, drab moths with fewer than 150 spe- cies described worldwide. This number, however, greatly underestimates the species richness, as there are hundreds of undescribed species represented in collections, especially from the Neotropics. Meyrick (1894) was the first to recognize the Blasto- basinae as a monophyletic group. Recent studies by Adamski and Brown (1989) and Hodges (1998) have corroborated this notion, and have established mono- phyletic groupings at the generic and familial levels within the Blastobasinae and Gelechioidea, respec- tively. Kornerup and Wanscher (1978) is used as a color standard for the description of the adult vestiture. Genitalia were dissected as described by Clarke (1941), except mercurochrome and chlorazol black were used as stains. Pinned specimens and genital preparations were examined with dissecting and com- pound microscopes. Measurements of wings and gen- italia were made using a calibrated ocular micrometer. Hypatopa tapadulcea Adamski, new species (Figs. 1-4) Diagnosis. Male with inner margin of gnathos slightly widened, forming a small broadened lobe, sac- culus subrectangular, aedeagus bulbous at base; fe- male with ostium elongate, ductus seminalis wide at base, ductus bursae spiralled. Description. Head: vertex and frontoclypeus with grayish-brown scales tipped with yellowish brown, fe- males mostly pale yellow brown; outer surface of labial palpus with grayish-brown scales tipped with yellowish brown intermixed with grayish-brown scales, and gray- ish-brown scales tipped with pale grayish brown, and yellowish-brown scales, inner surface mostly with yel- lowish-brown scales intermixed with few grayish- brown scales; some males with inner surface patterned similar to outer surface; females with mostly yellowish- brown scales on both surfaces; antennal scape and pedicel patterned as above; flagellum mostly brownish gray intermixed with few yellowish-brown scales; pro- boscis pale yellowish brown. Thorax: basal area of tegula and mesoscutum grayish brown, pale grayish brown distally. Legs with outer surface mostly brown intermixed with few yellowish-brown scales, yellowish- brown bands near midtibia, apices of femur, tibia, and tarsomeres, undersurface with mostly pale yellowish- brown scales intermixed with few brown scales. Forewing (Fig. 1): length 5.1-6.5 mm (n = 77), ground color yellowish brown intermixed with few brown scales; base posterior to CuP with a short dark-brown marginal streak; median fascia present, absent, or in- complete, usually distal 2/3 of wing darker than basal 1/3; discal spots absent; several similar-sized marginal brown spots on subapical and apical areas forming an irregular pattern; undersurface brown. Female paler than male. Venation (Fig. 2), cubitus 4-branched with M, and M, separate from M,, CuA, about 45 degree anes to mesic distal part of M, near parallel with M, and M,. Hindwing: brownish ty females paler. Ve- nation (Fig. 2), cubitus appearing 4-branched with CuA, and M, stalked beyond base of M,. Abdomen: Hrownich anny above, pale yellowish brown beneath, male with pale yellowish-brown scales on digitate pro- cess of upper part of valva, female pale yellowish brown beneath. Male genitalia (Fig. 3): uncus slightly broadened at base, narrowed distally, apical part hooked posteriorly, apex rounded and sparsely setose; gnathos medially narrowed, medially widened, form- ing a small broadened lobe; tergal setae present; vin- 30 Fic. 1. Holotype of Hypatopa tapadulcea Adamski. culum a thin band; juxta entire, not divided; lower part of valva with marginal setae, distal part setose, tapered to a pointed apex; upper part of valva with a subrect- angular sacculus; sacculus with mostly stout setae in- termixed with hairlike setae, a small cluster of mi- crotrichia near outer margin at base of long, setose digitate process; aedeagus bulbous at base, parallel sided to apex; anellus setose. Female genitalia (Fig. 4): ovipositor telescopic, in four membranous subdivi- sions; ostium elongate, delimited within membrane Fics. 2-3. Wing venation and male genitalia of Hypatopa tapadulcea Adamski. 2, Wing venation. Scale line = 1.0 mm. 3, Male genitalia. Genital capsule is figured above, aedeagus below. Scale line = 0.5 mm. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fic. 4. Female genitalia of Hypatopa tapadulcea Adamski. Scale line = 1.0 mm. near posterior margin of seventh sternum; antrum membranous, short, forming a common inception with ductus seminalis and ductus bursae; basal part of duc- tus seminalis as wide as posterior part of ductus bur- sae, ductus bursae spiralled from anterior end of antrum to corpus bursae; corpus bursae with a long hornlike signum. Types. Holotype: 4, “Est[acion] Pitilla, 700 ml[eters], 9 klilo]m[eters] S[outh] Sta[cion] Cecilia, PN. Gua- nacaste, Proviincia] Guan[acaste], COSTA RICA, C. Moraga, Se[p]t[iembre] 1991, L-N-330200, 380200”, “COSTA RICA, INBio, CR1000, 460377” [Bar code label]. Holotype is not dissected and is deposited in VOLUME 53, NUMBER 1 the Entomology Museum at Instituto Nacional de Bio- diversidad, Santo Domingo, Heredia, Costa Rica. Paratypes: 75 paratypes: 12 3d, 21 92, “Est Cacao, 1000-1400 m, Lado SO Vol Cacao, P.N.G. Prov Guan, COSTA RICA, C. Chaves, Abr 1991, L-N-323300, 375700”, two dissected males with the following label data, “INBio, Genitalia Slide by D. Adamski, No. 97, Sex 3”, “INBio, Genitalia Slide by D. Adamski, No. 98, Sex 6”, four dissected females with the following label data, “INBio, Genitalia Slide by D. Adamski, No. 99, Sex 2”, “INBio, Genitalia Slide by D. Adamski, No. 134, Sex 2”, “INBio, Genitalia Slide by D. Adamski, No. 135, Sex 2”, “INBio, Wing Slide by D. Adamski, No. 136, Sex 2”; 3 dd, 9 92, same label data as above ex- cept, “Set”; 6 29, same label data as above except, “25 Set—11 Oct 1990”; 2 92, same data as above except, “11 Set-11 Oct 1991”; 1 d, 2 92, same data as above except, “93 Oct—9 Nov 1990”; 1 d, same label data as above ex- cept, “Mar 1991”; “Est Pitilla, 700 m, 9 km S Sta Ce- cilia, P.N. Guanacaste, Prov Guanacaste, COSTA RICA, C. Moraga, Set 1991, L-N-330200, 380200”, “INBio, Wing Slide by D. Adamski, No. 137, Sex 2’, 5 22, same label data as above except, “Abr 1991”; 2 29, same label data as above except, “Set 1991”, one fe- male dissected, and with the following label data, “IN- Bio, Genitalia Slide by D. Adamski, No. 100, Sex 2”; 3 2, same label data as above except, “2-19 Mar 1992”, two specimens collected by P. Rios; “2 9°, same data as above except, “31-Mar-15 Abr 1992, P. Rios”; 1 4, same data as above except, “23 Oct-12 Nov 1992, C. Cano’; 1 d, 4 9°, “San Luis, Monteverde, Prov Punta, COSTA RICA, 1000-1350 m, Abr 1994, Z. Fuentes, L-N-449250-250850, #2845”. Two paratypes deposited in The Natural History Museum, London, England; ten paratypes in The National Museum of Natural History, Smithsonian Institution, Washington, D.C.; all other paratypes in Instituto Nacional de Biodiversidad, Santo Domingo, Costa Rica. Remarks. Hypatopa tapadulcea is probably more closely related to H. interpunctella (Dietz) of Utah, than other known species in the genus. This is based primarily on the similarity of the male genitalia be- cause the female genitalia are structurally very differ- ent and do not support close kinship between the two species. Male tapadulcea and interpunctella share a _ similarly shaped uncus and gnathos, however, in inter- punctella, the gnathos is notched. Both have the sac- culus longer than wide; in H. tapadulcea the distal 31 margin is obtuse while in H. interpunctella the outer margin is subtriangular. Hypatopa tapadulcea ranges from the high altitudes of the northwestern provinces along the Cordillera de Guanacaste southeast to the Cordillera de Tilaran in the Province of Puntarenas. The host for H. tapadul- cea is unknown. Etymology. The species epithet is derived from a brownish (similar in color to H. tapadulcea) crystalline, confectionary cake, tapa dulce, that is sold in market places throughout Costa Rica. This sweet is made from sugar cane and is broken into pieces and eaten like candy, or it is mixed with hot or cold water to make agua dulce. The costarriquenos consider agua dulce a national beverage. Because I have examined the type specimens of all the New World Hypatopa, and hundreds of specimens of New World Hypatopa representing at least 100 un- described species, I am confident that Hypatopa tapadulcea is the closest known relative of H. inter- punctella. Many, if not most, undescribed species of Blastobasinae from the New World belong to Hy- patopa, and with the large amount of museum speci- mens yet to be examined, the goal of hypothesizing nearest kin and phylogenetic relationships among all these taxa becomes problematic without a comprehen- sive alpha-taxonomic study. ACKNOWLEDGMENTS I thank Mario Camacho, Jessica Zamora Gonzalez, and Eugenie Phillips Rodriguez of The National Biodiversity Inventory Division, Instituto Nacional de Biodiversidad, Santo Domingo, Heredia, Costa Rica, for their help, cooperation, and warm hospitality during my visits to INBio; Carl Hansen of the Office of Imaging, Printing and Photographic Services for the photograph of the holotype of Hypatopa tapadulcea. LITERATURE CITED ADAMSKI, D. & R. L. BRowN. 1989. Morphology and systematics of North American Blastobasidae (Lepidoptera: Gelechioidea). Miss. Agr. For. Exp. Sta. Tech. Bull. 165 (Miss. Entomol. Mus No. 1):1-70. CLARKE, J. F. G. 1941. The preparation of slides of the genitalia of Lepidoptera. Bull. Brooklyn Entomol. Soc. 36:149-161. Hopces, R. W. 1998. Gelechioidea. In N. P. Kristensen (ed.), Handbuch der Zool. 494 pp. KORNERUP, A. & J. H. WANSCHER. 1978. Methuen handbook of colour. 2nd ed. Methuen and Co., Ltd., London. 243 pp. Meyrick, E. 1894. On a collection of Lepidoptera from upper Burma. Trans. Entomol. Soc. London 1894:1—29. Received for publication 20 June 1998 ; revised and accepted 2 Feb- ruary 1999. Journal of the Lepidopterists’ Society 53(1), 1999, 32-36 POPULATION DEMOGRAPHICS AND THE CONSERVATION STATUS OF THE UNCOMPAHGRE FRITILLARY BOLORIA ACROCNEMA (NYMPHALIDAE) Amy L. SEIDL Department of Biology, University of Vermont, Burlington, Vermont 04505, USA. ABSTRACT. Boloria acrocnema is an endangered relict arctic butterfly restricted to fewer than ten colony sites in southwestern Colorado, USA. Pollard transects were conducted from 1990-94 to establish relative population estimates for the butterfly. Results indicate a period of stability and increase for all three colony sites surveyed, and are a contrast to population estimates recorded in the 1980's. Collecting pressure, livestock grazing and local climate change are discussed as potential factors behind the butterfly’s original decline and its more recent stabiliza- tion and increase. Additional key words: alpine habitat, climate change, collecting activity, Pollard transect, Salicaceae. Boloria acrocnema Gall & Sperling (Nymphalidae) was discovered on Mt. Uncompahgre in the San Juan Mountains, Hinsdale County, Colorado, and subse- quently described by Gall and Sperling (1980) as a new species. The taxonomic relationship of Boloria acrocnema to Boloria improba (Butler) has been re- viewed (Gall & Sperling 1980, Britten & Brussard 1992) and some authorities recognize B. acrocnema as a valid species (Ferris 1981, Pyle 1981), whereas oth- ers consider it a subspecies of the more northern B. improba (Scott 1986). Since 1978, when B. acrocnema was discovered, its demography (Brussard & Britten 1989, Seidl 1995, Ellingson et al. 1996, Wasinger et al. 1997), genetic variation (Britten 1991, Britten & Brussard 1992), and adult and larval life history have been studied (Gall & Sperling 1980, Scott 1982, Gall 1984a, Britten & Riley 1994, Seidl 1995, 1996). Researchers found the spe- cies to be in decline in the early 1980’s and it was listed as Federally endangered in June 1991 (Opler 1990). Hypotheses for the butterfly’s decline include: exten- sive adult collecting pressure, disruption of larval mi- crohabitat by livestock grazing, and prolonged local drought conditions (Gall 1984a, 1984b, Brussard & Britten 1989, Britten 1991, Britten et al. 1994, Seidl & Opler 1994). The intent of this paper is to summarize and extend data on population size in three popula- tions of B. acrocnema located in the San Juan Moun- tains in southwest Colorado. Based on these data, I also evaluate the conservation status of the butterfly. MATERIALS AND METHODS Study Organism. Like some other alpine butter- flies, Boloria acrocnema is thought to have a biennial life cycle (but see Seidl 1996): each cohort overwinters twice and development occurs over three summers, thus creating odd- and even-year broods (Scott 1982, Brussard & Britten 1989). Eggs oviposited the first summer develop into first instar larvae that same sum- E-mail: aseidl@zoo.uvm.edu mer and overwinter. The following summer, develop- ment through third instar occurs. The third summer, larvae develop through fifth instar, pupate and eclose as adults, which live for approximately seven to nine days (Scott 1982). Britten and Brussard (1992) found no significant genetic differences between the two broods at two colony sites. Population estimates have therefore been characterized as either odd- or even- year broods (Gall 1984a, Brussard & Britten 1989). Boloria acrocnema specializes on snow willow, Salix reticulata L. ssp. nivalis (Hooker) Léve et al. (Sali- caceae), a common plant restricted to mesic alpine ar- eas in Colorado (Weber 1987). Females exhibit ovipo- sition preference for snow willow (Seidl 1996) and larvae followed in the field feed exclusively on it (Seidl 1995). However, in captivity B. acrocnema larvae will feed on Salix arctica (Scott 1982), a close relative to S. reticulata nivalis, despite no evidence of the use of this plant having been documented in the field. Study Sites. I conducted population demographic research from 1990—94 at three B. acrocnema colony sites: (1) a colony at Mt. Uncompahgre, the type local- ity (UP1); (2) a colony within four km of the type lo- cality (UP6); and (3) a colony on Redcloud Peak, ca. 20 km southeast of the type locality (RC1). All colonies are found in glacial cirques with a northeast exposure and range in altitude from 3800-4093 m, with UP1 being the highest in elevation and UP6 the lowest. Each colony site is located in alpine tundra habitat and each contains abundant patches of the butterfly’s wil- low host, S. reticulata nivalis. The area encompassed by each of the sites differs—RC1 is ca. 15 ha, UP1 is ca. 20 ha, and UP6 is less than 5 ha. All three study sites occur within public lands: UP6 and UP! are lo- cated in the Big Blue Wilderness Area of the Uncom- pahgre National Forest, and RC1 is within Bureau of Land Management lands. Hiking trails to both peaks pass through B. acrocnema habitat. Population Abundance. The Pollard transect tech- nique, a simple-to-use, non-intrusive method which identifies trends in butterfly abundance (Pollard 1977, VOLUME 53, NUMBER 1 33 TABLE 1. Relative population estimates and indexes of abundance for populations of Boloria acrocnema 1978-1996, Hinsdale County, Col- orado. Prior to 1988, capture-mark-recapture methods were used to estimate population size. Beginning in 1988, Pollard transect methods were used to derive an index of abundance from which a relative population estimate was determined. Colony Year RC1 UPI 1978 NA NA 1979 NA 6003 1980 NA 700 1982 1000 500 1987 250-300 4 1988 492 200. 1990 768 NA 1991 1624 0 1992 948 878 1993 2408 452 1994 3464 2918 1995 11773 6682 1996 3670 2163 UP6 Some NA Gall 1984a NA Gall and Sperling 1980 NA Gall 1984a NA U.S. Forest Service Files 3 Brussard and Britten 1989 2} Brussard and Britten 1989 NA this paper 20 this paper 414 this paper 2266 this paper 442 this paper 1979 Ellingson et al. 1995 130-200 Wasinger et al. 1997 1 1990-1993 estimates were originally derived without intercalculating missing days during the flight season (Seidl & Opler 1994). The data reported here have been reanalyzed such that missing days are calculated as the average of the previous day and the two following days (Cook et al. 1967, Watt et al. 1977). 2 Gall and Sperling (1980) state that several hundred individuals were seen on 30 July, 1978 but a population estimate for the brood was not calculated. 5 Gall (1984b) estimated 150-180 individuals at peak flight in 1979 at UP1. In 1980 peak flight estimates of 200 individuals were estimated and found to be asso- ciated with a final population estimate of ca. 700, i.e., 3.5 days at peak flight. With this in mind, I have tabulated an approximate population estimation for UP1 in 1979 to be ca. 600, i.e., 3.5 days * 165 individuals. + During the 1990 flight season I visited UP1 twice and UP6 a single time. These visits do not constitute a thorough search and therefore are specified as NA. Thomas 1983, Pollard & Yates 1993), was adopted in 1987 to assess trends in B. acrocnema abundance (Brussard & Britten 1989). This method is employed by the British Butterfly Monitoring Scheme (Pollard & Yates 1993) and enables researchers to assess popula- tion abundance with less intrusive sampling. The main criticism of the Pollard monitoring method is that ab- solute population estimates cannot be derived from the data since only trends in abundance are recorded and not the fate of individuals per se, as in capture-mark-re- capture methods (CMR). However, over a long-term monitoring period, the Pollard method can be equally as effective in assessing the condition of butterfly pop- ulations (Pollard & Yates 1993). The method includes laying fixed transect lines in a monitoring area that includes both suitable and less suitable butterfly areas (Pollard & Yates 1993). Since I was concerned that fixed transect lines would disturb B. acrocnema habitat, I set new transect lines each field season, placing 100 m long transects in areas of high butterfly density. Transects were walked in the morning and only under favorable conditions: low _ wind, relatively full sun, and no precipitation. If condi- tions turned unfavorable, counts were halted to allow poor weather to pass. Transects were walked at ap- proximately 10 min per 100 m transect. Individuals were included in the count if they occurred within five meters on either side of the transect line. Pollard transects were carried out at the RC1 colony site during the field seasons of 1990 through 1994 and at the UP1 and UP6 colony sites in 1991 through 1994. Searches for B. acrocnema were begun each year in mid to late June. Once butterflies were found, tran- sects were laid and counts were begun within three to five days. Three transects were laid each year at UP6, three to four transects were laid at UP1 each year, and five to eight transects were laid at RC1 each year. The number of transects laid at each site and each year var- ied with the number of high density butterfly areas found. If a day was missed during the flight season, in- terim day counts were estimated. These values were derived as the average of the preceding day and two days following the missing transect count after Cook et al. (1967) and Watt et al. (1977). To determine the relationship between Pollard tran- sect indices of abundance and population estimates, and to derive a relative population estimate for com- parison among years, CMR experiments were con- ducted simultaneously with Pollard transects in 1987 at RC1 (Brussard & Britten 1989). Pollard transect data were found to account for approximately 10% of the population estimates derived from CMR methods (Brussard & Britten 1989). Based on these calculations, each daily transect count (ti) was multiplied by 10 to derive the relative daily estimate (Ni) for a colony. In addition, Gall (1984b) derived a residency rate for B. acrocnema. A residency rate (r) is defined as the fraction of animals on day i that remain in the popula- tion on day i+1 and calculates the probability that an individual counted on day i is recounted the following 34 —fl___ odd-year brood —t— _even-year brood Relative population estimate 1980 1982 1984 1986 1988 1990 1992 1994 1996 Year Fic. 1. Relative population estimates for Boloria acrocnema at Redcloud Peak, colony site RCI, based on Pollard transect method. day (Gall 1984b). Residency rates for B. acrocnema at UP1 in 1980 were calculated as 0.46—0.70 (Gall 1984b). Therefore, transect counts in this study, and those in Brussard and Britten (1989), were multiplied by the average daily loss rate (1-r) or 0.4. Total population indices were summed across days to arrive at a relative population estimate for a colony for a given year. RESULTS The even-year population at RC1, initially estimated at maximally 1000 individuals (Gall 1984a), had de- clined to a count of 492 in 1988 (Brussard & Britten 1989). During this study however, the even-year popu- lation increased significantly and relative population estimates for the even-year population include: 768 in 1990, 948 in 1992, and 3464 in 1994 (Table 1, Fig. 1). Similarly, the odd-year population at RC1 increased from 1980's levels; in 1991 and 1993 estimates were 1624 and 2408 respectively (Table 1, Fig. 1). At UPI, the type locality for B. acrocnema, no indi- viduals were seen in 1991, despite a concerted search effort that lasted from July 1 through July 25. How- ever, in 1993 an estimate of 452 was calculated (Table 1, Fig. 2). For the even-year brood, relative population estimates of 878 and 2918 were recorded in 1992 and 1994 respectively (Table 1, Fig. 2). When the UP6 colony of B. acrocnema was discov- ered in 1988 only two individuals were recorded (Brus- sard & Britten 1989). An estimate of 20 individuals in 1991 supported the notion that this was a small colony (Table 1, Fig. 3). However, in 1992 and 1994 estimates of 414 and 442 respectively were recorded (Table 1, JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY —f__ odd-year brood —{— __even-year brood Relative population estimate 1977 1979 1981 1983 1985 1987 1989 1991 1993 1995 1997 Year Fic. 2. Relative population estimates for Boloria acrocnema at Mt. Uncompahgre, colony site UPI, based on Pollard transect method. Fig. 3). And in 1993 the odd-year brood rose to an es- timate of 2266 (Table 1, Fig. 3). DISCUSSION Population Biology. Even- and odd-year broods at all three colony sites surveyed in this study have ei- ther stabilized or increased when compared to 1980's estimates (Table 1). Original population estimates at RC1 and UPI were surpassed in 1993 and 1994 and the colony at UP6 grew substantially (Table 1). In addition to the population data reported here, more current estimates have also been calculated by Colorado Heritage Program researchers using similar methodology (Ellingson et al. 1996, Wasinger et al. 1997). In 1995, the odd-year estimate for RC1 was 11,773, an order of magnitude higher than any odd- year estimate calculated prior to 1995 at that site (Ellingson et al. 1996). Large population estimates were also found at Mt. Uncompahgre colony sites in 1995: 6682 at UP1 and 1979 at UP6 (Ellingson et al. 1996). Although the UP6 estimate is reasonably close to the previous odd-year brood, the estimate at UP1 is more than twelve times as large. While 1995 estimates were dramatically different from 1993 estimates, 1996 estimates derived by Col- orado Heritage Program researchers were more simi- lar to those found in 1994. For instance, the estimate at RC1 in 1996 was 3670 (Wasinger et al. 1997): in 1994 I found 3464 (Table 1). At Mt. Uncompahgre, the estimate at UP1 was 2163 in 1996 (Wasinger et al. 1997) and 2918 in 1994 (Table 1). Finally, the estimate at UP6 was 130-200 individuals in 1996 (Wasinger et al. 1997) and 442 in 1994 (Table 1). VOLUME 53, NUMBER 1 2500 2300 —__ odd-year brood 2100 —1—__even-year brood 1900 1700 1500 1300 1100 900 700 500 Relative population estimate 300 100 -100 1986 1988 1990 1992 1994 1996 Year Fic. 3. Relative population estimates for Boloria acrocnema at Mt. Uncompahgre, colony site UP6, based on Pollard transect method. Data from 1990-96 provide strong evidence that B. acrocnema is on a trajectory toward stabilization or in- crease. Yet the following caveats need to be men- tioned: 1) employing the CMR calibration calculation of 10% to transect counts may lead to inflated esti- mates since the CMR results from one colony site may be not transferable to other sites; and 2) residency rates may change between years and are largely deter- mined by the geographical area that the colony inhab- its. For instance, the UP6 colony site is one third the size of UP1 and RCI: three transects at UP6 include the majority of available S. reticulata nivalis habitat. Therefore, the likelihood of counting individuals on day i+1 that were present on day i is higher at UP6 than at RC1, due to the differences in area and assum- ing all else being equal. To remedy these problems in deriving present relative population estimates, CMR experiments and local residency rates should be calcu- lated for all colony sites. Those values should then be used to translate Pollard transect data into population estimates. Conservation Status. Collecting pressure in the early 1980's was reported to be intense and some col- lectors extracted more than 50 specimens at any given time (P. A. Opler, pers. comm.). Although there are no studies that show collecting to be the cause of local ex- _ tinction in Lepidoptera (Pyle et al. 1991), studies of vertebrate fauna demonstrate that overharvesting can reduce genetic variability and heterozygosity (Bonnell & Selander 1974, O’Brien et al. 1989, Barrowclough & Gutiérrez 1990). Conservation biologists assume that populations with greater genetic variability are better able to survive and evolve in changing environments (Avise 1994). 35 Livestock grazing is the second force proposed by some to have affected B. acrocnema (Seidl & Opler 1994). Sheep grazing in the alpine environments of the San Juan Mountains began in the mid 1800's (Lake City, Colorado Historical Society) and continues into the present. Foraging and trampling by sheep may re- sult in lower seedling survival of both nectar sources and host plants (Owens & Norton 1992). Both collecting and livestock grazing have been pro- hibited or suspended since B. acrocnema was federally listed in June 1991, presumably shielding the species from these pressures. However, local climate change and a consequent drying trend have not abated. Weather data collected over the last two decades indi- cate a trend of below-average snowfall and higher- mean ambient air temperatures for southwestern Col- orado (Colorado Avalanche Center, Denver, Colorado). It has been demonstrated that weather can influ- ence butterfly populations (Pollard 1988, Pollard & Yates 1993) and that catastrophic weather events may bring about local butterfly extinction (Ehrlich et al. 1972, Singer & Thomas 1996). Based on the unusual distribution and microhabitat requirements of B. acrocnema, its relict arctic history, specialist nature, and mesic habitat requirements, this species may be an excellent indicator of the effects of climate change in alpine environments. Although some species may migrate to cooler northern latitudes (Parmesan 1996), species with low vagility like B. acrocnema may be lim- ited in their dispersal capabilities (Dennis & Shreeve 1991, Murphy & Weiss 1992, Britten et al. 1994). Changes in host plant quality, phenology and distribu- tion are also likely as plants respond to changing abi- otic conditions (Dennis & Shreeve 1991). Butterfly abundance may be affected as host plants become less palatable or less available (Dennis & Shreeve 1991, Pollard & Yates 1993). Britten et al. (1994) suggested that B. acrocnema’s low population numbers in the late 1980's were due to human-induced factors including climate change and that the butterfly was headed for extinction. The data I have reported here suggest that the butterfly has re- covered from its low population numbers. However, I believe Britten et al. (1994) raised important questions concerning the effects of climate change on B. acroc- nema and other high altitude mesophilic butterflies. To address their concerns, careful analysis of how cli- mate change parameters (rising ambient tempera- tures, increased UVB, elevated CO, levels, etc.) affect the larvae and adults of B. acrocnema (or a close sur- rogate due to its endangered status) is needed. These experiments will provide essential evidence in identi- fying current and potential effects of climate change 36 on B. acrocnema and will help to predict the species’ future population dynamics and its probability of ex- tinction under a climate change model. ACKNOWLEDGMENTS I thank the Bureau of Land Management, the Colorado Mountain Club, Colorado State University’s Department of Entomology, the U.S. Fish and Wildlife Service, and the U.S. Forest Service for sup- porting this research. I thank A. Brody, A. Ellingson, B. Irwin, B. Kondratieff, P. Opler and D. Steingraeber for reviewing earlier drafts of this manuscript. I also thank D. Bowers, L. Gall and R. Pyle for of- fering many suggestions that improved the final quality of this paper. LITERATURE CITED AVISE, J.C. 1994. Molecular markers, natural history and evolution. Chapman and Hall, New York, New York. 511 pp. BARROWCLOUGH, G. F. & R. J. GUTIERREZ. 1990. Genetic variation and differentiation in the spotted owl (Strix occidentalis), Auk 107:737-744. BONNELL, M. L. & R. K. SELANDER. 1974. Elephant seals: genetic variation and near extinction. Science 184:908—909. BRITTEN, H. 1991. The conservation biology of the Uncompahgre fritillary and the related northern dingy fritillary. 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Evolutionary responses of a butterfly metapopulation to human- and climate-caused envi- ronmental variation. Am. Nat. 148:S9-S39. Tuomas, J. A. 1983. A quick method of estimating butterfly num- bers during surveys. Biol. Cons. 27:195—211. WASINGER, T., A. R. ELLINGSON, P. M. PINEDA & C. A. PAGUE. 1997. Population monitoring and inventory of the Uncompah- gre fritillary (Boloria acrocnema): 1996 field report. Colorado Natural Heritage Program, Fort Collins, Colorado. 62 pp. Watt, W. B., F. S. CHew, L. R. G. SYNDER, A. G. Watr & D. E. ROTHSCHILD. 1977. Population structure of Pierid butterflies. Oecologia 27:1—22. WeBER, W. 1987. Colorado flora: western slope. Colorado Assoc. Press, Boulder. 530 pp. Received for publication 8 April 1998; revised and accepted 31 Jan- uary 1999. Journal of the Lepidopterists’ Society 53(1), 1999, 37-44 PATTERNS IN THE SPATIAL AND TEMPORAL USE OF TEXAS MILKWEEDS (ASCLEPIADACEAE) BY THE MONARCH BUTTERFLY (DANAUS PLEXIPPUS L.) DURING FALL, 1996 WILLIAM H. CALVERT 503 East Mary Street, Austin, Texas 78704 ABSTRACT. In an attempt to better understand the fall migration of the monarch butterfly (Danaus plexippus L.) through Texas, the spatial and temporal distribution of milkweeds and immature monarchs were monitored along an 800 km transect. Important monarch host plants were found to be distributed unequally along the transect. Breeding monarchs were present at least one month before the main body of migrants appeared. The distribution of eggs and larvae did not spatially follow the distribution of milkweeds. Some milkweed species and locations were used more than others. Possible explanations for the distribution patterns observed include the circumscribed nature of the fall migratory pathway and the foraging efficiency of the polygyne version of the imported fire ant. Additional key words: natural regions, oviposition, fire ant, Solenopsis invicta, Asclepias, migration. Although the monarch butterfly (Danaus plexippus L.) is the best known of North America’s migratory in- sects, much remains to be learned about the spatial and temporal patterns of how it populates and then va- cates northern North America each spring and fall (Brower 1995). Our basic understanding of this pro- cess is as follows: During summer, the eastern group of monarchs spends two or more generations breeding in North America above latitude 35° (Malcolm et al. 1993). In late summer and early fall, the progeny of summer breeders migrate to central Mexico, crossing much of the North American continent (Urquhart 1987). They spend five months at the Mexican over- wintering sites, most of the time in a state of repro- ductive dormancy. After mating in late winter and early spring, they fly northward to once again exploit the milkweed flora (Cockrell et al. 1993) widely dis- tributed throughout North America (Woodson 1954). During spring and summer in their northern breeding grounds, they greatly increase their numbers, revers- ing the population decline that occurs during fall and winter. To understand more about monarch population dy- namics in Texas and the implications for the monarch population of North America, this study investigated the presence and abundance of monarch immatures on milkweed flora (Asclepiadaceae) in Texas during the fall of 1996. MATERIALS AND METHODS Milkweeds were examined along two routes be- tween 30°N and 32°N latitude. One route extended _east from Austin to Pineland in Sabine County in ex- treme eastern Texas near the Louisiana border and the other extended west from Austin to Ozona in Crockett County in West-central Texas (Fig. 1). A loop was made at the eastern end to insure that areas where sin- gle-queen (monogyne) fire ant colonies had been re- ported were well covered (Anonymous 1998; E. Vargo, pers. comm.). The two segments combined, referred to below as the cross-Texas transect, stretched about 60% of the way across Texas from 31.25°N, 93.985°W at Pineland to 30.00°N, 101.201°W at Ozona, a dis- tance of 801.66 km (ca. 900 road kilometers; see Mar- vin 1939, Table 92). The transect was traversed three times during the fall migratory season. The east and west segments of the first transect were conducted on 20-23 September and on 27-29 September respec- tively. The second transect was conducted on 6-7 and 11-13 October and the third on 12—13 and 12-13 No- vember. On the first transect to West Texas (27-29 September), the return route was slightly different from the two subsequent transects. Instead of pro- ceeding directly to Austin along Highway 290, a paral- lel route 30 km to the south of the regular route was taken. This route followed Interstate 10 to Comfort and then proceeded eastward along Highways 473, 281 and 165, rejoining Highway 290 at Henly. The last transect to East Texas (12-13 November) ended at Madisonville. The transect was truncated here be- cause no activity had been observed on the previous transect (6-7 October), and none had been observed prior to Madisonville. The scheduling of the transects was based on Texas Monarch Watch reports of the presence of monarchs in the state. Transect dates were chosen to cover peri- ods before the main body of migrants arrived, during the peak of the migration and after most monarchs had cleared the state. (The Texas Monarch Watch is an ed- ucational outreach service that solicits reports from volunteers concerning the presence and abundance of monarchs in the state (Calvert 1993-1997, Calvert & Wagner in press).) The relative abundance of milkweeds was assessed along the cross-Texas transect by counting stems whenever milkweeds were sighted. During September and October, many milkweeds were in flower and their presence was very conspicuous. Sampling stops were 38 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY fa) Piney Woods Oak Woods & Prairies MMB Blackland Prairie [ES Gulf Coast Prairies & Marshes GB Coastal Sand Plain (=I South Texas Brush Country 4 Edwards Plateau Rolling Plains #) High Plains ef Trans Pecos i enaee ase 4“ ess 7% jee ie PY Act Fic. 1. The cross-Texas transect route showing the location of milkweeds important to monarch butterflies along the 800 km transect from Pineland to Ozona. made when milkweeds were spotted in adjacent fields or roadside right-of-ways. Areas that appeared to be prime milkweed habitat such as short grass prairies and areas with poor soils were also searched. During November, when many fewer milkweeds were in flower, plants were located by revisiting areas where they were found abundant on previous transects. This study focused on the four abundant Asclepias species found in central Texas: Asclepias viridis Walt., A. oenotheroides Cham. & Schlecht, A. asperula Wood, and A. latifolia Raf. Other rarer species were examined as encountered, e.g., A. texana Heller and A. curassav- ica L. Milkweed vines in the genera Matelea, Sar- costemma and Cynanchum were also sampled, but only sporadically. Due to the very different growth form of these species and the difficulty of following a stem through the tangle of vines, the effort of examination cannot be considered the same as for the other species. Although every effort was made to keep the search method constant, initial search times were longer than later ones. It took less time to relocate milkweed patches after their initial discovery. Nonetheless, since the search method was always the same, the number of milkweeds counted should serve as a rough index of relative abundance of each host species in the genus Asclepias through space and time. During a sampling stop, the identity of the milk weed species was verified, the stem lengths were mea- sured, and the number of eggs and monarch larvae on the stems were counted. When available, 20 or more stems were examined. If fewer than 20 stems were present, all available stems were measured and exam- ined. Time of day, location, and the presence of adult monarchs were also noted. Geographical locations of sampling sites were determined from a U.S. Geologi- cal Survey Map of Texas (Anonymous 1985). A problem arose concerning the identification of eggs. At some locations in West Texas, both queen (Danaus gilippus L.) and monarch larvae were present at the same time during the fall. Since monarch eggs cannot be distinguished from queen eggs, eggs from both species were counted and are included the totals. Excluding the larvae present on M. reticulata, the host species that was used only by queens, 36% of the lar- vae encountered in West Texas were queens. No queen adult or larvae were encountered along the eastern segment of the transect. All eggs here were considered to be monarch. RESULTS Relative abundance and distribution of milk- weed species used by monarchs. Approximately VOLUME 53, NUMBER 1 300 200 Number of stems 150 100 50 0 - ve) (o) ve) ep) wn [co) Se ¢ © g€ ® @& ® Las io) a? (o>) op) Ay = = § © = r=) o 5 £ oS ue & iS i) c n lo) 2 5 o 2) 5 8 [= =) [= 8 6 w =. co Fic. 2. midway between Ozona to Pineland, the 800 mile transect crossed a major biogeographical barrier—the Balcones Fault (Fig. 1). At the latitude of the transect, the Balcones Fault is located at Austin (97.67°W). The transect west of the Balcones Fault lies entirely within the Edwards Plateau. Elevations here are 250—300 meters higher than the transect area to the east of the fault. East of this line rainfall is ample, >32 inches per year; west of it rainfall diminishes to ca. 16 inches per year at Ozona (101.2°W longitude; Arbingast, et al., 1973). Proceeding from east to west, the transect crossed four natural regions: Piney Woods, Oak Woods and Prairies, Blackland Prairie and Edwards Plateau (Anonymous 1978). The diminishing rainfall as one proceeds westward is conspicuous in the change in natural regions, the reduced density and stature of the trees, and the change in distribution of milkweeds. Above average rainfall during the fall of 1996 (Anonymous 1997) produced luxuriant crops of milk- weed giving ample opportunity to observe their distri- bution and relative abundance (Figs. 1, 2). In October, when the four major hosts (A. viridis, A. oenothero- _ ides, A. asperula and A. latifolia) were at their peak of flowering, A. viridis accounted for 41% of the milk- weeds along the transect followed by A. asperula (33%), A. oenotheroides (17%) and A. latifolia (4%). The distribution of these species along the transect varied with longitude. Asclepias oenotheroides had the widest longitudinal range (Figs. 1, 2). It was distrib- Austin 97.5 39 A. viridis [] Avenotheroides A. asperula BAA. latifolia fH other nn wy ice) w w w + w Oo @ ®&@ ww @®© & @ 8 [op) [op) aD [op) = o xe] oO > Cc > fod isi aaj x) 2 = ra) a < Longitude E The relationship between longitude and the occurrence and relative abundance of the principal monarch host plants. uted in patches throughout most of the transect, but was not found east of the Trinity River. It appears to be mainly a prairie species, reaching its highest density in the regions between Austin (97.67°W) and Midway (95.75°W). However, it was also consistently, but rarely, encountered in the Edwards Plateau to the west end of the transect at Ozona. Asclepias asperula was the dominant milkweed in the genus Asclepias on the Edwards Plateau (Figs. 1, 2). It was mainly present in patches of multiple- stemmed rosettes along roadsides and in pastures. It was found as far east as longitude 97.45°W in the Oak Woods and Prairies region between Bastrop and Austin, but east of Austin, it was rare. The highest den- sities were found between Austin and Johnson City. A. latifolia was encountered only in rare patches on the Edwards Plateau west of 100°W longitude. The Balcones Fault at Austin divides the ranges of A. viridis and A. asperula fairly well (Figs. 1, 2). Pro- ceeding eastward from Austin, A. viridis began a more or less continuous distribution around 97°W and, with two gaps, continued into the East Texas Piney Woods to the end of the transect at Pineland (93.99°W; Fig. 2). East of Austin, A. viridis reaches its greatest abun- dance in a zone extending along Highway 21 approxi- mately 65 km on either side of the Trinity River (95.70°W longitude). This area corresponds to an area of warmer than expected temperature that extends northward along the Trinity River (Fig. 1; Arbingast, et 40 Number of eggs and larvae A A wo No. stems 479 4 (oe) fo?) A. viridis A. asperula A oenotheroides A. latifolia JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 1 sth: 4th = o = i) A.texana w & A. curassavica M. reticulata Host species Fic. 3. Numbers of eggs and larval instars found on various milkweed hosts along the cross-Texas transect during the fall of 1996. M = mon- archs (Danaus plexippus); Q = queen (Dannaus gilippus). Numbers not followed by a letter are monarchs. al. 1973). Within this area the number of days of frost free weather are ca. 35% greater than regions above the Balcones Fault on the Edwards Plateau. The pres- ence of dense stands of A. viridis corresponded well with the occurrence of Blackland Prairie and with pockets of prairies within the Oak Woods and Prairies Natural Region. Although dense stands occurred in some places within the Piney Woods, A. viridis in this area was confined to roadsides. Because of this restric- tion, its overall abundance here must be considerably less than in the prairies. Two other milkweeds in the genus Asclepias were encountered along the transects. One stem of the non- native A. curassavica was found in a garden in Johnson City, and three stems of A. texana were found along Highway I 10 at ca. 99.0°W. Several milkweed vines were encountered in the western portion of the transects. These included Cy- nanchum barbigerum Shinners, Sarcostemma crispum Benth., S. cynanchoides Dene., and Matelea reticulata Woods. Both M. reticulata and S. cynanchoides achieve high biomass in the western portions of the transect. Stem counts of the four major milkweed species dur- ing the three months indicated that all species of milk- weeds except A. oenotheroides declined in abundance from October to November. Only A. oenotheroides be- came more abundant at the end of the three month period. The continued growth of A. oenotheroides may be explained by the apparently positive response of this species to the mowing of highway right-of-ways by the Texas Department of Transportation (Calvert, un- publ. obs.). Distribution of monarch eggs and larvae with respect to host species, location and time in the season. Prior reports to the Texas Monarch Watch in- dicated that in previous years, breeding monarchs had been present in low densities in Central Texas during September, and that the main mass of migrants did not arrive until the end of September. Migrants had mostly cleared the north and central parts of the state by the — end of October (Calvert 1993-1997, Calvert & Wag- ner in press). Cross-Texas transects commenced on 21 September and continued until 18 November. A total of 1422 milkweed stems were measured and examined for the presence of monarch and queen eggs and monarch larvae. Eggs and larvae per meter of stem were not equally distributed among milkweed species (Fig. 3). A. texana, A. curassavica and M. retic- ulata were infrequently encountered on the transects, but each of these species had eggs or larvae. Of the four major milkweed species encountered, leaves of Asclepias latifolia had the most eggs and monarch and queen larvae per meter of stem (0.73 eggs and lar- vae/m) and A. oenotheroides had the least (0.09 eggs and larvae/m). Asclepias asperula and A. viridis had 0.27 and 0.39 eggs and larvae/m respectively. Monarchs and queens used A. latifolia much more than any other milkweed species—almost three times more than A. viridis and ca. two times more than A. asperula. VOLUME 53, NUMBER 1 4] September Eggs and larvae per meter of stem Longitude _— wn lo) w fop) a) foo) te) Eeqgo OG ®@ GQ Ff ae oO a (op) ep) (op) © October Eggs and larvae per meter of stem uo 0.5 Se ®M © & @® ®M & (o) (s) on) CD: : (o) fon) ee) ~ SS Sy OS en) oy) or) Ss Longitude sth 4th Ei 3rd Z 2nd FB ist [1] Eggs Fic. 4. The relationship between numbers of eggs and larvae per meter of stem and the time of the transect. Breeding activity declined from a high in September to near zero in November. a, September; b, October; Most of the milkweed patches encountered were comprised of one plant species. Only rarely were two species found in the same area, e.g., A. oenotheroides and A. viridis in East-central Texas and A. latifolia and A. asperula in West Texas. Because of the spatial sepa- ration of host species, these data do not show oviposi- _tion preferences among the hosts, but rather, they show a presence in a specific geographic area. Cursory searches of vines in the genera Cynanchum and Sarcostemma revealed no eggs or larvae. Although it has been reported that Matelea is not used by mon- archs (P. Davis, pers. comm.), the importance of these other potential milkweed vine hosts needs to be inves- tigated. During the fall of 1996, the presence of eggs and larvae at both ends of the transect declined as the sea- son advanced. Eggs and larvae were encountered on milkweeds east of Austin (97.67°W) during the initial transect of 20-23 September, but not on the two sub- sequent transects of 6-7 October and 12-13 Novem- ber (Fig. 4a—c). In contrast, monarch and queen eggs or larvae were observed west of Austin during each of the three transects run from 27 September to 17 No- vember (Fig. 4a—c); however, the last transect of 17-18 November yielded no eggs and only one larva. The to- tal number of eggs and larvae for the complete tran- sect declined from a September high of 69, to 26 in October, to 1 in November. Breeding in East Texas 42 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY November =s § ti ‘5 2.5 M sth os NS4th = hS =| 3rd 2 1.5 : 8 2nd c 1 Ss A ist z 2) LO) = noenonenwaoenre nwo wn Se F®q"® aM*E OCG ® ® pee >" Gy) fon) fon) (>) fon) Longitude 6 All dates 5 5 7 - 1} 5th ° 5 4 4th o E Ei 3rd g 3 o 2nd iv 2 S 2 Ze BB ist = WH 5 » le ggs g 1 Y / Le?) a | (0) ams L am - wn S) w [op) w foo) w y w wo w w w T+ w Oe eS 896 7 Ff © GCF Boe 8s DW Es a iO) | Se OD or) fo) (>) a fo) a r% Longitude A. asperula ————e A. latifolia > ee ines soe ate Species Fic. 4. e, November; d, all dates combined. A conspicuous gap in breeding activity occurs in the central prairie area between and includ- ing 96°W and 97.5°W in spite of the presence of milkweed there. was likely over by the time the main mass of migrants arrived, but continued into November in West Texas (see discussion below). Monarch (and queen) eggs and larvae showed a bi- modal distribution along the 800 km transect (Fig 4d). The most eggs and larvae per meter of stem were found west of Austin between longitudes 98° and 101°W. The main concentration on the western end began about 15 miles east of Sonora (100.78°W) and continued to Ozona (100.96°W). Another area of con- centration was south of Johnson City (98.37°W). Con- spicuously and curiously absent were monarch eggs and larvae located in the center of the transect, from Austin (97.87°W) to Huntsville (95.55°W), the main prairie region of these latitudes in Texas. Traveling eastward, the number of eggs and larvae increased east of Livingston (94.63°W) and were especially concen- trated in patches in the Piney Woods between Broad- dus and Pineland (94.14°W). Averages for the fall of 1996 were 0.16 eggs and larvae/meter of plant stem east of Austin (n = 820 stems) and 0.47/meter of plant stem (n = 602) stems west of Austin. Most of the fall breeding activity occurred west of Austin, and most of it occurred on A. latifolia. DISCUSSION The distribution of milkweeds and monarch eggs and larvae with respect to the distribution of VOLUME 53, NUMBER 1 milkweeds. With the exception of one major gap be- tween Broaddus and Crockett and several minor gaps elsewhere (Fig. 2), milkweed species were continu- ously distributed throughout the transect. Monarch and queen eggs and larvae were not. The concentration of eggs and larvae in the eastern and western end of the transect and their absence in the middle prairies, in spite of the presence of milkweeds, requires expla- nation. Two possibilities are: 1) Monarchs migrating south during the fall largely avoid the east-central prairies, and therefore little oviposition occurs there. 2) The predacious activity of the imported fire ant, Solenopsis invicta Buren primarily determines the dis- tribution of eggs and larvae. The first hypothesis cor- responds well with data compiled from the Texas Monarch Watch. The fall migratory flyway is mainly lo- cated to the west of Austin and corresponds to the western end of the transect (Calvert & Wagner 1999). However, the migration did not reach Texas until after breeding had ceased in East Texas. Moreover, monarch eggs and larvae had already appeared in West Texas before the beginning of the migration. The second hypothesis best explains the bimodal distribution of monarch eggs and immatures during the fall of 1996. The absence or low densities of fire ants in West Texas may explain the relative abundance of eggs and larvae in that region, while the preponder- ance of the far less dense single-queened colonies in East Texas (Porter et al. 1991) may allow a relatively higher number of eggs and larvae to survive there. Other studies support the contention that fire ants are important in the decline of many species of Lepi- doptera in Texas. Long term records of the presence and abundance of Lepidoptera show that the abun- dance of lepidopterans in the vicinity of Austin has fallen to 50% of pre-fire ant levels. Especially hard hit were grass-feeding members of the Satyridae (C. Dur- den, pers. comm.). During the spring of 1995, a field in South central Texas near Luling containing an estimated 1250 monarch eggs failed to yield a single late instar monarch larva (Calvert 1996). This same field contained an estimated 1001 fire ant mounds. The high mound density and renowned foraging efficiency of the im- ported fire ant (Porter et al. 1991) suggested that this predator was the principal culprit in decimating the ‘monarch population. Finally, preliminary results from a study comparing larval growth inside exclusion zones, where fire ants densities were kept relatively low, to ar- eas outside the exclusion zones, showed the production of fifth instars inside the exclusion zones to be 13 times higher than outside. (Calvert, unpubl. data). The distribution of prairies in the mid-west and the pattern of the spring migration northward through 43 Texas suggests that the monarchs that breed on Cen- tral Texas prairies and plains are the progenitors of monarchs that will breed on the prairies of mid-west- ern states further north (Malcolm et al. 1993). The pattern of ample oviposition, combined with the fail- ure of larvae to develop into later instars, found during the spring of 1995 near Luling (Calvert 1996), suggests that monarchs breeding within the fire ant zones of Texas make only a small contribution to the North American monarch population. No evidence yet exists for similar effects in areas farther east where fire ants are also abundant, but eastern fire ants colonies are mostly monogyne and are not as dense as the multiply- queened (polygyne) variety on Texas prairies (Porter et al. 1991). The reproductive success of monarch mi- grants passing through in the fall may be diminished for the same reasons. Fall breeding in Texas. It has long been held that monarchs greatly increase their population size during summer months by breeding in the northern portion of their North American domain, especially in the lati- tudes of the Great Lakes and Northeastern States (Urquhart 1987; Malcolm et al. 1987). This is the area where the greatest biomass of one of their important milkweed hosts (A. syriaca L.) is found. At the peak of breeding (June and July), all females contain ovaries with numerous eggs in the oviducts and there is no communal roosting. During this period there are vir- tually no monarchs in Texas (Calvert 1993-1997). As the season advances into September, more and more females possess inactive ovaries and communal roost- ing increases. At the beginning of their southward mi- gration, most individuals show inactive ovaries and testes, but occasionally males and females with active reproductive organs are encountered (Brower 1985). It was recognized that some breeding did occur on the way south in states such as Texas, but the extent and importance of this was not known (Urquhart 1987). These data show that most fall breeding occurred early in the cycle before the main mass of migrants had arrived. Only in West Texas was there any breed- ing activity after the beginning of October (Fig. 4b). Prior to the arrival of the great masses of migrants, only an occasional monarch has been seen in Texas (Calvert 1993-1997), and communal roosting has not been observed. The early population, which may ar- rive as early as late August, may differ from the main migrant body in its breeding activity and non-commu- nal roosting. It may not be part of the migratory move- ment to Mexico. Instead these may be breeding but- terflies that have dispersed southward in the same manner that they disperse northward in the spring, stopping to lay eggs as the opportunity presents itself. 44 The presence of breeding monarchs in Texas in late August and early September was unexpected and re- quires a rethinking of the pattern of migration. Future studies may show that monarchs regularly breed on Texas milkweeds during September and perhaps Oc- tober, augmenting their numbers and adding a gener- ation to the monarch life history cycle. The origin of these late summer breeders is of yet undetermined. ACKNOWLEDGMENTS Many people from various agencies contributed to the success of this study. I gratefully acknowledge the following and apologize to any I have omitted: John Heron, Matt Wagner, Noreen Demude and Tonya Hunter at the Texas Parks and Wildlife Department's Nongame and Urban Wildlife Program; the Arldts at Manheim for permission to examine milkweed stems on their property; the Wray Trust for their support of the Texas Monarch Watch from which much background information for this study was derived; James Gillespie for help in identifying insects found on milkweeds; and Texas Monarch Watch volunteers for basic information about the monarch migration. Cynthis Banks of Texas Parks and Wildlife’s GIS Lab helped prepare fig. 1. I thank Lincoln Brower, Philip Russell, David Millard and two anonymous reviewers for many helpful sug- gestions that improved the manuscript. This study was supported by the Wildlife Division of Texas Parks and Wildlife Department. I thank them, especially Ron George, heartily for giving me the opportunity to study this most interesting migratory phenomenon. LITERATURE CITED Anonymous. 1978. Natural regions of Texas, In Preserving Texas’ Natural Heritage LBJ School of Public Affairs Research Project Report # 31. Compiled by Texas Parks and Wildlife Depart- ment, GIS Lab. . 1985. Map of the State of Texas. United States Depart- ment of the Interior Geological Survey. . 1997. Austin rainfall chart. Monthly precipitation records recorded by the National Weather Service at Robert Mueller Municipal Airport. Austin American Statesman, December 7, 1997. . 1998. Texas Department of Agriculture, Fire Ant Quaran- tine Map, TDA Q591D, revised 5-98. Texas Department of Agriculture, Austin, Texas. ARBINGAST, S. A., L. G. KENNAMER, R. H. RYAN, A. Lo, D. L. Kar- NEY, C. P. ZLATKOVICK, M. E. BONNIE & R. G. STEELE. 1973. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Atlas of Texas. Austin: Bureau of Business Research, University of Texas at Austin. Brower, L. P. 1985. New perspectives on the migration biology of the monarch butterfly, Danaus plexippus L., pp. 748-785. In M. A. Rankin (ed.), Migration: mechanisms and adaptive signif- icance. Contributions in Marine Science. Vol. 27. Marine Sci- ence Institute, University of Texas, Port Aransas, Texas. Brower, L. P. 1995. Understanding and misunderstanding the mi- gration of the monarch butterfly (Nymphalidae) in North America: 1857-1995. J. Lepid. Soc. 49:304—385. CALVERT, W. H. (1993-1997). Newsletter of the Texas Monarch Watch, Texas Parks and Wildlife Department, 4200 Smith School Road, Austin, Texas 78744. vols. 1-3. CALVERT, W. H. 1996. Fire ant predation on monarch larvae (Nymphalidae: Danainae) in a Central Texas prairie. J. Lepid. Soc. 50:149-151, CALVERT, W. H. & M. WAGNER, 1999. Patterns in the monarch but- terfly migration through Texas and Mexico—1993-1995. In Proceedings of the third Monarch Butterflies conference, No- vember, 1997, Morelia, Michoacan, Mexico. In press. COCKRELL, B. J., S. B. MALCOLM & L. P. BROWER. 1993. Time, temperature and latitudinal constraints on the annual recolo- nization of Eastern North America by the Monarch butterfly, pp. 233-252. In S. B .Malcolm & M. P. Zalucki (eds.), Biology and conservation of the Monarch Butterfly. Publication of the Los Angeles County Museum of Natural History, Los Angeles. MALCOLM, S. B., B. J. COCKRELL & L. P. BROWER. 1987. Monarch butterfly voltinism: effects of temperature constraints at differ- ent latitudes. Oikos 49:77—82. MALCOLM, S.B., B.J. COCKRELL & L.B. BROWER. 1993. Spring re- colonization of Eastern North America by the Monarch butter- fly: Successive brood or single sweep migration?, pp. 253-268. In S. B. Malcolm & M. P. Zalucki (eds.), Biology and conserva- tion of the Monarch Butterfly. Publication of the Los Angeles County Museum of Natural History, Los Angeles. MarvIN, C. F. (ED.). 1939. Smithsonian meteorological tables (1931 edition). Lord Baltimore Press, Baltimore. PorTER, S. D., A. BHATKAR, R. MULDER, S. B. VINSON & D. J. Ciair. 1991. Distribution and density of polygyne fire ants (Hymenoptera: Formicidae) in Texas. J. Econ. Entomol. 84:866—874. UrQuuakt, F. A. 1987. The Monarch Butterfly: international trav- eler. Chicago: Nelson-Hall. Woopson, R. E. JR. 1954. The North American species of Ascle- pias. Ann. Mo. Bot. Gard. 41:1-211. Received for publication 9 March 1998; revised and accepted 31 January 1999. BOOK REVIEWS Journal of the Lepidopterists’ Society 53(1), 1999, 45 Die NOCTUIDEN RUMANIENS, by L. Rakosy. 1996. Published by Staphia 46. 648 pp., 30 color plates, 11 habitat photographs, 68 text figures, 821 line drawings of genitalia, 651 distribution maps. Hard- back, 28/21 cm. Available from Apollo Books Aps., Kirkeby Sand 19, DK-5771, Stenstrup, Denmark, Tel. 45-62-26-3737, FAX: 45-62-26- 3780. Price DK 560.00 (approximately U.S. $90.00). Here is a remarkably fine work focusing on the rich noctuid fauna of a large eastern European area between the Carpathian Moun- tains, the lower Danube with its delta, and the Black Sea. The au- thor, who is one of Romania's leading experts on the Noctuidae, has done many years of research in the field and museum collections. This book is the pinnacle of all those years of work. It starts with an introductory section, written in cooperation with Dr. Eckbert Schneider, which presents a history of noctuid studies performed in the territory of present-day Romania since as early as the 18 century. In the following sections, Romania’s geography and landscape are presented and the country’s climate and vegetation are discussed. Several beautiful, colored illustrations complement the text, showing images of various biotopes from the high peaks of the Carpathian Mountains to the sand dunes on the Black Sea coast. Next, a brief section discusses the biogeography of the Romanian noctuid fauna. It has been discovered, among other things, that about 5% of Romania’s noctuids are holarctic in distribution, and so Romania shares over 30 noctuid species with North America (e.g., Scoliopteryx libatrix (L., 1758); Acronicta auricoma (F., 1787); Plu- sia putnami Grt., 1873; Calophasia lunula (Hufn., 1766); Xanthia to- gata (Esp., 1788); Agroperina lateritia (Hutn., 1766); Hydraecia mi- cacea (Esp., 1789); Cerapteryx graminis (L., 1758); the recently introduced Noctua pronuba (L., 1758)). The anatomy of the imago, egg, larva, and pupa is described in a concise and clear manner with very good illustrations. The author highlights the important features of the adult exoskeleton, genitalic structures, egg morphology, and larval chaetotaxy. The systematic part forms the bulk of the work. Each of the 670 species ever to be recorded on Romanian territory is discussed in detail. The author follows the systematic list of the European Noc- tuidae published by Fibiger and Hacker (1991). He gives a brief structural and genitalic description for each genus and treats each species by giving a list of selected synonyms with authors and years, biological data for the adult and the larva, general distribution and distribution within Romania. For many species there are very inter- esting parasitolgic mentions. It is worth mentioning here that the au- thor is describing two new subgenera (Synapamea for Apamea Ochsenheimer, 1816 and Denticucullus for Chortodes Tutt, 1897) and seven new subspecies of local/regional importance from en- demisms in Carpathian Mountains. The illustrations of the male genitalia for each species and the fe- male genitalia for many species are grouped together after the sys- tematic section. The author has made a tremendous effort in per- sonally drawing a total of 821 excellent illustrations of these important diagnostic tools. The distribution within the country is il- lustrated with the record/dot system and there is a map for each spe- cies with valid Romanian records. Bound together at the end of the text are 30 colored plates that show 882 excellent quality photographs of adults of each species dis- cussed in the work. The impeccable quality of these plates as well as of the photographed specimens make identification by superficial habitus possible even for the most difficult groups (e.g., Oligia Hbn., 1821; Cucullia Schran, 1802; Orthosia Ochs., 1816, etc.). The plates are followed by 3 more beautiful, folding plates that show 40 stun- ning photographs of live larvae. This work contains an extensive list of literature with 549 refer- ences on Romanian and general European papers and books dis- cussing noctuid related topics. The book ends with an Index that lists all genera, species, and synonyms with their authors and years of description. Unfortunately, the book lacks a species checklist, making the overall faunistic appreciation of the area and the com- parison with other areas somewhat difficult and time consuming. Because it covers 670 species of Noctuidae from Europe (over 50% of the whole continental fauna), this book is a landmark work. It is the first one to treat exhaustively a moth family (and of the mag- nitude of the Noctuidae!) in an eastern European country and with color photographs and genitalic illustrations for all listed species. Hacker did something similar when he published his book about the noctuids of Greece (1989, Die Noctuidae Griechenlands, Mit einer Ubersicht uber die Fauna des Balkanraumes (Lepidoptera: Noctu- idae), Herbipoliana 2:1—589) but he only illustrated a selected num- ber of adults and not all of them in color, with selective genitalic drawings. These two books, Hacker's and Rakosy’s complement each other very well by giving a very good idea of the composition of the noctuid fauna of Eastern Europe from the Mediterranean Sea to the Ukrainean Steppe. Although written in German, the text can be understood with ba- sic linguistic skills, making it an important source of information on the noctuids in general and a very good identification tool for the over 30 noctuid species shared by Romania and the U.S.A. Mr. Rakosy and his publisher, the Austrian house Staphia 46, are to be very highly praised for producing a book of the highest infor- mative and graphic standards, making of it a most valuable tool for the serious student of this large and heteromorphous family. VALERIU ALBU, 6 Kit Road, Charleston, West Virginia 25304 Journal of the Lepidopterists’ Society 53(1), 1999, 45-46 CONTRIBUTIONS TO THE KNOWLEDGE OF THE INSECTS OF THE PHILIPPINES, III. [Beitrage zur Kenntnis der Insekten der Philip- pinen, III], edited by Wolfgang A. Nassig, Colin G. Treadaway and Josef Settele. 1998. Nachrichten des Entomologischen Vereins Apollo e. V. Frankfurt am Main, Supplement 17, July 1998. Senck- enberganglage 25 D-60325 Frankfurt am Main. 576 pp., 48 color plates, text figures. ISSN 0723-9920. The Philippines is an archipelago of 7107 islands, many are very small in area with only about 500 islands with an area over one Km’. Only 2100 islands are actually inhabited by 65 million people com- posed of 60 ethnic groups. Not surprisingly, about one third of the islands are not listed by name in most reference books or maps. The Philippine Islands, with the highest mountain reaching 2954 m and with 17 active volcanoes, is geologically very complex. The zoogeo- graphical relationships to other areas of southeast Asia are manifold and complicated. Historically, the Philippines was noted for its extensive forest cov- erage. However, this has changed considerably over the past 30 years and now less than 10% remains of the original forest coverage present 50 years ago. Although there are 61 national parks and pro- tected areas, there will be no true forests shortly after the turn of the century if the current rate of deforestation continues. This will have a very strong impact on all forms of life in Philippine forests includ- ing insects. Human pressure on the global environment makes it critical that we acquire knowledge about biological diversity as fast as possible. An essential contribution to managing the biosphere intelligently is to discover, describe, and inventory its species. Southeast Asia is by no means an exception to these guidelines and several contributions in the form of national or regional faunal treatments of some groups of Lepidoptera have been published recently for example of Penin- sular Malaysia, Borneo, Thailand, Sulawesi, Sumatra, and Vietnam. On the same token, this volume is the third contribution to the knowledge of the insects of the Philippines with special emphasis on Lepidoptera within the Supplement series of “Nachrichten des En- tomologischen Vereins Apollo”, and it is an important addition to the taxonomy, nomenclature, and biogeography of the Lepidoptera. 46 This 576 page special issue is composed of 12 papers in English ranging from the physical description of the Philippines to anno- tated checklists of several lepidopteran groups and one of Tri- choptera and descriptions of several new taxa. Ten of the 12 papers are devoted to Lepidoptera, therefore this issue is of special interest to the lepidopterist. The following papers comprised the volume: “Short introduction to Philippine natural and geological history and its relevance for Lepidoptera” by C. G. Treadaway; “The Sphingidae (Lepidoptera) of the Philippines” by W. Hogenes and C. G. Treadaway; The Lasio- campidae (Lepidoptera) of the Philippines by V. V. Zohouhin, C. G. Treadaway and T. Witt; “The Saturniidae (Lepidoptera) of the Philip- pines” by W. A. Niassig and C. G. Treadaway; “The Brahmaeidae (Lepidoptera) of the Philippines” by W. A. Nassig and C. G. Tread- away; “Arguda sandrae n. sp., a new lasiocampid (Lepidoptera: Lasio- campidae) moth from Palawan, Philippines” by A. Zwick, “Samia treadawayi (Lepidoptera: Saturniidae), a new species from Palawan Island, Philippines” by S. Naumann; “Two new species of the genus Cyanosesia Gorbunov & Arita, 1995 (Lepidoptera: Sesiidae) from the Oriental Region” by O. G. Gorbunov and A. Kallies; “The genus Eoophyla Swinhoe, 1900 (Lepidoptera: Crambidae: Acentropinae) from the Philippine Islands” by W. Speidel; The Scopariinae and Heliothelinae stat. rev. (Lepidoptera: Pyraloidea: Crambidae) of the Oriental Region—a revisional synopsis with descriptions of new spe- cies from the Philippines and Sumatra” by M. Nuss; “New records of Cosmopterix Hiibner, [1825] (Lepidoptera: Cosmopterigidae) from the Philippines by W. Mey; “Contribution to the knowledge of the caddisflies (Insecta: Trichoptera a) of the Philippines. 2. The spe- cies of the Mt. Agtuuganon Range on Mindanao” by W. Mey. The issue starts with a brief, yet useful summary about the peo- ple, climate, forests, biogeography, and geological history of the archipelago. This general, introductory paper makes it easier to un- derstand the rest of the papers, specifically the biogeographic and faunistic sections. Four Lepidoptera families are thoroughly reviewed for the arch- ipelago for the first time: Sphingidae, Lasiocampidae, Brahmaeidae, and Saturniidae; as well as the genus Eoophyla (Crambidae). The first of these papers is an annotated checklist of the Sphingidae known from the Philippines; 116 out of 117 species are illustrated in 18 color plates, including two new species and one subspecies. There is an analysis of the number of species and endemic taxa for each subfamily, tribe, and genus of Philippine Sphingidae; an evalu- ation of the richness and endemicity for the nine largest islands; and an evaluation of Sphingidae endemicity for each of Vane-Wright’s faunal regions (R. I. Vane-Wright, 1990, The Philippines—key to the biogeography of Wallacea? Pp. 19-34. In Knight, W. J. and J. D. Holloway (eds.), Insects and the rain forests of South East Asia (Wallacea), London, Royal Entomological Society, iv+343 pp.) The distribution of taxa is summarized in 23 maps. The next paper is an annotated checklist of the Lasiocampidae. Sixty-one species are noted from the Philippines, including one new genus, 18 new spe- cies and 6 subspecies; all illustrated in 12 color plates. The distribu- tion of the species is figured in 34 maps, and the genitalia of most are illustrated in 13 black and white plates. In another contribution, 23 species of Saturniidae reported from the Philippine Islands (in- cluding 2 new species and 4 subspecies) are described, discussed, and illustrated in 13 color plates. In addition, pre-imaginal instars are depicted in 6 color plates. The male and female genitalia of most taxa are illustrated in 20 black and white plates; the known distribu- tion of Philippine Saturniidae is presented in 16 maps. The degree of endemicity for each island and zoogeographical region is dis- cussed. For the Brahmaeidae, the imaginal morphology, phenology, distribution, and variation of the only species present in the Philip- pines is thoroughly discussed. These papers will certainly be most often consulted by most lepidopterists, but the remaining six lepi- dopteran papers are providing important advances to our knowledge on Oriental Lepidoptera. In three brief papers, a new lasiocampid, a new saturniid, and two new sesiids are described from the Philippines, respectively. The genus Eoophyla (Crambidae) from the Philippines is reviewed, JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY and two new species are described. Similarly, in the Cosmopterigi- dae two new species are described and two new records are re- ported for the country. On a more ample basis, the Scopariinae and Heliothelinae (Crambidae) of the Oriental region are reviewed; 11 genera and 63 species (including 6 new species) are recognized. This paper includes diagnoses and phylogenetic remarks for the higher taxa. In the last paper, a checklist of the fauna of caddisflies of the Mt. Agtuuganon Range on Mindanao is presented. An amazing number of 63 species out of the 102 listed are described and male genitalia illustrated for the first time. It is important to note that special care was taken in order to in- sure accuracy in the localities used in the distribution discussions or maps. This is significant because specimens from commercial traders were usually collected or reared by local people on several is- lands, stored in the house of a Filipino trader with little or no data associated, imported to Europe, and then sold to customers. Thus, label data and the origin of most Philippine specimens in private col- lections (and probably in many museums) have been dubious as a result of the trade practices common in the area. Very few criticisms can be made to this series of papers which are apparently free of misspellings and typographical errors. Although the maps are small, they are very clear, but the font size used for the species names make the legends almost illegible. In some cases the color photographs are too small and some have shadows that may hamper character observation. This volume stresses the faunistics and taxonomy of Lepidoptera and contributes considerably to improve our knowledge of the Philippine insect fauna. This book is a must in the library of any in- dividual interested in Sphingidae, Saturniidae, and Lasiocampidae in particular, or Oriental Lepidoptera in general. It should be pres- ent in all libraries that maintain coverage in entomology worldwide. MANUEL A. BALCAZAR-LARA, Departamento de Zoologia, Insti- tuto de Biologia, UNAM, Apdo. Postal 70-153, 05410, Mexico, D.F. MEXICO Journal of the Lepidopterists’ Society 53(1), 1999, 46-47 THE AFROTROPICAL TIGER-Motus, by D. T. Goodger and A. Wat- son. 1995. Published by The Natural History Museum, London, and Apollo Books. 65 pp., 4 color plates of adults, 109 black-and-white photographs of genitalia. Softcover, 29.6 x 21.0 cm, ISBN 87-88757- 32-3. Available from Apollo Books Aps., Kirkeby Sand 19, DK-5771, Stenstrup, Denmark, Tel. 45-62-26-3737 FAX: 45-62-26-3780. Price DDK 200.00 (approximately U.S. $32.00). The purpose of this book, in the authors’ words, is to serve as an illustrated catalogue, with generic diagnoses and species distribution of the currently recognized and described afrotropical Arctiinae. The layout of the work is straightforward and easy to follow. There is a brief synopsis, introduction, comments on the structure of the cat- alogue entries, as well as a list of genera and species removed from the Arctiinae, followed by the main body of the catalogue, which oc- cupies some 20 pages. In the catalogue, generic entries are kept short and concise, and generally follow the pattern established in the well-known series Generic Names of Moths of the World. Information provided in- cludes the name, author, date of publication and pagination, fol- lowed by a similar statement on the type species. Also listed are ju- nior synonyms and homonyms. The entries on species include, again, the name, author, date of publication and pagination, and a statement (in parentheses) of the genus in which the taxon was originally published. Oddly, in the many cases where species were subsequently transferred to another genus, the authors do not indicate this by placing author and year in parentheses. The only explanation I can think of is that this was done in order not to interrupt the flow of text as the pagination is given immediately after the year (e.g., in an entry under Alpenus Walker: VOLUME 53, NUMBER 1 affiniola Strand, 1919:168 (Diacrisia)). Whatever the reasoning, this manner of citation is confusing and violates article 51(c) of the In- ternational Code of Zoological Nomenclature [Article 51(c) states: “Ifa species-group name is combined with generic name other than the original one, the name of the author of the species-group name, if cited, is to be enclosed in parentheses” |. Further information pro- vided in the species entries includes a brief statement about the na- ture of the type material and a similarly brief indication of distribu- tion. Inset appear junior synonyms, homonyms, incorrect spellings, or infrasubspecific names. A further irritation to me was the disregard shown to articles 31(b) and 34(b), dealing with the of late much-discussed problem of agreement in gender [Article 31(b) states: “A species-group name, if it is or ends in a Latin adjective or participle in the nominative sin- gular, or is latinized, must agree in gender with the generic name with which it is at any time combined, and its termination must be changed according to Latin inflection, . . .”; Article 34(b) states: “The termination of a Latin or latinized adjectival or participial species- group name must agree in gender with the generic name with which it is at any time combined; if the termination is incorrect it must be changed accordingly (the author and date of the species-group name remain unchanged Arts 50c(ii), 23c”]. This is especially obvious in cases such as the species-rich genus Spilosoma, which, despite its ending, is in fact neuter. I am well aware that many readers of these lines will argue that I am nit-picking and many of the rules govern- ing the use of the classical languages have become obsolete. I dis- agree, but even if I did not, the Code is quite unambiguous about the matter. Following these entries (said to cover 411) are listed the 20 gen- era and their constituent species removed from the Arctiinae. This section has the same layout as the main catalogue. Most of these genera are now placed in the Noctuidae, and it is a pity that no indi- cation is given of the subfamilies to which they are likely to belong, as this would have made making changes much easier. Given the more than doubtful monophyly of many noctuid subfamilies, how- ever, it seems quite possible that their affinities are simply not known. The text is complimented by a list of recorded hostplants of afrotropical tiger-moths and a bibliography with some 311 entries. The four color plates of adults are of high quality and depict mostly type species. Similarly, the black-and-white photographs il- lustrate the male genitalia mostly of type species. Here, depth of field is occasionally lacking, although this is hardly surprising given the frequently robust genitalia in this family. Allin all, this little book will serve as a useful introduction to, and overview of, this beautiful group of moths in the afrotropics. Its main strengths lies in its conciseness, but I feel it would have prof- ited from a little more attention to detail. MARTIN KRUGER, Lepidoptera Department, Transvaal Museum, P.O. Box 413, 0001 Pretoria, South Africa. Journal of the Lepidopterists’ Society 53(1), 1999, 47 THE Morus OF AMERICA NORTH OF MEXICco, fascicle 27.3, Noc- tuoidea, Noctuidae (part), Noctuinae (part Noctuini), by J. D. La- fontaine. 1998. Published by the The Wedge Entomological Re- search Foundation, Washington, D.C. 348 pp., 8 color plates. Soft cover, 8 x 11 inches, ISBN: 0-933003-09-9. Available from The Wedge Entomological Research Foundation, 85253 Ridgetop Drive, Eugene, OR, 97405, USA. $115.00+postage ($4.00 U.S., $5.00 else- where). Also available from Bioquip Products and Entomological Reprint Specialists. In October 1998, someone asked about the classification of the Lepidoptera via the Internet. Several persons replied: “There is no one universally agreed upon classification.” The classification of 47 Lepidoptera is not cast in stone, nor will it be anytime in the near fu- ture. Don Lafontaine’s superb work exemplifies the answer given above. The Board of Editors of The Moths of America North of Mex- ico series, affectionately known as MONA, deserves recognition for advancing classifications rather than casting one system in stone. This is the second fascicle of the MONA series, artfully written by Lafontaine, on part of the Noctuini with a discussion on the clas- sification of the tribe. He plans a third fascicle on the tribe Agrotini. Such facts would be simple except that a previously published fasci- cle of the MONA series by Robert W. Poole (1995, Noctuoidea, Noc- tuidae (part) in Dominick, R. B. et al., The Moths of America North of Mexico, fasc. 26.1:1-249) proposed a classification that signifi- cantly altered the definition of the subfamily Noctuinae. I don’t know if either classification is correct, but I am very pleased that the MONA series can be fodder for discussions about the classification of Lepidoptera, in addition to producing excellent monographic works. I seek knowledge, I love to learn, and the MONA series— well amore! Most afficionados of Lepidoptera are already familiar with the MONA series. It is recognized for its high quality, authoritative look at moths, and perhaps best of all, the fascicles introduce and illus- trate little known species, thus popularizing the study of moths. La- fontaine’s fascicle does not let us down. The volume starts with a morphological, systematic, and taxo- nomic overview of the group. One hundred sixty-nine species in 31 genera are included. Four new genera are proposed and 21 new species are described. Many new combinations are presented. Re- vised nomenclature abounds. Complete citations are provided for persons to fully understand Lafontaine’s overview and philosophy. Drawings supplement the morphological descriptions. The bibliog- raphy is rich with entries. A subtle digression from MONA’ perceived format is the inclu- sion of three species from Mexico. This important change allows the author to more fully describe and explain his groupings, and the reader will recognize these species if one day they are found to be part of the U.S. fauna. Lafontaine makes full use of keys to genera and species, a feature I find especially helpful when I want to know how the author differentiates taxa. Lafontaine is an excellent writer. His verbiage is succinct and lu- cid. Overviews and details are sufficient to allow a reader to under- stand the text, which is, as always, beautifully elucidated with illus- trations of genitalia, adults, larvae, and distribution maps. All of the illustrations are superior to what can commonly be found in scien- tific publications. My grammar teachers would disagree with the structure of only a couple of sentences. An extremely unfortunate trend in the book binding industry provides my only criticism of this volume. Breaking with tradition of the MONA series, this book is not Smythe sewn, rather it is perfect bound—which is in my opinion, highly imperfect. Smythe sewing is expensive, and as binders become convinced of the invincibility of glue, there are fewer binders who use the superior Smythe sewing. For persons who bind their books in buckram, as do I, perfect bind- ing creates cramped inner margins and books that will not lie flat open. Persons who do not bind their books will also notice that this volume will not gracefully lie flat when opened. I cannot predict that pages will fall out; time and use will test the glue. I don’t want anyone to miss the point that I give Lafontaine’s work, and this volume high marks. All Bee interested in the Noc- tuidae, on a worldwide basis, will need this information. This book will be a standard for a long time because it treats so many species of economic importance, i.e., cutworms. Other persons should be fa- miliar with this work so that they can explore the analysis, fully ex- plained by Lafontaine, employed in writing a revision of this magni- tude. I highly recommend it. Any person interested in moths should own a copy. Eric H. METZLER, 1241 Kildale Square North, Columbus, Ohio 43229-1306. 48 Journal of the Lepidopterists’ Society 53(1), 1999, 48 SATURNIIDAE MUNDI: SATURNIID MOTHS OF THE WORLD, Part 3, by Bernard D’Abrera. 1998. Published by Goecke & Evers, Sport- platzweg 5, D-75210 Keltern, Germany (email: entomology@ s-direktnet.de), in association with Hill House, Melbourne & Lon- don. 171 pages, 88 color plates. Hard cover, 26 x 35 cm, dust jacket, glossy paper, ISBN-3-931374-03-3, £148 (about U.S. $250), avail- able from the publisher, also in U.S. from BioQuip Products. Imagine a large book with the highest quality color plates show- ing many of the largest and most famous Saturniidae from around the world! Imagine that this book shows males and females of all the 14 known species of Attacus, in addition to the equally massive Cos- cinocera and Archaeoattacus. Imagine that all of the American Roth- schildia were depicted, as well as all of the African Epiphora. Imag- ine many of the species of Samia, some recently discovered in and described from Indonesia and China, to be included. Imagine that the North American Callosamia, Hyalophora, and Eupackardia were included to complete the tribe Attacini. Imagine that this book had all of the beautiful Actias, Argema, and Graellsia, with their del- icate green, yellow, and pink coloration, and tailed hindwings. Now suppose such a book included all species of the confusing and com- plex group from the Northern Hemisphere usually placed in Satur- nia, Caligula, Neoris, Agapema, Eriogyna, and Perisomena, all neatly sorted out and figured. Add to all these, numerous other sat- urniids in well-known genera like Rhodinia, Loepa, Copaxa, and Cricula, and lesser-known genera like Syntherata, Opodiphthera, Lemaireia, and Pararhodia. If you can imagine all of those saturni- ids shown as life-sized illustrations, then you can begin to visualize the satisfaction that this book will bring to everyone who owns or uses it. This is the book for which lepidopterists who love Saturni- idae have been waiting! The book shows many species that are absent from most museum and private collections. For example, from China is shown the rich maroon colored Samia watsoni, the massive peach-colored Loepa oberthuri, and Actias dubernardi, the most gracile of all the Actias. From East Africa is the unique Argema besanti, the smallest yet per- haps the most beautiful of its genus, and from Indonesia we see Samia yayukae, Samia naumanni, and Cricula hayatiae. Included also are several recently described species such as Neoris codyi from Pakistan, named for the American saturniid artist John Cody, Opodiphthera excavus from Queensland that forms its cocoons be- low ground to avoid sweeping fires, Samia treadawayi from the Philippines, Actias angulocaudata and Loepa obscuromarginata both from China, Agapema platensis from Texas, Copaxa evelynae from Guatemala, and Rothschildia renatae from Peru. None of the latter three species could be included in Claude Lemaire’s revision of the American representatives of the subfamily Satumiinae (1978, Les Attacidae Américains, Edition C. Lemaire, Neuilly-sur-Seine, France, 238 pp., 49 pls.), because they had not yet been discovered in 1978. I will provide some taxonomic commentary. In the book, D’Ab- rera describes two new species: Opodiphthera goodgeri and Pararhodia setekwa, both from New Guinea. The figure of Actias chapae from Chapa, Vietnam (near its northern border with China), shows that it is a very distinct species; I had not seen a figure or specimen of A. chapae until now. The figure of the Chinese Actias felicis still makes me think that this name is a synonym of A. gnoma from Japan and Siberia, an opinion I reached long ago. The taxo- nomic differences between Actias heterogyna and A. sinensis are not clear in the figures in this book. D’Abrera has shown good insight by sorting out some synonymies in Epiphora and Coscinocera. We now have a color figure of each pair of species of Opodiphthera, enabling JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY us to identify material from New Guinea in the sciron group, which includes several species that look much alike. Prior to this we only had a key published by E.-L. Bouvier (1936, Mem. Natl. Mus. Nat. Hist. Paris, 3:1—350), in which he called these species Neodiphthera. I agree with D’Abrera’s interpretation of the distribution of Attacus aurantiacus. As with D’Abrera’s similar books on Sphingidae and butterflies, this one is a pictorial guide to these moths, based largely on speci- mens in The Natural History Museum in London. In an effort to make the coverage as complete as possible, the author has done an exceptional job of gathering missing material to be photographed, receiving several loans and donations from Australia, Belgium, France, Germany, and the United States. He has largely succeeded; relatively few known species are missing. Missing also, but by de- sign, is the extensive genus Antheraea, because it will be treated in Part 2 along with the many African genera such as Imbrasia, Pseudobunaea, Gynanisa, Eochroa, Heniocha, Usta, Eustera, Ludia, and Microgone. Part 1 of Saturniidae Mundi was published in 1995 (reviewed in 1996, J. Lepid. Soc. 50(4):355—356) and included the magnificent moths in the genera CG opiopter yx, Arsenura, Automeris, Hemileuca, Citheronia, Eacles, and related groups. In Part 3, the text is not very extensive for each species, mainly giving the citation to the original description, diagnostic characters, and the distribution in general terms. An additional component is that for many species D’Abrera refers the reader to primary sources in literature where the life history and immature stages have been described and figured by other authors. D’Abrera does not use sub- genera and makes minimal use of the subspecies category, thereby preserving the most basic tenets of the binomial system of nomen- clature intended by Linnaeus. My taxonomic philosophy is in total agreement with his approach. D’Abrera’s taxonomy is both conser- vative and accurate. I found virtually no errors in spelling or fact. The book is extremely reliable as an authoritative source on these venerable moths. As in Part 1, the introductory section offers some photographic portraits of some of those who contributed significantly to our knowledge of Saturniidae in the past, and who are actively doing so in the present. These include Charles Oberthiir, Dr. Karl Jordan, Walter Rothschild, Dr. Stefan Naumann, Captain Ulrich Paukstadt, Laela Hayati Paukstadt, and Thierry Bouyer. All who consider Saturniidae to be among their special interests will want this book, but some will delay or avoid purchasing a copy because of the high cost. A book of this magnitude and physical quality simply cannot be sold these days for less money. Publishers and authors are not getting rich on these kinds of publications. Any- one who considers the price to be an obstacle should buy this one book instead of spending more money in the long run trying to buy several other books (some of which are out of print and hard to find) that still would not show as many color figures of saturniids. With a print run of only 1500 copies, I would not wait too long to buy one. Most of us will never be able to obtain most of the species shown in this book for our collections, no matter how intensively we collect, buy, and exchange specimens. However, we can have a complete collection of these moths in the form of fine color figures. For ama- teurs who rarely have the opportunity to see the foreign literature on Saturniidae, current and historical, this volume offers the best and really the only opportunity to become knowledgeable about the names of the many species that exist, where they are from, and what they look like. I highly recommend this book because it will prove to be both immensely useful and a true pleasure to own. It represents a milestone in the literature on Lepidoptera. RICHARD S. PEIGLER, Department of Biology, University of the In- carnate Word, 4301 Broadway, San Antonio, Texas 78209-6397 USA Date of Issue (Vol. 53, No. 1): 7 October 1999 EDITORIAL STAFF OF THE JOURNAL M. Deane Bowers, Editor Entomology Section University of Colorado Museum Campus Box 218 University of Colorado Boulder, Colorado 80309, U.S.A. email: bowers@spot.colorado.edu Associate Editors: Gerarpo Lamas (Peru), Kenetm W. Puiir (USA), Rosert K. Ropsins (USA), Feuix A. H. Spertinc (USA), Davin L. Wacner (USA), Curister WikLUND (Sweden) NOTICE TO CONTRIBUTORS Contributions to the Journal may deal with any aspect of Lepidoptera study. Categories are Articles, Profiles, General Notes, Technical Comments, Book Reviews, Obituaries, Feature Photographs, and Cover Illustrations. Obituaries must be authorized by the President of the Society. Requirements for Feature Photographs and Cover Illustrations are stated on page 111 in Volume 44(2). Journal submissions should be sent to the editor at the above address. Short manuscripts concerning new state records, current events, and notices should be sent to the News, Phil Schappert, Dept. Zoology, Univ. Texas, Austin, TX 78712. Requests to review books for the Journal and book review manuscripts should be sent to M. Alma Solis, Syst. Entomol. Lab, USDA, c/o National Museum of Natural History, MRC 127, Washington, DC 20560. 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Authors of Book Reviews and Obituaries are exempt from page charges. Correspondence: Address all matters relating to the Journal to the editor. PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A. CONTENTS CoNTRIBUTION TOWARDS THE STUDY OF THE PyraLINAE (PYRALIDAE): HISTORICAL REVIEW, MO NOMENCLATURE M. Alma Solis and Michael Shaffer... .:-oc:occoccc-ececeeeeeeteeeee nent POPULATION BIOLOGY AND WING COLOR VARIATION IN HELICONIUS ERATO PHYLLIS cae 2) QNOANGTEN Bic Preipasy ono BOOS es PETES Ns TENS Ss A HOSTPLANT-INDUCED LARVAL POLYPHENISM IN HYALOPHORA EURYALUS (SATURNIIDAE) Michael M. Collins aT A New Hyparopa FROM Costa Rica (GELECHIOIDEA: COLEOPHORIDAE: BLASTOBASINAE) David Adamski POPULATION DEMOGRAPHICS AND THE CONSERVATION STATUS OF THE UNCOMPAHGRE FRITILLARY Bouonia (Nympnanipsn): Amy LiSetdh © ou. Le vei eh ose ie eae le ee * (Danaus PLexippus L.) DURING FALL, 1996 William H. Calvert 0... ccc cee ceeceeceeeeve ee eeeeeeeeeeeeeeee Boox REviews TheAfrotropical'tiger-moths” Martin Katiger. so. > est ee ee eee The moths of America north of Mexico, fascicle 27.3, Noctuoidea, Noctuidae (part), Noctuinae (part Noctuini) Eric H. Metzler = Saturniidae Mundi: Saturniid moths of the world, Part 3 Richard S. Peigher ese: ec --eseeeee onsen eee cee eee eee MANUSCRIPT REVIEWERS), OOS p2 sea Pa pe oe ie a cen ee This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanance of Paper). Volume 53 1999 Number 2 ISSN 0024-0966 JOURNAL of the LEPIDOPTERISTS’ SOCIETY Published quarterly by THE LEPIDOPTERISTS’ SOCIETY Publié par LA SOCIETE DES LEPIDOPTERISTES Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN Publicado por LA SOCIEDAD DE LOS LEPIDOPTEROLOGOS 13 January 2000 THE LEPIDOPTERISTS’ SOCIETY EXECUTIVE COUNCIL Joun W. Brown, President Susan S. Borxin, Vice President Micuaet J. Smiru, Immediate Past President Mirna M. Casacranpe, Vice President Manuet A. Batcazar-Lara, Vice President Davin C, Irtner, Treasurer Ernest H. Winuiams, Secretary Members at large: Ronald L. Rutowski M. Deane Bowers George L. Balogh Felix A. H. Sperling Ron Leuschner Andrew V. Z. Brower Andrew D. Warren Michael Toliver Brian Scholtens EprroriaL Boarp Rosert K. Rossins (Chairman), Joun W. Brown (Member at large) M. Deane Bowers ( Journal) WituiaM E. Miniter (Memoirs) Puiu J. Scuaprert (News) Honorary Lire MEMBERS OF THE SOCIETY Cuar.es L. Remincton (1966), E. G. Munroe (1973), Ian F. B. Common (1987), Joun G. Franciemont (1988), Lincoin P. Brower (1990), Douctas C. Fercuson (1990), Hon. Miriam Rotuscui_p (1991), CLaupe Lemaire (1992), Freperick H. Rinpcr (1997) The object of The Lepidopterists’ Society, which was formed in May 1947 and formally constituted in December 1950, is “to promote the science of lepidopterology in all its branches, . . . to issue a periodical and other publications on Lepidoptera, to facilitate the exchange of specimens and ideas by both the professional worker and the amateur in the field; to secure cooperation in all measures” directed towards these aims. Membership in the Society is open to all persons interested in the study of Lepidoptera. All members receive the Journal and the News of The Lep- idopterists’ Society. Prospective members should send to the Assistant Treasurer full dues for the current year, together with their full name, address, and special lepidopterological interests. In alternate years a list of members of the Society is issued, with addresses and special interests. 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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, o Natural History Museum, 900 Exposition Blvd., Los Angeles, CA 90007-4057. Cover illustration: Antheraea montezuma (Sallé) (Saturniidae), male. A close relative of A. polyphemus, this species flies throughout much of Mexico. Pen and ink drawing by John T. Carrola Jr., University of the Incarnate Word, San Antonio, Texas. Ee a pe es Oe JOURNAL OF Tue LEPIDOPTERISTS’ SOCIETY Volume 53 Journal of the Lepidopterists’ Society 53(2), 1999, 49-54 1999 Number 2 A TECHNIQUE FOR EXTRACTION OF INTACT MITOCHONDRIAL DNA MOLECULES FROM LARVAE OF SATURNIID MOTHS (LEPIDOPTERA: SATURNIIDAE) FOR USE IN TAXONOMIC STUDIES RALPH M. CLARK Department of Biological Sciences, Plattsburgh State University of New York, 101 Broad Street, Plattsburgh, New York 12901, USA (e-mail: clarkrm@splava.cc.plattsburgh.edu) ABSTRACT. Analysis of mitochondrial DNA can yield information about evolutionary relationships. In this paper, a set of pro- cedures for the extraction and analysis of mtDNA from saturniid moths is described. Restriction fragment length polymorphisms re- veal differences in the mtDNA of the species investigated. The potential of this methodology to contribute to comparative studies of moth species is discussed. Additional key words: Hyalophora, Callosamia, RFLP, taxonomy, phylogeny. The use of mitochondrial DNA (mtDNA) in studies of taxonomic and evolutionary relationships in insect groups has become well established since Bultman and Laird (1973) extracted, purified and described the physical characteristics of mtDNA from Drosophila melanogaster Meigen. Brower and Boyce (1991), Sperling (1993), Sperling and Harrison (1994), and Brower's (1994) extensive taxonomic and phylogenetic studies on butterflies are examples of the application of mtDNA organization to phylogenetic studies of Lepidoptera. I became interested in carrying out mtDNA studies and began to learn the basic techniques of mtDNA ex- traction, purification and analysis. I chose to work with the genus Hyalophora Duncan (Lepidoptera: Saturni- idae) because I had been rearing and studying H. ce- cropia (L) and H. columbia gloveri Strecker for many years. The several species, subspecies and hybrid pop- ulations of the genus occur in geographic areas that are distinct from each other but overlap (Scriber & Grab- stein 1991, Collins 1997). They also show food prefer- ences and thus occupy slightly different niches (Ober- foell 1969, Scriber & Grabstein 1991, Collins 1984). The genus Hyalophora is presently considered to consist of three species: H. cecropia L., H. columbia Smith with the subspecies H. columbia gloveri Strecker and H. columbia columbia Smith; and H. eu- ryalus Boisduval with one subspecies, H. euryalus cedrosensis Cockerell. This last subspecies, found only on the Isla de Cedros, Baja California, Mexico, was thought to be extinct until rediscovered and described by Smith and Wells (1993). This organization of the genus was developed by Lemaire (1978) and reflects the studies of zones of hy- bridization between H. euryalus and H. columbia gloveri carried out by Sweadner (1937), repeated and extended by Collins (1973) and Kohalmi and Moens (1975, 1988). It is the classification used by Tuskes et al. (1996) and by Collins (1997). Molecular evolutionary studies of saturniids include those of Collins et al. (1993) whose analysis of the dis- tribution of 20 allozymes confirmed the hybrid nature of populations where H. euryalus and H. columbia gloveri came in contact with each other. Legge (1993) used DNA primers for the cytochrome oxidase II (COI) gene, in conjunction with polymerase chain re- action (PCR) amplification, to isolate COII genes form total genomic DNA of several species of Hemileuca. Legge (1993) was able to use the nucleotide sequences of COII genes to construct consensus trees for phylo- 50 genetic analysis of species of the genus Hemileuca. Friedlander et al. (1998) used the sequence informa- tion of two nuclear genes to construct consensus trees showing the phylogenetic relationships of species of the Attacini, including the three Hyalophora species, and the Saturniini. To my knowledge, no one has yet reported the iso- lation of complete mtDNA molecules from Saturniid species. Having the complete molecule would be an asset to anyone investigating phylogenetic relation- ships between moth species. It turns out that the moth tissue having the greatest quantity of mitochondria is the gut of mature larvae. Anderson and Harvey (1966) studied the fine structure of the H. cecropia midgut epithelium and found that both the microvillae which extended into the gut lumen and the channels formed by the deep infoldings of the apical and basal plasma membrane were packed with mitochondria. This rich source of mitochondria was used to initiate a compar- ative study of the mtDNA of Hyalophora species. In this paper I present the methodology developed and some initial results of this study. MATERIALS AND METHODS All glassware, Eppendorf tubes, pipet tips and solu- tions are autoclaved prior to mtDNA processing. Items that are going to be reused are washed in hot water containing Alconox ™ detergent, then rinsed in tap water, followed by a rinse in deionized water. These are air dried, placed in glass containers which are capped with aluminum foil, and autoclaved for 30 min at 120°C. Plastic gloves are worn during extraction procedures to reduce the chance for contamination of the mtDNA. Mitochondria are taken from the guts of healthy 4" or 5" instar larvae which have been cleaned of their contents. The larvae used for this study were reared from ova deposited by 15 H. cecropia, 8 H. columbia gloveri, 2 H. columbia columbia, 7 H. euryalus, and 2 Callosamia promethea Drury. The last were used as an outgroup. The sources of ova are listed in Table 1. Larvae, except as noted below, are laboratory reared in plastic sweater boxes on fresh leaves of cherry (Prunus sp.) or white oak (Quercus alba L.) which have been sprayed with an antibiotic solution (Riddiford 1967). The number of larvae per box is kept small and larvae are handled as little as possible. H. co- lumbia are reared on branches of Larix laricina (DuBoi) K. Koch. Mature larvae are anesthetized under carbon diox- ide gas, decapitated and severed near the posterior. The body wall is cut through along the ventral side and the larva is pinned out. Next, the gut is cut longitudi- JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TaBLE 1. Suppliers of the ova used in this research. Moth species Supplier H. cecropia + Ralph M. Clark, Plattsburgh, NY Larry Kopp, Klingerstown, PA + * James Oberfoell, Bowman, ND Mark Schmidt, Springboro, OH Scott Smith, Santa Rosa, CA + Dean Morewood, Victoria, BC, Canada + Norman Smith, Fresno, CA Scott Smith, Santa Rosa, CA Gardiner Gregory, Orland, ME William Kenny, Dixemont, ME ° Mark Schmidt, Springboro, OH + Ted Herig + * James Oberfoell, Bowman, ND Mark Schmidt, Springboro, OH Scott Smith, Santa Rosa, CA Mark Schmidt, Springboro, OH Larry Kopp, Klingerstown, PA Mark Schmidt, Springboro, OH H. euryalus H. columbia columbia H. columbia gloveri A. polyphemus C. promethea + Native moths reared by supplier from representatives of local populations. * Present address unknown. ° From stock supplied by W. Kenny. nally along its ventral side and the gut contents are re- moved. In the best of cases, the peritrophic membrane can be lifted and rolled forward to remove the gut con- tents cleanly. The tracheal trunks that serve the gut are severed. The gut is removed from the body cavity, washed thoroughly in deionized water and transferred to containers chilled in ice or dry ice. Most tissues are weighed and stored at —70°C within the hour. Each sample consists of 4 or 5 guts. These samples are processed to extract mtDNA as soon as possible. Tis- sues that are to be processed immediately are trans- ferred to a chilled Wheaton dounce homogenizer which contains 5 ml of a physiological buffer (0.44 M sucrose, 0.01 M Tris, 0.18 mM EDTA; pH 7.5). Tis- sues in the dounce, either fresh or frozen, are macer- ated and the mtDNA is extracted using a procedure described by Tamura and Aotsuka (1988), with the modifications given below. Differential centrifugation in a physiological buffer is followed by alkaline lysis of the mitochondrial frac- tion. A centrifugation step separates the mtDNA from cellular debris and the mtDNA is further purified by phenol extraction. The mtDNA is precipitated from an alcoholic solution, resuspended in T,,E, and treated with the enzyme, RNase, before storage at —20°C. Standard endonuclease digestions are carried out using a variety of endonucleases. A digest of lambda DNA with Hind III is used to generate a reference kilobase pair (Kbp) ladder against which to size re- striction fragments. The digests are loaded into the wells of 1% agarose (GIBCO/BRL) gels. Electro- phoresis is carried out in a Hoefer Scientific Instru- VOLUME 53, NUMBER 2 = een Swalene. os Bie Ow {50 [e) oO (Se oe) HOO 6 2a se fz ah Sy ee ae ee ih) Be f ——_— —_— — ol Hind III Xho | 5 A DNA Eco RI = Y) ja =: Hind III Pst | Xho | ZDNA Msp | Hpa | Ava | ADNA Eco RI Hpa II Xba | Fie. 1. ments unit set at 37V, 23 milliamps. TBE is the elec- trophoretic buffer. The gel slab is stained with ethidium _ bromide, then destained in 0.1 mM MgSO, before ex- amination under ultraviolet light. Gels are pho- tographed and common pins are positioned in the gel to mark the position of mtDNA fragments. Measures of distances of the pins from the wells are taken and used to construct gel replicas and to determine the sizes of the mtDNA fragments in kilobase pairs (Kbp). Representative electrophoretic patterns for A) H. columbia columbia; B) H. euryalus; C) H. cecropia; and D) C. promethea. RESULTS The photographs in Fig. la, b, c, d are examples of electrophoretic patterns obtained when mtDNA from larvel guts is digested with these endonucleases. A number of bands the size of the largest A-DNA frag- ment or larger is seen in Fig. 1c. These are taken as ev- idence of incomplete digestion; the presence of non- mt-DNA in the sample, and/or from complexes 52 TABLE 2. Number of endonuclease sites observed in mtDNA of four saturniid species for the listed enzymes. Enzyme Species Hind II Msp I (Hpa II) Hpa I Pst I H. cecropia 3 2 3 2 H. columbia 3 3 3 3 H. euryalus 3 2 2 3 C. promethea D} 3 3 3 formed when the DNA concentration in a sample is excessive. The patterns of cleavage for these species are very consistent. Not shown is the pattern for H. columbia gloveri which is identical to H. columbia columbia. Eight of the endonucleases tested to date yield infor- mation useful for phylogenetic analysis. These are Hind III, XhoI, PstI, MspI, EcoRI, Aval, Hpal and Hpall. The isoschizomers MspI and HpallI have iden- tical cleavage patterns indicating that the cytosines of 5-CCGG-3 sequences are not methylated. Xbal results are very inconsistent. This enzyme is known for aber- rant cleavage if exact reaction conditions are not fol- lowed (Gibco/BRL 1997-98 Catalog). Among the en- donucleases that do not cleave these saturniid mt DNAs are Bell, Haell, BglII and Bam HI (Fig. 1c). Species specific differences in cleavage patterns are observed when the photographs in Fig. 1 are com- pared (Table 2). There are no observed differences in the number of cleavage sites for the other endonucle- ases. The two small Hind III cleavage sites are about 2.2 and 1.7 kilobase pairs (Kbp) long and difficult to see as they stain faintly with ethidium bromide, and photograph poorly. The single band seen when Hyalophora mtDNA is cut by EcoRI (Figs. la, b, c) is judged to be about 920 Kbp long, based on its position relative to the 960 Kbp band of the A-Hind III standard. Since all my results indicate that moth mtDNA is between 14-19 Kbp in length (a size range common to animals), EcoRI ap- parently cuts the mtDNA in two places which yields two fragments of approximately equal size. The two fragments comigrate. The C. promethea mtDNA is also cut at two locations by Eco RI, but the product is two fragments of unequal size, thus there are two bands (Fig. 1d). The size of these two fragments totals to about 16 Kbp. This is one of two examples seen where RFLPs result from an internal reorganization of mtDNA, not from the gain or loss of the number of enzyme sites. A second example is seen when the Hpal digest pattern of C. promethea is compared to the hyalophorans (Fig. 1). JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TABLE 3. Haplotypes of species of Hyalophora and C. promethea based on electrophoretic patterns after digestion of mtDNA with the listed enzymes. H. cecropia’s haplotype is used as the basis for com- parison. RFLPs are indicated by letter changes. The two sub-species of H. columbia exhibit identical haplotypes. Enzyme Species Hind 111 MspI(Hpall) Hpal Pst I Ayal XhoI EcoRI H. cecropia A A A A A A A H. columbia A B A B A A A H. euryalus A A B B A A A C. promethea B B C B A B B Table 3 presents the haplotypes of the species stud- ied, based on RFLPs. If one elects the H. cecropia haplotype as the base against which to compare the others, then one can see that the H. cecropia restric- tion pattern differs from H. columbia and H. euryalus for two of seven enzymes, but not the same enzymes; from the C. promethea haplotype for 5 of 7 enzymes. H. columbia haplotypes show two mtDNA modifica- tions from H. euryalus and three from C. promethea. H. euryalus differs from C. promethea at five sites, but not the same five by which C. promethea and H. cecropia differ. The PstI digest pattern of H. cecropia is unique. Most of these polymorphisms are caused by the gain or loss of sites for enzyme attack, however the uniqueness of the C. promethea haplotype may result from RFLPs created by internal rearrangements of the mtDNA (Hpal and EcoRI digests). DISCUSSION The major reason for this paper is to present proce- dures for the extraction, partial purification and analy- sis of complete molecules of moth mtDNA. This has been accomplished by adapting procedures of others to the extraction of mtDNA from the guts of mature moth larvae. The results show that larval lepidoptera are good sources of mitochondrial DNA for use in studies of taxonomy and phylogeny. Even though the mtDNA obtained using this methodology was not highly purified, and even though the electrophoretic stain used was not the most sensitive, results are quite reliable. Distinct RFLPs and haplotypes of the four saturniid species were observed. Contamination of the mtDNA with nuclear DNA was not a problem. Tamura and Aotsuka (1988) stated that the purity of mitochondria as a result of differen- tial centrifugation is not important because the alka- line lysis procedure efficiently separates covalently closed circular mtDNA from linear DNA. Legges’ (1993) concern that mtDNA from gut tissues could be seriously contaminated by gut organisms are allayed by VOLUME 53, NUMBER 2 the results presented here. By using the procedures of Jones et al. (1988) greatly improved purification and fragment resolution is possible. The advantage of this method over that utilized by Legge (1993) and others to obtain and sequence the Coll gene is that the entire mtDNA molecule is ob- tained. This allows studies of RFLPs and the construc- tion of restriction maps of the mtDNA chromosome based on double-digest studies. Additionally, sequenc- ing of entire mtDNA molecules is possible. The results of such studies will reveal any internal rearrangement or mutations of the mtDNA molecule that is species or population specific. Two such internal rearrangements have been discovered during this study (Table 3). Se- quencing of the entire molecule may reveal any intro- gression of genetic sequences were hybrid zones exist. Kondo et al. (1990) presented clear evidence of het- eroplasmy in Drosophila and the occurrence of intro- gression of mtDNA. Introgression of mtDNA in Drosophila was also observed by Aubert and Solignac (1990). If introgression is found in natural hybrid zones it could reveal cross-overs between native and introduced mtDNAs and help clarify evolutionary re- lationships within Hyalophora. It could also reveal the existence or establishment of subpopulations within a population by revealing two or more maternal lineages that are established through reciprocal crosses. Table 3 is based on a system used by Avise and Nel- son (1989) to illustrate relationships between the genomes of seaside sparrows that were dispersed over a wide geographic area of the southeastern United States. I have adapted this system as it seems to illus- trate haplotype differences most clearly. The distinctness of the C. promethea mtDNA from that of Hyalophora is readily apparent as 3 of the 7 en- zymes used to digest the samples yield results that are unique to C. promethea. This pattern would suggest a separate line of evolution for the Callosamia and would corroborate the findings of Friedlander, et al. (1998) and Johnson, et al. (1996). One can infer that the identity of the digest patterns of the two sub- species of H. columbia indicates that these two are more closely related to each other than they are to the other Hyalophora. Also, the PstI digest pattern which is unique to H. cecropia suggests an evolutionary sep- aration from the others, with the PstI pattern of C. promethea possibly being more ancient and unchanged in columbians and euryalus. However one must be aware that both forward and reverse mutations occur. There is not enough information here to make fur- ther inferences as to the degree of relatedness and lines of evolution of the Hyalophora. However, noth- ing in these results contradict the findings of Sweadner 15 yo) (1937), Lemaire (1978), Tuskes, et al. (1996) or Collins (1973, 1997). It would be interesting to have samples of H. euryalus cedroensis mtDNA to study because of its potentially long-time isolation from the rest of the Hyalophora. The naming of the two H. columbia sub- species creates confusion because even though H. co- lumbia gloveri is believed to an ancestral to H. colum- bia columbia (Sweadner 1937, Collins 1997), the H. columbia name has priority. ACKNOWLEDGMENTS This research received financial support through in-house grants from the Office of Sponsored Research and Programs, and from the Department of Biological Sciences at Plattsburgh State University of New York, Plattsburgh, NY. I wish to ex- press my appreciation to the undergraduate students who as- sisted in the research, especially to Mary Guamieri, Kimberly Chesbrough, Lydia Grozzo, DruAnn Clark and Nicole Porter. Dean Morewood, Mark Schmidt, Scott Smith, and Ted Herig generously donated ova for this study. Thanks go to Dianna Seymour for help in preparing the paper and to my wife, Pa- tricia, who edited the paper and who put up with me. The edi- torial comments of Dean Morewood and Michael Collins are greatly appreciated. This paper is dedicated to my wife who passed away December 20, 1998, and to my daughter Theo, who gave loving care to her mother during the terminal weeks of her life. LITERATURE CITED ANDERSON, E. A. & W. R. Harvey. 1966. Active transport by the Cecropia midgut. II. Fine structure of the midgut epithelium. J. Cell. Biol. 31:107-134. AUBERT, J. & M. SoLicNac. 1990. Experimental evidence for mito- chondrial DNA introgression between Drosophila species. Evo- lution 44:1272-1282. AVISE, J. C. & W.S. NELSON. 1989. Molecular genetic relationships of the extinct dusky seaside sparrow. Science 243:646—648. Brower, A. V.Z. 1994. Rapid morphological radiation and conver- gence among races of the butterfly Heliconius erato inferred from patterns of mitochondrial DNA evolution. Proc. Nat. Acad. Sci. 91:6491—6495. BrRoweER, A. V. Z. & T. M. Boyce. 1991. Mitochondrial DNA varia- tion in monarch butterflies. Evolution 45:1281—1286. BuULTMAN, H. & C. D. Lairp. 1973. Mitochondrial DNA from Drosophila melanogaster. Biochem. Biophys. Acta 299:196—209. CoLiins, M. M. 1973. Notes on the taxonomic status of Hyalophora columbia (Saturniidae). J. Lepid. Soc. 27:225-235. . 1984. Genetics and ecology of a hybrid zone in Hyalophora (Lepidoptera: Saturniidae). U. Ca. Pub. in Ento- mology 104:1—-93. . 1997. Hybridization and speciation in Hyalophora (In- secta: Lepidoptera: Saturniidae): a reappraisal of W. R. Swead- ner’s classic study of a hybrid zone. Ann. Carnegie Museum 66:411—455. CoLiins, M. M., H. B. Britren & V. Rivers. 1993. Allozyme analysis of a known hybrid zone between Hyalophora euryalus and H. columbia gloveri (Lepidoptera: Saturniidae) in the Cali- fornia Sierra Nevada. J. Res. Lepid. 32:79-88. FRIEDLANDER, T. P., K. R. Horst, J. C. REGIER, C. MITTER, R. S. PEIGLER & Q. Q. FANG. 1998. Two nuclear genes yield con- cordant relationships within Attacini (Lepidoptera: Saturni- idae). Molec. Phylog. Evol. 9:131-148. JOHNSON, K. S., D. SNIDER & J. M. ScRIBER. 1996. Estimates of ge- netic differentiation among Callosamia species and Hyalophora cecropia (Saturniidae) using allozyme electrophoresis. J. Lepid. Soc. 50:217—225. 54 Jones, C. S., H. TEGELSTROM, D. S. LATCHMAN & R. J. BERRY. 1988. An improved rapid method for mitochondrial DNA iso- lation suitable for use in the study of closely related popula- tions. Biochem. Gen. 26:83-—88. KouHALMI, L., & P. MoENS. 1975. Evidence for the existence of an intergrade population between Hyalophora gloveri nokomis and H. columbia in Northwestern Ontario (Lepidoptera: Sat- urniidae). Can. Entomol. 107:793—799. . 1988. Patterns of inheritance of larval tubercle-colour polymorphism in Hyalophora columbia of northwestern On- tario (Lepidoptera: Saturniidae). Genome 30:307-312. Konpo, R., Y. Satra, E. T. MATsuuRA, H. IsHiwa, N. TAKAHATA & S. I. Cuicusa. 1990. Incomplete maternal transmission of mito- chondrial DNA in Drosophila. Genetics 126:657—663. LEGGE, J. T. 1993. Genetic differentiation of fen-dwelling Hemi- leuca (Lepidoptera: Saturniidae) within the H. maia species group. Master’s Thesis, Cornell University Graduate School, Ithaca, New York. 121 pp. LEMAIRE, C. 1978. Les Attacidae Americains. Attacinae. Edition C. Lemaire, Neuilly-sur-Seine, France. 238 pp. OBERFOELL, J. 1969. Occurrence of the cecropia and Rocky Moun- tain silk moths in southwestern North Dakota. Prairie Natur. 1:63-64. RippirorbD, L. M. 1967. Antibiotics in the laboratory rearing of Cecropia silkworms. Science 157:1451-1452. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY SCRIBER, J. M. & E. GRABSTEIN. 1991. Food plant processing adap- tations in four Hyalophora species (Lepidoptera: Saturniidae); regional and taxonomic specialization. Great Lakes Entomol. 24:109-120. SMITH, M. J. & R. E. WELLS. 1993. Rediscovery of Hyalophora eu- ryalus cedrosensis (Saturniidae), with descriptions of the adult and larval stages. J. Lepid. Soc, 47:114-124. SPERLING, F. A. H. 1993. Mitochondrial DNA variation and Hal- dane’s rule in the Papilio glaucus and P. troilus species groups. Heredity 71:227-233. SPERLING, F. A. H. & R. G. Harrison. 1994. Mitochondrial DNA variation within and between species of the Papilio machaon group of swallowtail butterflies. Evolution 48:408—422. SWEADNER, W. R. 1937. Hybridization and the phylogeny of the genus Platysamia. Ann. Carn. Mus. 25:163—243. Tamura, K. & T. AoTsuKA. 1988. Rapid isolation method of animal mitochondrial DNA by the alkaline lysis procedure. Biochem. Gen. 26:815-819. Tuskes, P. M., J. P. TurrLe & M. M. Couns. 1996. The wild silk moths of North America: a natural history of the Saturniidae of the United States and Canada. Cornell U. Press, Ithaca, New York. 250 pp. Received for publication 8 April 1998; revised and accepted 31 Jan- uary 1999, Journal of the Lepidopterists’ Society 53(2), 1999, 55-59 A NEW SPECIES OF LITHOPHANE (LEPIDOPTERA: NOCTUIDAE: CUCULLIINAE) FROM NORTHEASTERN NORTH AMERICA REGINALD P. WEBSTER 94 Millstream Drive, Charters Settlement, New Brunswick E3C 1X1, Canada AND ANTHONY W. THOMAS Natural Resources Canada, Canadian Forest Service—Atlantic Forestry Centre, P.O. Box 4000, Fredericton, New Brunswick E3B 5P7, Canada ABSTRACT: A new species of noctuid moth is described and illustrated. Lithophane thujae, new species, is known from two localities in New Brunswick, Canada, and one locality each in Michigan and Wisconsin, U.S.A. The probable larval host plant is northern white-cedar (ar- bor vitae), Thuja occidentalis L. (Cupressaceae). Notes on its biology are given and the last instar larva is figured. Additional key words: Winter moths, Lithophane lemmeri, New Brunswick, Michigan, Wisconsin. In late April 1990 a female specimen of an unusual Lithophane was collected at mercury vapor (M.V.) light by the authors a few km south of Harvey Station, York Co., New Brunswick. This specimen was tenta- tively identified as Lithophane species near lemmeri Barnes & Benjamin by Donald Lafontaine of the Biosystematics Laboratory, Ottawa, Canada. Another female of this species was collected at M.V. light by the senior author near Fredericton, York Co., N.B., in early June 1992. Eggs were obtained from this female and neonate larvae were given a choice of 21 species of woody shrubs and trees from the area where the moth was collected. The larvae refused to feed on the foliage of all species except northern white cedar, Thuja occi- dentalis L. (Cupressaceae). Larvae were reared on the foliage of T. occidentalis, and adults were produced in late September, 1992. Additional specimens were reared in 1996 from another female collected at the lo- cality near Fredericton in early May at M.V. light. An- other two specimens of this species from Michigan and one from Wisconsin were located in the respective collections of Mogens C. Nielsen and James C. Parkin- son. These had been identified as Lithophane new sp. near lemmeri by Dale F. Schweitzer. Comparison of the male genitalia of these moths to those of L. lem- meri, which occurs along the Atlantic coast of the U.S.A., demonstrated that this Lithophane was not conspecific with L. lemmeri. We therefore describe this insect as a new species. Lithophane thujae Webster and Thomas, new species (Figs. 1, 2, 6, 7) Description (Figs. 1 and 2). Forewing narrow, length 16.0-18.1 mm (mean = 17.2), width at tornus 6.0—7.5 mm (mean = 7.0) (n = 24) in males and length 16.0-18.5 mm (mean = 17.3), width at tor- nus 6.0—8.0 mm (mean = 7.2) (n = 46) in females. Anterior half of termen at nearly right angle with distal 1/3 of costa, then angled in- wards toward tornus at approximately a 135° angle; lower half of ter- men slightly emarginated above tornus. Forewing above brownish gray grading to light gray near the anterior margin basally and brown with pinkish hue along posterior margin becoming salmon pink near base of wing. Reniform spot indistinct, dirty salmon pink, basal por- tion contrasting with lighter gray anterior portion. Orbicular spot ab- sent. Basal dash consists of thin black line. Thicker black median streak between indistinct antemedial and postmedial lines, and well developed subreniform black line outlined anteriorly with white. Apical dash small, black and subterminal line indicated by series of 4 black dashes, most anterior connected to brownish apical streak. Hindwing above darker than forewing and grayish brown with salmon pink hue, discal dot faintly expressed. Fringes of both wings concolorous with adjacent portions of wing. Underside of fore- and hindwings uniformly light gray with strong salmon pink tint except for darker central area of forewing. Discal spot on underside of hindwing well developed. Color and pattern similar in both sexes and uniform among specimens examined. Dorsal side of thorax gray with mid-dorsal brown patch bisected by fine white line extending from collar to base of abdomen. Tegula elongated and gray, mar- gined with black above wing base. Anteriorly, mesothoracic vestiture terminates in V-shaped crest. Patagium concolorous with tegula, but bordered posteriorly towards mid-line by fine black line followed by white line. These lines on adjacent patagia form a V, opening formed by thoracic crest which emerges between arms of V. Head vestiture, dorsally, concolorous with patagia and tegulae. Palpi and ventral tho- rax gray with definite pinkish hue. Dorsally, abdomen gray with pinkish hue and without tufts; ventrally abdomen approaches dirty salmon pink. Prothoracic legs entirely dirty salmon pink. Outer lat- eral portions of femur and tibia of meso- and metathoracic legs black, remainder of legs salmon pink Male genitalia (Fig. 6). The interpretation of male genitalia was based primarily on Sibatani et al. (1954). Basal part of valve, com- posed of dorso-proximal costa and ventro-proximal sacculus, well developed. Heavily sclerotized costa forms right angle at junction of its dorso-proximal and proximal margins. Valvula, ventroapical re- gion of valve beyond membranous annelifer, narrower than basal part of valve, but of equal length. Valvula gradually tapers to simple narrowly rounded apex and lacking digitus. Relatively short harpe curved to form open half circle with its opening facing ventro-api- cally and inner diameter of 0.5 mm. Transtilla curved with its smaller diameter on its mesal surface. Vincula beyond ventral edge of sac- culus formV-shaped structure with length longer than width. Two vincula meet at acute angle. Juxta ends in two heavily sclerotized points forming shallow V, with depth of V equal to half width of juxta at level of base of V. Uncus terminates in widened tip with slight bi- furcation, Male genitalia is bilaterally asymmetrical, with apical por- tion of left valvula being broader than right. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY VOLUME 53, NUMBER 2 b Fic. 6. Male genitalia of Lithophane thujae. a, Genitalia with aedeagus removed; b, Aedeagus. A.W.T. Genitalia vial 95.x.30-#26. Paratype, New Brunswick, York Co., ecl. 22 September 1992, R. P. Webster. Scale bar = 1 mm. Female genitalia (Fig. 7). Similar to, but smaller than L. lemmeri (Fig. 8c). Major structural difference is in ventral edge of ostium. In L. thujae, sterigma is non-sclerotized and forms a straight, horizon- tal, edge. It is totally overshadowed by heavily sclerotized lamella an- tevaginalis which is deeply invaginated giving ostium an apparent V-shaped opening. In L. lemmeri, sterigma is distinct, sclerotized, and has wavy edge. Diagnosis. The forewing pattern of L. thujae (Figs. 1 and 2) is similar to that of L. lemmeri (Figs. 3 and 4), but the markings are bolder and more completely developed in L. thujae. In L. lemmeri, the wings are a uniform dirty brownish gray, the black median streak between the antemedial and postmedial lines is poorly developed, and the subreniform line is less developed and not outlined anteri- orly with white. Rubbed specimens may be mis-identified as L. lem- meri, but the forewing patterns of fresh specimens of the two spe- cies are distinctively different (Figs. 1-4). L. lemmeri has longer narrower wings: forewing length 20.0—20.5 mm, width 7.0 mm (n = 4). The male genitalia of L. thujae and lemmeri are also substantially different (Figs. 6 and 8). In L. lemmeri (Fig. 8a), valvula has an ex- panded costal bulge towards its apex to form a rounded protuber- 57 Fic. 7. Female genitalia of Lithophane thujae. A.W.T. genitalia vial 95.x30-#25. Paratype, New Brunswick, York Co., ecl. 19 Sep- tember 1992, R. P. Webster. Scale bar = 1 mm. ance on its mesal surface and then narrows to form a distinctively bi- fureate tip, with the dorsal branch twice as long as the ventral branch (digitus). In L. thujae the valvula (Fig. 6a) lacks a costal bulge and gradually tapers to a simple narrowly rounded tip, without a digitus. Types. Holotype male (Fig. 1): CANADA, NEW BRUNS- WICK, York Co., 5.3 km SW of Jct. of Hwy. 101 and Charters Set- tlement Rd. (45°50°38’N, 66°44°31’W), ex ovum from female col- lected at M.V. light 1 June 1992, reared on T. occidentalis, emerged 24 September 1992, R. P. Webster. Allotype female (Fig. 2): same lo- cality and data as male, emerged 16 September 1992. Paratypes: Fics. 1-5. Adults of Lithophane thujae, new species and Lithophane lemmeri; and larva of L. thujae. 1, Lithophane thujae, holotype male. 2, Lithophane thujae, allotype female. 3, Lithophane lemmeri, male, New Jersey, Atlantic Co., Egg Harbor Twp., ex. ovum 2 April 1994, reared on Juniperus virginiana, ecl. 1-4 Nov. 1994, Dale F. Schweitzer. 4, Lithophane lemmeri, female, same data as male. 5, Mature larva of Litho- phane thujae on Thuja occidentalis. Length 30 mm. Reared ex ovum from a female collected at M.V. light on 3 May 1996 at New Brunswick, York Co., R. P. Webster. 58 Fic. 8. Male and female genitalia of Lithophane lemmeri. a, Male genitalia with aedeagus removed; b, Aedeagus. A.W.T. geni- talia vial 95.x.30-#23. New Jersey, Atlantic Co., Egg Harbor Twp., ex ovum 2 April 1994, reared on Juniperus virginiana, ecl. 1-4 Noy. 1994, Dale Schweitzer; c, Female genitalia. A.W.T. genitalia vial 95.x.30-#24. Same data as male. Scale bar = 1mm. 27 males and 47 females as follows: CANADA, NEW BRUNS- WICK: York Co., 3.5 km. S. of Jct. Hwy 3 & 4, Jct. Hwy 3 & Davis Brook, 27 April 1990 (1 female), A. W. Thomas & R. P. Webster; same locality and data as Holotype, emergence dates 16-30 Sep- tember 1992 (8 males, 14 females), R. P. Webster (2 males and 1 fe- male dissected, A. W. Thomas genitalia vials 95.x.30-#21 (male), 95.x.30-#26 (male), 95.x.30-#25 (female)); same locality as Holo- type, 8 October 1994, at M.V. light (1 female), R. P. Webster; same locality as Holotype, ex ovum from female collected at M.V. light 3 May 1996, reared on T. occidentalis, emergence dates 4-17 Sep- tember 1996 (17 males, 29 females), R. P. Webster; same locality as Holotype, 13 October 1996, at M.V. light (1 female), R. P. Webster. U.S.A., MICHIGAN: Otsego Co., T29N R2W Section 15, 27 April 1974 (1 male), 13 May 1994 (1 male), both M. C. Nielsen. WIS- CONSIN: Florence Co., T38N R19E Section 10, 11 October 1980 at light (1 female), J. C. Parkinson. Disposition of types. Holotype (no. CNC-22575) and allotype in the Canadian National Collection, Ottawa, Ontario; paratypes at the American Museum of Natural History, New York; Canadian Na- tional Collection, Ottawa, Ontario; Florida State Collection of Arthropods, Gainesville, Florida; National Museum of Natural His- tory, Washington, D.C.; Los Angeles County Museum of Natural History, Los Angeles, California; The Natural History Museum, London, England; Insect Reference Collection, Natural Resources Canada, Canadian Forest Service - Atlantic Forestry Centre, Fred- ericton, New Brunswick; and private collections of Henry H. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Hensel, Edmunston, New Brunswick; Mogens C. Nielsen, Lansing, Michigan; James C. Parkinson, Mosinee, Wisconsin; Dale F. Schweitzer, Port Norris, New Jersey; Anthony W. Thomas, Freder- icton, New Brunswick; Jim Troubridge, Agassiz, British Columbia; Martin N. Turgeon, St. Basile, New Brunswick; Reginald P. Webster, Charters Settlement, New Brunswick. Type locality. The type locality is within a small, partially wooded, residential area 5.3 km SW of the Jet. of Hwy. 101 and Charters Settlement Rd, about 8 km SW of Fredericton, New Brunswick, Canada. The forested area where the L. thujae speci- mens were collected is a mature second growth mixed conifer forest with a small brook passing through it. Dominant species of trees are T. occidentalis, Abies balsamea (L.) Mill. with scattered Salix sp., Be- tula allegheniensis Britt., B. papyrifera Marsh., Acer rubrum L., and Fraxinus americana 1. T. occidentalis is most abundant near the shaded stream. Openings along the stream are dominated by Alnus incana (1.) Moench. Etymology. The specific epithet thujae is named after the name of the genus of the probable host plant of this species. Biology. Lithophane thujae was reared on T. occidentalis on two occasions. In the initial attempt to rear this species, the larvae were offered 21 species of woody plants from the general area where the adults were collected. The larvae accepted only T. occi- dentalis and completed development on this host. T. occidentalis is common at the type locality where most specimens of this species have been collected to date and is present at all other known sites for this species and thus, we feel that T. occidentalis is the likely host plant for this species. The related L. lemmeri also feeds on a mem- ber of the Cupressaceae (Juniperus virginiana L.) (Dale F. Schweitzer, pers. comm. _). Lithophane thujae has five larval instars. A mature last instar of L. thujae is shown in figure 5. The mature larvae are 33.0 to 35.0 mm in length and range in color from green to dark green with a se- ries of longitudinal rows of white spots as shown in the figure. This pattern makes the larvae very difficult to locate on the foliage of their host plant. The second through fourth instar caterpillars exhibit a similar color pattern. A female collected on May 3, 1996, and confined to a 2-liter plastic container laid yellowish white eggs (0.89 mm diam., 0.69 mm high) near the tips of the underside of the foliage of T. occidentalis. The description that follows is based on observations of larvae main- tained on cuttings of T. occidentalis between 12° and 18°C. The eggs hatched 12-15 days after they were laid. The tan-colored first instar larvae gradually tured to light green as they fed near the tips of the new growth. The molt to second instar took place about six days later. The second instar larvae developed the color pattern that per- sisted through the last instar. The duration of the second, third, and fourth instars was each about six days under this temperature regime. The duration of the last instar was nine to eleven days. All larval instars fed near the tips of the new growth and usually con- sumed between 1 and 2 mm of the tip of the needle before moving to an adjacent needle. Last instar larvae also fed on the previous year’s growth. When development was completed (mid June to early July) the last instar larvae formed cocoons in the leaf litter. However, the larvae did not pupate until late August. Adults emerged in mid September under laboratory conditions. Flight season. Lithophane thujae has been collected in Octo- ber (3 freshly emerged specimens) and during April, May, and early June. Specimens collected in the spring were worn suggesting that L. thujae, like other Lithophane species, overwinter as adults and will be present in the fall and the following spring. Geographical distribution. South central New Brunswick, northern Michigan and northern Wisconsin. Probably has a broader distribution and will likely be found in intervening areas where T. oc- cidentalis is common. DISCUSSION Lithophane thujae is a member of an assemblage of Lithophane species which feed during the larval stage VOLUME 53, NUMBER 2 on members of the Cupressaceae. These species in- clude L. lemmeri from the Atlantic coastal plain of the U.S.A. from Connecticut southward (Dale F. Schweitzer, pers. comm.); L. subtilis Franclemont, L. tarda (Barnes and Benjamin), and L. longior (Smith) from southwestern North America (Franclemont 1969): and the Palaearctic L. leautieri Boursin (Bretherton et al. 1983). None of these species has a combination of wing pattern and male genitalia similar to L. thujae (Bames and Benjamin 1929, Boursin 1971, Bretherton et al. 1983, Franclemont 1969). Although the wing pat- tern of the more northern L. thujae and L. lemmeri are similar, the genitalia of the two are very different. Very few field-collected specimens are currently known of L. thujae. The reason for this apparent rarity is unclear. Possibly L. thujae is less attracted to light or bait than other Lithophane species, or has very specific habitat requirements. The current distribution in- cludes only New Brunswick, Michigan, and Wisconsin. However, it is likely that this species will be found in the intervening areas in Maine, northern New Hamp- shire, Quebec and Ontario once more collecting is done in habitats with T. occidentalis. ACKNOWLEDGMENTS Many thanks are due to Dale Schweitzer, Port Norris, New Jer- sey, for sharing unpublished data on the biology of L. lemmeri, sup- 59 plying specimens of L. lemmeri, and providing invaluable informa- tion on the location of specimens of L. thujae in eastern North American institutional and private collections obtained during a sur- vey of the distributional records of L. lemmeri. We thank Donald Lafontaine, Biosystematics Laboratory, Ottawa, Canada, for supply- ing information on the location of possible L. thujae specimens in The Canadian National Collection and other Canadian institutions. We thank Martin Honey for supplying information on L. leautieri and the location of possible L. thujae in The Natural History Mu- seum, London, England. We thank Mogens C. Nielsen, Lansing, Michigan, and James C. Parkinson, Mosinee, Wisconsin, for loaning specimens of L. thujae. We thank P. Z. Goldstein and two anony- mous reviewers for helpful suggestions on the manuscript. LITERATURE CITED BARNES, W. & E. H. BENJAMIN. 1929. Lepidopterological contribu- tions. I. Three new species of Phalaenidae from the New Jersey pine barrens. Bull. Brooklyn Entomol. Soc. 24:164-166. BoursIn, C. 1971. Lithophane lapidea Hb. et Lithophane leautieri Bsd. Entomops 20:129-135. BRETHERTON, R. F., B. GOATER & R. I. LORIMER. 1983. The moths and butterflies of Great Britain and Ireland. In John Heath and A. Maitland Emmet (eds.). Volume 10 Noctuidae (Cuculliinae to Hypeninae) and Agaristidae. Harley Books, Colchester, En- gland. 459 pp. FRANCLEMONT, J. G. 1969. Two new species of Lithophane from California (Noctuidae, Cuculliinae). J. Lepid. Soc. 23:10-14. SIBATANI, A., M. OcaTA, Y. OKADA & H. OKAGAKI. 1954. Male gen- italia of Lepidoptera: morphology and nomenclature. I. Divi- sions of the valvae in Rhopalocera, Phalaenidae (=Noctuidae) and Geometridae. Ann. Entomol. Soc. Am. 47: 93-106. Received for publication 10 May 1998; revised and accepted 8 April 1999. Journal of the Lepidopterists’ Society 53(2), 1999, 60-64 A NEW GENUS OF TORTRICID MOTHS (TORTRICIDAE: EULIINI) INJURIOUS TO GRAPES AND STONE FRUITS IN CHILE JOHN W. BROWN Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, % National Museum of Natural History, Washington, D.C. 20560-0168, USA (e-mail: jbrown@sel.barc.usda.gov) ABSTRACT: Accuminulia, new genus, is described and illustrated from Chile. The new genus includes two species: A. buscki, new spe- cies (type species), and A. longiphallus, new species. Accuminulia buscki has been reared from the fruit of grape (Vitis sp.; Vitaceae), plum (Prunus domestica; Rosaceae), apricot (Prunus armeniaca; Rosaceae), and peach (Prunus persica; Rosaceae) in Chile; the oldest specimen ex- amined is an adult intercepted at the port of New York in cargo (grapes) from Chile in 1926. Several specimens of A. buscki have been collected recently (1983) in traps baited with Proeulia-lure. The new genus is assigned to Euliini on the basis of its putative phylogenetic relationship to Proeulia Clarke. Additional key words: _ pest species, Neotropical, Euliini, systematics, Vitis, Prunus. In 1926 an adult of an undescribed tortricid moth was intercepted at the port of New York in cargo (grapes) that originated from Chile. The late August Busck, a lepidopterist at the National Museum of Natu- ral History, recognized the moth as representing a new genus and species, but did not describe it, probably ow- ing to the lack of sufficient material. Over the 70 years since that interception, numerous specimens of the spe- cies have accumulated—both sexes, the pupa, and larval food plants now are known. A second undescribed con- gener also is known from Chile. This paper describes and illustrates the new genus and its two species, and presents information on the biology of one of them. MATERIALS AND METHODS Taxonomic material was obtained from the follow- ing institutions: The Natural History Museum (BMNH), London, England; Mississippi Entomologi- cal Museum (MEM), Mississippi State, Mississippi, U.S.A.; Essig Museum of Entomology, University of California, Berkeley (UCB), California, U.S.A.; and National Museum of Natural History (USNM), Smith- sonian Institution, Washington, D.C., U.S.A. Dissec- tion methodology follows that summarized in Brown & Powell (1991). Illustrations of genitalia and wing ve- nation were drawn with the aid of a Ken-A-Vision mi- croprojector (model X1000-1). Forewing measure- ments were made with an ocular micrometer mounted in a Leica MZ12 dissecting microscope. Terminology for wing venation and genitalic structures follows Ho- rak (1984). Abbreviations and symbols are as follows: FW = forewing; HW = hindwing; DC = discal cell; n = number of specimens examined; X = mean; ca. = circa (approximately). SYSTEMATICS Accuminulia J. Brown, new genus Type species.—Accuminulia buscki J. Brown, new species. Description. Adult. Head: Antennal cilia ca. 0.5—0.8 times fla- gellomere diameter in male, ca. 0.1 times flagellomere diameter in female. Labial palpus (all segments combined) ca. 1.5 times hori- zontal diameter of compound eye, segment II weakly upturned, slightly expanded distally by scaling, segment III 0.2—0.3 times as long as II, smooth-scaled, exposed. Maxillary palpus rudimentary. Dorsal portion of frons with short overhanging tuft of scales. Ocelli present. Chaetosema present. Proboscis ca. as long as segment II of labial palpus, presumably functional. Thorax: Smooth-scaled. Legs unmodified, male foreleg without hairpencil. Forewing (Fig. 1): Length ca. 1.8 times width; length of DC ca. 0.68 times FW length; width of DC ca. 0.18 times DC length; CuA, originates ca. 0.70 along length of DC; all veins separate beyond DC; chorda weak but present; M-stem absent; CuP absent. Upraised scale tufts present (A. longiphallus) or absent (A. buscki); male without costal fold. Hindwing: Sc+R and Rs approximate; Rs to costa before apex; Rs and M, stalked ca. 0.4 distance; M, and M, approximate; M, and CuA, stalked ca. 0.3 distance; CuP present; M-stem absent; tuft of hairlike scales along 1A+2A (cubital pecten), originating near base of wing; male with (A. longiphallus) or without (A. buscki) modified sex scaling on basal portion of wing. Abdomen: Dorsal pits absent; no modified corethrogyne scaling in female. Male genitalia (Figs. 2-3): Uncus simple, rodlike, weakly curved. Socius moderate in size, ca. 0.7 length of gnathos arms, pendant, rounded; not fused to gnathos. Gnathos complete, arms narrow, joined distally into expanded trian- gular plate with densely spined venter. Subscaphium and hami ab- sent. Transtilla a moderately broad, arched plate, with a sclerotized posterior band bearing a dense row of short, fine, spinelike teeth. Valva moderately slender, parallel-sided, rounded apically; sacculus simple, well defined at base, without free apical process(es); costa weakly differentiated. Pulvinus absent. Vinculum complete, well de- veloped. Juxta a sclerotized subrectangular plate with lateral pointed processes at dorsum. Aedeagus broad, moderately short, with scle- rotized, attenuate, thornlike process distally; phallobase simple, rounded; vesica with variable number of mostly lanceolate cornuti. Female genitalia (Figs. 4—5): Papillae anales slender. Apophyses an- teriores and posteriores long, slender, posteriores slightly longer. Sterigma a slender, weakly sclerotized band. Antrum large, broad, membranous, with slender, sclerotized dorsal band; accessory pouch weakly developed on right side, either as a slightly expanded area with an irregular line of sclerotization or as a triangular flap. Ductus bursae broad, moderately long, with longitudinal creases of sclero- tization, twisted about two-thirds distance from antrum to junc- tion with corpus. Corpus bursae rounded, finely punctate; signum lacking. Pupa (Figs. 6-7). Description and illustrations based on recon- structions of 3 pupal shells (i.e., adults eclosed) of A. buscki. Typi- cally tortricine; head without apical projection; no conspicuous sculpturing on dorsum of T3 or Al—2 (similar to Anopina Obraztsov and Chileulia Powell, and in contrast to Dorithia Powell and Cuproxena Powell & Brown); abdomen with two rows of spines dor- sally on A2-7, one row on A8—9; rows on A2 well developed in con- VOLUME 53, NUMBER 2 Fic. 1. Wing venation of Accuminulia buscki. trast to that of other euliine pupae examined; cremaster short and broad, with 4 pairs of long, hooked setae. The pupa of Accuminulia differs from that of Dorithia and Cuproxena (both Euliini) in the fol- lowing: 1) absence of ornate sculpturing on the dorsum of abdomi- nal segments T3 and Al—2 (see Brown & Powell 1991 for compari- son); 2) the anterior row of spines on the dorsum of segments A3—7 extends from spiracle to spiracle across the dorsum (in Dorithia and Cuproxena the row is restricted to approximately the middle 0.6 of the dorsum); and 3) the cremaster is short and broad compared with that of Dorithia and Cuproxena. Diagnosis. Superficially, adults of Accuminula are similar to Apotomops Powell and Bonagota Razowski on the basis of size and pattern of the forewing, size and shape of the labial palpus, and length of the antennal cilia. However, the only feature of the geni- talia reminiscent of those two genera is the weakly developed acces- sory pouch in the female; a well defined accessory pouch is one of several convincing synapomorphies demonstrating the sister rela- tionship of Apotomops and Bonagota (Brown & Powell 1991). The male genitalia of Accuminulia are similar to those of Varifula Ra- zowski in the possession of a thornlike sclerite at the distal end of the aedeagus and a densely spined transtilla. In contrast, the facies of Accuminulia are remarkably dissimilar to those of Varifula: forewing length in Varifula varies from 12-15 mm, that of Accuminulia from 6-7 mm; forewing color and pattern in Varifula are simple with mostly yellow and pale tan, those of Accuminulia are complex, mot- tled gray, black, and white; and the labial palpi are extremely elon- gate (ca. 2.2 times compound eye diameter) and nearly porrect in Varifula, while short (ca. 1.5 times compound eye diameter) and weakly upturned in Accuminulia. Autapomorphies for Accuminulia include the enlarged, triangular, ventrally spined distal portion of the gnathos, the narrow, parallel-sided valvae, and the partially twisted ductus bursa. Although documented food plants of Accuminulia are similar to those of Chileulia stalactitis (Meyrick), a Chilean species also known to feed on grapes and various Prunus species (see Powell 1986, Brown & Passoa 1998), the two genera have little in common mor- phologically. Etymology. The generic name is a combination of parts of three _ words: “accumulate,” “in,” and “Eulia” Hiibner; it is interpreted as masculine in gender. Phylogenetic relationships. Based on the large, broad aedeagus with long, strong cornuti in the male and corresponding large, variably sclerotized antrum and ductus bursae of the female, Accuminulia appears to belong to a group of genera that includes Proeulia 61 Clarke, Argentulia Brown, Varifula, Inape Razowski, and Subtranstillaspis Razowski. Accuminulia shares with Argentulia, Varifula, and several species of Proeu- lia a slender, sclerotized, attenuate process along the lateral margin of the distal portion of the aedeagus. It shares with Varifula an extremely similar, densely spined transtilla (see Razowski 1995 for comparison). Although the spined transtilla appears to represent a convincing synapomorphy for Accuminulia and Vari- fula, their considerable phenotypic difference, their dissimilar cornuti, and their extremely dissimilar fe- male genitalia shed doubt on their possible sister group relationship. Superficially, Varifula looks like Proeulia, and its female genitalia are extremely similar to those of Argentulia. The assignment of Accuminulia to the tortricid tribe Euliini is based on the hypothesized phylogenetic relationship with Proeulia, which possesses the charac- teristic euliine foreleg hairpencil in the male (Brown 1990). The presence of a gnathos excludes it from Cochylini, which it resembles superficially and in the possession of an extremely broad aedeagus. KEY TO MALES OF ACCUMINULIA 1. Hindwing without modified sex scales in basal one-half, mostly dingy white with pale gray mottling (Fig. 8)..... buscki 1’. Hindwing with basal one-half covered by modified sex scales, cream-white in contrast to pale gray-brown of remainder of hindwing (Fig. 10)................ longiphallus Accuminulia buscki J. Brown, new species Figs. 1, 2, 5-9 Description. Male. Head: Frons smooth-scaled below mid-eye, whitish; roughened above, red-brown, pale yellow, and white. Labial palpus whitish yellow and brown mesally; brown mixed with tan lat- erally. Antennal scaling pale tan. Thorax: White, with brown and tan at anterior portion. Forewing (Fig. 8): Length 6.5-8.0 mm (x = 6.9 mm; n = 8). Upper side whitish tan, with irregular gray, brown, and cream overscaling and irrorations; gray rhomboidal or semicircular patches at costa ca. 0.3 and 0.5 distance from base to apex; ill de- fined, transverse, reddish brown band in distal 0.33, bordered basally by a white band; white band well defined at dorsum, becom- ing less defined toward costa; small black spot at apex of DC. Under side uniform dark tan with faint indication of upperside markings. Hindwing: Upper side dingy white, with pale gray overscaling and mottling. Under side light gray-brown with darker mottling. Ab- domen: Light cream. Genitalia: As in Fig. 2 (drawn from USNM slide 88447; n = 8). Uncus, socius, gnathos, transtilla, and valva as described for the genus. Aedeagus extremely short, broad, with 5-6 large, narrow-triangular cornuti, with broad bases. Female. FW length 6.0-7.0 mm (x = 6.6 mm; n = 5). Superfi- cially as in male (Fig. 9), except forewing with basal 0.5 mostly whitish with grayish overscaling, pattern slightly more defined; an- tennal cilia shorter. Genitalia: As in Fig. 5 (drawn from USNM slide 68618; n = 3). Essentially as described for the genus. Holotype. 3, Chile, Santiago Province, 29 Mar 1954, reared from grape (fruit), emerged 12 Apr 1954, 54-3351 (M. J. Ramsay, USNM). 62 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 2—5. Genitalia of Accuminulia; males with valvae spread, aedeagus removed. 2. Male of A. buscki; 3. Male of A. longiphallus; 4. Fe- male of A. longiphallus; 5. Female of A. buscki. VOLUME 53, NUMBER 2 Fics. 6-7. Pupa of Accwminulia buscki. 6. Ventral aspect; 7. Dorsal aspect. Paratypes. 10 dd, 17 92 as follows: CHILE: Aconcagua Province: Los Molles, ca. 10 km S Pichidangui, 1 9, 15/17 Nov 1981 (D. & M. Davis, USNM). Coquimbo Province: Fray Jorge National Park, ca. 70 km W Ovalle, 12, 6/9 Nov 1981 (D. & M. Davis, USNM); Nague, J1 km N Los Vilos, 20 m, 1 2, 4/5 Nov 1981 (D. & M. Davis, USNM): Cta. Cavilolen, NE Los Vilos, 5 9°, 5 Feb 1986 (L. Petia, USNM); El Naranjo, S Caimanes, 1 2, 7 Feb 1986 (L. Petia, USNM); La Viluma, SE Melipilla, 350 m, 1 2, 15/17 Dec 1987 (L. Pefia, USNM); Co- quimbo, 1 d, 12 Mar/14 May 1884 (Walker 3214, BMNH). Nuble Province: Alto Tresgualemu, ca. 20 km SE Chovellen, 1 2, 1/3 Dec 63 1981 (D. & M. Davis, USNM). Santiago Province: Santiago, 1 2, 5 Apr 1954, reared from grapes, 4-3161 (M. J. Ramsay, USNM), 12,5 Apr 1954, reared from peach, 54-3165 (F. Rotundo, USNM), 1 2, 24 Feb 1954, reared from peach, 54-2979 (M. J. Ramsay, USNM), 1 2, 9 Feb 1954, reared from peach, 54-2233 (Damos, USNM), 1 °, 22 Mar 1954, emerged 2 Apr 1954, reared from plum (fruits), 54-3352 (M. J. Ramsay, USNM), 1 d, Noy 1955, reared from apricot fruit (G. Olalquiaga, USNM), 14, 17 Feb 1954, reared from peach (M. Ram- say, USNM); Santiago, ex-Proeulia bait, 1 6, 5 Sep 1983 (P. Alvarez, UCB), 2 dd, 3 Sep 1993 (R. Gonzales, UCB), 1 3, Sep 1993 (D. Cepeda, UCB), 1 d, 8/12 Nov 1993 (R. Gonzales, UCB); Rio Col- orado, ca. 40 km SE Santiago, 1100 m, 1 9, 29/31 Oct 1981 (D. & M. Davis, USNM); Valparaiso, 1 d, 30 Sep/8 Oct 1883 (Walker 3070, BMNH). USA: New York: In cargo [packaged grapes] from Chile, on ship, 1 6, NY #57651, A. Busck gen. prep. 22 May 1926 (USNM). Remarks. In the female paratype reared from grapes, the ac- cessory pouch is considerably more developed, represented by a conspicuous triangular flap lacking sclerotization. Although it is pos- sible that this specimen is not conspecific with the remaining series, its conformity in phenotype and all other morphological features to other specimens, in additional to its food plant, suggest that the un- usual pouch represents infraspecific variation. Etymology. This species is named in honor of the late August Busck, one of the most prolific early microlepidopterists at the U.S. National Museum of Natural History. Distribution and biology. Accuminulia buscki is known only from Chile; captures range from 20 to 1100 m elevation. Specimen records suggest an adult flight period from October through April. It has been reared from the fruit of grape (Vitis sp.; Vitaceae), Fics. 8-11. Adults of Accuminulia. 8. A. buscki (male); 9. A. buscki (female); 10. A. longiphallus (male); 11. A. longiphallus (female). 64 plum (Prunus domestica L.; Rosaceae), apricot (Prunus armeniaca L..; Rosaceae), and peach (Prunus persica (L.) Batsch; Rosaceae). Captures in native habitat suggest that it is not an introduced pest in Chile, but a native species that has expanded its food plant range to include exotic (i.e., agricultural) plants. Although most larval Tortricinae are leaf-rollers, a few genera are known to bore into the fruit of their food plants. In Euliini these include Proeulia Obraztsoy, Chileulia Powell, and Accuminulia, all from Chile (see Brown & Passoa 1998). Accuminulia buscki was first recorded as an inter- ception at the port of New York in cargo (grapes) from Chile. Nearly all current interceptions of Lepidoptera at United States ports of entry are larvae. Because the larva of Accuminulia remains unknown, it is impossi- ble to determine whether this species currently is in- tercepted in fruit from Chile. Because grapes from Chile routinely are fumigated at U.S. port of entry, fo- liage-feeding insects are eliminated (J. Cavey pers. comm.). However, larvae feeding within fruit may be unaffected by such treatments. Recent collections (1983) of this species have come from traps baited with Proeulia-lure. Several species of Proeulia are pests of crops in Chile, including grapes, citrus, kiwis, and various stone fruits. Accuminulia longiphallus J. Brown, new species Figs. 3-4, 10-11 Description. Male. Head: Frons smooth-scaled below mid-eye, white mixed with red-brown; roughened above, dark gray-brown. Labial palpus white and gray mesally; mostly brown laterally. Anten- nal scaling red-brown on scape, whitish on flagellomeres. Thorax: Mixed white, red-brown, and tan. Forewing (Fig. 10): Length 6.5—-7.8 mm (x = 7.2 mm; n = 2). Upper side mostly gray, with irreg- ular tan, black, and cream overscaling and irrorations; distal 0.25 with moderately dense red-brown overscaling; patch of upraised cream scales at dorsum near base; an irregular, lustrous white band from near middle of dorsum, terminating just basad of apex of DC; diffuse black patch of overscaling distad of termination of white line. Under side uniform dark tan with faint indication of upperside markings. Hindwing: Upper side with basal 0.5 covered by modified sex scaling, cream-white; patch of sex scales just below costa near base; distal 0.5 of wing pale gray-brown. Under side light gray- brown with darker mottling. Abdomen: Light cream. Genitalia: As in Fig. 3 (drawn from RLB slide 1087; n = 2). Uncus, socius, gnathos, transtilla, and valva as described for the genus. Aedeagus compara- tively long, curved in distal 0.4, with a patch of 4 short, external teeth near tip; 10-12 short, spinelike cornuti in vesica. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Female. FW length 6.1 mm (n = 1). Superficially as in male (Fig. 11), except forewing mostly whitish in basal 0.5; antennal cilia shorter. Genitalia; As in Fig, 4 (drawn from RLB slide 1085; n = 1). As described for the genus, except accessory pouch poorly defined, represented by a pair of spinelike sclerites (possibly deciduous cor- nuti?), Holotype. 3, Chile, Santiago Province, 6 km W Tiltil, 15 Dec 1982 (R. L. Brown, MEM). Paratypes. 1d, 19 as follows: CHILE: 14, same data as holotype. Aconcagua Province: 10 km E San Felipe, 19, 14 Dec 1982 (R. L. Brown, MEM). Etymology. The specific epithet refers to the comparatively long aedeagus of this species. Distribution and biology. Accuminulia longiphal- lus is known only from Chile. It has been recorded only from Santiago and Aconcagua, provinces in which A. buscki also has been collected. Nothing is known of the biology; the three adults were collected in December. ACKNOWLEDGMENTS I thank the following for helpful comments on the manuscript: William Miller, University of Minnesota, St. Paul, Minnesota; Jerry Powell, University of California, Berkeley; Kevin Tuck, The Natural History Museum, London; Douglass Miller, USDA, Systematic En- tomology Laboratory, Beltsville, Maryland; and David Smith, USDA, Systematic Entomology Laboratory, Washington, D.C. I thank Kevin Tuck (BMNH), Richard Brown (MEM), and Jerry Powell (UCB) for the loan of material in their care. I thank Josef Razowski for allowing me to describe the species A. buscki, which we discovered virtually simultaneously. I thank Joseph Cavey, USDA, Animal and Plant Health Inspection Service, Riverdale, Maryland, for information re- garding current actions on fruit imported from Chile. LITERATURE CITED Brown, J. W. 1990. Taxonomic distribution and phylogenetic sig- nificance of the male foreleg hairpencil in the Tortricinae (Lep- idoptera: Tortricidae). Entomol. News 101:109-116. Brown, J. W. & S. Passoa. 1998. Larval foodplants of Euliini (Lep- idoptera: Tortricidae): from Abies to Vitis. Pan-Pacif. Entomol. 74:1-11. Brown, J. W. & J. A. POWELL. 1991. Systematics of the Chrys- oxena group of genera (Lepidoptera: Tortricidae: Euliini). Univ. Calif. Publ. Entomol. 111. 87 pp. Horak, M. 1984. Assessment of taxonomically significant struc- tures in Tortricinae (Lep., Tortricidae). Mit. Schweiz. Entomol. Gesel. 57:3-64. POWELL, J. 1986. Synopsis of the classification of Neotropical Tor- tricinae, with descriptions of new genera and species (Lepi- doptera: Tortricidae). Pan-Pacif. Entomol. 62:372—398. RAZOWSKI, J. 1995. Proeulia Clarke, 1962, the western Neotropical Tortricidae genus (Lepidoptera), with descriptions of five new species and two allied genera. Acta Zool. Cracov. 38:271—283. Received for publication 15 June 1998; revised and accepted 5 April 1999. Journal of the Lepidopterists’ Society 53(2), 1999, 65-71 PRELIMINARY ESTIMATES OF LEPIDOPTERA DIVERSITY FROM SPECIFIC SITES IN THE NEOTROPICS USING COMPLEMENTARITY AND SPECIES RICHNESS ESTIMATORS MICHAEL G. POGUE Systematic Entomology Laboratory, P. S. I., Agricultural Research Service, U.S. Department of Agriculture, % Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0168, USA ABSTRACT. Lepidoptera were collected and species richness and complementarity or uniqueness were compared between two rainfor- est sites: Pakitza, Peru and Beni, Bolivia. The total number of species collected from both sites was 1,879 of which 60 were shared resulting in a complementarity of 96.8%. Non-parametric equations and species accumulation curves of Hemiceras Guenée (Lepidoptera: Notodontidae) were used to compare species richness between three rainforest sites, Pakitza and Tambopata, Peru and Reserva Ethnica Waorani, Onkone Gare, Ecuador. Cluster analysis, using complementarity values for selected sites was used to determine altitudinal relationships between sites in Costa Rica; relationships between forest types in Brazil; and faunal differences among sites in western Amazonia using Hemiceras. Additional key words: Biodiversity, Notodontidae, Hemiceras, Chao, jackknife. Biodiversity as defined by E. O. Wilson (Reaka- Kudla et al. 1997) is “everything”. Biodiversity encom- passes the genes within a single local population or species, the species within a local community, and communities comprising the diverse ecosytems of the world. Life on earth is supported by the interactions and products produced by all other life on earth. With- out biodiversity there would be no life on earth as we know it. Therefore, it is essential that biologists begin to document and record biodiversity, whether it be how many species of insects are in your backyard to how many species of trees in a forest to how many for- est types in the world. Anyone with an interest in natural history can begin to study their local biodiversity and to document it. One basic element of biodiversity is to know how many species are present at a particular site. This pa- per will outline how anyone can begin to document lo- cal biodiversity by gathering data on species richness or the number of species present at a site at a particu- lar point in time. Knowing what species are present at a site is essential because it is the first step in under- standing the interactions between the species docu- mented and their interactions within the local commu- nity and ecosystem. Presently, no one has much of an idea exactly how many species are present today on earth. Estimates of the number of species worldwide range from 3 to 100 million (Erwin 1982, 1983, Stork 1988, Hodkinson & Casson 1991, May 1992, Raven and Wilson 1992). The study of species richness and complementarity, or how different species composition is between sites, is es- sential to assessing global biodiversity patterns. To address the question of world insect diversity one must get accurate estimates of site-specific species richness for a variety of taxa (Colwell & Coddington 1994), and then to compare these species lists to mea- sure relative levels of overlap and richness of these taxa around the globe. After a site has been sampled, species richness estimates are used to predict how many species were missed during the sampling pro- cess, thus arriving at an estimated number of species based on the actual number observed plus the number missed. By using species lists, either generated by sampling at a site or from museum collections, species composition among sites can be compared. These comparisons can then be used in setting policy and making informed conservation and management deci- sions. The goals of this paper are 1) to emphasize the im- portance of adequate sampling, 2) to assess whether inadequate sampling can still be useful in predicting species richness and complementarity between study sites, and 3) to provide a method of using complemen- tarity to compare faunal relationships between sites. To accomplish these goals, I compared overall species richness and complementarity of Lepidoptera from two rainforest sites, one in Peru and the other in Bo- livia. In addition, I estimated species richness from species accumulation curves and non-parametric esti- mators to compare the Hemiceras Guenée (Notodon- tidae) fauna between 2 sites in Peru and 1 in Ecuador. Finally, museum specimens of Hemiceras were used for faunal comparisons among sites in Costa Rica, Brazil, and western Amazonia, using complementarity and cluster analysis. MATERIALS AND METHODS Lepidoptera complementarity in SW Amazo- nia. Lepidoptera were collected at two rainforest sites: Beni, Bolivia and Pakitza, Peru. The Beni study site at 14°49’ S, 66°28’ W is 40 km E of San Borja, at 250 me- ters elevation. Pakitza is located on the Rio Manu at 11°56’47’S, 71°1700°W within the large drainage basin of the Rio Madre de Dios in southeastern Peru, at 356 m elevation, approximately 550 km NW of Beni. 66 40 35 30 25 20 No. of Species Trap Nights JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY —e-Pakitza | |——Tambopata | [eoiione Cee Fic. 1. Species accumulation curves of Hemiceras at Pakitza and Tambopata, Pert, and Onkone Gare, Ecuador. Samples were collected between August 26—Septem- ber 15, 1987 (Beni) and from September 27—October 5, 1987 (Pakitza). Adult moths were collected by UV light traps, spread, identified to family, sorted to mor- pho-species, and counted. Voucher specimens have been deposited in the National Museum of Natural History, Smithsonian Institution, Washington, D.C. Collecting effort was defined as the number of trap nights: 12 and 8 trap nights in Pakitza and Beni, re- spectively. Other studies have included the number of person-hours spent collecting (Coddington et al. 1991, Robbins et al. 1996), number of collecting days (Lou- ton et al. 1996), or trap nights (approximately 12 hours in length). The Lepidoptera faunas of Pakitza and Beni were compared using complementarity (Colwell and Cod- dington 1994). In comparing two sites, j and k, the first site has a species richness of S. and the second site has S, If the number of species in common be- tween both sites is Vee then the total species richness for both sites is $= 5,+5,-V, (1) jie and the number of species unique to both sites (U,) is WSS (2) The complementarity between the two sites is the proportion of the unique species to the pooled rich- ness, or Cis. (3) Hemiceras species richness in western Amazo- nia. The genus Hemiceras (Lepidoptera: Notodonti- dae), representing 245 species, was used as an indica- tor group for estimating species richness at three rainforest sites: Pakitza and Rio Tambopata Research Station in southeastern Peru, and Onkone Gare, in Ecuador. Rio Tambopata Research Station, at 14°14, 69°11W, is located on the Rio Tambopata, 30 air km SE of Puerto Maldonado, Madre de Dios, Peru, at 290 m elevation. Onkone Gare, at 00°38’S, 76°36W, is a research station within the Reserva Ethnica Waorani, Ecuador, at 220 m elevation. Tambopata had a total of 29 trap nights from November 2—25, 1979 and Sep- tember 16-21, 1990. The 11 trap nights at Onkone Gare were January 10, 12, 13-18, 25; June 20; and July 16, 1994. Species accumulation curves (Fig. 1) were used to plot the cumulative number of new species collected over unit effort (number of trap nights) at each of the VOLUME 53, NUMBER 2 67 Fic. 2. Localities of Costa Rican sites used in cluster analysis of dissimilarity matrix in Table 3. Fic. 3. Localities of South American sites used in cluster analyses of dissimilarity matrices in Tables 4 and 5. three sites. To extrapolate total species richness from the species accumulation curves at each site, four dif- ferent nonparametric equations were compared: 1) Chao 1, 2) Chao 2, 3) first-order jackknife, and 4) sec- ond-order jackknife. Chao (1984) developed a simple estimator of the true number of species at a given site based on the number of rare species in the pooled sample j. This is considered an abundance-based estimator because it is based on the number of species that are only repre- sented by only 1 or 2 individuals to estimate overall species richness. Colwell and Coddington (1994) called this Chao 1, Chao 1 =S,, + a°/2b, (4) where S_,. is the observed number of species in a sam- ple, a is the number of species that are only repre- sented by one specimen in the pooled sample (single- tons), and b is the number of species represented by two specimens in the pooled sample (doubletons). This estimator works well when the samples contain a large number of rare species (Chao 1984), which fre- quently occurs when sampling diverse groups such as insects. A related estimator is Chao 2 (Colwell & Codding- ton 1994), which is based on the incidence of rare spe- cies among samples, Chao 2 = S$, + L?/2M, (5) where L is the number of species that occur in only one sample, and M is the number of species that occur in exactly two samples. Jackknife estimators (Burnham & Overton 1978, 1979) also use the distribution of species among sam- ples (Colwell & Coddington 1994). The first-order jackknife estimator of species richness is based on the number of species that occur in only one sample L, 1 jackknife = S,,. + L(n- I/n), (6) where n is the number of samples. The second-order jackknife (Burnham & Overton 1978, 1979) is like the Chao 2 estimator where L is the number of species that occur in only one sample and M is the number of species that occur in exactly two samples, 2 jackknife = Sos i ip [L(2n 3 3)/n =) M(n = 2)?/n(n = 1)], (7) where n is the number of samples. Hemiceras faunal comparisons between three tropical regions. Species lists of Hemiceras were com- piled from specimens in the National Museum of Natu- ral History, Smithsonian Institution, Washington, D.C.., and were used to examine faunal relationships among sites in Costa Rica, Brazil, and western Amazonia. In Costa Rica, six sites were chosen to see how altitude af- fected species composition. The sites chosen were Juan Vinas (1500 m), Tuis (732 m), Turrialba (634 m), Guapiles (259 m), La Selva (40 m) and Sixaola River (0 m) (Fig. 2). Six sites in Brazil were chosen to examine the effect of forest type: the lowland Amazonia forest in- cluded the sites of Sao Paulo de Olivenga (Amazonas) and Porto Velho (Rondonia), and the Atlantic coast for- est sites were Baixo Guandu (Espirito Santo), Campo Bello (Rio de Janeiro), Hansa Humboldt (Santa Cata- rina), and Santa Catherines (Santa Catarina) (Fig. 3, sites 1-6). Six sites were chosen to determine faunal re- lationships in western Amazonia. They included Neblina, Venezuela; Onkone Gare, Ecuador; S40 Paulo 68 TABLE 1. Comparison of % complementarity and number of spe- cies of Lepidoptera between Beni, Bolivia and Pakitza, Peru. Family % Complementarity Number shared species Microlepidoptera Cosmopterygidae 100.0 50 (0 Tineidae 100.0 113 (0 Gelechiidae 98.8 167 (2 Oecophoridae 97.8 324 (7 Pyralidae/Crambidae 96.8 281 (9 Macrolepidoptera Noctuidae 98.6 296 (4 Notodontidae 95.2 63 (3 Geometridae 93.3 213 (14) Arctiidae 91.5 189 (16) de Olivenga and Porto Velho in Brazil; and Pakitza and Tambopata in Peru (Fig. 3, sites 1-2, 7-10). Complementarity was calculated between sites. These values were used to produce dissimilarity matri- ces which were converted, for easier interpretation, into dendrograms using cluster analysis (SYSTAT 1992). RESULTS Lepidoptera complementarity in SW Amazonia. Some 38 families of Lepidoptera were collected from both sites, although only nine families had numbers of species sufficient to illustrate trends in complementarity. Five of these families were Microlepidoptera and the remaining four were Macrolepidoptera. A total of 1748 specimens representing 933 species were collected at Beni and 1731 specimens representing 1006 species from were collected at Pakitza. The pooled species richness for both sites (S..) was 1879, the total number of unique species (U, i was 1819, resulting in a com- plementarity of 96. 8%. The Microlepidoptera families had higher complementarity values (100—96.8%), than the Macrolepidoptera families (95.2—91.5%), with the exception of the Noctuidae (98.6%) (Table 1). Hemiceras species richness in western Amazo- nia. Comparing the four nonparametric equations for estimating species richness of Hemiceras at Pakitza and Tambopata, Peru, and at Onkone Gare, Ecuador resulted in Chao 1 estimating the highest number of species. The incidence-based estimators (Chao 2 and the Jackknifes) consistently estimated total richness TABLE 3. Dissimilarity matrix of Costa Rican altitudinal sites. Juan Vinas (1500 m) Tuis Turnialba Guapiles La Selva Tuis (732 m) 0.389 Turrialba (634m) 0.771 0.758 Guapiles (259m) 0.897 0.829 0.960 La Selva (40 m) 0.718 0.730 0.897 0.769 Sixaola River (0m) 0.794 0.781 0.909 0.667 0.571 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TABLE 2. Estimated total species richness of Hemiceras species for 4 nonparametric estimators. Species richness Pakitza Tambopata Onkone Gare estimator Peru Peru Ecuador Observed 36 24 22 Chao 1 43 29 40 Chao 2 39 25 24 1 Jackknife 41 24 24 2 Jackknife 39 25 23 closer to the observed values than did Chao 1, on abundance-based estimates (Table 2). Hemiceras faunal comparisons between three tropical regions. Costa Rica: The high altitude sites of Juan Vinas and Tuis (1500 m and 732 m) were clus- tered at a dissimilarity value of 0.389, and the low alti- tude sites (0 m and 40 m) of Sixaola River and La Selva were clustered at a dissimilarity value of 0.571. Guapiles (259 m) was clustered with the low sites at a dissimilarity value of 0.667. The high altitude and low altitude sites were clustered at a dissimilarity value of 0.718. Turrialba (634 m) showed the greatest faunal dissimilarity between all other sites at 0.758 (Table 3; Fig. 4). Brazil: The two Amazonian sites of SAo Paulo de Olivenga and Porto Velho were clustered at a dissimi- larity value of 0.883. Within the Atlantic Coast sites Hansa Humboldt and Santa Catherines were least dis- similar (0.600), Campo Bello was most similar to Hansa Humboldt + Santa Catherines (0.724), and Baixo Guandu was most dissimilar to the previous At- lantic Coast Forest sites (0.833). Dissimilarity between the Amazonia and Atlantic Coast Forests was 0.951 (Table 4; Fig. 5). Western Amazonia: Pakitza and Tambopata in southeastern Peru were clustered at a dissimilarity value of 0.766. Neblina, Venezuela and Onkone Gare, Ecuador were clustered at a dissimilarity value of 0.793. These four sites were clustered at a dissimilarity value of 0.806. The fauna of Sao Paulo de Olivenga had a dissimilarity value of 0.815 compared to the previous four sites and Porto Velho’s fauna was the most dissim- ilar (0.833) (Table 5; Fig. 6). TaBLE 4. Dissimilarity matrix of comparison of Brazilian Amazo- nia and Atlantic Coast Forest. Sao Paulo de Porto Baixo Campo St. Olivenga Velho Guandu Bello Catherines Porto Velho 0.883 Baixo Guandu 0.982 0.951 Campo Bello 0.927 0.930 0.833 St. Catherines 0.945 1.000 0.867 0.724 Hansa Humboldt 0.958 1.000 0.870 0.792 0.600 VOLUME 53, NUMBER 2 Turrialba 634 m (1) Tuis 732 m (2) Juan Vinas 1500 m (3) La Selva 40 m (4) Sixola River O m (5) Guapiles 259 m (6) 0.389 69 0.667 0.758 0.571 0.718 Fic. 4. Dendrogram for cluster analysis of dissimilarity matrix (Table 3) of Costa Rican sites. Numbers refer to localities on Fig. 2. Altitudes are indicated for each site. The scale indicates dissimilarity value. DISCUSSION Lepidoptera complementarity in SW Amazo- nia. The lower complementarity found in most Macrolepidoptera, compared to the Microlepidoptera, may be due to sampling bias; because the collecting was done by UV light, larger moths may be coming from a larger collecting universe than the smaller moths, because the the larger species are better able to disperse than the smaller ones. The relatively high complementarity of the Noctuidae is curious given that they are generally strong fliers, medium to large moths, and many species are known for their migra- tion and wide ranging dispersal abilities. The answer may lie in their diversity; because these moths are the most speciose lepidopteran family, the sampling time possible in this preliminary study may be inadequate to accurately assess the ranges of many noctuid spe- cies, resulting in a higher complementarity value. Hemiceras species richness in western Amazo- nia. To accurately estimate the total number of species of a target taxon (such as Hemiceras) at a particular site the species accumulation curve (or the curve of a Sao Paulo de Olivenca (1) Porto Velho (2) Baixo Guandu (3) Campo Bello (4) St. Catherines (5) Hansa Humboldt (6) suitable richness estimator) should reach an asymptote or remain constant over time with additional sampling. Of the three sites, the species accumulation curve reached an asymptote, only at Pakitza (Fig. 1), sug- gesting that the species estimate there should be the most accurate. Although Tambopata is known for the high species richness of various insect groups (Fisher 1985, Paulson 1985, Pearson 1985, Wilkerson & Fairchild 1985, Robbins et al. 1996). The Hemiceras fauna there is poor, considering that there are 245 species in the neotropics. Although samples taken at Tambopata and Onkone Gare were insufficient to ac- curately estimate richness, as shown by the non- asymptotic species accumulation curves (Fig. 1), it ap- pears from all the nonparametric estimates, and the steeper accumulation curve, that Pakitza is even richer in Hemiceras than is Tambopata. That contrasts with a study of the faunal relationships between the Ci- cadoidea (Homoptera) (Pogue 1996) and Odonata (Louton et al. 1996) at Pakitza and Tambopata, in which the species richness was greater at Tambopata. Thus, either the preliminary estimate for Hemiceras at Tambopata is inaccurate, or this study demonstrates 5 0.883 0.600 0.724 0.833 0.951 Fic. 5. Dendrogram for cluster analysis of dissimilarity matrix (Table 4) of Brazilian Amazonian and Atlantic Coast Forest sites. Numbers refer to localities on Fig. 3. The scale indicates dissimilarity value. 70 Porto Velho (2) S&o Paulo de Olivenca (1) Pakitza (7) Tambopata (8) Neblina (9) Onkone Gare (10) FIc. 6. The scale indicates dissimilarity value. that different taxa, with divergent biologies, have dif- ferent centers of diversity. At the Onkone Gare site in Ecuador it is not clear whether the species accumulation curve is approach- ing an asymptote (Fig. 1) and there are more rare spe- cies (12 singletons) than at the other sites, so the pre- diction of 40 species by Chao 1 (Table 3) may be accurate. Based on the species richness data of Hemiceras, the following are recommended for follow- up studies: 1) collecting effort must be adequate for the species accumulation curve, or the estimate curve, to reach an asymptote, 2) if there is a preponderance of rare species, (singletons) Chao 1 should give the highest, and perhaps best estimate, and 3) if the num- ber of rare species is low, Chao 2 or the second-order jackknife may give a better estimate. It is also impor- tant to choose your target taxon carefully. To assess taxon richness, target taxa can be any cat- egory, an order, family, subfamily, tribe, or genus. The taxon should be chosen with care. For example, one that is too speciose requires too much time to process and extrapolate the needed data, one with too few spe- cies could result in insufficient data. I have found in the Neotropics that a target taxon of 200-400 species seems to be large enough so that the species accumu- lation curve reached an asymptote after 20-30 sam- ples (trap nights, in this case). The data for the target TABLE 5. Dissimilarity matrix of Amazonian sites. Onkone Sao Paulo de Porto Neblina Gare Olivenga Velho Pakitza Onkone Gare 0.793 S. P. De Olivenca 0.918 0.873 Porto Velho 0.947 0.886 0.883 Pakitza 0.860 0.863 0.831 0.833 Tambopata 0.806 0.821 0.815 0.915 0.766 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 6 0.806 0.833 0.766 0.7930.815 Dendrogram for cluster analysis of dissimilarity matrix (Table 5) of western Amazonian sites. Numbers refer to localities on Fig. 3. taxon can come from collecting, or from using mu- seum collections to obtain faunistic data from specific sites. An advantage of using museum collections are the data available from sites that are no longer pristine, such as those in Amazonia that were collected more than 50 to 100 years ago. Today, with the destruction of the rain forest, these sights will no longer have the same biota. The target taxon has to be common throughout the study area so it can be easily sampled and there should not be a dominance of rare species. Hemiceras faunal comparisons between three tropical regions. The six sites in Costa Rica show a broad altitudinal range from 0—1500 meters. Altitude seems to influence species composition among sites Costa Rica more than distance between sites. Juan Vinas (1500 m) and Tuis (732 m) are the highest sites and are clustered at a dissimilarity value of 0.389, the lowest of any pair. If distance was the significant limit- ing factor of faunal composition, one would expect that Tuis and Turrialba would be clustered. The same is true for the lower altitudinal sites, with La Selva (40 m) and Sixaola River (0 m) having the lower dissimi- larity value (0.571), even though La Selva is closer to Guapiles than the Sixaola River site (Table 3) (Fig. 4). Analysis of the six sites within Brazil were used to show if there was a faunal difference between Amazo- nia and the Atlantic Coast forest. The Atlantic Coast sites were clustered (Baixo Guandu to Hansa Hum- boldt) and were quite distinct from the Amazonian fauna (Sao Paulo de Olivenga and Porto Velho) which were also clustered. Within the Atlantic Coast Forest distance seems to be influencing the faunal relation- ships. Hansa Humboldt and Santa Catherines were most faunistically similar and closest in distance. Campo Bello shows similarity with Hansa Humboldt + Santa Catherines and is closer than Baixo Guandu, which is the most dissimilar and furthest from these VOLUME 53, NUMBER 2 two sites. Cluster analysis indicates that there is a fau- nal difference between Amazonia and the Atlantic Coast Forest (Table 4) (Fig. 5). Amazonia is often treated as one large biogeo- graphical area, but just how different are sites within Amazonia? Among the six Amazonian localities dissim- ilarities (Table 5) seemed strongly affected by distance and habitat. Pakitza and Tambopata, both in south- eastern Peru clustered, as did Onkone Gare, Ecuador and Neblina, Venezuela which are similar in habitat despite being approximately 1160 km distant from each other. Taken together these western Amazonia sites appear to form a region (Fig. 3), perhaps because they lie along the eastern edge of the Andes. Porto Velho was less faunistically similar to Sao Paulo de - Olivenga than to Pakitza. These studies from Costa Rica to Bolivia demon- strate that site-specific data analysis is a prerequisite to a thorough understanding of regional biodiversity pat- terns. The methods presented above were useful in as- sessing biodiversity on a site by site basis, and once similar data from other studies and other organisms are pooled, it may be possible to predict complemen- tarity between and among sites and to predict species numbers at other sites. Complementarity, or distinct- ness of species assemblages among sites can be used with cluster analysis to predict complementarity be- tween a wide variety of parameters such as biogeo- graphic, habitat differences, or host plant specificity. Using species richness estimates, complementarity val- ues, and species lists generated from biodiversity in- ventories can be useful for biologists and conservation- ists to make more informed decisions about land use and conservation. ACKNOWLEDGMENTS I would like to thank Terry L. Erwin for his support of this proj- ect and for providing time in the field for collecting and processing material. For critically reviewing a draft of this paper, I thank Marc E. Epstein and Jerry A. Louton, Smithsonian Institution, Washing- ton, D.C., and M. Alma Solis, Stuart H. McKamey and David R. Smith, Systematic Entomology Laboratory, Washington, D.C., Eric H. Metzler, Columbus, Ohio, and Deane Bowers, University of Col- orado, Boulder, Colorado. Funding was provided by the Smith- sonian Institution's BIOLAT Program. LITERATURE CITED BuURNHAM, K. P. & W. S. OvERTON. 1978. Estimation of the size of a closed population when capture probabilities vary among ani- mals. Biometrika 65:623—633. . 1979. Robust estimation of population size when capture probabilities vary among animals. Ecology 60:927—936. Cuao, A. 1984. Non-parametric estimation of the number of classes in a population. Scand. J. Statistics 11:265—-270. Wl CoppinctTon, J. A., C. E. GriswoLp, D. Siva Davia, E. PE- NARANDA & S. F, LARCHER. 1991. Designing and testing sam- pling protocols to estimate biodiversity in tropical ecosystems, pp. 44-60. In E. C. Dudley (ed.), The unity of evolutionary bi- ology: Proceedings of the Fourth International Congress of Sys- tematic and Evolutionary Biology. Dioscorides Press, Portland, Oregon. 1048 pp. COLWELL, R. K. & J. A. CoppINcTON. 1994. Estimating terrestrial biodiversity through extrapolation. Philos. Trans. R. Soc. Lon- don 345:101-118. ErRwIN, T. L. 1982. Tropical forests: their richness in Coleoptera and other arthropod species. Coleopts. Bull. 36:74—75. . 1983. Beetles and other insects of tropical forest canopies at Manaus, Brazil, samples by insecticidal fogging, pp. 59-75. In S. L. Sutton, T. C. Whitmore & A. C. Chadwick (eds.), Trop- ical rain forest ecology and management. Oxford, Blackwell Sci- entific Publications. 498 pp. FISHER, E. M. 1985. A preliminary list of the robber flies (Diptera: Asilidae) of the Tambopata Reserved Zone, Madre de Dios, Pert. Rev. Peru. Entomol. 27:25—36. Hopkinson, I. D. & D. Casson. 1991. A lesser predilection for bugs; Hemiptera (Insecta) diversity in tropical rain forests. Biol. J. Linn. Soc. 43:101—109. LoutTon, J .A., R. W. Garrison & O. S. FLintT. 1996. The Odonata of Parque Nacional Manu, Madre de Dios, Peru; natural his- tory, species richness and comparisons with other Peruvian sites, pp. 431-449. In D. E. Wilson & A. Sandoval (eds.), La biodiversidad del sureste del Peru: Manu (Biodiversity of Southeastern Peru). Editorial Horizonte, Lima, Peru. 679 pp. May, R. M. 1992. How many species inhabit the Earth? Sci. Am. 267(4):42—48. PAULSON, D. R. 1985. Odonata of the Tambopata Reserved Zone, Madre de Dios, Pert. Rev. Peru. Entomol. 27:9-14. PEARSON, D. L. 1985. The tiger beetles (Coleoptera: Cicindelidae) of the Tambopata Reserved Zone, Madre de Dios, Pert. Rev. Peru. Entomol. 27:15—24. PocuE, M. G. 1996. Biodiversity of Cicadoidea (Homoptera) of Pakitza, Manu Reserved Zone and Tambopata Reserved Zone, Pert: a faunal comparison, pp. 313-325. In D. E. Wilson & A. Sandoval (eds.), La biodiversidad del sureste del Peru: Manu (Biodiversity of Southeastern Peru). Editorial Horizonte, Lima, Peru. 679 pp. RAVEN, P. H. & E. O. WiLson. 1992. A fifty-year plan for biodiver- sity surveys. Science 258:1099—1100. ReAkKA-KupDLa, M. L., D. E. WILSON & E. O. WiLson. 1997. Biodi- versity IJ: understanding and protecting our biological re- sources. The National Academy of Sciences. John Henry Press, Washington, D.C. 551 pp. ROBBINS, R. K., G. Lamas, O. H. H. MIELKE, D. J. HARVEY & M. M. CASAGRANDE. 1996. Taxonomic composition and ecological structure of the species-rich butterfly community at Pakitza, Parque Nacional del Manu, Pert, pp. 217-252. In D. E. Wilson & A. Sandoval (eds.), La biodiversidad del sureste del Peru: Manu (Biodiversity of Southeastern Peru). Editorial Horizonte, Lima, Peru. 679 pp. StorK, N. E. 1988. Insect diversity: facts, fiction and speculation. Biol. J. Linn. Soc. 35:321—337. SYSTAT. 1992. Statistics, version 5.2 edition. SYSTAT Inc., Evanston, Illinois. 724 pp. WILKERSON, R. C. & G. B. FAIRCHILD. 1985. A checklist and generic key to the Tabanidae (Diptera) of Pert with special ref- erence to the Tambopata Reserved Zone, Madre de Dios, Pert. Rey. Peru. Entomol. 27:37—53. Received for publication 24 July 1998; revised and accepted 9 April 1999. Journal of the Lepidopterists’ Society 53(2), 1999, 72-73 LETHAL AND NON-LETHAL PARASITOIDS OF PLATYPREPIA VIRGINALIS (ARCTIIDAE) Additional key words: Tachinidae, tritrophic interactions, development rate, parasitism. By definition, a parasitoid is supposed to kill its host (Borror et al. 1989, Ricklefs 1990, Godfray 1994). Previously we reported that Platyprepia virginalis Byd. (Lepidoptera: Arctiidae) caterpillars in- fected with parasitoids of Thelaira americana Brooks (Diptera: Ta- chinidae) (formerly called T. bryanti) often survived the emergence of their flies (English-Loeb et al. 1990, 1993, Karban & English- Loeb 1997). At our study site at the Bodega Marine Reserve, Sonoma County, California (38°19.06N, 123°4.20W), caterpillars survived approximately 50% of the time although rates of survival depended upon the host plants and behavior of the caterpillars (En- glish-Loeb et al. 1993, Karban and English-Loeb 1997, Karban 1998). Most early instar P virginalis caterpillars use Lupinus ar- boreus at this study site although Coniwm maculatum is preferred by later instar caterpillars that are parasitized (Karban & English-Loeb 1997). Caterpillars that survive their parasitoids take longer to de- velop and pupate at slightly smaller weights than unparasitized caterpillars, although they are reproductively viable (English-Loeb et al. 1990, Karban & English-Loeb 1997). We most commonly ob- serve non-lethal parasitism when we rear caterpillars in the field in large sleeve cages; lab rearings are much more likely to be fatal. Because non-lethal parasitism is an unusual phenomenon (other examples have been reported for tachinids and caterpillars, e.g., Richards & Waloff 1948, DeVries 1984), we conducted a compara- tive study to determine what features of this system allowed both the caterpillar and its tachinid parasitoid to survive. We found popula- tions of P. virginalis in other locations where caterpillars used dif ferent host plants to determine if T. americana was ever non-lethal under those circumstances. We also reared 316 individuals of P. vir- ginalis outdoors in sleeve cages to determine if other, less common, parasitoid species were ever non-lethal to our host population at the Bodega Marine Reserve. During 1994 we found populations of P. virginalis in riparian habitats of the Trinity Alps (along Rush Creek in Trinity County (40° 46.80°N, 122°51.14°W) and along French Creek (40°41.75'‘N, 122° 38.27W) and Water Gulch (40°40.08°N, 122°42.20°W) in Shasta County, California). We returned to these sites during April 1995, 1996, and 1997 and caged individuals (50 caterpillars in 1995, 75 in 1996, and 44 in 1997) of this inland race on several host plants that were being used naturally by caterpillars at those locations. Our field rearing techniques are described in detail elsewhere (Karban & En- glish-Loeb 1997). Of 22 caterpillars that produced adult T. americana flies from these inland samples, 10 survived to become adults. This 45% rate of survival is indistinguishable from the 37% survival rate of cater- pillars parasitized by T. americana during the same three seasons at Bodega (Fisher's exact test, n = 60, p = 0.59). Survival of both cater- pillars and flies from this inland (Trinity Alps) population occurred when caterpillars were reared on C. maculatum (2 individuals), Lupinus albicaulis (1 individual), and especially Lupinus albifrons (7 individuals). Caterpillars were also reared on Rubus ursinus, a com- monly used host plant along Rush Creek; of 3 parasitized individuals reared on this host plant, none survived. Caterpillars were found on other host plants including species of Plantago, Phacelia, Nemophila, Plagiobothrys, and Rumex, although these were not tested as host plants because single individual plants were not large enough to support the complete development of a caterpillar. These observations indicated that the populations of P. virginialis and T. americana at the Bodega Marine Reserve were not unique in ex- hibiting non-lethal parasitism. In addition, species of lupine other than arboreus could serve as the sole host plants of late instar cater- pillars that survived emergence of parasitoids. Late instar caterpillars (already containing parasitoid larvae) were reared in field cages at the Bodega site in 1987, 1989, 1990, 1991, 1993, 1994, 1995, 1996, and 1997, and at the Trinity Alps sites in 1995, 1996, and 1997. Parasitoids that they contained pupated in these field cages and were collected. Rates of parasitism at Bodega have ranged from 6% in 1996 to 71% in 1990. Most of the para- sitoids were T. americana, although occasionally other species were recovered. We have 5 rearings of Carcelia reclinata (A&W), a sec- ond tachinid, from P. virginalis in the Trinity Alps. This parasitoid was common at our populations in the Trinity Alps in 1994, although we did not rear caterpillars in outdoor cages that season. C. reclinata has also been reared from our Bodega population at least 1 time. We reared Leschenaultia adusta (Wulp), a very large tachinid, from in- dividuals in our Bodega population on 5 different occasions. We also reared eight individuals of a large ichneumonid wasp, Ichneumon sp. from caterpillars at Bodega and from Shasta and Trinity counties. In no case (n = 18), did caterpillars recover to pupate successfully af- ter emergence of C. reclinata, L. adusta or Ichneumon sp. During this same period of time (1995-97), 45% of caterpillars with T. americana survived to become viable adult moths (n = 22). Given the same rate of survival after emergence of T. americana, we would have expected 8 caterpillars to have recovered after emergence of the other parasitoids. The likelihood that recovery rates are as high following emergence of these other parasitoids (considered to- gether) as it is following T. americana is 0.001 (Fisher's exact test). These results suggest that there is something unique about the in- teraction between P. virginalis and T. americana that allows for non- lethal parasitism. Of these three less common parasitoids, C. reclinata seems the most likely candidate to be non-lethal. Because of the low sample size (n = 5) of observations of this parasitoid, the likelihood that re- covery rates following emergence of C. reclinata differ from those of T. americana are only 0.12 (Fisher's exact test). More observations of C. reclinata parasitizing P. virginalis are necessary to determine whether C. reclinata can be non-lethal. What factors could allow T. americana but not the other para- sitoids to be non-lethal to P. virginalis? The ichneumonid is much larger than T. americana and remains in the host for longer, often (3 of 8 cases) emerging from the pupa. Lepidopteran larvae sometimes live for days or weeks after emergence of hymenopteran parasitoids although they invariably perish before successfully reproducing (Clausen 1962, Strand et al. 1988). (Hymenopteran parasitoids are much better studied than dipteran parasitoids [Feener and Brown 1997]). Recovering after emergence of L. adusta, another tachinid, seems about as unlikely as recovering after the ichneumonid. Like the ichneumonid, L. adusta is very much larger than T. americana and emerges later, often when the host is beginning to spin its co- coon (3 of 4 cases). C. reclinata is a smaller tachinid and can com- plete its development relatively rapidly. However, in 3 of the 5 cases in which we observed C. reclinata, this parasitoid emerged from a caterpillar that was spinning its cocoon. We have noted that the chances that P. virginalis will survive the emergence of T. americana decrease the later in the development of the caterpillar that para- sitoid emergence occurs. One difference between T. americana and the other parasitoids that are always lethal is that T. americana often completes development during one of the middle stadia of the host larva. It may be informative to consider the differences from the per- spective of the parasitoid’s life history traits. Belshaw (1994:149) de- scribes two developmental strategies for tachinids. Some species, perhaps including the ichneumonid and L. adusta, delay their own development and only kill their host close to pupation. Selection on these species presumably favors individuals that maximize their size at pupation by consuming most or all of the host. These species will always be lethal. Belshaw describes other tachinids that develop rapidly, often restricting their attack to late instars. C. reclinata may fall into this category. Selection on this species presumably favors speed of development or early emergence at the expense of not ex- ploiting all of the host. T. americana does not fit this category per- fectly because it develops quickly although it attacks early instars. This suggested that T. americana that develop rapidly may be more fit than those that take longer. A reanalysis of pupal weights of T. americana that developed in P. virginalis at Bodega Bay (methods VOLUME 53, NUMBER 2 and data in Karban & English-Loeb 1997) revealed that those that completed development earlier in the season were markedly heavier than those completing development later (F46= 25.037, p < 0.001). This result is consistent with the hypothesis that selection for rapid development at the expense of incomplete host exploitation may sometimes produce non-lethal parasitism. It would be informative to examine whether other tachinids that develop rapidly are also non- lethal. In conclusion, there does not seem to be anything special about the populations of P. virginalis, T. americana or the host plants used at Bodega Bay. Non-lethal parasitism was observed for other popu- lations of these insects. However, T. americana does appear to be unique among the four parasitoids that attack P. virginalis in allow- ing the host to recover and eventually reproduce. Hopefully these comparisons can be used in the future to elucidate the nature of the interactions that determine lethal and non-lethal parasitism. ACKNOWLEDGMENTS -We thank Alison Brody, Claire Karban, Anurag Agrawal, and Jennifer Thaler for help collecting and rearing caterpillars. Steve Hawthorne and Art Shapiro helped us locate new populations. Monty Wood and Paul Arnaud identified tachinids, Steve Heyden and Clement Dasch identified ichneumonids, and Ellen Dean iden- tified plants. The mansucript was improved by Deane Bowers and Michael Strand. We thank the UC Bodega Marine Reserve, Shasta- Trinity National Forest, and Whiskeytown National Rcreation Area for permission to work on their land. We were supported, in part, by UC intercampus research grants. LITERATURE CITED BrELsHAw, R. 1994. Life history characteristics of Tachinidae (Diptera) and their effect on polyphagy, pp. 145-162. In B. A. Hawkins & W. Sheehan (eds.), Parasitoid community ecology. Oxford Univ. Press, Oxford UK. BorreR, D. J., C. A. TRIPLEHORN & N. F. JOHNSON 1989. An intro- duction to the study of insects. Sixth edition. Saunders College Publishing, New York. 73 CLAUSEN, C. P. 1962. Entomophagous insects. Hafner, New York. DEVRIES, P. J. 1984. Butterflies and tachinidae: does the parasite always kill its host? J. Nat. Hist. 18:323-326. ENGLISH-LOEB, G. M., A. K. BRopy & R. KARBAN. 1993. Host- plant-mediated interactions between a generalist folivore and its tachinid parasitoid. J. Anim. Ecol. 62:465—471. ENGLISH-LOEB, G. M., R. KARBAN & A. K. Bropy. 1990. Arctiid larvae survive attack by a tachinid parasitoid and produce viable offspring. Ecol. Entomol. 15:361—362. FEENER, D. H. & B. V. BRowN. 1997. Diptera as parasitoids. Annu. Rev. Entomol. 42:73-97. Goprray, H. C. J. 1994. Parasitoids. Behavioral and evolutionary ecology. Princeton University Press, Princeton. KarBAN, R. 1998. Caterpillar basking behavior and non-lethal par- asitism by tachinid flies. J. Insect Behav. 11:713-723. Karsan, R. & G. ENGLISH-LOEB. 1997. Tachinid parasitoids affect host plant choice by caterpillars to increase caterpillar survival. Ecology 78:603—611. RICHARDS, O. W. & N. WALoFF. 1948. The hosts of four British Ta- chinidae (Diptera). Entomol. Monthly Mag. 84:127. RICKLEFS, R. E. 1990. Ecology. Third Edition. W. H. Freeman. New York. STRAND, M. R., J.A. JOHNSON & J. D. CuLin. 1988. Developmental interactions between the parasitoid Microplitis demolitor (Hy- menoptera: Braconidae) and its host Heliothis virescens (Lepi- doptera: Noctuidae). Ann. of the Entomol. Soc. 81:822—830. RICHARD KARBAN. Department of Entomology, University of California, Davis, California 95616, USA, AND GREGORY ENGLISH- LoEB. Department of Entomology, Cornell University, New York Agricultural Experiment Station, Geneva, New York 14456, USA. Received for publication 3 June, 1998; revised and accepted 10 Feb- ruary, 1999. Journal of the Lepidopterists’ Society 53(2), 1999, 74-75 A NEW SYNONYMY IN DICHRORAMPHA THAT REVEALS AN OVERLOOKED IMMIGRANT RECORD FOR NORTH AMERICA (TORTRICIDAE) Additional key words: D. vancouverana, D. gueneeana, Grapholitini, Olethreutinae, holarctic. The holarctic genus Dichrorampha currently includes nine rep- resentatives in North America (Miller 1983, Powell 1983) and sev- eral dozen in Eurasia (Obraztsov 1958, Razowski 1996). In revising Palaearctic Dichrorampha, Obraztsov (1953) proposed the new name D. gueneeana for a long-known species to which no available name Clearly applied. Male D. gueneeana have a distally upturned valva, a small sharp-pointed process on the ventroposterior edge of the cucullus, and a long, distally curved aedeagus, these characters together forming a diagnostic suite (Figs. 35, 15, respectively, in Obraztsov 1953, 1958; Fig. 4 in Roberts 1991). The sharp-pointed process on the cucullus is sometimes partially obscured in genital slide preparations. Unknown to Obraztsov, McDunnough (1935) had earlier de- scribed the same moth as D. vancouverana based on one male cap- tured on Vancouver Island, BC. The D. vancouverana holotype male and its genitalia (Fig. 5 in McDunnough 1935) match illustra- tions and specimens of both Palaearctic and Nearctic D. gueneeana in all respects. The new synonymy is formalized in the following nomenclatural review. Dichrorampha vancouverana McDunnough Dichrorampha vancouverana McDunnough (1935). D. gueneeana Obraztsov (1953). New Synonymy. This new synonymy and specimen examinations resulting from it have four implications beyond taxonomy. (1) D. vancouverana is a previously unsuspected immigrant in North America. (2) The year of McDunnough’s description of D. vancouverana, 1935, is more than five decades earlier than the first published record of D. van- couverana in North America (Roberts 1991); Roberts found adults at two Maine sites in 1988 and 1991. (3) The moth occurs in western as well as eastern North America; besides the type specimen from Vancouver, D. vancouverana is now recorded from four counties in Washington State, as shown by the specimens enumerated below. These specimens suggest that the species was established in Wash- ington in the 1940s. The moth was not seen there again for half a century; it reappeared during the 1990s in surveys for exotic pests in western Washington (LaGasa 1998, LaGasa et al. 1998). Powell (1988) reported a similar collection hiatus for Clepsis consimilana (Hiibner), another Pacific Northwest tortricid immigrant. (4) The synonymy brings to 12 the number of North American immigrant tortricids first detected in coastal British Columbia (W. E. Miller un- publ.), a number that spotlights the area as a premier immigrant en- try portal. Larval foodplants of D. vancouverana in North America have not been reported, but in Britain and Europe, the larva feeds in rootstocks of milfoil (Achillea millefoliwm L.) and ox-eye daisy (Chrysanthemum leucanthemum L.) (both Asteraceae) (Bradley et al. 1979, Kuznetsov 1987). The latter plant has become naturalized in North America (Fernald 1950), and the former also according to some authors (Fernald 1950), but others argue that American Achil- lea millefolium is actually the very similar native A. lanulosa Nutt. (Woodward & Rickett 1979). D. vancouverana is widely distributed in the Palaearctic (Kuznetsov 1987, Razowski 1996). In the Nearctic, its presently known distribution by states and provinces is British Columbia (Mc- Dunnough 1935), Maine (Roberts 1991), New Hampshire (W. Kiel in Winter 1993), New York (R. L. Brown pers. comm.), and Wash- ington State (LaGasa 1998, specimens listed below). As suggested by Roberts (1991), further searching of North American collections might uncover additional specimens and locality records because adults superficially resemble and could be mixed with more com- mon congeners such as D. sedatana (Busck). Indeed, R. L. Brown (pers. comm.) recently found D. vancouverana specimens captured in 1975 at Ithaca, New York, thus pushing back the earliest eastern North American record by more than a decade. Depository abbreviations used in the specimen enumerations be- low are as follows: CNC, Canadian National Collection of Arthropods, Ottawa, ON; USNM, National Museum of Natural History, Washing- ton, DC; WDA, Washington State Dept. of Agriculture, Olympia. I thank P. T. Dang, Ottawa, ON, J. W. Brown, Washington, DC, and E. LaGasa for the opportunity to examine specimens in their care. I also thank J. A. Powell, D. P. Prowell, and an unnamed reader for useful manuscript reviews, and R. L. Brown and M. A. Roberts for helpful information. Specimens examined. D. vancouverana holotype male (CNC); 1 male, “Achil. [probably referring to the Achillea foodplant] 4-16- [18]98, Hamfelt Collection” [of known European origin], male genit. slide #2, 2-13-34, C[arl]. H[einrich]., USNM slide 72540; 1 male, “Collection O. Hofmann” [of known European origin], genit. slide # 1, 9-14-22, C[arl]. H[einrich]., USNM slide 72543; 1 male, Sumas, Whatcom Co., WN, 7-12-44, W. H. Baker, Truck Crop No. 5276, genit. slide 12-27-44, Clarl]. H[einrich]., USNM slide 71298; 2 males, Drayton Harbor, Whatcom Co., WN, 7.16.44, W. H. Baker, Truck Crop No. 5278 (diagnostic parts of genitalia revealed by descaling) (all USNM); 1 male, 3.2 km NW Tenino, Thurston Co., WN, 7-9-96, E. LaGasa, genit. slide M 18; 2 females, Port of Seattle, King Co., WN, 7-17 to 28-98, M. Allen (genitalia not examined); 1 male, Shelton, Mason Co., WN, 7-15-98, M. Allen (genitalia not ex- amined) (all WDA). LITERATURE CITED BRADLEY, J. D., W. G. TREMEWAN & A. SmiTH. 1979. British tortri- coid moths. Tortricidae: Olethreutinae. Ray Society, London, England. 336 pp. FERNALD, M. L. 1950. Gray's manual of botany. Ed. 8. 1632 pp. Dioscorides Press, Portland, Oregon. [1987 reprint]. KUZNETSOY, V.I. 1987. Family Tortricidae (Olethreutidae, Cochyli- dae)—tortricid moths, pp. 279-956. In Medvedev, G. S. (ed.), Keys to the insects of the European part of the USSR. Vol. 4. Lepidoptera, Part 1. U.S. Dept. of Agriculture & (US) National Science Foundation, Washington, D.C. [translated from Russian]. LaGasa, E. 1998. Database record summary: Lepidoptera species collected in Washington State. Washington State Dept. Agr. Econ. Ins. Coll. & Entomol. Surv. database (EICES). 4 pp. LaGasa, E., M. Hircucox, T. UNRUH & T. Boyp. 1998. 1997 up- date-survey and methods development for the apple tortrix- Archips fuscocupreanus (Lepidoptera: Tortricidae) in Washing- ton State. Washington State Dept. Agr. Lab. Serv. Div. 1997 Entomol. Proj. Rept. 6 pp. McDUuNNOUGH, J. 1935. New Canadian eucosmids with notes (Lep- idoptera). Can. Entomol. 67:140-149. MILLER, W. E. 1983. New synonymies in Nearctic Dichrorampha (Lepidoptera: Tortricidae). Proc. Entomol. Soc. Wash. 85: 727-733. OsRAZTSOV, N. 1953. Systematische Aufstellung und Bemerkungen tiber die palaearktischen Arten der Gattung Dichrorampha Gn. (Lepidoptera, Tortricidae). Mitt. Miinch. Entomol. Ges. 43: 10-101. . 1958. Die Gattungen der palaearktischen Tortricidae. II. Die Unterfamilie Olethreutinae. Tijd. Entomol. 101: 229-261. POWELL, J. A. 1983. Tortricidae, pp. 31-41. In R. W. Hodges, T. Do- minick, D. R. Davis, D. C. Ferguson, J. G. Franclemont, E. G. Munroe & J. A. Powell (eds.), Check list of the Lepidoptera of America north of Mexico including Greenland. E. W. Classey & Wedge Entomological Research Foundation, London, England. . 1988. Records of the Palearctic tortricid, Clepsis consimi- lana, in the Pacific Northwest: can an urban moth be over- looked for half a century? Pan-Pac. Entomol. 64:98—99. VOLUME 53, NUMBER 2 RAZOWSKI, J. 1996. Tortricidae, pp: 130-157. In O. Karsholt, &. J. Razowski (eds.), The Lepidoptera of Europe: a distributional checklist. Apollo, Stenstrup, Denmark. RogeErRts, M. A. 1991. Two Palaearctic species of Dichrorampha discovered in Maine (Tortricidae). J. Lepid. Soc. 45:169-171. WINTER, W. D. (ED.). 1993. The Northeast, pp. 46—48. In Season summary 1992. News Lepid. Soc. 1993, No. 2. WoopwakbD, GC. H. & H. W. RICKETT. 1979. Common wild flowers of the northeastern United States. Barron’s, Woodbury, New York. 318 pp. WILLIAM E. MILLER, Department of Entomology, University of Minnesota, St. Paul, Minnesota 55108, USA. Submitted for publication 10 June 1998; revised and accepted 24 February 1999. Journal of the Lepidopterists’ Society 53(2), 1999, 76 “ARGYRESTHIA VISALIELLA CHAMBERS 1875” (ARGYRESTHIIDAE), A NOMEN NUDUM Additional key words: While checking the type localities of species names attributed to Vactor T. Chambers for an annotated checklist of Kentucky Lepi- doptera, I encountered a name that stands in synonymy of Ar- gyresthia apicimaculella Chambers 1874 (Argyresthiidae; type local- ity: “Kentucky”). The name is “visaliella Chambers 1875,” as the two names are listed as entry #2438 in Hodges et al. (1983). This same synonymy appears as #6456 in Dyar (“1903” [1904] ) along with a page reference to Chambers’ presumed original de- scription; however the publication date is quoted erroneously by Dyar as “1874.” In that 1875 paper we find not an original descrip- tion of visaliella but a reference to it in the discussion of “Ar- gyresthia goedartella Auct.” in which Chambers states, after de- scribing a specimen presumed to be that species, “It is a more handsome species than A. andereggiella, next after which as to beauty I would place A. visaliella Cham.” In searching for the original description of “A. visaliella’ I found only one “visaliella’—that of Chambers 1873, p. 113, described in Cyane Chambers, but listed in Hodges et al. (1983) #307 as Choro- pleca vesaliella |sic] (Cham. 1873). The type locality of this name is Visalia, Kentucky, which is in Kenton County not far south of Cham- bers’ home town of Covington. Chambers wrote “Several specimens captured in June resting on forest trees at Visalia, Kentucky.” This constitutes a rare specific type locality published by Chambers. Thus it appears that “A. visaliella Chambers” is a nomen nudum. Also the name Choropleca visaliella (Cham. 1873) is misspelled in Hodges et al. (1983) and was also misspelled in Forbes (1923). Lepidoptera, North America, taxonomy. I thank Ronald W. Hodges and Paul A. Opler for reviewing this manuscript. LITERATURE CITED CuHaMBerS, V. T. 1873. Micro-Lepidoptera. Can. Entomol. 5:12-15, 44-50, 72-75, 85-91, 110-115, 124-138, 147-152, 173-176, 185-190, 229-232, CHAMBERS, V. T. 1875. Tineina from Canada. Can. Entomol. 7;124-128, 144-147, 209-213. Dyar, H. G. “1903” [1904]. A List of North American Lepidoptera and key to the literature of this order of insects. Bull. U.S. Nat. Mus. 52, 722 pp. Forbes, W. T. M. 1923. The Lepidoptera of New York and Neigh- boring States. Part 1. Cornell Uniy. Agr. Exp. Station Bull. 23, 729 pp. Hopces, R. W. 1983. Check List of the Lepidoptera of America North of Mexico, E. W. Classey Ltd. and The Wedge Entomol. Res. Found., Washington, D.C. 284 pp. CHARLES V. COVELL JR., Department of Biology, University of Louisville, Louisville, Kentucky 40292-0001, U.S.A. Received for publication 1 August 1999; revised and accepted 30 Au- gust 1999. ERRATA STUDIES IN THE GENUS HYLEPHILA BILLBERG, I. INTRODUCTION AND THE IGNORANS AND VENUSTA SPECIES GROUPS (HESPERIIDAE: HESPERIINAE) In the above paper by C. D. MacNeill and J. Herrera G. (Journal of the Lepidopterists’ Society 52(3):277-317) there were two typographical errors in the text: pp. 283-287. The legends for Figures 4, 5, 6, 7, and 8 should read “.. . (descaled to show caudal array of tuberculate bristle-sockets at 70x with inset enlarged below), . . .” pp. 316, lines 7 and 8. The reference should read: . 1940. Hesperioidea XI. Especies nuevos para nuestra fauna y anotaciones sobre otros. Rev. Soc. 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CONTENTS A TECHNIQUE FOR EXTRACTION OF INTACT MITOCHONDRIAL DNA MOLECULES FROM LARVAE sea DOPTERA: SATURNIIDAE) FOR USE IN TaxoNOMIC sTuDIES Ralph M. Clark 020-2 A NEw species OF LirHopHANE (LEpiporTERA: NocTuIDAE: CUCULLIINAE) FROM NORTHEASTERN Noxri \MER Reginald P. Webster and Anthony W. Thomas ... ES a Res TR sn ere kT IF A NEW GENUS OF TORTRICID MOTHS (TORTRICIDAE: os. INJURIOUS TO GRAPES AND STONE FRUITS IN Joh Wo Brown sipe 26 sk! (cas FEA GW RS IR Re ere 2d ok 2a 4 GENERAL NOTES Lethal and non-lethal parasitoids of Platyprepia virginalis (Arctiidae) Richard Karban and Gregory English-Loeb_..... aa ntnn ann This paper meets the requirements of ANSI/NISO Z239.48-1992 (Permanance of Paper). q@ L- sA| LS3q ENT Volume 53 1999 Number 3 JOURNAL of the LEPIDOPTERISTS’ SOCIETY Published quarterly by THE LEPIDOPTERISTS’ SOCIETY Publié par LA SOCIETE DES LEPIDOPTERISTES Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN Publicado por LA SOCIEDAD DE LOS LEPIDOPTEROLOGOS 16 June 2000 ISSN 0024-0966 THE LEPIDOPTERISTS’ SOCIETY EXECUTIVE CoUNCIL Joun W. Brown, President Susan S. Borxin, Vice President Micuae J. Smita, Immediate Past President Mirna M. Casacranpde, Vice President Manuex A. Batcazar-Lara, Vice President Davip C. Irtner, Treasurer Ernest H. Wii.taMs, Secretary Members at large: Ronald L. Rutowski M. Deane Bowers George L. Balogh Felix A. H. Sperling Ron Leuschner Andrew V. Z. Brower Andrew D. 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Ink and watercolor by Eowyn Burke, University of Colorado, Boulder, Colorado. —_— i ee JOURNAL OF Toe LepiporTeRiIstTs’ SOCIETY Volume 53 1999 Number 3 Journal of the Lepidopterists’ Society 53(3), 1999, 77-89 DREPHALYS: DIVISION OF THIS SHOWY NEOTROPICAL GENUS, PLUS A NEW SPECIES AND THE IMMATURES AND FOOD PLANTS OF TWO SPECIES FROM COSTA RICAN DRY FOREST (HESPERIIDAE: PYRGINAE) JOHN M. Burns Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0127, USA AND DANIEL H. JANZEN Department of Biology, University of Pennsylvania, Philadelphia, Pennsylvania 19104, USA ABSTRACT. Mainly on the basis of many male and female genitalic characters, Drephalys splits cleanly into two subgenera: Drephalys (Drephalys) Watson 1893 (=Paradros Watson 1893), with at least 16 species, and Drephalys (Paradrephalys) Burns, new subgenus (type species Hesperia dumeril Latreille 1824), with at least 7 species. Although these showy, diurnal, neotropical pyrgine hesperiids generally are rare in collections, two species have been reared repeatedly in the tropical dry forests of the Area de Conservacion Guanacaste (ACG) in north- western Costa Rica: Drephalys (Drephalys) kidonoi Burns, new species, and D. (D.) alcmon (Cramer). In the ACG, larvae of D. kidonoi (N = 236) eat Roupala montana Aublet (Proteaceae), and larvae of D. alemon (N = 70) eat Hirtella racemosa Lamarck (Chrysobalanaceae). In Panama, as well, D. alemon eats Hirtella racemosa; but in Pard, Brazil, it eats the Hirtella relatives Couepia and Parinari (Chrysobalanaceae), whereas D. (D.) eous (Hewitson) eats Vochysia vismiaefolia Spruce (Vochysiaceae). As far as we can tell (admittedly not far), different species of Drephalys (Drephalys) seem to be specializing on food plants in taxonomically unallied families. Larvae of these three Drephalys (Drephalys) species share a basically similar color pattern, which is distinctive. With a development time of 45-55 days from newly-eclosed larva to prepupa, D. kidonoi is one of the slower-growing of some 190 species of pyrgines reared in the ACG. Adults of D. kidonoi apparently breed chiefly dur- ing the first half of the dry season, when other dry-forest skippers have emigrated or are sexually dormant. While D. kidonoi still is known only from Guanacaste, Costa Rica, D. alemon ranges far more widely—from eastern Peru and central Brazil all the way to Guatemala, at least (it has not previously been reported from Central America). Despite close genitalic similarities that mark D. kidonoi as the sister species of D. helixus (the type species of Drephalys), D. kidonoi departs sharply in color pattern from it and all other species of Drephalys, apparently to mimic sev- eral common species of the silver-spotted skippers Epargyreus with which it is sympatric. We illustrate comparatively the larvae, pupae, adults, and genitalia of species of Drephalys (Drephalys) that are central to this paper. Additional key words: Drephalys (Drephalys) alemon; Drephalys (Drephalys) kidonoi Burns, new species; Drephalys (Paradrephalys) Burns, new subgenus; Hirtella racemosa (Chrysobalanaceae), Roupala montana (Proteaceae). Once more, taxonomy and ecology join forces for genitalic form (in both sexes), by a unique feature of the greater good. the labial palpus, and by shared larval food plants (in A several-decade inventory of Lepidoptera larvae in the Malpighiaceae and Combretaceae) (Burns 1996). the tropical dry forests of the Area de Conservacion The fact that some species of Cephise have long hind- Guanacaste (ACG) in northwestern Costa Rica (Janzen wing tails while others do not, may explain why these & Hallwachs 1998) currently is focusing on the Hes- skippers had never been seen as related, much less periidae or skipper butterflies (Burns & Janzen in congeneric; but presence or absence of tails, though prep.). Of some 190 species of pyrgine hesperiids now striking, can be taxonomically trivial. The second reared, at least two are new. The first, Cephise nuspesez reared new species, Drephalys kidonoi, debuts here. Burns, joined a number of “known” species—which Because one ought not describe a new species with- were buried in the wrong genus or in synonymy—to out considering its taxonomic setting, Burns examined swell Cephise from a monotypic to a polytypic neotrop- the genus Drephalys in as much detail as available ma- ical genus, tightly characterized by many aspects of terial would allow. Unfortunately, this neotropical 78 genus of showy, medium large skippers is remarkably rare in collections. To illustrate, when Evans (1952) treated Drephalys (the most recent overall review), he included two taxa that were not represented in the vast hesperiid holdings of The Natural History Museum (BMNH); and he described three new species from single (or, in one case, a pair of ) specimens. Evans also described what he called a new subspecies (D. orian- der oria, from Honduras) from just 3 specimens (oria Evans is actually a separate species, distinct from oriander |Hewitson] though related to it). One of the two new species of Drephalys that Mielke (1968) de- scribed from far southeastern Brazil (Santa Catarina) was based on a single male. One of the two new species that Austin (1995) described from westcentral Brazil (Rondénia) was based on 4 specimens. In re- viewing a meager accumulation of Drephalys to de- velop a context for D. kidonoi, Burns already has dis- covered five more new species, represented by scant samples of 8, 6, 3, 1, and 1. In sharp contrast, we have a huge sample of D. ki- donoi—the type series amounts to 53 adults from Guanacaste—obtained almost entirely by rearing from 236 larvae found in nature. (Adults are far fewer than larvae because, in the course of rearing, many larvae were lost to parasitoids, fungi, and other diseases; and because some healthy reared adults were released.) Even though the larvae can be located with fair ease, only one adult has been caught in the wild. This, along with the general scarcity of Drephalys in collections, suggests adult behavior that tends to keep the skippers and their potential human captors apart. However, Drephalys is definitely diurnal, not crepuscular and/or nocturnal like some tropical skippers; and males are known to hilltop, though only for a very limited and specific period in any given day (Mielke 1968, pers. comm., Casagrande & Mielke 1992). Watson (1893:34) proposed the genus Drephalys (with the type species helixus Hewitson) but did not define it well. Eleven genera later in the same paper, Watson (1893:39) also proposed the genus Paradros (with the type species phoenice Hewitson). Godman and Salvin (1894:349) immediately set these genera side by side, observing “There can be no doubt that Drephalys should be placed next Paradros, and the only question that arises is whether these two genera ought not to be merged into one.” While Mabille and Boullet (1919) still kept them separate but adjacent, Evans (1952:23) correctly listed Paradros as a syn- onym of Drephalys, remarking that “Typically these 2 genera appear very different, but they are connected by intermediate species and the genitalia conform to a general pattern.” Although his statement is partly true, JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Drephalys as treated by Evans is gravely polyphyletic. Burns (1999) addressed this problem by removing morphologic misfits to the new and unrelated genus Pseudodrephalys. Any lingering questions about the generic limits of Drephalys have no bearing on the inclusion of kidonoi for the simple reason that kidonoi is the sister species of helixus. Since helixus is the type species of Drephalys, the sister species kidonoi must also go in Drephalys. It will be treated together with helixus to document their similarity. Drephalys heraclides Bell (1942:1), which is yet an- other species described from a single male (this time from “Peru”), was said in the original description to be “extremely like helixus.” Though Bell further noted that “The form of the male genitalia is similar in the two species, but the details materially differ,” he did not elaborate. Still known only from the holotype, heraclides is one of those taxa Evans (1952) never saw; and, as a result, he conservatively called it a subspecies of D. helixus. This is wrong. Study of the holotype of heraclides (borrowed from AMNH) and of Bell’s (1942:fig. 1) illustration of its genitalia (because the slide of its genitalia has been lost) shows not only that D. heraclides is a distinct species of Drephalys but also that it is morphologically farther from D. helixus than is D. kidonoi. In connection with five species of Drephalys occur- ring in Rond6nia, Brazil, Austin (1995:127) com- mented that “There appear to be a number of species groups in Drephalys with quite different genitalia of both sexes.” Even after Burns’s (1999) transfer of a pair of incredibly misplaced species—atinas (Mabille) and hypargus (Mabille)—from Drephalys to the dis- tant new genus Pseudodrephalys, Drephalys is genital- ically complex. This complexity can be resolved into two readily characterizable groups that are highly dis- tinct. Despite their differences, each is apparently the other’s nearest relative; so at this point in the analysis of hesperiid biodiversity, it is better to treat them as subgenera than as separate genera. A similarly cau- tious approach was taken in recognizing—but not overemphasizing—two useful, valid, well-differenti- ated divisions of the dusky wing skippers of the genus Erynnis: Erynnis (Erynnis) and Erynnis (Erynnides) (Burns 1964). Drephalys (Drephalys) Watson, 1893:34 =Paradros Watson, 1893:39. Type species. Eudamus helixus Hewitson (1877:320). Male genitalia. Uncus with pair of caudally projecting prongs that form U (Figs. 1, 3) or V in dorsal or ventral view. In lateral view, juxta either at level of vinculum or considerably anterior to it (Figs. 2, 4). If present, any dorsal projection from sacculus (i.e., sclero- VOLUME 53, NUMBER 3 TABLE 1. Species of Drephalys (Drephalys). alemon (Cramer) olva Evans eous (Hewitson) olvina Evans helixus (Hewitson) opifex Evans heraclides Bell kidonoi Burns miersi Mielke mourei Mielke phoenice (Hewitson) phoenicoides (Mabille & Boullet) 4 undescribed species tized, anteroventral portion of inner lamina of valva) arises from proximal part of sacculus (Figs. 2, 4); apart from this projection, sac- culus slopes sharply downward from anterior to posterior in lateral view (Figs. 2, 4). At least slightly dentate process arising from distal end of valva curves mostly mediad (Figs. 1, 3), but sometimes also dorsad (Figs. 2, 4) or caudad (or even cephalad so as to be recurved). Below this dentate, medially curved process, ventrodistal comer of valva usually rounded and extended slightly (Figs. 2, 4) to moder- ately caudad, but sometimes prolonged into blunt point. Above the dentate, medially curved process, dorsodistal corner of valva ranges from completely undeveloped or slightly developed (Figs. 1-4) to a small to large process, variable in orientation and degree of denta- tion (rarely none). Aedeagus with very short to very long subtermi- nal to terminal (Figs. 1-4) titillator on left side. Vesica long, delicate, fingerlike, with 1-14 needlelike cornuti (almost always in cluster) at distal end (Figs. 1, 3). In dorsal or ventral view, anterior end of sac- cus pointed (Figs. 1, 3), keeled, or narrowly rounded. Female genitalia. Posterior portion of ductus bursae, which is well-sclerotized, extending well anteriad of sterigma (Figs. 5-8) and flattened dorsoventrally, at least anteriad (Figs. 6, 8). Thereafter, ductus bursae both narrow and long (Figs. 5-8). Altogether, bursa copulatrix takes rather indirect course from posterior to anterior (Figs. 5-8) (except in some individuals of D. alemon [Cramer] and presumably also D. mourei Mielke [female unknown]). Lamella an- tevaginalis and lamella postvaginalis distinct from each other (Figs. 5-8) rather than fused and inseparable. Lamella antevaginalis a more or less simple, narrow band which may (Figs. 5-8) or may not be sclerotized midventrally; it forms conspicuous, paired, more or less caudally pointing, sharp projections immediately lateral to os- tium bursae (Figs. 5-8) (except in D. alemon and presumably D. mourei). Lamella postvaginalis a ventrally convex plate with mid- ventral U or V in its posterior margin (Figs. 5-8). Costal fold of male. Well-developed to vestigial or absent. Wingshape. Hindwing elongate (Figs. 9-20), appreciably longer than wide (more so in males [Figs. 9, 10, 13, 14, 17, 18] than in fe- males [Figs. 11, 12, 15, 16, 19, 20]), and clearly lobed at end of vein 1b (Figs. 9-16, 19, 20 [lobes of male in Figs. 17, 18 damaged)]). Included species. N = 16 (Table 1). Drephalys (Paradrephalys) Burns, new subgenus Type species. Hesperia dumeril Latreille (1824:757). Male genitalia. Uncus truncate, with no long caudally project- ing prongs in dorsal or ventral view. In lateral view, juxta begins at level of vinculum and extends posteriorly beneath aedeagus. Tall, dorsal projection arises from distal part of sacculus (i.e., sclerotized, anteroventral portion of inner lamina of valva); apart from this pro- jection, sacculus nearly or quite uniform in height from anterior to posterior in lateral view. Process arising from middle part of dorsal margin of valva extends mostly caudad, usually becoming at least slightly dentate distally. Narrow, slightly dentate process arising from distal end of valva extends mostly dorsad to overlap aforementioned process. Valva at its ventrodistal corner so totally undeveloped as to be “chinless” in lateral view. Aedeagus short, relatively stout, and de- void of titillators (although sometimes very finely dentate ventrolat- erally or midlaterally, on one or both sides, slightly before its poste- rior tip). Short, fat vesica everts caudally but especially to right and sports numerous (11-33) needlelike cornuti in several loose assem- 79 TABLE 2. Species of Drephalys (Paradrephalys). talboti (Le Cerf) tortus Austin 1 undescribed species croceus Austin dumeril (Latreille) oria Evans oriander (Hewitson) blages. In dorsal or ventral view, anterior end of saccus expanded or broadly rounded. Female genitalia. Posteriormost portion of ductus bursae a cylindrical tube, well-sclerotized—except for broad, middorsal, clear strip—and so short that it extends only slightly anteriad of sterigma. Sterigma reflects major fusion between lamellae antevaginalis and postvaginalis. At posterior end of ductus bursae, parts of lamella an- tevaginalis that are closely appressed to its sides meet midventrally to form (in ventral view) a large U or V (which opens caudally) just ventrad of ostium bursae. Posterior to this, lamella postvaginalis forms another large (and similarly oriented) midventral U or V. All of the above framed laterally by paired, (essentially) parallel, large, thin, vertical plates extending ventrad from sterigma (so as to be seen on edge in ventral view). Although these long, thin plates closely flank the central equipment, they leave a deep fissure on ei- ther side of it. Immediately dorsal and lateral to all of above—about halfway up sides of sterigma—a conspicuous, somewhat fingerlike and caudally pointing, unsclerotized area extends two-thirds dis- tance from anterior to posterior margin of sterigma. Seen ventrally, entire sterigma tends to look long, narrow, and more or less rectan- gular. Immediately anteriad of point where posterior sclerotized tube of ductus bursae becomes membranous, ductus swells to large sac (which may have folded, sclerotized plate in its walls) and then constricts sharply before expanding anteriad into corpus bursae. Al- together, bursa copulatrix takes rather direct course from posterior to anterior. Costal fold of male. Well-developed. Wingshape. Hindwing roundish, nearly as wide as long (more so in females than in males), and barely lobed at end of space 1b. Included species. N = 7 (Table 2). The helixus/kidonoi species pair of Drephalys (Drephalys) Male genitalia. Dorsal projection arising from proximal part of sacculus uniquely long from anterior to posterior and finely dentate dorsally (Figs. 2, 4). In dorsal or ventral view, saccus tapers anteriad to extremely delicate, sharp point (Figs. 1, 3). In lateral view, distal end of valva roughly truncate, with no major development of dor- sodistal corner, but with narrow, finely dentate process (which ex- tends mediad, dorsad, and caudad) arising closer to ventrodistal cor- ner of valva than to dorsodistal corner (Figs. 2, 4). Female genitalia. Anteriad, where the well-sclerotized poste- rior portion of the ductus bursae abruptly becomes membranous and sharply decreases in diameter, a large, blind, membranous sac extends at least slightly to the right; and the narrowed, membranous ductus bursae angles dorsad, perpendicular to its sclerotized course, before continuing anteriad to the corpus bursae (Figs. 5-8). Costal fold of male. Narrow (helixus) or almost vestigial (ki- donoi). Number of antennal nudum segments. High (Table 3). Al- though there are too few specimens of other species of Drephalys to include the conspicuously variable nudum in the preceding sub- generic characterizations with confidence, there are enough to indi- cate that the mean number of nudum segments is greater in helixus and kidonoi than it is in other species of Drephalys. Moreover, the mean number of nudum segments is clearly greater in females than it is in males (Table 3). Evans (1952:6) said for Drephalys “Nudum typically 16/15, arcuate or hooked”; but his total of 31 segments is a little too low to be typical for this genus. Palpus. Third segment of palpus (which in Drephalys and its rel- atives is not centered on the second segment but, instead, shifted 80 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY wets - Fics. 1,2. Male genitalia of Drephalys (Drephalys) kidonoi (paratype) from the Area de Conservacion Guanacaste, Guanacaste, COSTA RICA (D. H. Janzen & W. Hallwachs rearing voucher no. 92-SRNP-445) (J. M. Burns genitalic dissection no. X-3422) (USNM); scale = 1.0 mm. 1, Tegumen, uncus, gnathos, and aedeagus [all stippled] with everted vesica and cornuti (plus cut, everted sperm duct), as well as both valvae, vinculum, juxta, and saccus [all outlined] in dorsal view. Rotating the genitalia until the top of the tegumen/uncus is about flat makes the un- derlying structures look shorter than they do in the lower figure (but the scale is the same in both figures). 2, Complete genitalia (minus right valva and everted vesica and cornuti) in left lateral view. conspicuously laterad—see Burns 1999:figs. 19, 20) unusually short, protruding less far anteriad of second segment than in other species of Drephalys. (Note that in Drephalys generally, the third segment of the palpus is shorter in males than it is in conspecific females.) TABLE 3. Number of antennal nudum segments in sister species of Drephalys (Drephalys). Species Sex N Range Mean D. helixus 3 18 33-37 35.3 es Q 1 40 40 D. kidonoi fo) 27 30-42 35.0 Z 2 22, 3451 40.8 Size. Among the larger species of Drephalys. As usual in hes- periids, males average smaller than coexisting females (Table 4). Al- though helixus from Panama looks a little larger than kidonoi (Table 4), the difference may not be real: all specimens of helixus are wild- caught whereas all measured specimens of kidonoi are reared, with the likely result that most are appreciably stunted. Drephalys (Drephalys) kidonoi Burns, new species Male genitalia. Aedeagus shorter than that of helixus, owing mainly to titillator. Titillator (i.e., caudal extension of left side of aedeagus, beginning where vesica emerges) shorter by half, as well as stouter (Figs. 1, 2), than that of helixus (Figs. 3, 4) and not tumed slightly up at distal end. Saccus also shorter (a condition almost cer- tainly correlated with the reduced aedeagal length). Body of aedea- VOLUME 53, NUMBER 3 81 —<—<— SS! Fics. 3,4. Male genitalia of Drephalys (Drephalys) helixus from Rodman, 8°58’N 79°35’W, Canal Zone, PANAMA, 22 December 1972 (G. B. Small) (J. M. Burns genitalic dissection no. X-4254) (USNM); scale = 1.0 mm. 3, Tegumen, uncus, gnathos, and aedeagus [all stippled] with everted vesica and cornuti (plus cut, everted sperm duct), as well as both valvae, vinculum, juxta, and saccus [all outlined] in dorsal view. Rotating the genitalia until the top of the tegumen/uncus is about flat makes the underlying structures look shorter than they do in the lower fig- ure (but the scale is the same in both figures). The oblique position of the aedeagus is unnatural. 4, Complete genitalia (minus right valva and everted vesica and comnuti) in left lateral view. gus, in lateral view (Fig. 2), bowed more dorsad than in helixus (Fig. 4). Process arising from distal end of valva a little less conspicuously dentate (Figs. 1, 2) than in helixus (Figs. 3, 4) and straight along its dorsal margin (Figs. 1, 2) rather than slightly convex as in helixus (Figs. 3, 4). TABLE 4. Forewing length (mm) in sister species of Drephalys (Drephalys). Species Locality Sex N Range Mean 3 22.7-23.9 23.20 15 =. 20.6-22.2 21.49 23.6 23.6 28 17.0-22.0 20.22 24 19.0-24.7 22.22. D. helixus Brazil i Panama uw u D. kidonoi Costa Rica uw 40 OL 10 AQ — Female genitalia. Lamella postvaginalis deeply notched mid- ventrally in its posterior margin and grooved midventrally along its entire length (Fig. 5) rather than shallowly notched and grooveless as in helixus (Fig. 7). Immediately farther anteriad, at posterior end of ductus bursae, at least a small midventral notch (Fig. 5) not pres- ent in helixus (Fig. 7). Sclerotized portion of ductus bursae flattened dorsoventrally in its anterior half (Fig. 6) instead of its anterior two- thirds as in helixus (Fig. 8). Paired, caudally pointed projections from lamella antevaginalis (immediately lateral to ostium bursae) shorter, less delicate, and originating farther posteriad (Figs. 5, 6) than in helixus (Figs. 7, 8). Costal fold of male. Present but exceedingly narrow (much narrower than that of helixus)—almost vestigial. Facies/mimicry. Unique among species of Drephalys: kidonoi (Figs. 9-12) departs sharply from a more usual Drephalys appear- ance (as, for example, in helixus [Figs. 13-16]) to suggest several 82 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 5,6. Female genitalia of Drephalys (Drephalys) kidonoi (paratype) from the Area de Conservacion Guanacaste, Guanacaste, COSTA RICA (D. H. Janzen & W. Hallwachs rearing voucher no. 92-SRNP-480) (J. M. Burns genitalic dissection no. X-3423) (USNM)); scale = 1.0 mm. 5, Ovipositor lobes, sterigma, and bursa copulatrix in ventral view. 6, The same, plus part of the ductus seminalis, terminal abdominal tergite, and posterior apophysis, in right lateral view. common species of the silver-spotted skippers Epargyreus with which it is sympatric. On the forewing, this involves (a) eliminating (Figs. 9-12) the spot in space 1b (Figs. 13-20); (b) reducing to tiny points or, more often, completely eliminating (Figs. 9-12) the small subapical spots in spaces 6, 7, and 8 (Figs. 13-20); (c) reducing (Figs. 11, 12) or eliminating (Figs. 9, 10) the outer cell spot (Figs. 13-20); and (d) suppressing (Figs. 10, 12) the yellow coloration along the proximal half of the ventral costal margin (Figs. 14, 16, 18, 20). On the hindwing, this involves (dorsally) more or less suppress- ing (Figs. 9, 11) what is normally a conspicuous double row of spots (Figs. 13, 15); and (ventrally) shortening and making less regular (Figs. 10, 12) a conspicuous central white stripe (Figs. 14, 16). On both wings, this involves (dorsally) intensifying the color of proximal scales—to an orangy yellow—to heighten the contrast with more distal ones (Figs. 9, 11); and (ventrally) adding lavender overscaling broadly along the outer margins (Figs. 10, 12). The total effect is one VOLUME 53, NUMBER 3 83 Fics. 7,8. Female genitalia of Drephalys (Drephalys) helixus from Rodman Naval Station, Canal Zone, PANAMA, 9 January 1972 (S. S. Nicolay) (J. M. Burns genitalic dissection no. X-4263) (USNM); scale = 1.0 mm. 7, Ovipositor lobes (by chance, wider apart than in Fig. 5), | sterigma, and bursa copulatrix in ventral view. 8, The same, plus part of the ductus seminalis, terminal abdominal tergite, and posterior apoph- ysis, in right lateral view. of apparent mimicry. Because the hindwing is a little less elongate in Types. Holotype: COSTA RICA, PROVINCIA GUANACASTE, kidonoi (Figs. 9-12) than it is in helixus (Figs. 13-16), kidonoi even Area de Conservacion Guanacaste, Sector Poco Sol, Poco Sol, 270 approaches the wingshape of Epargyreus. m, Janzen & Hallwachs rearing voucher 91-SRNP-2736, adult emer- Larval food plant. Roupala montana Aublet (Proteaceae). gence date 20 Feb 1992, 4, genitalia no. X-4243 J. M. Burns 1997; 84 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 17 18 . 19 20 Fics. 9-20. Males and females of Drephalys (Drephalys) from the Area de Conservacion Guanacaste, Guanacaste, COSTA RICA (reared) and from Rodman, Canal Zone, PANAMA (wild-caught) in dorsal (odd-numbered) and ventral (even-numbered) views (x0.85) (all USNM). 9, 10, D. kidonoi 4, holotype, COSTA RICA (D. H. Janzen & W. Hallwachs rearing voucher no. 91-SRNP-2736) (J. M. Burns genitalic dissection no. X-4243). 11, 12, D. kidonoi 2, paratype, COSTA RICA (D. H. Janzen & W. Hallwachs rearing voucher no. 92-SRNP-345). 13, 14, D. helixus 4, PANAMA, 28 November 1974, G. B. Small. 15, 16, D. helixus 9, PANAMA, 9 January 1972, S. S. Nicolay (J. M. Burns genitalic dissection no, X-4263). 17, 18, D. alemon 3, COSTA RICA (D. H. Janzen & W. Hallwachs rearing voucher no. 96-SRNP-1047) (J. M. Burns genitalic dissec- tion no. X-4248). 19, 20, D. alemon °, COSTA RICA (D. H. Janzen & W. Hallwachs rearing voucher no. 96-SRNP-1090) (J. M. Burns genitalic dissection no. X-4249). deposited in National Museum of Natural History, Smithsonian In- stitution (USNM). Paratypes N = 52 (27 4, 25 2). Reared Paratypes (with emergence dates): COSTA RICA, PROVINCIA GUANACASTE, Area de Con- servacion Guanacaste: Sector El] Hacha, Casa Oeste, 420 m, 93- SRNP-8708, 21 Jan 94, °; 93-SRNP-8712, 17 Mar 94, d; 93-SRNP- 8738, 22 Jan 94, ?; 93-SRNP-8740, 17 Jan 94, 3: 94-SRNP-19, 5 Mar 94, 5; 94-SRNP-20, ?, 2, X-4489: 94-SRNP-22, 31 Jan 94, 9; 94- SRNP-25, 24 Jan 94, 3; 94-SRNP-27, 11 Mar 94, 6; 94-SRNP-29, ?, d; 94-SRNP-31, 22 Feb 94, d: 94-SRNP-32, 5 Mar 94, 6. Sector El Hacha, Vado Rio El Hacha, 290 m, 94-SRNP-879, 2 Jun 94, 9; 94- SRNP-882, 11 Jul 94, 3, X-4246. Sector Orosi, Estacion Maritza, 520 m, 95-SRNP-378, 14 Mar 95, 6, X-4491. Sector Poco Sol, Poco Sol, 270 m, 91-SRNP-2737, 21 Feb 92, 2: 92-SRNP-361, 20 Mar 92, 6; 92-SRNP-379, 21 Mar 92, 9: 92-SRNP-380, 5 Mar 92, °, X-4244. 99- SRNP-385, 27 Mar 92, ?: 92-SRNP-386, 18 Apr 92, 9; 92-SRNP- 400, 21 Mar 92, °, X-4247: 92-SRNP-403, 23 Mar 92, 6; 92-SRNP- 410, 5 Mar 92, ?; 92-SRNP-419, 14 Mar 92, 9; 92-SRNP-426, 15 Apr 92, 6: 92-SRNP-433, 10 Mar 92, d, X-4245: 92-SRNP-435, 21 Mar 92, 6: 92-SRNP-437, 24 Mar 92, 6: 92-SRNP-438, 15 Apr 92, 9; 92- SRNP-441, 12 Mar 92, 6, X-4487; 92-SRNP-444, 18 Mar 92, d; 92- SRNP-445, 29 Feb 92, 6, X-3422: 92-SRNP-450, 18 Mar 92, 2, X-4488; 92-SRNP-455, 15 Mar 92, 9; 92-SRNP-458, 5 Apr 92, d; 92- SRNP-466, 30 Mar 92, 6; 92-SRNP-470, 21 Apr 92, °; 92-SRNP- 473, 5 Apr 92, °; 92-SRNP-477, 2 Mar 92, 3, X-4278: 92-SRNP-480, 6 Mar 92, 2°, X-3423-; 92-SRNP-498, 15 Apr 92, d: 92-SRNP-639, 29 Apr 92, °. Sector Poco Sol, Quebrada Aserradero, 160 m, 92-SRNP- 354, 7 Apr 92, 2; 92-SRNP-366, 13 Mar 92, d; 92-SRNP-370, 3 Mar 92, 2; 92-SRNP-377, 23 Mar 92, 6: 92-SRNP-698, 9 Jun 92, 6; 94- SRNP-641, 8 May 94, d. Sector Santa Rosa, Cruz de Piedra, 290 m, 92-SRNP-345, 29 Feb 92, °. Sector Santa Rosa, Porton de Los Per- ros, 300 m, 94-SRNP-870, ?, 2, X-4490. Wild-caught paratype: COSTA RICA, PROVINCIA GUA- NACASTE, Comelco, 8 km N Bagaces, 50 m, 24 May 1972, 2, P. A. Opler (CAS). Etymology. Named in honor of Dr. Hiroshi Kidono of the Japan International Cooperation Agency who is an enthusiastic and dedi- cated supporter of the INBio and ACG parataxonomists’ research on the caterpillars of the Hesperiidae of Costa Rica. NATURAL HISTORY OF DREPHALYS KIDONOI AND DREPHALYS ALCMON As stated above, just one adult of D. kidonoi has been collected in nature, although larvae have often been found eating both new and mature leaves of Roupala montana (Proteaceae), the only known host plant. This shrubby tree is abundant on the poor soils and rocky pastures in the central portion of the ACG at 100-500 m elevation (Figs. 31, 32). Since R. mon- tana occurs throughout the dry forest remnants in Pa- cific coastal Mesoamerica, since it “ranges from Vera- cruz, Mexico, to Peru, Bolivia, and Brazil” (Burger 1983:14), and since adults of D. kidonoi elude collec- tors, we infer that this skipper is more widespread. In this connection, note that its sister species, D. helixus, is represented (a) in the USNM by 144 1 2, all taken at a single, hilltop locality (Rodman) in the former Canal VOLUME 53, NUMBER 3 Zone of Panama (on 12 different days in 5 different years), (b) in the AMNH by 1 d from Balboa in the for- mer Canal Zone of Panama, and (c) in the BMNH by 6 d, all from Panama (Evans 1952)—which suggests a geographically limited species. But it most certainly is not: Mielke, Miers, and Casagrande (Mielke pers. comm.) have caught 28 d of D. helixus (only 1 or 2 on any given day, and always on hilltops) very far away in the southern Brazilian states of Santa Catarina (24 d at Joinville) and Sao Paulo (3 6 at Morro do Diabo, Teodoro Sampaio) and in the city of Rio de Janeiro (1 4) (specimens in UFPR, 3 of them donated to USNM and examined by Burns); and C. Callaghan has caught 1 d (on a hilltop at km 500 of the Belo Horizonte-Brasilia highway) in Minas Gerais, the next Brazilian state to the north (specimen seen by Burns). Drephalys kidonoi larvae (Figs. 21-23) are unlikely to be confused with any other hesperiid larvae known from the ACG. The dorsal and lateral part of the body is sharply, and rather narrowly, banded black on a whitish to greenish white background, while the head (which is slightly rugose) is pale to dark orange and de- void of markings. The black bands are broken by the whitish ground color just anterior to the spiracles on all segments but the first two and the last three. This banded color pattern starts to appear in the second in- star; the first instar larva is green (once it has fed) with a black head. When the larva changes to a prepupa, the ground color becomes creamy, and the black bands, light beige (Fig. 23). The head may become lighter orange at this time. Drephalys alemon (Cramer) (Figs. 17—20), the only other species of Drephalys known from the ACG, ranges widely from Guatemala (1 ° from Cayuga in USNM) through Central and South America to Brazil and southeastern Peru (1 2 from°30 km SW Puerto Maldonado in USNM)—as well as the island of Trinidad (Cock 1984). Brazilian specimens come from northern and central states—Roraima: Ilha de Maraca, Alto Alegre (UFPR); Rond6nia: Fazenda Rancho Grande, Cacaulandia (Austin 1995, UFPR), Fazenda Urupa, Candeias do Jamari (UFPR); Paré (Evans 1952): Belém (Moss 1949), 15 km S Itaituba (USNM), Obidos (UFPR), Santarém (UFPR); Mato Grosso: Alto Rio Arinos, Diamantino (USNM), Barra dos Bu- gres (UFPR), Fazenda Parana, Brasnorte (UFPR); ' Goids: Goids Velho (UFPR), Ilha do Bananal (UFPR): Pernambuco: Camaragibe, Recife (UFPR); and Es- pirito Santo: Linhares (UFPR). The latitude of the southernmost record (Linhares, Brazil, 19°25’S) is slightly higher than that of the northernmost record (Cayuga, Guatemala, 15°32’N). However, D. alcmon 85 probably occurs as far north as southern Mexico (the distribution of its larval food plant would allow this— see below). When he recorded D. alemon from Rondonia, Brazil, Austin (1995:127) gave for the rest of its range only “northeastern South America.” In his list of BMNH holdings, Evans (1952:27) included, be- sides specimens from the Guianas and Amazonian Brazil, “1 2 “Honduras”; but the quotation marks were his way of questioning the accuracy of a locality label. Adults of D. alemon (Figs. 17-20) are superficially and morphologically well removed from the sister species D. kidonoi (Figs. 9-12) and D. helixus (Figs. 13-16). Although the larva of D. alemon shares a gen- eral color pattern with that of D. kidonoi, its black bands are so broad that the white ground color is re- duced to thin white bands connected to a thin, white ventrolateral line (Figs. 24, 25). Viewed from the front (Fig. 25), its orange (and slightly rugose) head bulges less than does that of D. kidonoi (Fig. 22). The larvae of D. kidonoi (N = 236) apparently feed only on Roupala montana (Proteaceae). In the ACG, the larvae of D. alemon (N = 70) apparently feed only on Hirtella racemosa Lamarck (Chrysobalanaceae) (Burns & Janzen in prep., Janzen & Hallwachs 1998), a quite unrelated plant—but a common one that spans the neotropics from central and southern Mexico to southeastern Peru, Bolivia, and northeastern Sado Paulo, Brazil (Prance & Campbell 1988, Prance 1989). In 1998, Aiello (pers. comm.) reared an adult male of D. alcmon from a larva found on H. racemosa in Loma del Rio, Arraijén, Panama province, Panama (Aiello Lot 98-9). But in the vicinity of Belém, Para, Brazil, Moss (1949:59) reared D. alcmon from larvae found on two food plants that were incompletely determined as “Parinarium or Couepia, Rosaceae.” “Parinarium’ is presumably Parinari; and both Parinari and Couepia are now in the Chrysobalanaceae, along with Hirtella. Indeed, Couepia and Hirtella are sister genera, ex- tremely closely related (Prance pers. comm.). Moss (1949:59) reared one other species of Drephalys, D. eous (Hewitson), whose “larval shelters [he] commonly observed in the forest on the leaves of . . . Vochysia vismiaefolia Spruce” (Vochysiaceae). So far (but ad- mittedly it is not very far), different species of Drephalys seem to be specializing on food plants in taxonomically unallied families (Proteaceae, Chryso- balanaceae, and Vochysiaceae). The drawings of last instar Drephalys larvae in Moss (1949:pl. I) are both black-and-white and small. Still, his D. eous larva (fig. 9) closely resembles our D. ki- donoi larva (Figs. 21-23). However, his D. alemon larva (fig. 11) is one in which every other vertical, thin, white band of ground color fails to reach the horizon- 86 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 21-25. Larvae of two species of Drephalys (Drephalys) from the Area de Conservacion Guanacaste, Guanacaste, COSTA RICA. 21, D. kidonoi last instar larva in dorsolateral view (93-SRNP-8715, 11 January 1994). 22, D. kidonoi last instar larva in anterior view (93-SRNP- 8715, 11 January 1994). 23, D. kidonoi prepupal larva in dorsolateral view (94-SRNP-147, 11 January 1994). 24, D. alemon penultimate instar larva in dorsolateral view (94-SRNP-8577, 10 October 1994). 25, D. alcmon last instar larva in anterior view (96-SRNP-1091, 22 May 1996). tal, thin, white ventrolateral line, something that hap- In all instars, the larva of D. kidonoi forms its shelter pens only once, near the anterior end, in our examples by silking together two halves of a leaflet (R. montana of D. alemon (Figs. 24, 25). has pinnately compound leaves), thus folding the For both D. eous and D. alcmon, Moss (1949:59) leaflet along its midrib so that its upper surface is in- notes that “the pupa squeaks audibly when touched.” side the shelter. Search for larvae is greatly facilitated VOLUME 53, NUMBER 3 Fics. 26-30. Pupae of two species of Drephalys (Drephalys) from the Area de Conservacion Guanacaste, Guanacaste, COSTA RICA. 26, D. kidonoi in ventral view (93-SRNP-8708, 11 January 1994). 27, D. kidonoi in lateral view (93-SRNP-8708, 11 January 1994). 28, D. kidonoi in anterior view (93-SRNP-8708, 11 January 1994). 29, D. alemon in ventral view (96-SRNP-1090, 22 May 1996). 30, D. alemon in anterodor- sal view (96-SRNP-1090, 22 May 1996). by looking for shelters on young saplings and sucker shoots, where the folded leaflets are easier to see and where they may also be more abundant. However, the larvae are occasionally encountered at all heights above the ground and on leaves of all ages. The larva walks out of the shelter to feed at night and remains within it by day. The larva of D. alemon lightly silks to- gether two leaves to form its shelter. The pupa of D. kidonoi (Figs. 26-28) is largely ivory white. It has a pair of conspicuous pink/orange/brown- colored thoracic spiracles (false eyespots) and a black/brown “mustache” between the ivory-colored 88 ag Fics. 31, 32. Roupala montana (Proteaceae)—larval food plant of Drephalys (Drephalys) kidonoi—in the Santa Elena Sector of the Area JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY de Conservacion Guanacaste, Guanacaste, COSTA RICA, during the dry season. 31, Sapling (1 m tall) at the age commonly used as larval food (15 January 1986). 32, Sapling (lower arrow) and adult tree (upper arrow) in their general habitat (8 April 1985). true eyes (Fig. 28). The true eyes turn red several days before eclosion and become dark brown the day be- fore. The pupa of D. alemon (Figs. 29, 30) is likewise ivory white with a pair of conspicuous pink/brown (false eye-like) thoracic spiracles, but it lacks the dark “mustache” (Fig. 30). The striking “face” on the ante- rior end of both species’ pupae is part of a pupal de- fense against diurnal vertebrate predators that is com- monplace in ACG hesperiids (Janzen in prep.). The pupa of D. kidonoi rests in the whitish and densely silked pupation chamber that is constructed from the last larval shelter. The last instar larval skin remains in the pupal chamber and lodges close to the point where the cremaster attaches to the silked walls. In captivity, the larva takes 45-55 days to develop from a newly-hatched first instar to prepupa. This puts D. kidonoi among the slower-growing pyrgine larvae that have been reared in the ACG (Janzen & Hall- wachs 1998). Slowness of growth probably is related to feeding on leaves that range from newly expanded to very old and tough. The last instar larva remains 3-4 days in the prepupal stage and 16-18 days in the pupal stage. Such durations are normal for a pyrgine hes- periid of this body weight. There is no hint of prepupal or pupal dormancy, either in the wet season or in the (very hot and dry) ACG dry season. More than 90% of the 236 larvae collected between 1991 and 1997 were found in the first half of the ACG dry season (late December through March) (Janzen 1993). Four pupae were found in the wild in mid-Feb- ruary 1992, and adults eclosed from them a week later. However, a few larvae were also found in April, May, July, August, and November. It would appear that D. kidonoi breeds mainly during the first half of the dry season. At this time, almost all other species of Hes- periidae that breed in this dry forest are sexually dor- mant adults on site, are dormant prepupae (very rarely), or have migrated out of these dry forests into nearby riparian bottomlands or the more distant ever- green, montane, cloud or rain forests to the east of the ACG dry forests. The ACG habitat currently occupied by the food plant, Roupala montana, is extensive, deforested, windswept, highly insolated, and dry (Fig. 32). Most, if not all, of this habitat has been generated by centuries of logging and burning, which have left large areas as rocky plains and knolls with low, sparse, native grasses and three species of widely scattered, stunted, and rel- VOLUME 53, NUMBER 3 atively fire-resistant trees: Curatella americana L. (Dilleniaceae), Byrsonima crassifolia (L.) DC (Malpighiaceae), and Roupala montana (Janzen 1988:fig. 26). In some of the old pastures, R. montana is the only species of tree present. All three of these fire-tolerant tree species are vertebrate-dispersed. Originally, the D. kidonoi population may well have persisted on a fragmented and low density R. montana population growing on cliff faces, ravine banks, and rocky outcrops scattered throughout the original old growth dry forest blanketing the ACG landscape. ACKNOWLEDGMENTS The following individuals and institutions generously provided material: F. H. Rindge, American Museum of Natural History. (AMNH), New York, New York; P. R. Ackery, The Natural History Museum (BMNH), London, England; C. D. MacNeill, California Academy of Sciences (CAS), San Francisco, California; G. Lamas, Museo de Historia Natural, Universidad Nacional Mayor de San Marcos (MUSM), Lima, Peru; O. H. H. Mielke, Departamento de Zoologia, Universidade Federal do Parana (UFPR), Curitiba, Parana, Brazil; National Museum of Natural History, Smithsonian Institution (USNM), Washington, D.C., and S. S. Nicolay. E. A. Klafter, D. J. Harvey, and P. Gentili dissected genitalia, and Y. T. Sohn drew them. C. C. Hansen photographed adults and digitally collated his color photos. A. Aiello, Lamas, Mielke, and G. T. Prance provided useful information. S. N. Burns helped in many and vari- ous ways. D. L. Pawson and the Research Opportunities Fund, and R. B. Simons, gave financial support to JMB. The caterpillar/host/parasitoid inventory that sparked this study got financial support from NSF grants BSR 90-24770, DEB 93- 06296, DEB-94-00829, and DEB-97-05072 to DHJ, plus financial, administrative, and logistic support from INBio, the government of Costa Rica, the Area de Conservacion Guanacaste, CONICIT of Costa Rica, Keidanren of Japan, and the Japan Childrens’ Rainfor- est. W. Hallwachs, C. Camargo, and A. Masis offered their expertise. Caterpillar collection and husbandry was conducted by the team of R. Moraga, G. Sihezar, G. Pereira, L. Rios, M. Pereira, O. Espinosa, E. Cantillano, M. Pereira, R. Franco, and H. Ramirez of the Area de Conservacion Guanacaste, Costa Rica. A. Aiello, B. Scholtens, and J. A. Shuey reviewed the manuscript. We are grateful to all. LITERATURE CITED AusTIN, G. T. 1995. Hesperiidae of Rond6nia, Brazil: Drephalys, with descriptions of two new species (Lepidoptera: Hesperi- idae: Pyrginae). Trop. Lepid. 6:123-128. BELL, E. L. 1942. New genera and new species of Neotropical Hesperiidae (Lepidoptera: Rhopalocera). Amer. Mus. Novit. 1205:1-9. BurGER, W. 1983. Proteaceae, pp. 8-14. In Burger, W. (ed.), Flora Costaricensis, Fieldiana, Botany, New Series, No. 13. Burns, J. M. 1964. Evolution in skipper butterflies of the genus Erynnis. Univ. Calif. Publ. Entomol. 37:1—-217. . 1996. Genitalia and the proper genus: Codatractus gets mysie and wvydixa—in a compact cyda group—as well as a hys- 89 terectomy, while Cephise gets part of Polythrix (Hesperiidae: Pyrginae). J. Lepid. Soc. 50:173-216. . [1999] “1998.” Pseudodrephalys: a new genus comprising three showy, neotropical species (one new) removed from—and quite remote from—Drephalys (Hesperiidae: Pyrginae). J. Lepid. Soc. 52:364-380. Burns, J. M. & D. H. JANZEN. In prep. Larval food plants of pyrgine and pyrrhopygine Hesperiidae (the dicot-eaters) in the tropical lowland dry forest of the Area de Conservacion Gua- nacaste, northwestern Costa Rica. CASAGRANDE, M. M. & O. H. H. MIELKE. 1992. Borboletas (Lepi- doptera) ameacadas de extingao no Parana. Revta bras. Zool. 9:75-92. Cock, M. J. W. 1984. The skipper butterflies (Hesperiidae) of Trinidad. Part 3 Pyrginae (first section). Living World. J. Trinidad & Tobago Field Naturalists’ Club, 1983-1984:38-42. Evans, W. H. 1952. A catalogue of the American Hesperiidae indi- cating the classification and nomenclature adopted in the British Museum (Natural History). Part II. Pyrginae. Section 1. British Museum, London. 178 pp., pls. 10-25. GopmaN, F. D. & O. SALvIN. 1879-1901. Biologia Centrali-Ameri- cana; Insecta; Lepidoptera-Rhopalocera. Vol. 2, 782 pp.; Vol. 3, 113 pls. Hee W. C. 1877. Descriptions of twenty-three new species of Hesperidae from his own collection. Ann. Mag. Nat. Hist. (4)20:319-328. JANZEN, D. H. 1988. Guanacaste National Park: tropical ecological and biocultural restoration, pp. 143-192. In Cairns, J. J. (ed.), Rehabilitating damaged ecosystems. Vol. II. CRC Press, Boca Raton, Florida. . 1993. Caterpillar seasonality in a Costa Rican dry forest, pp. 448-477. In Stamp, N. E. & T. M. Casey (eds.), Caterpillars. Ecological and evolutionary constraints on foraging. Chapman and Hall, New York. . Inprep. Larval and pupal defense through eye mimicry by Costa Rican Lepidoptera. JANZEN, D. H. & W. Hatiwacus. 1998. Janzen and Hallwachs caterpillar rearing database. . Filemaker Pro 4.0 database accessible through Netscape and other web browsers. LATREILLE, P. A. & J. B. Gopart. 1819-1824. Encyclopedie méthodique. Histoire naturelle des insectes. Vol. 9, 828 pp. MABILLE, P. & E. BOULLET. 1919. Essai de revision de la famille des hespérides. Ann. Sci. nat. Zool. (10)2:199-258. MIELKE, O. H.H. 1968. Duas espécies novas de Drephalys, prove- nientes de Santa Catarina (Lepidoptera, Hesperiidae). Atas Soc. Biol. Rio de Janeiro 12:129-133. Moss, A. M. 1949. Biological notes on some “Hesperiidae” of Para and the Amazon (Lep. Rhop.). Acta Zool. Lilloana 7:27-79, pls. LV. PRANCE, G. T. 1989. Chrysobalanaceae. Flora Neotropica, Mono- graph 9S:1—267. PRANCE, G. T. & D. G. CAMPBELL. 1988. The present state of trop- ical floristics. Taxon 37:519-548. Watson, E. Y. 1893. A proposed classification of the Hesperiidae, with a revision of the genera. Proc. Zool. Soc. London 1893(1):3-132, pls. LIII. Received for publication 16 November 1998; revised and accepted 15 July 1999. Journal of the Lepidopterists’ Society 53(3), 1999, 90-98 NEW PRONOPHILINE BUTTERFLIES FROM THE VENEZUELAN TEPUYES (NYMPHALIDAE: SATYRINAE) ANGEL L. VILoRIA! Biogeography & Conservation Laboratory, The Natural History Museum, Cromwell Road, London SW7 5BD, Great Britain AND TOMASZ W. PYRCZ Muzeum Zoologiczne Instytutu Zoologii Uniwersytetu Jagiellofiskiego, Ingardena 6, 30-060 Krakéw, Poland ABSTRACT. Four new species of satyrine butterflies collected in four Venezuelan tepuyes are described and illustrated: Protopedaliodes ridouti from the Roraima-Tepui, Protopedaliodes profauna and Pedaliodes terramaris from the Auyén-Tepui, and Pedaliodes yutajeana from the Cerros Yutajé, Yavi, and Marahuaka. Taxonomic considerations and discussion on affinities are presented. Additional key words: Pantepui, Pedaliodes, Pleistocene, Protopedaliodes, Venezuela. Venezuelan scientists involved in research on the tepuyes (most of them currently within national parks) have made major efforts to decrease and limit the so- called “ecological tourism” in this region. This is due mainly to their concern over the dramatic degradation of the fragile environments at the tops of these moun- tains. Propaganda recently generated by science fic- tion films (often recalling the imaginary Lost Worlds of Conan-Doyle and Crichton) and persons attempting to set new Guinness’ records in the tepuyes generate public clamor and still more visitor interest in the area. The accumulated result of such misguided publicity, together with the general paranoia about commercial collecting of wild animals and plants anywhere in the national territory, has resulted in the thoughtless en- forcement of rigorous laws that virtually prohibit bio- logical research in most of the Venezuelan protected areas. Consequently, there is now an exceedingly com- plex bureaucy to deal with in applying for insect col- lecting permits in the tepuyes (and elsewhere in Venezuela). Generally speaking, three separate ap- plications must be prepared for the consideration and approval, respectively, of the National Council for Scientific and Technological Research (CON- ICIT), the National Parks Institute (INPARQUES), and finally the Service for the Fauna of the Ministry of Environment and Natural Resources (PRO- FAUNA). They have to be submitted synchronously and well in advance, taking into account the fact that the delivery of the third of these depends on the ap- proval of the second, and so on. Even if these end- less requirements were all satisfactorily met at the initial submission, we have found that there is no guarantee of receiving such permission, and when "Current address: Museo de Biologia, Facultad Experimental de Ciencias, La Universidad del Zulia, Apdo. 526, Maracaibo 4011, Zu- lia, Venezuela. given, permits are often so badly delayed that are out-of-date and useless. To face this problem has become an essential worry for Venezuelan scientists (let alone foreigners), who find themselves handicapped in their field work, even if they are entirely innocent of any involvement with film and TV productions, or commercial dealing with biological specimens. As “illegal” procedures are being sternly punished with confiscation of material, finan- cial penalties, and menace of imprisonment, it is rather frustrating to find that we are virtually forbidden to study our own biota while at the same time extensive gold mining (never controlled as are insect collecting activities) is quickly devastating large areas of pristine forests in marginal regions of the Venezuelan territory. These include all National Parks south of the Orinoco. Massive “ecotourism” and all its undesirable conse- quences however continues with no problems of per- mission (just a local application, approved on the day of submission) in the Canaima National Park and all of the tepuyes, even in the remotest Cerro de La Neblina. This scenario leads us to believe that it will take sev- eral decades for Venezuelan and international scien- tists to be able to study good series of entomological samples from the tepuyes, which can satisfy the ac- cepted paradigm of having two or more individuals to proceed to a satisfactory taxonomic description. In the meantime, we strongly feel that a few butterflies we know as undescribed, collected in four of the 54 tepuyes existing in Venezuela (all potentially populated by these insects), deserve to be described as part of a major revisionary work of the group currently being undertaken by the senior author. Two of these unde- scribed taxa are known from single male individuals. One is so distinctive within an endemic and hitherto monobasic genus (which was erected by ourselves in 1994) that we would not hesitate in providing a new place for it in the increasing list of Neotropical butter- VOLUME 53, NUMBER 3 fly names; the other one, although externally undistin- guished, is also a new member of a small group of species which may be unmistakably recognized by the contortion of the tip of the male genitalic valvae. Four descriptions are presented here. All type spec- imens, except for the solitary individual of Protope- daliodes ridouti, new species, which is held by The Natural History Museum (BMNH) in London, are de- posited in the Museo del Instituto de Zoologia Agri- cola of the Universidad Central de Venezuela (MIZA) in Maracay, as required by the Venezuelan environ- mental authorities. Protopedaliodes ridouti Viloria & Pyrcz, new species (Figs. I 7) Description. Male. Forewing length: 30 mm (n = 1). Eyes dark brown, hairy. Palpi twice as long as head, covered with long dark- brown hair. Antennae to over half costa. Thorax and abdomen all dark brown; walking legs (second pair) same length as in P. kukenani Viloria & Pyrez (third pair broken in holotype). Forewing subtrian- gular, apex and tornus obtuse, outer margin convex and smooth; hindwing oval, outer margin smooth. Upperside all dark brown, blackish in basal and postbasal areas; faint, barely visible ocellus in cell M1, black with white pupil. Underside forewing ground color dark brown, sparsely sprinkled with lighter brown or silvery scales on apex and upper one third of submarginal area; ocellus in cell M1 large, as wide as cell, black, pupilled with white, circled with faint orange framing. Hindwing ground color similar to forewing but lib- erally sprinkled with lighter brown scales over the entire surface; lighter area between postmedian and submarginal lines, forming a 5-8 mm wide band, being the narrowest in cell M2 and towards tor- nus on vein 1A (3 mm), its inner and outer edge do not merge; two large ocelli in cells R5 and Cul, of same shape, color and size as on forewing. Genitalia illustrated in Fig. 7. Female. So far unknown. Types. Holotype. Male, Mt. Roraima, 8000 ft, Venezuela, 12-ix- 1974, B. V. Ridout, B. M. 1974-650, BM(NH) Rhopalocera vial number 4198. Etymology. We dedicate the name of this species to its collector, Dr. B. V. Ridout. Distribution. This species is only known from the summit of the Roraima-Tepui (=Mount Roraima, Fig. 11), where it probably flies sympatrically with Protopedaliodes kukenani, and another pronophi- line species (presumably of the genus Lymanopoda Westwood; Orellana, pers. comm.). For a description of the habitats, vegetation, and general geographic aspects of this region, see Brewer (1984). Protopedaliodes profauna Viloria & Pyrcz, new species (Figs. 2, 3, 8) Description. Male. Forewing length: 32-32.5 mm, mean = 32.25 (n = 2). Eyes coffee brown, covered by short black hairs. Palpi twice as long as head, pale brown; outer ventral long hairs coffee ' brown, inner hairs shorter and light brown. Antennae reaching half of costa, black, except for ventral region of club, which is brown. Body dorsally coffee brown with very shiny hairs, ventrally pale brown, lighter towards abdomen, in general very hairy (including leg femorae), but hairs denser and shorter than on dorsal surface. Forewing triangular, apex and tornus slightly rounded, outer margin slightly convex; hindwing subtriangular, tornus somewhat truncated, outer margin moderately scalloped. Dorsal ground color of wings 91 dark coffee brown, very shiny; creamy-light brown scales between veins in fringes of both wings; brownish sheen on hindwing anal margin region; forewing exhibiting very thick androconial patches in discal region; hindwing very hairy on basal two thirds. Underside ground color of wings similar to upperside; lighter postdiscal bands on both wings, laterally limited (except near tornus) by darkening of ground color. Forewing band anteriorly broadened, dusted with red- dish scales; sparse white scales on subapical and apical regions, more densely in inner border of band near costa; sparse short white hairs along costal region. Hindwing covered by short creamy-white setae, more densely towards basal region, anal margin, and on discal band; reddish scales dusted over basal region, in space anterior to cell. Genitalia in Fig. 8. Female. Forewing length: 33.5 mm (n = 1). The only known fe- male is worn and differs from male in the following features: general color much paler; dorsal wing color rather paler towards distal third, particularly on forewing. Types. Holotype. Male, Auyén-Tepui, 1700 m, Bolivar, Venezuela, 5°58N, 62°32/W, 14/19-ii-1994, J. L. Garcia, A. Chac6n. Paratypes. 1 male same data; 1 female, Auyén-Tepui, 1800 m, Bolivar, Venezuela, 5°51’N, 62°35’W, 4/10-ii-1988, L. J. Joly & A. Chacon. Etymology. This butterfly bears the name of one of the Venezuelan environmental institutions mentioned in the introduc- tion. We do that because of the resulting euphony. Distribution. Apparently endemic to the Auyan-Tepui (Fig. 11), an extensive table mountain massif in southeastern Venezuela. Gen- eral accounts of the geography and ecological aspects of this region were presented by Brewer (1978) and Fundacion Terramar (1993). P. profauna seems to fly in a different altitudinal zone located above the species described below. Pedaliodes terramaris Viloria & Pyrcz, new species (Figs. 4, 9) Description. Male. Forewing length: 27 mm (n = 1). Eyes black, covered by short black hairs. Palpi twice as long as head, hairy, dor- sally and ventrally black, laterally creamy-white. Body dorsally cov- ered by dark, bright, brown hairs, ventrally pale brown (including hairs covering femorae), somewhat reddish on anterior part of tho- rax. Forewing triangular, apex and tornus softly rounded, outer mar- gin more or less linear; hindwing suboval, outer margin excavated between veins. Wing upperside ground color chocolate brown, very dark in discal region (of both wings), lighter towards basal region and distal third, except in marginal region; hindwing also lighter in marginal region. Forewing upperside bearing six androconial patches in discal region, the two elongated ones in cell Cu2 not as distant as in P. yutajeana, new species. Wing underside groundcolor chocolate brown, postdiscal bands lighter, bordered distinctly but ir- regularly with darker lines that never reach tornus; marginal region reddish chestnut, flanked by fine dark chocolate brown lines on both sides; forewing basal third, region adjacent to costa, and inner mar- gin, almost as light as postdiscal band; basal half of wing densely cov- ered by short brownish hairs; some white scales over costal portion of band inner border; discal cell finely sprinkled with dark chocolate brown scales. Hindwing underside sprinkled with brown and red- dish scales (the latter less conspicuous on postdiscal band), anal re- gion suffused with brick-orange, and dusted with yellow scales; two submarginal white dots within band in cells Cu2 and M3, respec- tively; basal region very hairy. Genitalia illustrated in Fig. 9. Female. So far unknown. Types. Holotype. Male, Auyan-Tepui, 1500 m, Bolivar, Venezuela, 5°57’N, 62°39’W, 19/24-ii-1994, A. Chacon. Etymology. The specific name, terramaris, is a derivation from the name of the Fundacién Terramar, a private Venezuelan organi- zation that has been responsible for much of the recent biological exploration of the tepuyes. Distribution. Only known from the slopes of the Auyan-Tepui, where it flies in lower cloud forest. 92 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY VOLUME 53, NUMBER 3 Pedaliodes yutajeana Viloria & Pyrez, new species (Figs. 5, 6, 10) Description. Male. Forewing length: 29 mm (n = 1). Eyes black covered by black hairs (with reddish sheen). Palpi twice as long as head, hairy, dorsally and ventrally black, laterally yellowish white. Antennae reaching approximately half of costa, dorsally dark brown, ventrally reddish. Thorax dorsally black, covered by very bright dark coffee hairs, as well as rest of body, ventrally lighter. Forewing trian- gular, apex and tornus slightly rounded, outer margin very softly sin- uate; hindwing subtriangular, outer margin convex and moderately excavated between veins. Wing upperside ground color dark coffee brown, shiny, slightly lighter towards distal quarter (particularly in forewing), some light creamy-brown scales between veins in fringes; androconial patches on forewing discal region contiguous, two of them lengthened, and running parallel in cell Cu2. Wing underside ground color dark chocolate brown, lighter towards postmedial re- gion. Forewing also lighter in basal quarter; some white scales dusted over costal region of band; reddish scales dusted over sub- apical region; six tiny submarginal white dots in cells R4 to Cul; dense, short, reddish hairs on anterior portion of basal region. Hind- wing postmedial band less distinct than on forewing; reddish suffu- sion in tornus and anal margin area; yellow scales dusted within band, from tornus along its inner margin, to costa; basal third of wing hairy; one submarginal white dot in cell Cul, another one (ves- tigial) in M3; costal and marginal area with chestnut tone. Genitalia illustrated in Fig. 10. Female. Forewing length: 30-32 mm, mean = 31 mm (n = 2). In general bigger than male, with less bright coloration. Dorsally with light postdiscal bands, which on hindwing containing a suffusion of brick-reddish in costal region near apex. Ventral pattern similar to male, but ground color with general speckling of reddish chestnut scales (almost imperceptible within bands); marginal region reddish; forewing submarginal white dots variable, sometimes missing; hind- wing exhibiting a contrasting “marble pattern” as a result of reddish and chocolate brown speckling over plain brown ground color; yel- lowish scales uniformly dusted over entire surface, especially con- centrated in inner border of postdiscal band; series of five submar- ginal white dots in cells R5 to A2; anal region with reddish suffusion as in male. Types. Holotype. Male, Cerro Marahuaka, 2470 m, Parque Na- cional Duida-Marahuaka, Amazonas, Venezuela, 3°37'N, 65°22’W, 3/6-ii-1992, Exp[edicion]. Terramar, J. Clavijo, A. Chac6én. Paratypes. 1 female, Cerro Yutajé, 1750 m, Amazonas, Venezuela, 5°45’/N, 65°08'W, 12/17-ii-1995, J. Clavijo A., Exp[edicion]. Terramar; | fe- male, Cerro Yavi, 2200 m, Amazonas, Venezuela, 5°43’N, 65°54’W, 24/28-ii-1995, J. L. Garcia, Exp[edicién]. Terramar. Etymology. The name of the species is derived from one of the original localities, the Cerro Yutajé. Distribution. This species is distributed in an extensive, discon- tinuous, montane area of northern Amazonas State, from Cerro Yavi and Yutajé to Cerro Marahuaka (Fig. 11). This range implies that it may also be found in the intervening mountains: the Sierra de Maigualida, Jaua-Sarisarifiama massif, and certainly the Cerros Huachamacari and Duida, which are adjacent to the Marahuaka. Geographical and ecological aspects of the area, as well as recent bi- ological discoveries, are discussed by Michelangeli et al. (1988) and Fundacion Terramar (1989, 1993). — 93 DISCUSSION Protopedaliodes. The genus Protopedaliodes was recently erected for a species (P. kukenani Viloria & Pyrcz, 1994) from the upper cloud forest on neighbor- ing table mountains in South-Eastern Venezuela, Kukenan-Tepui and Roraima-Tepui. Further research in 1995 in the Natural History Museum (BMNH) re- vealed the existence of a small collection of butterflies made by B. V. Ridout on the top of Mount Roraima. This material comprises 16 males and two females of P. kukenani (plus one male collected on the north ridge of Roraima, on the Guyana side, at 7400 ft., by Adrian Warren). This large series agrees with the original de- scription of P. kukenani and no modifications of the specific diagnosis are required. However, among the Ridout material a single male was readily recognized to represent the second species of the genus (P. ri- douti). In 1996 we examined the material recently col- lected on the tepuyes by staff members of the Museo del Instituto de Zoologia Agricola of the Universidad Central de Venezuela (MIZA), and found both P. pro- fauna and the two species of Pedaliodes Butler also de- scribed in this paper. The above mentioned specimens of Protopedaliodes, plus the type series of P. kukenani (in MIZA) and five additional individuals of this species (three males, two females) obtained by the American mammalogist G. H. Tate in the summit of Roraima in 1927 (deposited in the American Museum of Natural History, New York [AMNH)]), are to our knowledge the only ones existing in scientific collections. Protopedaliodes ridouti is easily distinguished from its allies, P kukenani and P. profauna, by its wing shape and quite different hindwing underside pattern, especially the well developed ocelli in cells R5 and Cul. Although some specimens of P. kukenani also have faint ocelli in cell R5 of the underside of the forewing, in P. ridouti the ocelli are very well developed in forewing cell M1 and in cells R5 and Cul of the hindwing. The wing pattern of P. ridouti is unusual for pedaliodine butterflies, and is reminiscent, but perhaps not homol- ogous to, that of the genus Praepronophila Forster (1964) (see also Miller 1986). It places P. kukenani well apart from other members of the tribe. The type specimen of P. ridouti (30 mm) is slightly smaller than average sized P. kukenani (mean 32.8 Fics .1-3. 1, Protopedaliodes ridouti Viloria & Pyrez, new species, Holotype. Male, Mt. Roraima, 8000 ft, Venezuela, 12-ix-1974, B. V. Rid- out, B. M. 1974-650, BM(NH) Rhopalocera vial number 4198; right upperside, left underside. 2, P. profauna Viloria & Pyrez, new species, Holo- type. Male, Auyan Tepui, 1700 m, Bolivar, Venezuela, 5°58’N, 62°32’W, 14/19-ii-1994, J. L. Garcia, A. Chacon; right upperside, left underside. 3, P. profauna Viloria & Pyrez, new species. Female paratype, Auyan Tepui, 1800 m, Bolivar, Venezuela, 5°51’N, 62°35’W, 4/10-ii-1988, L. J. Joly & A. Chacén; right upperside (forewing discal white mark represents rubbing of the scales), left underside. 94 JOURNAL OF THE LEPIDOPTERISTS * SOCIETY VOLUME 53, NUMBER 3 mm). Its wing shape differs, the hindwing apical and tornal corner being more angular and giving the wings of P. ridouti a slightly square appearance. Venation is the same for all three species of Protopedaliodes, but the wing fringes of P. ridouti are shorter. The male genitalia of P. ridouti show certain very characteristic features common to this species and to P. kukenani, such as the extremely long, straight and toothed aedeagus, and the deep saccus. On the other hand, subunci are nearly atrophied in P. ridouti, its un- cus is even longer than in P. kukenani, and its valvae are devoid of any secondary process. The wing pattern of P. profauna is simple and resembles P. kukenani. The genitalia of P. profauna is structurally characteris- tic of the genus, but the aedeagus is three times as broad as those of the two other species, and it lacks the tooth at the tip; the uncus is relatively short, thick- ened, and remarkably bifurcated at the extremity (so far a unique feature in the tribe it belongs to, the Pronophilini); the saccus is as deep as in the two other species, although curved downwards; the valvae re- semble those of P. kukenani, but are more stylized. Common features among the species of Protope- daliodes, such as the ground color of the upperside, the lack of any androconial patch on the forewing upper- side, similar head and leg morphology, and the charac- teristic male genitalia, confirm the validity of the genus when compared to other American Pronophilini. The existence of further species of Protopedaliodes in the Guayana shield area (one certainly sympatric with P. kukenani) directly implies that some radiation occurred within the Pantepui which, until the present time, has been poorly researched, as compared to avian or mammalian faunas or high altitude floras. The cloud forest fauna of the upper slopes of the table mountains in the Pantepui seems to be impoverished compared to similar montane habitats in the Andes. This is possibly due to the isolation of this region and relatively small area of suitable cloud forest habitats. Most butterfly species (including Pedaliodes, see be- low) reported to date from the cloud forests of the Pantepui are endemic, but apparently offshots of the Andean fauna (see Strand 1912, Brown 1932, Viloria [1995], 1998, Viloria & Pyrez 1995, Pyrez 1995, Neild 1996), at least in those cases where affinities of the species can be recognized. — Protopedaliodes cannot yet be related with certainty to any pronophilines known from the Andes. We pre- viously suggested that it has possible affinities with the less derived lineage of Pedaliodes (sensu lato) (i.e., Praepronophila, Parapedaliodes Forster). This as- sumption was based, among other characters, on male genitalic morphology (which is also reminiscent of that of Praepronophila). It seems unlikely that Protopedaliodes originated from modern Andean “colonizers”, and we favor the hypothesis that it is derived from older elements proper to the Pantepui. A fast adaptive radiation of Protopedaliodes could also be involved as a factor ob- scuring its phyletic origins. Pedaliodes. Pedaliodes Butler sensu stricto is cer- tainly one of the most speciose genera of Satyrinae in the world. We recognize 132 described valid species plus nine subspecies, and 105 species (excluding the two described here) plus 22 subspecies confidently identified as new, undescribed taxa, which are de- posited in seven major entomological collections in America and Europe (Viloria unpubl.). This makes a grand total of 270 taxa, most of which are highly en- demic to restricted montane areas of the tropical An- des. Only 15 species are known to occur out of the An- des, five of them being restricted to the mountains of the Pantepui (i-e., P. roraimae Strand, 1912 (Gran Sa- bana, Roraima-Tepui and Kukenan-Tepui, 1280-1900 m), PB demarmelsi Viloria, [1995] (Cerro de La Neblina, 1690-2100 m), P. chaconi Viloria, 1998 (Ser- rania de Tapirapec6, 1300 m), P. terramaris (Auyan- Tepui, 1500 m), and P. yutajeana (tepuyes of northern Amazonas State, from Cerro Yavi to Cerro Duida, 1750-2470 m)). Other alleged records from the Guayana region, such as Pedaliodes prytanis (Hewit- son) (Adams & Bernard 1979:109) and P. manis (C. & R. Felder) (d’Abrera 1988:852), are incorrect. The first one is based on two old specimens mislabelled as be- ing from Corosita, Caura Valley (in the BMNH), which were purchased by J. J. Joicey from Klages. This is obviously wrong as P. prytanis is endemic to the highest elevations of the Cordillera de La Costa, where Klages obtained part of his collections. The sec- ond case represents misidentifications of four males of P. roraimae (from Mount Roraima, Venezuela) in the same institution. Fics. 4-6 4, Pedaliodes terramaris Viloria & Pyrez, new species, Holotype. Male, Auyén Tepui, 1500 m, Bolivar, Venezuela, 5°57’N, 62°39°W, 19/24-ii-1994, A. Chacén; right upperside, left underside. 5, P. yutajeana Viloria & Pyrez, new species, Holotype. Male, Cerro Marahuaka, 2470 m, Parque Nacional Duida-Marahuaka, Amazonas, Venezuela, 3°37’N, 65°22’W, 3/6-ii-1992, Exp. Terramar, J. Clavijo, A. Chacén; right upperside, left underside. 6, P. yutajeana Viloria & Pyrcz, new species, Female paratype, Cerro Yutajé, 1750 m, Amazonas, Venezuela, 5°45’N, 65°08’W, 12/17-ii-1995, J. Clavijo A., Exp. Terramar; right upperside, left underside. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 7-10. Male genitalia of the species herein described; in each case aedeagus and left valva have been removed from natural positions. 7, Protopedaliodes ridouti. 8, P. profauna. 9, Pedaliodes terramaris. 10, P. yutajeana. The species of Pedaliodes found south of the Orinoco River are all closely related, as deduced by the strong similarities in wing pattern and genitalia. With the exception of P. demarmelsi (which exhibits a very distinctive color pattern), they are, in fact, almost indistinguishable from each other in facies. Pedaliodes terramaris and P. yutajeana, however, can be told apart by subtle differences in size and wing shape (compare Figs. 4 and 5), and by the differences in the distribution of the male androconial patch on forewing cell Cu2 (see descriptions above). The shape and extent of the male forewing scent patches have proved to be most useful in identifying Pedaliodes species with few wing markings, and using these characters we have been able to recognize seven “black” species occurring sym- patrically in the Colombian Cordillera Occidental (Pyrez & Viloria 1999b), all previously misidentified under one or two names (Adams 1986). When compared, the valvae of P. roraimae and P. chaconi are shorter and more robust than those of P. terramaris and P. yutajeana. Differences between P. roraimae and P. chaconi were discussed in a previous publication (Viloria 1998). The Pantepuian Pedaliodes belong to a group that is not restricted to the Guayana biogeographical region. On the contrary, this is the most widely distributed clade within the genus, ranging from Mexico to Bolivia (see below). Viloria [1995] pointed out the structural and superficial similarities between the Pantepuian P, demarmelsi, and the Mesoamerican species P. dejecta (Bates) and P. napaea (Bates). This observation was at that time surprising, because of the apparently huge distributional gap between Mesoamerica and the Cerro de La Neblina, especially for these montane in- sects. However, a better understanding of the mor- phology, taxonomy, and distribution of a number of closely related species that occur at lower altitudinal levels (the lowest possible for the genus) in the cloud forest of almost every mountain range in between these range extremes has led us to believe that the group probably started diverging in isolation only geo- logically very recently, perhaps from a single wide- spread lowland ancestor. Although Neotropical cloud forest satyrines are ex- ceedingly sedentary and do not migrate (De Marmels et al. 1996, Viloria et al. in prep.), it is possible that past global climatic fluctuations may have led to the lowering and conjugation of cloud forests, enabling the ancestor of these butterflies to spread between cur- rently isolated cloud forest “islands.” Results of paleoclimatic studies in Venezuela (Rull 1996) and other areas of the northern Neotropics VOLUME 53, NUMBER 3 66° 67° 68° 69° 70° Caribbean 7° 72° 73° 74 Fic. 11. Localities of the Pantepui region mentioned in the text: 1, Cerro Yutajé. 2, Cerro Yavi. 3, Sierra de Maigualida. 4, Jaua-Sarisar- iflama massif. 5, Cerro Huachamacari. 6, Cerro Marahuaka. 7, Cerro Duida. 8, Cerro de La Neblina. 9, Serrania de Tapirapecd. 10, Auyan- Tepui. 11, Kunenan-Tepui. 12, Roraima-Tepui (modified from Steyermark 1986, Fundacién Terramar 1993 and Huber 1995). (Schubert 1987), indicate much lower average temper- atures at the end of the Pleistocene, and although con- ditions were also much drier in some areas, this does not rule out the possibility that the floristic equivalent of cloud forest could have been present in others. We speculate that not only the Pantepuian Pedaliodes, but also all allied congeners’ elsewhere in the Neotropics, may be derived from an ancestor which was wide- spread at lower elevations during the Pleistocene. ‘The members of this group of allied congeners outside the Pan- tepui are: Pedaliodes croizatorum. Viloria and Camacho (Serranja del Turimikire, northeastern Venezuela, 1500-2300 m); P. pisonia (Hewit- son) (Venezuelan Cordillera de La Costa, 1100-1700 m); P. manneja | Thieme (Cordillera de La Costa and Sierra de Perijé, 1800-2300 m); P, montagna Adams & Bemard (Andes from Venezuelan Cordillera de Mérida to Bolivian Yungas, 1050-3000 m); P. ereiba (C. & R. Felder) (Cordillera Oriental, Colombia, ca. 1800 m); Pedaliodes canela (=Ped- aliodes canela) Pyrcz and Viloria (Cordillera Occidental, Colombia, 1000-3300 m); P. phrasiclea Grose-Smith (Andes of Colombia to Bo- livia, 1000-2250 m); P. pomponia (Hewitson) (Andes of southeastern Ecuador, 450-1400 m); Pedaliodes balnearia Pyrcz and Viloria (1999a) ACKNOWLEDGMENTS We thank the trustees of the BMNH for allowing us to study the material deposited in this institution; P. Ackery for locating the Rid- out collection in the BMNH in 1995; J. Reynolds for his kind assis- tance in the Museum; D. C. Lees, R. I. Vane-Wright, G. Lamas, and G. W. Beccaloni for reviewing, discussing and editing several drafts of this paper; and J. Wojtusiak for his relentless support for our stud- ies on the satyrines. We also thank T. Emmel, M D. Bowers and an anonymous referee for critical reading and editorial improvements to this text, and J. De Marmels for his encouragement and the gen- erous loan of the specimens from the MIZA, which most of this (southeastern Ecuador, 2000-2600 m); Pedaliodes, new species (Zamora valley, 1000-1300 m); Pedaliodes, new species, Lamas & Vilo- ria, MS (southern Ecuador to northem Bolivia, 1800-2300 m); P. phra- sis Grose-Smith (Andes of southern Peru and Bolivia, 750-3000 m); P prosa Staudinger (southeastern Peru and Bolivia, 1000-3000 m); P. de- jecta Bates (mountains of Panama, Costa Rica and Guatemala, 850-2300 m): P. cremera Godman & Salvin (Irazti volcano, Costa Rica, ca. 2000 m); P. napaea Bates (mountains of Guatemala and southern Mexico [Chiapas], 1000-1700 m); P. circumducta Thieme (Mexico, 1100-1450 m); Pedaliodes sp. [nov.] Luis-Martinez and Llorente (1993) (Mexico, Puerto Los Mazos, Jalisco, Michoacén Mountains and Sierra de Atoyac in Guerrero, ca. 1000-2000 m). 98 study is based on. Our gratitude is also expressed to J. M. Gonzalez (Fundacién Terramar) for successfully locating the Tate collection of Venezuelan satyrines in the AMNH. The junior author acknowl- edges A. Mee and A. Neild for their hospitality in London, for their company and for editing preliminary manuscript notes. This re- search has been supported by The British Council, CONICIT, and La Universidad del Zulia (PhD grant to ALV), and the Institute of Zoology of the Jagiellonian University (intemal grant DS/IZ 1995/01, to TWP). LITERATURE CITED Apams, M. J. 1986. Pronophiline butterflies (Satyridae) of the three Andean Cordilleras of Colombia. Zool. J. Linn. Soc. 87:235-320. ApAMS, M. J. & G. I. BERNARD. 1979. Pronophiline butterflies (Satyridae) of the Serrania de Valledupar, Colombia-Venezuela border. Syst. Ent. 4:95-118. BREWER, C. 1978. La vegetacién del mundo perdido. Fundacién Eugenio Mendoza, Caracas. 223 pp: . 1984. Roraima, la montamia de cristal. Editorial Arte, Cara- cas. 153 pp. Brown, F. M. 1932. Pieridae from the regions of Mt. Duida and Mt. Roraima. Am. Mus. Nov. 572:1-7. D’ABRERA, B. 1988. Butterflies of the Neotropical Region. Part V. Nymphalidae (Conce.) and Satyridae. Hill House, Victoria. Pp. [viii] + 679-877. DE MARMELS, J., J. A. CLAvljo & M. E. Cuacin. 1996. A new sub- species of Xylophanes tersa (Sphingidae) from Venezuela. J. Lepid. Soc. 50:303-308. Forster, W. 1964. Beitriige zur Kenntnis der Insektenfauna Bo- liviens XIX. Lepidoptera III. Satyridae. Veroff. zool. Staatssam. Miinchen 8:51-188, pls. 27-35. FUNDACION TERRAMAR (ED.). 1989. Bidsfera del Marahuaka y zonas adyacentes (Territorio Federal Amazonas-Venezuela). Acta Ter- ramaris 1:1—107. . 1993. Informe técnico sobre los tepuyes, Formacién Ro- raima, Venezuela. Acta Terramaris 6:1—74. Huser, O. 1995. Geographical and physical features, pp. 1-61. In Berry, P. E., B. K. Holst & K. Yatskievych (eds.), Flora of the Venezuelan Guayana. Volume 1: Introduction. Missouri Botan- ical Garden/Timber Press, St. Louis/Portland. LuIs-MARTINEZ, A. & J. LLORENTE-BOUSQUETS. 1993. Mari- posas, pp. 307-385 In Luna, I. & J. Llorente (eds.), Histo- ria natural del Parque Ecolégico Estatal Omiltemi, Chilpancingo, Guerrero, México. Conabio-Universidad Na- cional Auténoma de México, Ediciones Técnico Cientificas, México. MICHELANGELI, A., F. MICHELANGELI, R. S. BORGES, W. SMITTER, A. SUBERO & K. JAFFE (EDS.). 1988. Marahuaka. Editorial Ar- mitano, Caracas. 352 pp. MILLER, L. D. 1986. The relationship between Pedaliodes per- perna and petronius (Satyridae), with the description of a new subspecies. J. Lepid. Soc. 39:187-195. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY NEILD, A. F. E. 1996. The butterflies of Venezuela. Part 1: Nymphalidae I (Limenitidinae, Apaturinae, Charaxinae). Meridian Publications, Greenwich. 144 + [ii] pp., 32 pls. Pyrcz, T. W. 1995. A new species of the genus Memphis Hiibner (Nymphalidae: Anaeini) from the Pantepui. Trop. Lepid. 6:11-13. Pyrcz, T. W. & A. L. Vitoria. 1999a. Contribution to the knowl- edge of the Ecuadorian Pronophilini, Second Part; new ped- aliodines from Ecuador (Lepidoptera: Nymphalidae: Satyri- nae). Genus (Wroclaw), 10:117—150. . 1999b. Mariposas de la tribu Pronophilini de la Reserva Forestal Tambito, Cordillera Occidental, Colombia. Primera Parte. Convergencia de los patrones de coloracién en mariposas andinas: siete nuevas especies del género Pedaliodes Butler, 1867 (Lepidoptera: Nymphalidae, Satyrinae). SHILAP, Revista de lep- idopterologia (Madrid) 27(106):173-187. RuLL, V. 1996. Late Pleistocene and Holocene climates of Venezuela. Quaternary Int. 31:85-94. SCHUBERT, C. 1987. Climatic changes during the last glacial maxi- mum in northern South America and the Caribbean: a review. Interciencia 13:128—-137. STEYERMARK, J. A. 1986. Speciation and endemism in the flora of the Venezuelan Tepuis, pp. 317-373. In Vuilleumier, F. & M. Monasterio (eds.), High altitude Tropical biogeography, Oxford University Press, New York. STRAND, E. 1912. Zwei neue Satyriden von Roraima. Gesammelt von Herrn Botaniker E. Ule. Fauna Exotica. 2:47— 48. Vitoria, A. L. [1995]. Description of a new species of Pedaliodes (Lepidoptera: Satyridae: Pronophilini) from the Cerro de La Neblina, Venezuela. Atalanta 25:525-529, pl. XVIIa. . 1998. Un nuevo Pedaliodes Butler, 1867 de la Serrania de Tapirapec6é, Venezuela (Lepidoptera: Nymphalidae, Satyrinae, Pronophilini). SHILAP Revista de lepidopterologia (Madrid). 96:13-18. VitoriA, A. L., M. J. ADAms, T. W. Pyroz & F. ROMERO. Noticia hist6rica sobre satfridos venezolanos coleccionodos por Karl Moritz (1797-1866) y discusi6n de la identidad taxonémica y la distribucién de Pedaliodes pisonia (Hewitson, 1862) (Lepi- doptera: Nymphalidae, Satyrinae). Unpublished MS. Vitoria, A. L. & J. CAMACHO. 1999. Three new pronophiline but- terflies from the Serrania del Turimiquire, eastern Venezuela, and type designation for Corades enyo enyo Hewitson (Lepi- doptera: Nymphalidae, Satyrinae). Fragmenta Entomologica (Roma). 31:173-188. Vitoria, A. L. & T. W. Pyrcz. 1994. A new genus, Protopedaliodes and a new species, Protopedaliodes kukenani from the Pan- tepui, Venezuela (Lepidoptera, Nymphalidae, Satyrinae). Lam- billionea 94:345-352. . 1995. Notes on Pedaliodes roraimae Strand, a little known satyrid from south-eastern Venezuela (Lepidoptera: Nymphali- dae: Satyrinae). Lambillionea 95:584—586. Received for publication 30 June 1998; revised and accepted 20 Au- gust 1999. Journal of the Lepidopterists’ Society 53(3), 1999, 99-103 AN EXAMINATION OF INTRASEASONAL VARIATION IN THE INCIDENCE OF MELANISM IN PEPPERED MOTHS, BISTON BETULARIA (GEOMETRIDAE) BRUCE S. GRANT Department of Biology, College of William and Mary, P.O. Box 8795, Williamsburg, Virginia 23187-8795, USA AND CyriL A. CLARKE Emeritus Professor of Medicine, University of Liverpool, Liverpool L69 3BX, UK ABSTRACT. We examined daily catch records of peppered moths (Biston betularia) taken over a forty-year period to determine whether melanic and pale forms of this species fly at the same or different times within seasons. We also compared the emergence rates of the two forms from reared broods to determine whether differences in developmental rates might contribute to intraseasonal flight patterns. Although melanic phenotypes develop slightly faster than their typical siblings in some laboratory broods, the field records show no consistent pattern for one phe- notype being more common than the other early or late in the same summer. The work is discussed in the context of industrial melanism for which this species is the classic example. Additional key words: cognataria, f. carbonaria, industrial melanism, f. swettaria. Industrial melanism is a familiar textbook example of observable evolution brought about by natural se- lection. The term pertains to increases in the frequen- cies of genetically determined melanic versus pale phenotypes in populations living in habitats modified by regional industrial development and urbanization. The phenomenon has been well documented in many species of Lepidoptera; however, attention has focused primarily on the peppered moth, Biston betularia (L) (Geometridae). The broad aspects of the subject have been reviewed recently by Majerus (1998), Sargent et al. (1998), and Grant (1999). The corpus of experimental work to date is consis- tent with the interpretation that selective predation on the moths by birds is the primary, though not exclu- sive, force driving the changes in the frequencies of peppered moth phenotypes (Majerus 1998). Ket- tlewell (1955, 1956) provided the first quantitative ev- idence that birds eat the different color phases of pep- pered moths according to their conspicuousness on different backgrounds. His mark-release-recapture ex- periments also demonstrated that the melanic pheno- types fared better than the pale forms in soot-black- ened woodlands; whereas, the pale forms fared better than the melanics in unpolluted woodlands. Kettlewell (1973) entertained other possibilities be- side selective predation that might contribute, at least in part, to the high incidence of melanism in moth populations living in the vicinities of British industrial centers. He speculated that larvae developing in the early part of the summer feed on leaves that are less contaminated by industrial pollutants than are the older leaves that larvae feed on later in the summer. From his observations of larval developmental rates, he proposed that pale peppered moths, as fast devel- opers, avoided pollution, and/or that the melanics, as slow developers, “may be capable of getting rid of toxic substances.” He cited no reference to support his statement that “slow feeding and a capacity for excret- ing noxious materials has been demonstrated . . . out- side the Lepidoptera” (Kettlewell 1973:85). Unfortunately, Kettlewell’s developmental analysis of peppered moths was limited to one brood that was partially consumed by mice, and a second brood that provided “no corroboration of the earlier results.” He also acknowledged that he could provide no evidence from samples of wild populations that industrial melanic forms change in frequency during the flight period within single seasons. However, he did discuss intraseasonal changes in phenotype frequencies for several other moth species which show what he called “ancient” (=stable polymorphism) melanism. For ex- ample, pale Amathes glareosa increase late in the sea- son (the melanics appear early on); whereas, Cleora repandata melanics increase, relative to pale forms, as the season progresses. In analyses of seasonal catch records of other moth species polymorphic for melanic forms, Bishop et al. (1978) concluded that melanic Gonodontis bidentata emerged later than pale forms, and Sargent (1983) re- ported slight increases in melanism during the second half of seasons in Phigalia titea. As only field data were available from these studies, clear distinctions between selection on the adults and developmental differences in emergence schedules between the phenotypes were not possible. Equally problematical, S. Poitout (cited 100 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TABLE 1. June (=A) and July (=B) catch records of B. betularia phenotypes (Mel. = melanics, T+I = typicals and f. insularia) at Caldy Com- mon between 1959 through 1998. The phenotypic distributions during June and July of each year are compared by G-tests of independence. Year Mel T+I G Year Mel T+I G 59A 174 15 69A 349 25 59B 90 4 1.46 69B 216 20 0.67 60A 178 11 70A 658 68 60B 34 2 — 70B 112 10 0.18 61A 161 18 TIA 109 8 61B 240 12 4,45* 71B 79 11 1.75 62A 159 17 72A 45 3 62B 588 38 2.56 72B 202 26 1,26 63A 725 67 73A 102 14 63B 185 18 0.03 73B 181 21 0.21 64A 238 25 74A 99 14 64B 335 37 0.03 74B 178 22 0.14 65A 361 40 75A 78 16 65B 70 7 0.06 75B 155 20 1.60 66A 163 18 76A 129 29 66B 110 8 0.93 76B 242 46 0.05 67A 123 10 TIA 68 12 67B 243 21 0.02 77B 360 50 0.46 68A 209 26 78A 91 28 68B 216 26 0.01 78B 205 31 0.92* *p < 0.05 *p < 0.01 by Bishop et al. 1978) showed in laboratory strains of Spodoptera exigua that melanic genotypes had a longer total development than non-melanic genotypes; thus melanic individuals tended to emerge from pupae later than the non-melanics, but no field data were re- ported. The above studies serve to illustrate that generalities are not obvious regarding developmental differences and/or in intraseasonal variations in flight patterns be- tween melanic and pale forms of polymorphic moth species. Therefore, we have analyzed the flight patterns in B. betularia directly, and compared the emergence rates of melanic and pale phenotypes. To determine if melanic and pale phenotypes fly at different times during the summer, we examined the daily catch records of peppered moths taken at a single location over a forty-year period. To assess potential differ- ences in emergence schedules, we recorded the eclo- sion sequences of melanic and pale phenotypes from three large broods produced from controlled crosses. MATERIALS AND METHODS Field studies of intraseasonal flight patterns. Biston betularia were collected near Caldy Common, Year Mel T+I G Year Mel T+I G 79A 387 111 89A 52 114 79B 111 23 1.72 89B 13 40 0.91 80A 379 115 90A 36 68 80B 103 40 1.30 90B 15 35 0.33 81A 118 46 91A 102 337 81B 178 65 0.08 91B 139 355 2.93 82A 43 ifs) 92A 227 750 82B 45 19 0,22 92B 32 iLL 0.05 83A 219 111 93A 4l 146 83B 226 133 0.88 93B 22 65 0.37 84A 172 105 94A 39 196 84B 43 32 0.56 94B 26 87 2.01 85A 189 144 95A 33 149 85B 271 256 2.34 95B 13 66 0.11 86A 213 198 96A 5 78 86B 169 225 6.78** 96B 12 106 1,12 87A 141 201 97A 12 165 87B 4] 61 0.03 97B 5 56 — 88A 44 66 98A 14 113 88B 29 AO 0.07 98B 12 87 0.07 West Kirby, England over a period of 40 years, begin- ning in 1959. The original purpose of the study was to assess frequency changes in the melanic, pale, and in- termediate phenotypes over the course of years; these annual changes have been reported elsewhere (see Clarke et al. 1985, 1994, Grant et al. 1996, 1998). To determine if the different phenotypes fly at different times during the same season, we reexamined the daily catch records for each of the 40 years moths were trapped at this location. Each season a mercury vapor (MV) light trap was operated nightly from 1 June through 31 July. When virgin females were available, an assembling (phero- mone) trap was also used. No difference in the pro- portions of the phenotypes caught by one trapping method or the other has ever been observed (Clarke et al. 1994), but the incidental use of the assembling trap does increase the total catch size. For our analysis, therefore, we subdivided each season into early and late halves based on calendar date (June versus July) rather than by the mid-point (median) of the total numbers of the moths caught within seasons. We then tabulated the numbers of each phenotype caught dur- ing the first and second halves of each season. The null VOLUME 53, NUMBER 3 % Melanic 0 59 62 65 68 71 74 77 80 83 86 89 92 95 98 ear Fic. 1. The decline in the frequency of melanic B. betularia be- tween 1959 through 1998 at Caldy Common, West Kirby, England, plotted from June samples (solid symbols) and from July samples (open symbols). hypothesis is that there should be no difference in the percentage of melanics caught in the early and in the late samples taken during the same summer. Laboratory experiments on differential emer- gence. To determine if differential emergence (time to eclosion) of melanic versus pale phenotypes exists, we crossed known heterozygous melanic B. betularia cognataria to their pale siblings. (The expected ratio of pales to melanics among the progeny from such mat- ings is 1:1.) The stock material was produced by a melanic female crossed to a pale male, both of which were caught at the same location in Pennsylvania in 1996, and the crosses used in our experiments were made using their progeny which emerged in the spring of 1997. In all, three sets of siblings were crossed, and their broods were subdivided to avoid overcrowding. The caterpillars were housed inside plastic “garbage cans” and were provided a continuous supply of fresh leaves from a single, large Chiswell crab apple tree (Malus spp.). The caterpillars pupated “at will” in moistened beddings in the bottoms of their containers. The pu- pae were then stored in containers lined with moist- ened paper towels. All storage containers were checked daily to remove newly emerged adults until the experiment was terminated several weeks after emergences ceased entirely and no living pupae re- mained. For each brood, each moth to emerge was _ identified by phenotype and sex and the date of its emergence. RESULTS Field studies. From 1959 through 1998, 18,255 Biston betularia were collected near Caldy Common. 101 The year-to-year sample sizes varied widely, ranging between 122 to 1120, with an average annual catch of 456 + 257. The early and late halves (June vs. July) of seasons also varied widely with respect to sample sizes, with mean catches at 259 + 206 and 197 + 143, re- spectively. The differences between the early and late sample sizes, though large, are not statistically signifi- cant by paired samples t-tests (t = 1.622, df = 39, p = 0.113), nor by the non-parametric Wilcoxon’s signed- ranks test (p = 0.226). The complete set of catch records, subdivided by early (A = June) and late (B = July) catches, are sum- marized by phenotype in Table 1. In the table, the melanics (f. carbonaria) are separated from the other phenotypes (pale = f. typica or “typicals” and interme- diates = f. insularia). The intermediates have re- mained rare at Caldy Common (for complete data through 1993 see Clarke et al. 1994), therefore the combined category (T+I) is essentially “typical” (pale), or non-melanic. Sample sizes permitting, the numbers of melanics and non-melanics collected during the first half and second half of each season were compared using 2 x 2 contingency G-tests of independence. The G statistics are listed in Table 1. Of the 38 comparisons made, only three showed significant differences in the phenotypic proportions between the early and late halves of the same season; in two instances the melanics increased significantly (p < 0.05) in the second half of the season, and in one instance the melanics declined very signifi- cantly (p < 0.01) during the second half of the season. In 35 of the 38 comparisons, no significant differences in the proportions of melanics between the early and late subsamples within seasons were observed. To test for the possibility that small but consistent differences might exist within seasons (differences too slight to be detected by G-tests), we analyzed the en- tire 40-year record by Wilcoxon’s signed-ranks test. The null hypothesis is that the differences in the per- centages of melanics collected during first and second halves of seasons are random. The data show that the percentage of melanics increased during the second half of summers 23 times, and decreased 17 times (Fig. 1), but the differences are not significant by Wilcoxons signed-ranks test (p = 0.55). Between 1977 and 1997 the annual incidence of melanism declined rapidly at Caldy Common (Fig. 1), and for several years during this same period, there ap- peared to be a short run of seasons in which the per- centage of melanics decreased in the second half of summers compared to the first half. However, the pat- tern is not consistent throughout the period of rapid annual decline in melanism, and the differences in 102 TABLE 2. Early and late emergences from reared broods of B. betularia cognataria from crosses expected to produce 1:1 ratios of melanic and pale (typical) phenotypes. The phenotypic distributions between the early and late groups within each brood are compared by G-tests of independence. Brood Melanics Typicals iG A Early 53 46 Late 49 36 0.31 B Early 30 23 Late 16 29 4.37* C Early 27 Di Late 23 30 0.47 *™*D < 0.05 melanic frequencies between first and second halves of seasons are not significant by Wilcoxons signed- ranks test (p = 0.82). Differential emergence. We recorded the emer- gence date, sex and phenotypes of all adults produced by the three crosses of heterozygous melanics mated to their pale, homozygous siblings. Eclosion patterns within broods generally begin with one or a few adults emerging from puparia on the initial day, then the number of daily emergences increases quickly and peaks a few days later, after which the daily emer- gences tail off until finally many days may separate the stragglers from the main group and from each other. The null hypothesis of our experiment is that the dif- ference in emergence sequence within broods is not related to the color phenotypes; therefore, the propor- tions of melanics and typicals should be the same in the first half of a brood to emerge as in the second half. We divided each brood into early and late groups using the median. All of the moths emerging up to and including the day the first half of the total emergences for the brood was reached were assigned to the early group, and all of the moths emerging after that day were assigned to the late group. Table 2 lists the early and late emergences by phenotype (melanic and typi- cal) for each of the three broods. The numbers of melanics and typicals emerging during the early and late periods are compared within broods by 2 x 2 con- tingency G-tests of independence. The G statistics are also listed in the table. In brood B, melanics are signif- icantly more common in the early than in the late group (p < 0.05), but no significant emergence differ- ences are apparent in the other two broods. DISCUSSION The data from Caldy Common provide little support for the idea that the different phenotypes of Biston betularia fly at different times of the season. That three of the 38 contingency tests indicated significant JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY intraseasonal differences between the phenotypic proportions is consistent with type-I error rates ex- pected when large numbers of statistical comparisons are made (Sokal & Rohlf 1981). Furthermore, only two of the three observed significant deviations from random expectations were skewed in the same direc- tion which further supports that these exceptional years can be attributed to chance. Of these three ex- ceptional years (1961, 1978, 1986), only 1986 shows a very pronounced deviation in phenotypic proportions between the early and late halves of a season, and in this instance the significant decline in the proportion of melanics occurred during the brief run of intra- seasonal declines in melanism that occurred during those years of the steepest declines in the annual inci- dence of melanism. Here, at least, it seems reasonable to suggest that selection at the adult stage rather than developmental differences in emergence schedules might account for the drop in melanism within that season. Differences in emergence schedules between melanic and pale peppered moths are slight, at best. Of the three broods we examined, only one showed significantly accelerated emergence of the melanics versus their typical siblings. Different broods, no doubt, emerge throughout the normal season of sev- eral months, and if developmental differences, how- ever slight, between melanics and pale forms are con- sistent, then, in the absence of offsetting selection at the adult stage, we should expect to observe a consis- tent bias favoring melanics earlier in the summer. Clearly this did not happen over the 40 years of obser- vations at Caldy Common. Our developmental studies in the laboratory em- ployed the North American subspecies, B. betularia cognataria, and our field studies centered on the British subspecies of peppered moths, B. betularia be- tularia. Direct comparisons between the two must be qualified. American “pale” or typical forms are gener- ally much darker than British typicals; however, the melanics (called f. carbonaria in Britain and f. swet- taria in America) are phenotypically indistinguishable, and are caused by alleles at the same locus (Grant & Clarke in prep.). The melanics, both in America and Britain, have also shown parallel increases (Owen 1962) and decreases (Grant et al. 1995, 1996) in fre- quencies associated with environmental modifications related to industrial development and urbanization. The reductions in melanism on both continents are now widespread (Grant et al. 1998). The phylogenetic relationship (Rindge 1975, Clarke et al. 1993), and the genetics of melanism in these sub- species (West 1977, Grant & Clarke in prep.), and the VOLUME 53, NUMBER 3 ecological events coincidental to the parallel evolu- tionary changes in phenotype frequencies (Grant et al. 1996, 1998) combine to suggest that we are observing the same basic phenomenon in peppered moth popu- lations on both sides of the Atlantic. While we must re- main cautious in extrapolating what we learn from the study of one subspecies to draw conclusions about the other, we also recognize that what we learn from one subspecies might be instructive in our study of the other. In this instance, there again appears to be agree- ment: No consistent intraseasonal flight pattern differ- ences between pale and melanic British peppered moths are apparent, nor are the differences in emer- gence schedules of pale and melanic American pep- pered moths consistent among broods. Our study does not address one of Kettlewell’s (1973) suggestions that differences in developmental rates might exist among the larvae of different geno- types that produce pale and melanic adult phenotypes. No clear relationship between emergence rates be- tween melanic and pale peppered moths and the time the forms spend as larvae has ever been established; therefore, our study on emergence sequences does not bear directly on the question of the seasonal condi- tions early and late larvae might experience. However, there is evidence that B. betularia cognataria is bivol- tine (Owen 1962, Manley 1981); therefore, larvae of those genotypes producing melanic and pale adults in the first versus the second generations of summers must endure late and early summer conditions, re- spectively, as they feed and develop. Yet, there is no evidence that the proportions of melanic and pale adults change consistently between the first and sec- ond broods of summers (Owen 1962, Manley 1981). These observations do not support Kettlewell’s specu- lations that selection at the larval stage influences melanism at the adult stage. We do not consider this a “negative results” paper. There has been much speculation about the relevant ecological factors and the putative selective agents re- sponsible for industrial melanism, and now its decline (recently critiqued by Majerus 1998, Sargent et al. 1998, and Grant 1999). The occasional but recurring query that intraseasonal variation in the frequencies of the forms may offer some insights can, in our view, be laid to rest. Part of the process in identifying what is important from what is not involves the process of elimination. We hope we are making progress. 103 ACKNOWLEDGMENTS We thank Jean Butler, Winifred Cross, Sally Thompson, and An- gela Urion for assistance with population sampling at Caldy Com- mon, and the Nuffield Foundation for funding the fieldwork. We also thank Annie Harvilicz for assistance in feeding caterpillars. Deane Bowers, Laurence Cook, Theodore Sargent, Ward Watt, and Lawrence Wiseman helped us improve this report by their com- ments on an earlier draft. We dedicate this paper to the memory of Lady Féo Clarke. LITERATURE CITED Bisuop, J. A., L. M. Cook & J. MUGGLETON. 1978. The response of two species of moths to industrialization in Northwest England: I. Polymorphism for melanism. Philos. Trans. R. Soc. Lond. B. Biol. Sci. 281:489-442. CLARKE, C. A., G. S. MAnI & G. WYNNE. 1985. Evolution in re- verse: clean air and the peppered moth. Biol. J. Linn. Soc. 26:189-199. CLARKE, C. A., F. M. M. CLARKE & B. GRANT. 1993. Biston betu- laria (Geometridae), the peppered moth, in Wirral, England: an experiment in assembling. J. Lepid. Soc. 47:17-21. CLARKE, C. A., B. GRANT, F. M. M. CLARKE & T. ASAMI. 1994. A long term assessment of Biston betularia (L.) in one UK locality (Caldy Common near West Kirby, Wirral), 1959-1993, and glimpses elsewhere. Linnean 10:18-26. Grant, B.S. 1999. Fine tuning the peppered moth paradigm. Evo- lution 53:980—984. Grant, B., D. F. OWEN & C. A. CLARKE. 1995. Decline of melanic moths. Nature 373:565. . 1996. Parallel rise and fall of melanic peppered moths in America and Britain. J. Hered. 87:351-357. GRANT, B. S., A. D. Cook, C. A. CLARKE & D. F. Owen. 1998. Geographic and temporal variation in the incidence of melanism in peppered moth populations in America and Britain. J. Hered. 89:465-471. KETTLEWELL, B. 1973. The evolution of melanism. Clarendon Press, Oxford. KETTLEWELL, H. B. D. 1955. Selection experiments on industrial melanism in the Lepidoptera. Heredity 9:323-342. . 1956. Further selection experiments on _ industrial melanism in the Lepidoptera. Heredity 10:287-301. Majerus, M. E. N. 1998. Melanism: evolution in action. Oxford Univ. Press, Oxford. MANLEY, T. R. 1981. Frequencies of the melanic morph of Biston cognataria (Geometridae) in a low-pollution area of Pennsylva- nia from 1971 to 1978. J. Lepid. Soc. 35:257-265. Owen, D. F. 1962. The evolution of melanism in six species of North American geometrid moths. Ann. Entomol. Soc. Amer. 55:695-703. RINDGE, F. H. 1975. A revision of the New World Bistonini (Lepi- doptera: Geometridae). Bull. Am. Mus. Nat. Hist. 156:69-155. SARGENT, T. D. 1983. Melanism in Philgalia titea (Cramer) (Lepi- doptera: Geometridae): a fourteen-year record from Central Massachusetts. J. N. Y. Entomol. Soc. 91:75-82. SARGENT, T. D., C. D. MILLAR & D. M. LamBERrT. 1998. The clas- sical explanation of industrial melanism. Evol. Biol. 30:299-322. SOKAL, R. K. & F. J. RonF. 1981. Biometry, 2nd. ed., Freeman, San Francisco. West, D. A. 1977. Melanism in Biston (Lepidoptera: Geometri- dae) in the rural Central Appalachians. Heredity 39:75-81. Received for publication 11 November 1998; revised and accepted 16 December 1999. Journal of the Lepidopterists’ Society 53(3), 1999, 104-107 URIC ACID DEPOSITION IN LARVAL INTEGUMENT OF BLACK SWALLOWTAILS AND SPECULATION ON ITS POSSIBLE FUNCTIONS SUSANNE TIMMERMANN AND May R. BERENBAUM Department of Entomology, 320 Morrill Hall, University of Illinois, 505 S$. Goodwin, Urbana, Illinois 61801-3795, USA ABSTRACT. From the first through third instar, larvae of the black swallowtail, Papilio polyxenes Fabr., display a distinctive color pattern characterized by an irregular circle of white pigment on the dorsum. This white spot is surrounded by brown pigmentation, creating the im- pression of a bird dropping. It has long been assumed that this color pattern evolved as a defense mechanism against avian predators. We report here that the source of the white color is accumulated uric acid. Although uric acid has traditionally been viewed as an excretory product, it also can act in biological systems as a powerful antioxidant. Thus, the possibility exists that the white spot serves a protective function not only against predators in a high fidelity mimicry system, but also against oxidative stress generated by the phototoxic allelochemicals that characterize most hostplants of Papilio species. Additional key words: Papilio polyxenes, bird dropping, furanocoumarin. Early instars of a range of Lepidoptera species are brownish in color with a white saddle traversing the middle of the abdomen; this color pattern, from the human perspective, bears a remarkable resemblance to a bird dropping and thus has been characterized as an example of homotypism, or protective resemblance to an object considered inedible by a predator (Ed- munds 1974). Although this pattern has been reported in at least one nymphalid genus (and typifies the larvae of the viceroy Basilarchia archippus archippus (Cramer), the red-spotted purple Basilarchia arthemis astyanax (Fabr.), and the white admiral Basilarchia arthemis arthemis (Drury)), it is practically universal among early stages of species in the genus Papilio (Lepidoptera: Papilionidae) (Munroe 1961). Among the many substances contributing to larval lepidopteran pigmentation patterns is uric acid. Al- though uric acid is the major nitrogenous waste prod- uct of terrestrial insects (Cochran 1985), in many species it is retained and deposited in body tissue. This mechanism of dealing with excess nitrogen is known as storage excretion (Wigglesworth 1942, 1965, 1987) and provides a source of pigmentation, particularly in larval stages. Mauchamp and Lafont (1975) demon- strated that most of the uric acid in young Pieris bras- sicae (L.) caterpillars lies in the integument and is ac- cumulated in the fat body before pupation, and Buckner and Newman (1990) determined that uric acid deposition in the integument is principally re- sponsible for the appearance of white stripes that con- trast with the green abdominal base color. Uric acid, presumably generated by larval metabolism, also can be found as a pigment in the yellow scales of Papilio xuthus L. (Tojo & Yushima 1972) and in the wings of male Pieris brassicae (Lafont & Pennetier 1975). In that birds are uricotelic, producing uric acid as the principal form of waste nitrogen, the white color of bird droppings is due to the presence of uric acid. We examined the integument of larvae of Papilio polyx- enes Fabr., the black swallowtail butterfly, a species that displays the typical Papilio early stage “bird drop- ping” morphology, in order to determine if in fact the white saddle results from the accumulation of uric acid, the substance that provides the model for the mimetic resemblance. We also examined the integu- ment of ultimate (fifth) instars, to determine if uric acid contributes to color patterns in mature larvae as well, and in adult male butterflies, to ascertain whether sequestered uric acid is retained through metamorphosis. MATERIALS AND METHODS The eastern black swallowtail butterfly, Papilio polyxenes asterius Stoll., is found throughout eastern North America, ranging from southern Canada to Florida; it also occurs west along the eastern Rockies into northern Mexico (Opler and Krizek 1984). The larval stages feed almost entirely on herbaceous repre- sentatives of the families Apiaceae and, to a more lim- ited extent, Rutaceae, and are found in a variety of open habitats. Larval development requires from two to three weeks, depending on temperature and host- plant (Blau 1981). Larvae in the first three stadia are primarily black, with a characteristic white saddle across the dorsal midsection; fourth and fifth instars are greenish-white to green, with black bands running horizontally across each segment, interrupted by a se- ries of yellow to orange spots (Fig. 1). In central Illi- nois, there are two to three generations each year. Principal natural enemies of the black swallowtail in- VOLUME 53, NUMBER 3 cm 2 -€ Fic. 1. Third (left) and fifth (right) instar Papilio polyxenes caterpillars. clude spiders, wasps, predaceous bugs in the families Nabidae, Reduviidae, Coreidae, and Pentatomidae (Blau 1981, Feeny et al. 1985), ants, and possibly birds. Caterpillars defend themselves with an eversible osmeterial gland, the constituents of which change de- velopmentally from primarily mono- and sesqui-ter- penes in early instars to aliphatic acids and their esters in fourth and fifth instars (Berenbaum et al. 1992). Gravid female P. polyxenes were collected in Cham- paign County, Illinois, and allowed to oviposit on fo- liage of parsley, Petroselinum crispum (Mill.); eggs col- lected in this manner were used to found a colony from which larvae were taken for experimental use. Caterpillars were reared on potted parsley or on parsnip, Pastinaca sativa L., plants in a greenhouse at 27°C (day): 21°C (night) under a 16L:8D photoperiod. As caterpillars reached either third or fifth instar, they were collected for chemical analysis. Male adults col- lected earlier from a laboratory colony initiated with wild-caught butterflies from Champaign County, Illi- nois, were stored at —80°C prior to chemical analysis. 105 Prior to all larval tissue collections, we removed the gut and Malpighian tubules. Epidermis of the ab- domen of the third instar was divided into portions corresponding to the white saddle area and the re- maining brown portion; each portion was analyzed separately. The final instar was divided according to apparent integument coloration. Black bands, green ground color, and yellow spots were cut out with iri- dectomy scissors and analyzed separately. The wings of the males were divided into black parts and yellow spots by cutting out the spots and each tissue type was analyzed separately. All tissues were collected directly into dry ice-chilled microcentrifuge tubes (1.8 ml) and kept in a freezer at 80°C prior to the uric acid assay. We homogenized tissue samples with a tissue tearor (Biospec Products, Inc., Bartlesville, Oklahoma) and used a chloroform rinse to free the solution of lipophilic compounds. The integument was eluted in lithium carbonate (1%) to dissolve uric acid and after centrifu- gation (12,200 g) an aliquot of the supernatant was used for uric acid determination (Van Handel 1975). Each aliquot was brought up to a volume of 2 ml with distilled water. To each sample we added 1 ml of reagent (copper sulfate 0.05%, glycine 1.6%, sodium carbonate 4%) and 0.05 ml neocuproine reagent (Sigma, St. Louis, Missouri). Optical density was determined at 450 nm wavelength (Perkin Elmer Lambda 3B spectrophotometer). Because ascorbic acid can inter- fere in the uric acid assay (25 ug shows the same opti- cal density as 0.9 ug uric acid), we corrected uric acid readings according to the ascorbic acid content of the sample (Omaye et al. 1979). To quantify ascorbic acid, 0.5 ml of supernatant was added to 0.5 ml of ice-cold 10% trichloroacetic acid, mixed thoroughly, and cen- trifuged for 5 minutes (12,200 g); 0.5 ml of super- natant was mixed with 0.1 ml of DTC (thiourea, CuSO,,.5H,0, 2,4-dinitrophenylhydrazine in 9N H,SO.,: Omaye et al. 1979) and incubated for 3 hours at 37°C, to form the 2, 4-dinitrophenylhydrazone. After incu- bation, the test tube was removed from the water bath and placed into ice water; 0.75 ml of ice-cold H,SO, (65%) was added and the solution mixed well. Solutions stood at room temperature for 30 more min- utes, after which time we measured the absorbance at 520 nm. RESULTS AND DISCUSSION Integument from ten third instars and nine fifth in- stars and wing scales from three adult male butterflies were analyzed for their uric acid content and distribu- tion. All parts of the integument of third instar P. polyxenes contain measurable amounts of uric acid (Table 1). The white saddle, however, contains over 106 TABLE 1. Distribution of uric acid in the variously colored in- teguments of third and fifth instar and male wings of Papilio polyx- enes. Values are given as the mean (standard deviation). Life stage N Integument color Uric acid (g/mg) Third 10 Brown 57.2 (16.1) Third 10 White 116.0 (57.6) Fifth i) Black 24.9 (6.9) Fifth f°) Green 49.1 (20.3) Fifth 9 Yellow 45.5 (9.2)¢ Male 3 Black 2.2 (0.5) Male 3 Yellow 15.0 (2.5)4 * Significantly different from brown integument, paired ¢ = 3.72, p = 0.005 » Significantly different from black integument, paired ¢ = 3.2, p = 0.013 © Significantly different from black integument, paired t¢ = 5.7, p = 0.0001 4 Significantly different from black scales, paired t = 8.6, p = 0.01 twice the amount of uric acid in brown-colored in- tegument (paired t test, t = 3.72, p = 0.005). Integu- ment that appears light in color in fifth instars also contains significantly greater amounts of uric acid than does adjacent integument that is black in color. Yellow spots contain almost twice as much uric acid (t = 5.65, p = 0.0001) as black stripes; the green ground color also contains almost twice the uric acid content of the black stripes (t = 3.19, p = 0.013). The accumulation of uric acid in the yellow wing scales in male butterflies is sevenfold higher than the amount of uric acid in the black wing scales (¢ = 8.6, p = 0.01). Over all life stages examined, uric acid content is highest in the white sad- dle of the third instar. To a large extent, caterpillars rely on coloration to avoid visually orienting natural enemies. Although some species rely on crypsis (matching their back- ground) and concealment, many can remain in plain sight by virtue of aposematism and associated un- palatability or by mimetic resemblance to inedible ob- jects or substances. Birds have been identified as im- portant predators of caterpillars in general (Morris 1972, Holmes et al. 1979, Atlegrim 1989, Marquis & Whelan 1994) and the body coloration of many species is thought to reflect selection pressures exerted by avian predation. Resemblance to a bird dropping may be common in that it is likely such a color pattern would be unappetizing to a wide variety of birds. In P. polyxenes, the appearance of the white saddle of the “bird dropping” pattern is largely due to the accumu- lation of uric acid, the same substance that creates the white, shiny appearance of authentic bird droppings. The fidelity of the visual mimetic resemblance to a bird dropping is no doubt enhanced by use of a sub- stance identical to that found in the “model.” Resemblance to a bird dropping is at least in part re- sponsible for the ability of early stage P. polyxenes to forage in full view of predators. Such foraging is ad- JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY vantageous in that caterpillars can process food more quickly and efficiently at higher temperatures associ- ated with daylight hours (Ali et al. 1990). Such behay- ior, however, is not without attendant risks. Exposure to sunlight (particularly ultraviolet wavelengths) can cause oxidative stress; this stress may be exacerbated by the presence of photosensitizing allelochemicals in foliage. The apiaceous hosts of P. polyxenes (indeed, of most Papilio species—Berenbaum 1983) characteristi- cally possess furanocoumarins, photosensitizers that can cause oxidative damage to DNA (Berenbaum 1991). Although all stages of P. polyxenes are capable of rapid and efficient metabolism of these compounds (Harrison et al. submitted), early instars in general have higher relative consumption rates than late in- stars (Slansky & Scriber 1985) and thus may encounter greater quantities of furanocoumarins relative to their body weight. While melanin in the dark parts of P. polyxenes integument may function as a neutral den- sity filter, eliminating photoactivating wavelengths, the white saddle could potentially leave caterpillars vul- nerable to ultraviolet light exposure. Uric acid may be selectively retained by P. polyx- enes, and Papilio species in general, not only because of its color but because of its powerful antioxidant and radical-scavenging properties (Becker 1993). The function of uric acid as an antioxidant in insects has al- ready been established. Souza et al. (1997) observed greatly increased urate concentrations in the he- molymph of Rhodnius prolixus following a blood meal and suggested an antioxidant protective function of urate against prooxidant activity generated during the hydrolysis of hemoglobin. In Drosophila melanogaster, the antioxidant properties of urate have been demon- strated by the sensitivity of urate-null mutants to ex- perimentally induced oxidative stress (Hilliker et al. 1992). The role of uric acid in epidermis as a protec- tive pigment has also been suggested. In Anopheles mosquito larvae, the white dorsal pigmentation may represent protective coloration against solar radiation. Anopheline larvae are confined to shoal biotopes, liv- ing at the air/water interface and lying horizontally im- mediately below the water surface. Under these cir- cumstances, the antioxidant properties of uric acid could effectively reduce possible damage by UV radia- tion (Benedict et al. 1996). Other antioxidants (partic- ularly antioxidant enzymes) are known to occur in the integument of swallowtails (Lee and Berenbaum 1992), and foliage of swallowtail hostplants has been demonstrated to produce singlet oxygen at the leaf surface (Berenbaum and Larson 1988), which may present particular risks to integumentary tissues. Thus, the high concentrations of uric acid in the VOLUME 53, NUMBER 3 white saddle of the “bird dropping” morph of young black swallowtail caterpillars may serve multiple pur- poses—contributing to a compelling mimetic/protec- tive resemblance to an inedible object in the environ- ment while at the same time scavenging free radicals generated by ultraviolet light exposure and ingestion of photosensitizers. Such physiological economy may be enhanced further by the fact that accumulation of uric acid, a waste product generated as a consequence of processing food, does not divert nitrogen away from other physiological needs, as would synthesis of such nitrogenous pigments as pteridines or ommochromes. ACKNOWLEDGMENTS This project was funded by the Swiss National Science Founda- tion to S. E. Timmermann and National Science Foundation grant DEB96-28977 to M. R. Berenbaum and A. R. Zangerl. We thank Dr. Arthur Zangerl for comments on the manuscript and an anony- mous reviewer for helpful references. LITERATURE CITED Aut, A., R. J. LUTTEREL & J. C. SCHNEIDER. 1990. Effects of tem- perature and larval diet on development of the fall armyworm. Ann. Entomol. Soc. Am. 83:725-733. ATLEGRIM, O. 1989. Exclusion of birds from bilberry stands: impact on insect larval density and damage to the bilberry. Oecologia 79:136-139. BECKER, B. F. 1993. Towards the physiological function of uric acid. Free Rad. Biol. Med. 14:615-631. BENEDICT, M. Q., A. COHEN, A. J. CORNEL & D. L. BRUMMETT. 1996. Uric acid in Anopheles mosquitoes (Diptera: Culicidae): effects of collarless, stripe, and white mutations. Ann. Entomol. Soe. Am. 89:261—265. BERENBAUM, M. R. 1983. Coumarins and caterpillars: a case for co- evolution. Evol. 37:163-179. . 1991. Coumarins, pp. 221-249. In G. Rosenthal and M. Berenbaum (eds.), Herbivores Their interactions with sec- ondary plant metabolites. Vol.1. 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Gilbert (eds.), Comprehensive insect physiology, biochemistry and pharmacol- ogy. Pergamon Press, New York.. SouZA, A. V. G., J. H. PETRETSKI, M. DEmasI, E. J. H. BECHARA & P. L. OLIveIRA. 1997. Urate protects a blood-sucking insect against hemin-induced oxidative stress. Free Rad. Biol. Med. 22:209-214. Tojo, S. & T. YusHIMA. 1972. Uric acid and its metabolites in but- terfly wings. J. Insect Physiol. 18:403-422. VAN HANDEL, E. 1975. Direct determination of uric acid in fecal material. Biochem. Med. 12:92-93. WIGGLESWORTH, V. B. 1942. The storage of protein, fat, glycogen, and uric acid in the fat body and other tissues of mosquito lar- vae. J. Exp. Biol. 19:56—-77. . 1965. The principles of insect physiology. Methuen & Co. LTD., London. . 1987. Histochemical studies of uric acid in some insects. 2. Uric acid and polyphenols in the fat body. Tissue & Cell 19 (1):93-100. Received for publication 1 December 1998; revised and accepted 19 November 1999. Journal of the Lepidopterists’ Society 53(3), 1999, 108-113 FEEDING PREFERENCE OF HELICONIUS ERATO (LEP.: NYMPHALIDAE) IN RELATION TO LEAF AGE AND CONSEQUENCES FOR LARVAL PERFORMANCE DANIELA RODRIGUES AND GILSON RUDINEI PIRES MOREIRA! Departamento de Zoologia, Instituto de Biociéncias, UFRGS, Av. Paulo Gama, 40, 90046-900, Porto Alegre, RS, BRAZIL ABSTRACT. In South Brazil, female Heliconius erato phyllis (Fabricius) lay isolated eggs on branch tips of Passiflora suberosa Linnaeus, leading to early larvae first contacting young host plant tissues. These feed on young tissues, but the adaptive meaning, if any, for such an asso- ciation is still unknown. In this paper, we determine what leaf ages are usually consumed by each instar, the existence of feeding preference in relation to leaf age, and the performance of H. erato when fed either on young or old leaves of P. suberosa under laboratory conditions. Freshly- hatched larvae were transferred to the terminal portion of isolated, intact shoots, and observed daily for order of leaf choice until pupation. Feeding preference in relation to leaf age was evaluated for all instars through choice tests using leaf disks from young and mature leaves. Ad- ditionally to evaluate induction of feeding preference, larvae were reared through fourth instar on either young or mature leaves and then tested using the same choice procedure. Growth performance was assessed by progressively transfering larvae, according to instar number, from intact shoots to sections where apical portion was cut off. Larvae fed starting at the terminal bud (young tissues) and moving progressively to lower leaves of increasing age (mature leaves). Larvae of all instars consumed more young leaf disks, and preference for old leaves could not be in- duced. Larvae reared initially on young tissues had greater survivorship and grew faster than those on mature, and this may explain the oviposi- tion site selection and larval feeding preference of this butterfly. Additional key words: host-plant selection, heliconian butterflies, passion vines, leaf age effects. Heliconius erato phyllis (Fabricius) (Nymphalidae) is common in the forests of southern Brazil (Brown & Mielke 1972). It uses a number of passion vine species as larval hosts, a few being preferred in a given locality (Benson et al. 1976, Brown 1979, 1981, Ramos & Frei- tas 1999). Isolated eggs are laid primarily on terminal buds of Passiflora shoots, young tendrils and young leaves (Benson 1978, Perico 1995, Mugrabi-Oliveira & Moreira 1996a). Larvae feed initially on young leaves near the apical portion of shoots (Alexander 1961, Benson 1978). Ovipositing females reject shoots de- prived of or with a damaged apical portion, and prefer to lay on large shoots compared to small (Mugrabi- Oliveira & Moreira 1996a). They also assess egg- and larval-load, and reduce daily oviposition rates under low host-plant availability (Mugrabi-Oliveira & Mor- eira 1996b). There are at least three hypotheses to explain Heli- conius oviposition and early larval feeding on young Passiflora tissue, which are not necessarily mutually exclusive. First, Benson et al. (1976) suggested that Heliconius oviposition on passion vine tendrils would help to prevent egg-predation, because eggs on ten- drils are less likely to be attacked by ants. In addition, Benson (1978) hypothesized that the feeding pattern of heliconian larvae that use young tender growth, such as H. erato, was in part shaped by interspecific competition with species specialized on older leaves. Also, it is known that host-plant selection by phy- tophagous insects is linked to plant suitability as larval food (Jones 1991, Bernays & Chapman 1994). Thus, it is expected that H. erato phyllis oviposition on the api- ‘From whom reprints should be requested. cal portion of host-plants is also related to higher suit- ability of the corresponding tissues as larval food. In this paper, we address the latter hypothesis from a behavioral ecology perspective. Feeding of H. erato phyllis larvae and its consequences are evaluated re- garding variation in leaf age on shoots of Passiflora (Plectostemma) suberosa. This widely spread passion vine is a primary host of H. erato phyllis, and one of the most abundant passion vine species in southern Brazil (Menna-Barreto & Aratijo 1985, Perico & Araujo 1991, Perico 1995). Information on the phenol- ogy of P. suberosa and corresponding effects on H. er- ato phyllis oviposition site selection are provided by Mugrabi-Oliveira & Moreira (1996a). Specific goals of the current study are 1) to characterize the natural feeding pattern of H. erato phyllis larval instars in rela- tion to P. suberosa leaf age, 2) to evaluate the feeding preference of each instar regarding leaf age, and 3) to determine the consequences of feeding on leaves of variable age on survivorship, growth rate and adult size. MATERIALS AND METHODS Insects and plants. Larvae used in the experi- ments came from eggs obtained from an H. erato phyllis outdoor insectary maintained at the Zoology Department of Federal University of Rio Grande do Sul, Porto Alegre, RS (see Mugrabi-Oliveira & Mor- eira 1996a). P. suberosa plants were obtained from cut- tings that were transplanted into plastic pots. The plants originated from a population at Aguas Belas Ex- perimental Station, Viamao County. The plastic pots were provided with 50 cm high wooden frame support for shoot growth, covered with a fine mesh cloth and maintained in an outdoor screened cage. Prior to the VOLUME 53, NUMBER 3 tests, plants were standardized for size by removing all branches except the main shoot in each pot. Experi- ments were conducted in a large laboratory chamber, adopting the prevailing abiotic conditions for the re- gion during the summer (photoperiod 14L:10D, tem- perature 25 + 1°C, humidity 75 + 5%). Feeding pattern. To determine larval feeding pat- tern in relation to leaf age, twenty freshly hatched H. erato phyllis larvae were individually placed on the terminal bud of healthy growing shoots bearing 12 open leaves. Additional studies that have been carried out in our laboratory (S. S. Borges & G. R. P. Moreira unpubl. data) showed that in all cases when young lar- vae are put on any mature leaf, they move to the tip of P. suberosa shoots, always starting feeding on the ter- minal bud. Larvae were inspected for molting daily until pupation. At each inspection, we noted the age of leaf being consumed. To make sure that molts were not overlooked, larvae were gently marked with small colored dots of enamel paint (Testors) on the dorsal part of the penultimate abdominal segment. Leaf age was determined by leaf position in relation to the ter- minal bud (made up of apical meristem, leaf primordia and unopen leaves) (Fig. 1A). Choice experiments. Feeding preference in rela- tion to leaf age was evaluated through leaf disk choice tests. Trials were conducted in plastic pots, following the methodology described in Hanson (1983). The same twenty larvae were used over all instars; corre- sponding rearing procedure followed that descrived above. Unless noted, two leaf age categories were adopted: young (consisting of the first and second open leaves on actively growing branch) and mature (sixth and seventh open leaves). Disks were cut with a cork borer (diameter of 6.75 mm; 35.78 mm? in area), and a fixed number offered (3, 3, 6, 12 or 18 disks of each age per larvae tested from first to fifth instar, re- spectively). Young leaf disks alternated with mature ones in the pot. To keep leaf disks from drying out, pots were covered with a plastic film. Leaves were col- lected from different plants every time. Total leaf area offered a given instar was adjusted to double the aver- age leaf area consumed by a given instar in 5 hours (C. A. Barcellos & G. R. P. Moreira unpubl. data). All tri- als lasted for 5 hours, after which larvae were returned to their original rearing plants. To evaluate possible induction of feeding prefer- ence in relation to leaf age, additional 40 larvae were reared up to third instar on young leaves. After molt- ing to fourth instar, half of them were transferred to cuttings containing only mature leaves; the remaining 20 larvae were kept on cuttings bearing young leaves. They were all then individually tested in the fifth instar 109 Fic. 1. Schematic representation of Passiflora suberosa shoots used to determine influence of leaf age on Heliconius erato larval performance. A, intact, with terminal bud and ten open leaves; B, lower section, with five mature leaves (terminal bud and five apical open leaves were cut out). TB, terminal bud; L, open leaf. Numbers indicate position in relation to shoot apex. Tendrils and stipules associated with open leaves were not drawn. using the leaf-disk choice test described above (18 disks of each age per larvae). Feeding was measured as area of disk eaten, recorded by placing the disks against graph paper at the end of each feeding trial and counting the number of square millimeters corresponding to the missing leaf area. If only young leaves were damaged, or over twice the area of young as mature leaves was con- sumed, larvae were scored as having discrimated in favour of young leaves. Conversely, if only mature leaves were consumed, or over twice the area of ma- ture as young leaves was missing, then larvae were scored as having chosen mature leaves. Feeding trials were recorded as neutral when neither leaf-age cate- gory had more than twice the damage of the other (see Thomas 1987). To test the hypothesis that consump- tion of young leaves was greater than mature, scores were tested using one-tailed sign tests, following the procedure described in Conover (1980). Larval performance and adult size. To deter- mine leaf age effects on performance, larvae were in- 110 TABLE 1. Experimental scheme adopted to test influence of Pas- siflora suberosa leaf age on Heliconius erato larval performance. Treaments correspond to numbers of instar staying on intact shoots. Shoot type offered as food Treatment Intact Lower section 0 all instars 1 instar I instars II-V y) instars I and II instars IIJ-V 3 instars I-III instars IV and V 4 instars J-IV instar V 5 all instars dividually reared on potted plants receiving one of the following shoot types: 1) intact, with ten leaves and terminal bud; 2) lower section, with five mature leaves (terminal bud and five apical open leaves were cut out) 60 4 Peete

40 L 6 c 0 a (ee ; c 40 LZ 40 L 8 0 —— ee 40 Eg) i fas 40 L10 | eit == | 40 111 V 1 2 3 4 5 6 Le RS FED 10 VW age of larvae (days) 12°13 «#14 «215 Fic. 2. Sequential feeding pattem of Heliconius erato larvae (n = 20) in relation to Passiflora suberosa leat age and instar number. TB, ter- minal bud; L, open leaf. Arabic numbers designate position of leaf in relation to shoot apex. Roman numbers represent larval instars. Closed circles and associated bars denote the median and corresponding interval of leaf age eaten by a given instar. VOLUME 53, NUMBER 3 TABLE 2. Selection by different instars of Heliconius erato larvae (n = 20) for young vs. old leaves of Passiflora suberosa. Data show number of larvae falling into each preference category. See text for description of discrimination categories. Preferred leaf category Instar Younger Neutral Older ip I 20 — — < 0.001 I 19 1 — < 0.001 Ill 18 1 1 < 0.001 IV 18 ] 1 < 0.001 Vv 9 9 2 < 0.04 * probabilities calculated from sign tests. were log transformed for the analysis of variance, fol- lowing the criteria described in Sokal and Rohlf (1981). Differences among treatments were tested us- ing Fisher's Protected Least Significant Difference multiple comparison tests, when analyses of variance were significant. To determine specific effects of leaf age on adult size, 40 additional larvae were tested. These were reared on young leaves up to third instar, and after molt- ing for the fourth instar, half of them were kept feeding on young leaves while the remaining fed on mature leaves until pupation. After emergence, adult forewing length was measured, and data were compared through unpaired, two-tailed t-tests. There was no indication that size of males and females differed in these exper- iments, and thus data were grouped. Unless noted measurements are given as mean + standard error. RESULTS In general, the larvae first ate the terminal bud, fol- lowed by the first open leaf and progressively de- voured older leaves, as they grew (Fig. 2). They con- sumed all P. swberosa shoot parts, including tendrils, stipules and stem. In general, the terminal bud pro- vided enough food to complete the first instar. During this instar some larvae ate part of the first open leaf as well. Leaf damage extended to the third, fourth and fifth open leaves during instars two, three and four, re- spectively (Fig. 2). There was considerable variation in fifth instar consumption rates. This variation in leaf consumption was expected, because leaves vary in size. About half the larvae required seven open leaves to complete larval development; however, some con- sumed up to ten leaves before pupation. Larvae reared on intact P. suberosa shoots discrimi- nated between the two leaf categories tested (Table 2). All instars preferred young over mature leaves in the leaf-disk choice test. Type of leaf age offered during fourth instar did not influence choice of larvae during fifth instar (one-tailed sign tests, alpha = 0.05); they JLILIL survivorship (%) I ll Ml IV Vv Cc instar of transfer to mature leaves Fic. 3. Heliconius erato larval stage survivorship when trans- ferred from young to mature leaves of Passiflora suberosa shoots. C, control = treatment 5. preferred young over mature leaves when reared ei- ther on young (p < 0.002) or mature (p < 0.005) leaves of P. suberosa. Leaf age affected H. erato survivorship during early instars (Fig. 3). There was a significant association be- tween the age of transfer to mature leaves and percent survivorship (Spearman’s rank correlation test, Rho = 0.975, n = 5, p < 0.05). When transferred to mature leaves during first and second instars only 30% and 60% of larvae respectively survived until pupation (Fig. 3). When transferred to mature leaves during third, fourth and fifth instars, larvae had similar sur- vivorship to those kept on intact shoots (Fig. 3). Larval development rates were significantly longer when lar- vae were transferred to shoots containing only mature leaves as first or second instars, but there was no effect on later instars (Fig. 4). 18 - @ es a S 16 - Aen il E -_ 2 144 ® b 5 ; Cc {e) a 7 FF | Py ® 10 -— — - = I mM IV Vv Cc instar of transfer to mature leaves Fic. 4. Heliconius erato larval stage development time when transferred from young to mature leaves of Passiflora suberosa shoots. C control = treatment 5. Values followed by the same letter are not significantly different (Anova — log transformed data, n = 48, p < 0.001; Fisher's Protected Least Significant Difference multiple comparison tests, p < 0.001). 112 The size of adults that emerged from larvae trans- ferred during fourth instar to cuttings containing ma- ture leaves (36.97 + 0.42 mm) was not significantly dif- ferent from that of those kept feeding on young leaves (38.08 + 0.37 mm) (unpaired, two-tailed t-test, n = 36, p > 0.06). Similarly, the size attained in the adult stage was not affected by transference imposed on all instars in the experiment designed to test effects of leaf age on larval performance (one-way Anova, n = 35, p > 0.37). DISCUSSION Our data confirmed field observations in that H. erato phyllis larvae feed continuously from the termi- nal bud (young tissues) to the basal portion (mature leaves) of P. suberosa shoots. Young tissues are pre- ferred by all instars, and this preference is related to larval survivorship and development rates. As already pointed out, young larvae move to the terminal bud when placed on any mature leaf. Thus, larvae start feeding on the apical portion of the plant not simply because that is where the female lay the eggs. Larvae begin feeding on the apical portion of P. swberosa shoots because the corresponding tissues are more suitable for development of first larval instars. Larvae feed on a wider range of leaf ages in the later instars. A progressive loss in selectivity as larvae age has been found in some insects, however the underlying mech- anisms for such an ontogenetic changes still not being clearly understood (Lewis & van Emden 1986). This ontogenetic change in feeding preference may be re- lated to changes in nutrition, plant chemistry or me- chanical barriers offered by young and mature P. suberosa leaves. Later instars may be better able to overcome defenses of older leaves and differences in food quality have most effect on the early instars; thus, the necessity to make the correct choices may be greatest for early instars (Reavey 1993). The choice tests showed that later instar larvae pre- fer young leaves independent of the experience in the previous instar. Thus, they are expected to be found mostly on the youngest surviving portions of host- plants under field conditions. Results also indicated that the age of leaves consumed in a given instar does not influence choices made by larvae during subse- quent instar. Even fifth instar larvae, which were com- paratively less selective in relation to leaf age, showed preference for young leaves when fed mature leaves during fourth instar. Thus, H. erato phyllis larval pref- erence could not be induced in relation to P. suberosa leaf age. Induction of feeding preference, as a result of previ- ous experience regarding particular plant species of- fered as larval food, has been demonstrated through choice tests performed with several lepidopterans JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY (Hanson 1983, Bernays & Chapman 1994; but see Bernays & Weiss 1996). Contrary to most of these lep- idopterans that are polyphagous, H. erato is oligopha- gous, using a few Passiflora species as larval hosts in southern Brazil (Perico & Arattijo 1991, Perico 1995). In an additional study carried out in our laboratory, Kerpel (1999) failed to show induction of larval feed- ing preference regarding the two preferred hosts (Pas- siflora misera and P. suberosa) of H. erato phyllis in Rio Grande do Sul State. The data showed that P. suberosa leaf age strongly affects H. erato phyllis larval performance. Although it is not known to what degree mechanical and nutri- tional factors contribute to these effects, they may con- stitute the proximate cause for the maintainance of at least two egg-laying behaviors of H. erato. First, it may explain why females selectively oviposit on terminal portion of intact P. suberosa shoots (Lopes 1991, Mu- grabi-Oliveira & Moreira 1996a); females that eventu- ally oviposit on shoots lacking or with damaged termi- nal portion may be in disadvantage compared to those laying on intact shoots due to lower larval survivorship and perhaps inferior performance. Second, it may elu- cidate why females lay only isolated eggs on P. suberosa shoots (Mugrabi-Oliveira & Moreira 1996b). By avoiding multiple oviposition they may reduce in- traspecific competition for the limited amount of leaves available on many host plants. H. erato larvae are cannibalistic, which is also an adaptation to host- plant size limitation (Brower 1997). Mugrabi-Oliveira and Moreira (1996a) found for a P. suberosa popula- tion of Viam&o County, RS, that in more than half of H. erato eggs are laid on plants whose total leaf area is less than that required for larval development. Food shortage in the fifth larval instar has profound effects on survivorship and size attained in the adult stage (D. Rodrigues & G. R. P. Moreira unpubl. data). Thus, we confirm the importance of variation in in- traspecific attributes of P. suberosa plants, more pre- cisely leaf age, in relation to preference and perfor- mance of H. erato phyllis larvae. Feeding preferences for young Passiflora auriculata tissues had already been demonstrated through choice experiments by Denno and Donnelly (1981) for larvae of Heliconius sara in Costa Rica. The physiological basis for such a feeding specialization in earlier instars remains unde- termined. It is known that several chemical and physi- cal traits of leaf tissues, such as pubescence, hardness, water content, nutrients and secondary metabolic compounds, vary according to age. It is also known that these leaf characteristics can play a major role in host-plant selection or performance of herbivorous in- sects (e.g., Scriber 1984, Slansky 1993, Bernays & Chapman 1994, Fernandes 1994), including flea bee- VOLUME 53, NUMBER 3 tles that use Passiflora as host plants (Thomas 1987). Smiley and Wisdom (1985) found a significant correla- tion between leaf nitrogen content of several sym- patric Passiflora species and larval growth rates of He- liconius ismenius and Heliconius melpomene in Costa Rica. They, however, failed to find any deleterious ef- fects of potential toxins (including alkaloids, tannins and cyanogenic compounds) on larval growth rates and survivorship. Although not quantified precisely yet, there is a substantial increase in tissue toughness with advance in age in P. suberosa leaves. In consequence, ontogenetic changes in leaf age selection could also be associated with morphological constraints on H. erato phyllis feeding apparatus. Early instars of some grasshoppers and lepidopterans do not feed upon fa- vored tissue or mature leaves because the mandibles are either not hard enough for effectively chewing, or their gape is not wide enough to grasp thick tissues (Bernays 1991). ACKNOWLEDGMENTS We thank Elna Mugrabi-Oliveira for supplying the plants and larvae used in the experiments. Cassiano S. Moreira and André Silva Gomes assisted in building the wooden frame supports. We are especially grateful to Woodruff W. Benson (UNICAMP) for significant improvements in the final version of manuscript made possible by his comments. We also wish to thank, Andrew V.Z. Brower (Oregon State University), Gilberto Albuquerque (UNFla), Lenice Medeiros (UNIJUI) and Nicoleta T.N. Sabetzki (EPAGRI), and two anonymous reviewers, for suggestions that improved earlier drafts of the manuscript. Financial support for the study came from a FAPERGS grant, number 94/50941.7, to G.R.P. Moreira. This is contribution number 315 of the Zoology Department of Federal University of Rio Grande do Sul. LITERATURE CITED ALEXANDER, A. J. 1961. A study of the biology and behavior of the caterpillars, pupae and emerging butterflies of the subfamily heliconiinae in Trinidad, West Indies. Part I. Some aspects of larval behavior. Zoologica 46:1—24. BENSON, W. W. 1978. Resource partitioning in passion vine butterflies. Evolution 32:493-518. BENSON, W. W., K. S. BRown JR. & L. E. GILBERT. 1976. Coevolution of plants and herbivores: passion vine butterflies. Evolution 29:659-680. Bernays, E. A. 1991. Evolution of insect morphology in relation to plants. Phil. Trans. R. Soc. Lond. B 333:257-264. Brernays, E. A. & R. F. CHAPMAN. 1994. Host-plant selection by phytophagous insects. Chapman & Hall, New York. BERNays, E. A. & M. R. WeIss. 1996. Induced food preferences in caterpillars: the need to identify mechanisms. Ent. Exp. App. 78:1-8. BroweER, A. V. Z. 1997. The evolution of ecologically important characters in Heliconius butterflies (Lepidoptera: Nym- phalidae): a cladistic review. Zool. J. Linn. Soc. 119:457-472. Brown, K.S. Jr. 1979. Ecologia geografica e evolugao nas florestas neotropicais. Universidade Estadual de Campinas, Campinas. . 1981. The biology of Heliconius and related genera. Ann. Rev. Entomol. 26:427-456. Brown, K. S. Jr. & O. H. H. MIELKE. 1972. The heliconians of Brazil (Lepidoptera: Nymphalidae). Part II. Introduction and general comments, with a supplementary revision of the tribe. Zoologica 57:1—40. 113 Conover, W. J. 1980. Practical nonparametric statistics. 2nd. ed., Wiley, New York. DENNO, R. F. & M. A. DONNELLY. 1981. Patterns of herbivory on Passiflora leaf tissues and species by generalized feeding insects. Ecol. Entomol. 6:11—16. FERNANDES, G. W, 1994. Plant mechanical defenses against insect herbivory. Revta. bras. Ent. 38:421—-433. Hanson, F. E. 1983. The behavioral and neurophysiological basis of food plant selection by lepidopterous larvae, pp. 3-23. In S. Ahmad (ed.), Herbivorous insects: host-seeking behavior and mechanisms. Academic Press, New York. Jones, R. E. 1991. Host location and oviposition on plants, pp. 139-171. In W. J. Balley & J. Ridsdill-Smith (eds.), Reproductive behavior of insects: individuals and populations. Chapman & Hall, New York. KERPEL, S. M. 1999. Mecanismos ecolégico-comportamentais envolvidos na selecao da planta hospedeira por Heliconius erato phyllis (Lepidoptera: Nymphalidae). Unpublished MSc. Dissertation, Instituto de Biociéncias, Universidade Federal do Rio Grande do Sul, Porto Alegre. Lewis, A. C. & H. F. VAN EMDEN. 1986. Assays for insect feeding, pp- 95-119. In J. R. Miller & T. A. Miller (eds.), Insect-plant interactions. Springer-Verlag, New York. Lopes, F. S. 1991. Padrées sazonais e evolucao do uso de plantas hospedeiras de larvas por Heliconius erato phyllis (L.) (Lepidoptera, Nymphalidae) na Serra do Japi, Sao Paulo. Unpublished Ph.D. Dissertation, Instituto de Biologia, Universidade Estadual de Campinas, Campinas. MENNA-BARRETO, Y. & A. M. ARAUJO. 1985. Evidence for host plant preferences in Heliconius erato phyllis from southern Brazil (Nymphalidae). J. Res. Lepid. 24:41—-46. MucraBl-O.iverra, E. & G. R. P. Morera. 1996a. Size of and damage on shoots of Passiflora suberosa (Passifloraceae) influence oviposition site selection of Heliconius erato phyllis (Lepidoptera: Nymphalidae). Revta bras. Zool. 13:939-953. . 1996b. Conspecific mimics and low host plant availability reduce egg laying by Heliconius erato phyllis (Lepidoptera: Nymphalidade). Revta bras. Zool. 13:929-937. PERICO, E. 1995. Interac&o entre quatro espécies de Heliconiini (Lepidoptera: Nymphalidae) e suas plantas hospedeiras (Passifloraceae) em uma floresta secundaria do Rio Grande do Sul, Brasil. Biociéncias 2:3-18. Perico, E. & A. M. ArRAUjo. 1991. Suitability of host plants (Passifloraceae) and their acceptableness by Heliconius erato and Dryas iulia (Lepidoptera: Nymphalidae). Evol. Biol. 5:59-74. Ramos, R. R. & A. V. L. Freitas. 1999. Population biology and wing color variation in Heliconius erato phyllis (Nymphalidae). J. Lepid. Soc. 53:11-21. REAVEY, D. 1993. Why body size matters to caterpillars, pp. 248-279. In N. E. Stamp & T. M. Casey (eds.), Caterpillars: ecological and evolutionary constraints on foraging. Chapman & Hall, New York. SCRIBER, J. M. 1984. Host-plant suitability, pp. 159-200. In W. J. Bell & R. T. Cardé (eds.), Chemical ecology of insects. Chapman & Hall, London. SLANSKY, F. JR. 1993. Nutritional ecology: the fundamental quest for nutrients, pp. 29-91. In N. E. Stamp & T. M. Casey (eds.), Caterpillars: ecological and evolutionary constraints on foraging. Chapman & Hall, New York. SMILEY, J. T. & C. S. WisDoM. 1985. Determinants of growth rate on chemically heterogenous host plants by specialist insects. Biochem. Syst. Ecol. 13:305-312. SOKAL, R. R. & F. J. ROLHF. 1981. Biometry. 2nd. ed. W. H. Freeman and Company, New York. Tuomas, C. D. 1987. Behavioural determination of diet breadth in insect herbivores: the effect of leaf age on choice of host species by beetles feeding on Passiflora vines. Oikos 48:211-216. Received for publication 28 December 1998; revised and accepted 10 December 1999. Journdl of the Lepidopterists’ Society 53(3), 1999, 114-125 FIVE NEW SPECIES OF ARGYROTAENIA (TORTRICIDAE: ARCHIPINI) FROM MEXICO AND THE SOUTHWESTERN UNITED STATES JOHN W. BROWN Systematic Entomology Laboratory, Plant Sciences Institute, Agricultural Research Service, U.S. Department of Agriculture c/o National Museum of Natural History, Washington, DC 20560-0168, U.S.A. (e-mail: jbrown@sel.barc.usda.gov) AND ASHLEY CRAMER Department of Biology, University of New Mexico, Albuquerque, New Mexico 87131, U.S.A. ABSTRACT. Based on an examination of 187 specimens, five new species of Argyrotaenia are recognized from Mexico and the south- western United States. A hypothesis of the phylogenetic relationships among the species is derived using Hennig86 to find the most parsimo- nious solution to the distribution of 14 morphological characters (5 characters of the forewing, 7 of the male genitalia, and 2 of the female gen- italia). Argyrotaenia spinacallis Brown & Cramer, new species, from the State of Veracruz, A. unda Brown & Cramer, new species, from the states of Mexico and Morelos, and A. octavana Brown & Cramer, from the states of Puebla and Veracruz, appear to form a monophyletic group with A. ponera (Walsingham), from Puebla. The last is redescribed and illustrated. Although superficially similar, A. coconinana Brown & Cramer, new species, from Arizona and New Mexico, and A. bialbistriata Brown & Cramer, from Arizona (Cochise Co.) and Durango, Mexico, may not be members of the “ponera group” owing to their considerable divergence in male and female genitalia. Additional key words: phylogeny, ponera group, morphology, genitalia. With the exception of a single widespread Palaearc- tic species, the tortricid genus Argyrotaenia Stephens is restricted to the New World (Razowski 1997), with about 64 described species occupying portions of the Western Hemisphere from Canada to Argentina. Field work over the past four decades has revealed a surpris- ingly large number of undescribed species in Central and South America. The purpose of this paper is to de- scribe three of them from central Mexico that appear to form a monophyletic group with Argyrotaenia pon- era (Walsingham), and two from the southwestern United States and northern Mexico that are superfi- cially similar to A. ponera. Although the “ponera group” may be confined to Mexico, it is clearly tem- perate/boreal rather than tropical in origin, restricted to the higher elevations of the Mexican states of Mex- ico, Morelos, Puebla, and Veracruz. The five new species described herein underscore the considerable alpha-level taxonomic work that remains to be done in the New World south of the United States border. MATERIALS AND METHODS Material examined. We examined 187 pinned specimens of adult moths obtained from or studied at the following institutions: American Museum of Natural History (AMNH), New York, New York; The Natural History Museum (BMNH), London, United Kingdom; Canadian National Collection (CNC), Ot- tawa, Ontario, Canada; Ray B. Nagle private collection (RNC), Tucson, Arizona; Essig Museum of Entomol- ogy (UCB), University of California, Berkeley; and Na- tional Museum of Natural History (USNM), Smith- sonian Institution, Washington, DC. Specimens were sorted by geographic location and examined for differ- ences in the male and female genitalia. Genitalia preparations of representative individuals were made following the methodology summarized in Brown and Powell (1991). Male genitalia were photographed us- ing a SONY DKC5000 digital camera and enhanced using Adobe Photoshop. Illustrations of female geni- talia were drawn with the aid of a microprojector. Un- less indicated otherwise, genitalia illustrations are of a single preparation. Forewing measurements were made with a transparent millimeter ruler under low power of a Leica MZI12 dissecting microscope. Forewing length was measured in a straight line from the base of the wing to the apical region, including the fringe. Forewing width was measured at the widest place on the wing perpendicular to the line measuring length. Where available, 10 individuals of each sex were measured. Photographs were taken with a Wild M 400 microscope with camera attachment. Terminol- ogy for wing venation and genitalic structures follows Horak (1984). Abbreviations are as follows: FW = forewing; HW = hindwing; DC = discal cell; ca. = circa (approximately); n = number of individuals or prepara- tions examined; X = mean; Mtns = Mountains; N, S, E, W = compass points. Phylogeny. A phylogenetic analysis was conducted on the 6 species of Argyrotaenia suspected to com- prise the “ponera group” (plus an out-group). The analysis was based on 14 morphological characters (8 binary and 6 multi-state), including 5 characters of the forewing, 7 of the male genitalia, and 2 of the female VOLUME 53, NUMBER 3 115 TABLE 1. Morphologic characters for cladistic analysis (0 = plesiomorphic state; 1-3 = apomorphic states). 1. FW L:W ratio 0 - 2.3-2.7 5 1 - 2.8-3.0 2. FW length 0 - 9.0-10.5 mm 1 - 11.0-13.0 mm. 0 - Without white spot at end of DC 3. FW pattern elements 1 - With diffuse white spot at end of DC 4. FW pattern elements 0 - Without pale longitudinal streak 1 - Pale longitudinal streak weak, extending to ca. end of DC 2 - Pale longitudinal streak well-developed, extending beyond end of DC 5. FW pattern elements 0 - Without dark longitudinal streak below distal two-thirds of DC 1 - With dark longitudinal streak below distal two-thirds of DC D> . Shape of valva ~] . Base of sacculus 0 - Somewhat rounded-triangular 1 - Subrectangular 0 - Unmodified 1 - With small rounded lobe 2 - With moderate, partially free, rounded lobe 3 - With large, free, rounded lobe . Venter of sacculus 0 - Smooth Go 1 - Slightly bumpy/warty 2 - With spinelike teeth 0 - Unmodified, moderately broad 9. Uncus 1 - Extremely broad 10. Uncus 0 - Unmodified, uniform in width 1 - Slightly tapered distally 11. Aedeagus 0 - Smooth 1 - With 2-4 spinelike teeth at distal end 2 - With 4-6 spinelike near middle 3 - With irregular rows of spinelike teeth from middle to end 12. Aedeagus 0 - About 4 times as long as wide, weakly curved 1 - Greater than 6 times as long as wide, strongly curved 13. Antrum 0 - Small, poorly defined 1 - Well developed, funnel shaped 2 - Large, cup shaped 14, Signum 0 - Lacking sclerotized projections from region bearing capitulum 1 - With sclerotized projection extending anterad from area bearing capitulum. 2 - With sclerotized projection extending posterad from area bearing capitulum. genitalia. Character state polarity was determined pri- marily using the out-group method. Although sister- group relationships are virtually unknown for Argyro- taenia, recent work by Landry et al. (1999) has identified a monophyletic species group comprised of A. franciscana (Walsingham), A. citrana (Fernald), A. citrana isolatissima Powell, A. citrana insularis Powell, A. niscana (Kearfott), and an undescribed species; this group was used as the out-group. Descriptions of character states. The character states used in the analysis are described below. Their putative plesiomorphic and apomorphic states are pre- sented in Table 1, and their distribution among the taxa is presented in Table 2. Forewing (Characters 1-5). Most species of Argyro- taenia, including the out-group, have a rather broad forewing with a pattern that typically includes one or more oblique fascia and a small, flattened, triangular patch at the costa about two-thirds the distance from the base to the apex. All of the species in the putative “ponera group” have a more narrow forewing that lacks the typical Argyrotaenia pattern. A forewing length-to-width ratio of 2.3-2.7, present in the out- group, was considered the plesiomorphic condition (character state 1.0); a forewing length-to-width ratio of 2.8—3.0 was considered the apomorphic condition (character state 1.1). A comparatively small forewing length (9.0-10.5 mm) was assumed to represent the plesiomorphic condition (character state 2.0), and a larger forewing length (11.0-13.0 mm) the apomor- phic state (character state 2.1). A forewing without a white spot at the distal end of discal cell was assumed to represent the primitive condition (character state TABLE 2. Distribution of character states for cladistic analysis (“P” = missing data). 000 000 000 000 00 ponera 111 200 310 021 ?? spinacallis 101 000 210 131 11 unda 101 100 110 111 21 outgroup octavana 101 100 110 111 20 coconinana 110 211 121 000 02 bialbistriata 100 211 121 000 01 116 3.0), and a forewing with a diffuse white spot at the end of the discal cell, characteristic of A. ponera, A. spinacallis, A. unda, and A. octavana, the apomorphic state (character state 3.1). The absence of a pale longi- tudinal streak through the discal cell of the forewing was interpreted as plesiomorphic (character state 4.0); the presence of an ill-defined or narrow, pale longitu- dinal streak extending from the forewing base to about the end of the discal cell, as is present in A. unda and A. octavana, was interpreted as derived (character state 4.1); and the presence of a well-developed pale longitudinal streak extending from near the base of the forewing to beyond the end of the discal cell, present in A. ponera, A. coconinana, and A. bialbistriata, was considered the most derived state (character state 4.2). The absence of a dark longitudinal streak below the distal two-thirds of the discal cell of the forewing was considered the plesiomorphic condition (character state 5.0), and the presence of a dark longitudinal streak in this region was considered the apomorphic condition (character state 5.1). Male genitalia (Characters 6-12). A short, some- what rounded-triangular valva, present in many species of Argyrotaenia including the out-group, was considered plesiomorphic (character state 6.0), and a subrectangular valva apomorphic (character state 6.1). An unmodified base of the sacculus was considered plesiomorphic (character state 7.0); the presence of a small, rounded lobe the first step in the transformation series (character state 7.1); the presence of a well-de- veloped, partially free, rounded lobe the next advance- ment (character state 7.2); and the presence of a large, free, rounded lobe the final step in the series (charac- ter state 7.3). The venter of the sacculus is relatively smooth in most species of Argyrotaenia (character state 8.0). A slightly bumpy or warty ventral margin was considered derived (character state 8.1), and the presence of spinelike teeth along the margin, charac- teristic of A. coconinana and A. bialbistriata, was con- sidered the most derived state (character state 8.2). A moderately broad uncus (character state 9.0) of uni- form width (character state 10.0) is present in most species of Argyrotaneia. The extremely broad uncus of A. coconinana and A. bialbistriata was considered the apomorphic condition (character state 9.1). An uncus that is slightly tapered distally also was considered an apomorphic state (character state 10.1). The moder- ately slender, slight curved aedeagus lacking external spines, present in A. coconinana and A. bialbistriata, was considered the plesiomorphic state (character state 11.0). An aedeagus with 24 spinelike teeth at the distal end (character state 11.1) was considered the first step in a transformation series leading to 4-6 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY spinelike teeth near the middle of the aedeagus (char- acter state 11.2) and irregular rows of spinelike teeth from near the middle to the distal tip of the aedeagus (character state 11.3). A comparatively short (about 4 times as long as wide), weakly curved aedeagus, present in A. coconinana and A. biablistriata, was considered the plesiomorphic state (character 12.0), and a rela- tively long (greater than 6 times as long as wide), slen- der, strongly curved aedeagus (character state 12.1), present in A. ponera, A. spinacallis, A. unda, and A. octavana, was considered the apomorphic state. Female genitalia (Characters 13-14). A small, poorly defined antrum was considered plesiomorphic (character state 13.0); a well developed, funnel-shaped antrum apomorphic (character state 13.1); and a large, cup-shaped antrum the most derived state in the transformation series (character state 13.2). In most Argyrotaenia the signum is typical of other members of the tribe Archipini, with a well developed, elongate, interior spine and an external capitulum (see Horak 1984). Because of the extremely variable configuration and development of internal sclerites extending dorsad and posterad from the area bearing the capitulum, we selected the most simple condition, i.e., sclerites ab- sent, as the plesiomorphic state (character state 14.0). The presence a sclerotized projection extending an- terad of the capitulum (character 14.1) was interpreted as apomorphic, and the presence of an additional scle- rotized projection extending posterad (character state 14.2) was considered the most derived state. RESULTS The data set was subjected to parsimony analysis us- ing Hennig86 version 1.5 (Lipscomb 1994), employing the “mhennig*” command. This algorithm constructs trees, each by a single pass through the data, by adding the taxa in a different sequence each time, and then applies branch-swapping to each of the trees. The analysis generated one most parsimonious tree with a length of 27, a consistency index of 0.81, and a reten- tion index of 0.77. The resulting cladogram (Fig. 1) in- dicates that A. unda, A. ponera, A. spinacallis, and A. octavana form a monophyletic group supported by the following characters: 1) diffuse white spot near end of the discal cell of the forewing (character 3); 2) rounded lobe at the base of the sacculus (character 7); 3) spinelike teeth on the aedeagus (character 11); 4) a long, slender, curved aedeagus (character 12); and 5) antrum well developed (character 13). The basal posi- tion of A. coconinana plus A. bialbistriata, with a com- parable set of characters supporting their monophyly, suggests that they may not be members of the “ponera group’ as currently defined. The cladogram (Fig. 1) is VOLUME 53, NUMBER 3 concordant with the geographic distribution of the species (Fig. 2), i.e., A. unda, A. spinacallis, A. octa- vana, and A. ponera occur in close geographic proxim- ity, while A. coconinana and A. bialbistriata occur rel- atively far to the north. All of the species treated herein are distinguished from other Argyrotaenia by the absence of the typical semicircular or triangular subapical patch of dark scales on the forewing characteristic of most species, and the presence of a diffuse white dot near the termi- nation of the discal cell or a pale streak through the discal cell. The forewing is more elongate or narrow than most species in the genus: forewing length: width ranges from 2.8-3.0 (X = 2.9; n = 10) for the in- cluded species in contrast to 2.3-2.8 (k = 2.6; n = 10) for members of the franciscana group. According to Powell (pers. comm.), the latter feature is characteris- tic of cold temperature microlepidoptera in various families (e.g., Elachistidae (Ethmiinae), Tortricidae). The short, somewhat rounded-triangular valvae and broad uncus of the species treated herein are similar to many other congeners, e.g., A. franciscana and A. am- atana (Dyar), and the large, cup-shaped antrum of A. unda, new species, is highly reminiscent of that of A. dichotoma (Walsingham), of which no males are known. The dichotomous key below is based on maculation and size of forewing. However, genitalia are much more reliable than facies for species-level identifications, and dissections are recommended. Although males of each species possess at least one diagnostic feature, shapes of the antrum and the signum in the female exhibit con- siderably more conspicuous and reliable differences. KEY TO THE SPECIES OF THE PONERA GROUP 1. Forewing with at least one white or pale yellow longi- tudinal streak extending through discal cell ............... 2 V’. Forewing without pale longitudinal streak in discal cell (LSA. 2B) sa cy ee eee eM mea GR te a spinacallis 2. Pale longitudinal streak extending along upper portion of discal cell arising at or near base of forewing .............. 4 2’, Pale longitudinal streak extending along upper portion of discal cell arising approximately one-fourth distance from base to apex of forewing, bordered below in distal two- thirds by a distinct, narrow, dark brown streak ............. 3 3. Female with second white streak along lower edge of discal cell; male forewing ground color usually red brown; Du- rango, Mexico, southeastern Arizona (Fig. 8)..... bialbistriata 3’. Female without second white streak along lower edge of discal cell; male forewing ground color usually pale brown; Arizona, New Mexico (Fig. 7) .................. coconinana 4, Forewing length greater than 11.0 mm (Fig. 3) ....... ponera 4”, Forewing length less than 11.0mm ..................... 5 5. Forewing with distinct rust-brown dash along costal edge of discal cell in distal one-third; diffuse whitish spot at end of Cell(icn ois ee eme nn eine emer hohe. Toh unda 5’. Forewing lacking distinct rust-brown dash along costal edge of discal cell; fine white dot at end of cell (Fig. 6) ... . octavana IG Outgroup coconinana bialbistriata spinacallis ponera unda octavana Fic. 1. Hypothesis of phylogenetic relationships among Argyro- taenia ponera, A. spinacallis, A. unda, A. octavana, A. bialbistriata, and A. coconinana. Numbers on the left refer to characters (1—14); numbers on the right refer to character states (e.g., 0 = plesiomor- phic, 1-3 = apomorphic) (see Tables 1 and 2). SYSTEMATICS Argyrotaenia ponera (Walsingham) (Figs. 3, 9) Tortrix ponera Walsingham, 1914:279. Argyrotaenia ponera: Obraztsov 1961:38; Powell et al. 1995:147. Redescription. Male. Head: Frons with sparse, smooth scaling below mid eye, pale red brown; vertex roughened above, light brownish red. Labial palpus pale red brown mesally, light brownish red laterally. Antennal scaling brown; scape pale red brown, frosted with whitish scales. Thorax: Light brownish copper, lighter mediodorsally. Forewing (Fig. 3): Length 11.0-13.0 mm (x = 12.2; n = 10). Upper side light brown; a conspicuous silver-white longitudi- nal streak extending from base, ending in a distinct silver-white dot just beyond end of DC, more diffuse and ill-defined from spot to termen; an ill-defined, darker brownish copper longitudinal streak immediately above white longitudinal streak; a small patch of darker brownish copper scales below silver-white streak beyond end of DC, accentuating silver-white dot near end of DC. Fringe pale gray. Un- der side pale gray. Hindwing: Upper side pale gray. Fringe slightly lighter than ground color. Under side pale gray. Genitalia: As in Fig. 9 (photograph of JWB slide 1035, Puebla, Mexico; 3 preparations examined). Uncus moderately broad, uniform in width, with trun- cate tip. Socius short, fused with tegumen. Arms of gnathos narrow, 118 600 MILES KILOMETERS JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fic. 2. Geographic distribution of Argyrotaenia ponera (star), A. spinacallis (open square), A. unda (closed triangles), A. octavana (closed squares), A. bialbistriata (open circles), and A. coconinana (closed circles). uniform in width, united distally into an elongate pointed tip. Transtilla a simple, sclerotized, narrow band. Valva short, somewhat rounded-triangular, costa undifferentiated; sacculus moderately long, slender, curved, ending ca. 0.8 distance from base to apex; base of sacculus with large, free, semicircular lobe; venter of distal 0.5 of sacculus with small bumps. Aedeagus long, slender, curved, with 4-6 spinelike teeth near the middle along right side; comuti not evident (deciduous and lost); phallobase moderate, slightly swollen distad to attachment point of juxta. Female. Unknown. Holotype 4, Mexico [Puebla], Popocatapetl Park, 13,000’ [4000 m], 6047 Wlsm 1908 (Wm. Schaus, USNM). Paratypes. Same data as holotype, 1 4, “co-type,” 33946 (BMNH), 1d, “co-type,” 33947 (BMNH). VOLUME 53, NUMBER 3 Additional specimens examined. MEXICO: Puebla: Popocat- apetl Park, 13,000’ [4000 m], 1 5 (USNM), 1 4, “co-type” Wlsm 6048 (USNM); Tlamacas, Volcan Popocatapetl, 3600-3660 m, 7d, 24 Au- gust 1987 (J: Brown & J. Powell, UCB). Diagnosis. Argyrotaenia ponera is superficially most similar to A. coconinana, from which it can be distinguished by the following features: 1) ground color of the forewing brownish copper rather than pale brown, lacking a distinct dark brown streak below silver- white streak; 2) large, well-defined lobe at the base of the sacculus, less defined in A. coconinana; 3) ventral edge of the distal 0.5 of the sacculus bumpy rather than dentate; and 4) aedeagus longer and more evenly curved, with 4-6 spinelike teeth near the middle. Remarks. Walsingham (1914:279) had 4 specimens when he de- seribed Tortrix ponera: one was designated as the holotype [“HT”] (USNM) and two as paratypes [“PT”] (BMNH), although the latter two are labeled as “co-types.” The fourth specimen (6048 Wlsm) also is labeled “co-type” but was not designated as a paratype in the orig- inal description—it is referred to only after the locality data: “Four specimens.” There is one additional specimen (USNM) with identi- cal collecting data that was not mentioned (or seen?) by Walsingham. Obraztsov’s (1961:35) illustration of A. ponera includes a photograph of an aedeagus, which is not that of the holotype of A. ponera. Argyrotaenia spinacallis Brown & Cramer, new species (Figs. 4, 11, 15) Description. Male. Head: Frons with sparse, smooth scaling be- low mid eye, pale red brown; vertex roughened above, pale red brown. Labial palpus pale yellow mesally, pale red brown laterally. Antennal scaling light brown; scape brown, with pale red-brown scales. Thorax: Light brown. Forewing (Fig. 4): Length 9.5-10.5 mm (xX = 10.0; n = 10). Upper side with costal one-half mostly cop- per brown, lower one-half pale yellow orange; distal one-third faintly reticulated; occasionally with ill-defined, thin, pale red-brown longi- tudinal streak along lower edge of DC; a pair of diffuse, small, dark- brown spots near distal end of DC, separated by diffuse white streak. Fringe pale red brown with basal row of slightly darker red- brown scales. Under side pale gray. Hindwing: Upper side pale gray. Fringe concolorous with hindwing. Under side pale gray. Genitalia: As in Fig. 11 (photograph of JWB slide 1063, Veracruz, Mexico; 3 preparations examined). Uncus moderately broad, with truncate tip. Socius short, fused to tegumen. Gnathos united distally into moder- ately long, pointed tip. Transtilla a simple, sclerotized, narrow band. Valva short, somewhat rounded-triangular, costa undifferentiated; sacculus moderately long, curved, attenuate distally, ending in a short free tip ca. 0.85 distance from base to apex; sacculus with moderately large, rounded, partially free lobe at base; venter of sacculus weakly dentate in distal 0.25. Aedeagus moderately long, slender, curved near middle; irregular rows of spinelike teeth from near middle to distal end of aedeagus. Cornuti not evident (deciduous and lost). Female. FW length 9.3-9.5 mm (x = 9.4; n = 2). Essentially as described for male, except hindwing with pale gray-brown overscal- ing. Genitalia: As in Fig. 15 (drawn from JWB slide 1036, Veracruz, Mexico; 2 preparations examined). Papillae anales unmodified, broad. Sterigma a slender, strongly sclerotized, weakly undulate band; ostium bursae moderately broad; antrum large, funnel shaped, with long ventral tube, strongly sclerotized. Ductus bursae moderately long, with distinct junction between corpus bursae and ductus bursae. Corpus bursae ovoid; signum with sclerite extending posterad of capitulum greatly reduced, portion extending anterad well developed. ; Holotype 3, Mexico, Veracruz, Cafion de las Minas, 13 km NE Perote, 2150 m, 18 August 1987 (J. Brown & J. Powell, UCB). Paratypes. Same data as for holotype, 20d, 2 2, 18/19 August 1987 (J. Brown & J. Powell, UCB), 1 3, 19 August 1987 (J. Doyen, UCB). Additional specimen examined. MEXICO: Mexico: La Mar- quesa, 1 d, 13 July 1966 (O. Flint & A. Ortiz, USNM). Diagnosis. Argyrotaenia spinacallis differs from all other species in the group by the absence of the pale longitudinal streak 119 on the forewing. Average male forewing length (10.0 mm) is less than that of A. ponera (12.2 mm) and A. coconinana (12.2 mm) and slightly greater than that of A. unda (9.4 mm) and A. octavana (9.4 mim). The female genitalia of A. spinacallis are distinguished easily from all other species by the strongly sclerotized, funnel-shaped antrum. In comparison with A. ponera, the male genitalia have slightly shorter, more rounded valvae; a slightly less well-developed rounded lobe at the base of the sacculus; and a slightly longer aedea- gus, with irregular rows of spinelike teeth from near the middle to the distal end. The somber forewing pattern, spinelike teeth of the aedeagus in the male, and funnel-shaped antrum in the female eas- ily separate this species from other members of the ponera group. Remarks. The single male from La Marquesa is similar to A. spinacallis in general appearance, but has a larger forewing length (10.8 mm) and a darker brown ground color. Because the genitalia slide (Razowski no. 11072) apparently is lost, comparison of these structures is impossible; hence the specimen is not included in the type series. In addition to the locality and slide labels, the specimens bears a label indicating “Holotype” and one indicating “Arygyrotae- nia Ppovera Wlsm.,” probably referring to A. ponera. Etymology. The species name refers to the irregular rows of spinelike teeth on the aedeagus. Argyrotaenia unda Brown & Cramer, new species (Figs. 5, 10, 16) Description. Male. Head: Frons with sparse, smooth scaling be- low mid eye, copper brown; vertex roughened above, yellowish white. Labial palpus pale red brown mesally, light copper laterally. Antennal scaling pale red brown; scape copper orange, with pale red-brown frosting. Thorax: Copper orange. Forewing (Fig. 5): Length 9.0-10.0 mm (x = 9.4; n = 10). Upper side rust to light cop- per, rust brown apically; conspicuous thin, silver-white longitudinal streak from base to beyond middle of DC; an ill-defined pale-brown line immediately below silver-white streak; conspicuous rust brown streak along distal one-third of costal edge of DC, bordered above by diffuse white scaling, terminating in a diffuse silver-white spot. Fringe pale red brown. Under side light gold. Hindwing: Upper side white. Fringe yellowish white. Under side yellowish white. Geni- talia: As in Fig. 10 (photograph of JAP slide 6271, Mexico, Mexico; 3 preparations examined). Uncus moderately broad, slightly tapered distally, with truncate tip. Socius, gnathos, and transtilla as in ponera and spinacallis. Valva short, costa slightly arched dorsally, with a lightly sclerotized subrectangular lobe near base; sacculus moder- ately long, curved, attenuate distally, ending in a short free pointed tip; base of sacculus with an ill-defined, semicircular lobe. Aedeagus moderately long, not as strongly bent as in ponera and spinacallis; distal end with 24 small spinelike teeth; cornuti 12-15 in a dense fascicle. Female. FW length 9.0-10.0 mm (x = 9.4; n = 6). Essentially as described for male, except hindwing with gray-brown overscaling. Genitalia: As in Fig. 16 (drawn from JAP 6272 slide, Mexico, Mex- ico; 3 preparations examined). Papillae anales unmodified, broad. Sterigma moderately heavily sclerotized; ostium bursae moderately small, somewhat crescent shaped; antrum extremely large, cup shaped, strongly sclerotized. Ductus bursae moderately long, with distinct junction between corpus bursae and ductus bursae; poste- rior one-thrid of ductus bursae broad, strongly sclerotized. Corpus bursae ovoid; signum with sclerite extending posterad of capitulum reduced; anteriorly-directed sclerite extremely elongate, attenuate, extending nearly to anteriormost wall of corpus bursae as narrow line. Holotype 4, Mexico, Mexico, 7 air km WSW Juchitepec, 2750 m, 24 August 1987 (J. Brown & J. Powell, UCB). Paratypes. MEXICO: Mexico: same data as for holotype, 10 ¢, 3 2, 24/25 August 1987 (J. Brown & J. Powell, UCB); 10 air km SE Amecameca, 2720 m, 1 d, 23 August 1987 (J. Brown & J. Powell, UCB). Morelos: Lag. Zempoala, 3°, 10/11 July 1965 (O. Flint & A. Ortiz, USNM). Diagnosis. Argyrotaenia unda is superficially and genitalically most similar to A. octavana. In facies most specimens of A. unda can 120 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 3-8. Adults of Argyrotaenia. 3. A. ponera; 4. A. spinacallis; 5. A. unda; 6. A. octavana; 7. A. coconinana; 8. A. bialbistriata. VOLUME 53, NUMBER 3 be distinguished from the latter by a more isolated, narrow, dark rust-brown dash along the costal edge of the discal cell in the distal one-third, although this character is not always obvious. Likewise, male genitalia are not easily separated. In contrast, the female geni- talia are separated easily from those of A. octavana by the large, cup- shaped antrum and by the extremely elongate, slender, anteriorly-di- rected sclerite from the base of the area that bears the capitulum. Comments. The female genitalia of A. unda are similar to those of A. dichotoma (Walsingham) (illustrated in Obraztsov 1961:30) in the presence of a large, cup-shaped antrum. However, the adult of A. dichotoma, a large, dark-brown moth, with an ill-defined grayish spot near the end of the discal cell and a white hindwing, is superfi- cially quite dissimilar to A. wnda. It is possible that A. dichotoma is a member of the ponera group, based on the similarity of the female genitalia to those of A. unda and the pale spot at the end of the dis- cal cell, but in the absence of a male (i.e., the species is known only from two females from Guatemala), its assignment to the group would be speculative. While its forewing is narrow like other mem- bers of the ponera group (i.e., L:W = 2.88), the slightly undulate costa is not consistent with members of this group. It is unusual that Obraztsov (1961) figured the female genitalia of A. dichotoma but did not discuss it or formally propose a new combination in the text. Etymology. The species name is Latin for “wave.” Argyrotaenia octavana Brown & Cramer, new species (Figs. 6, 12, 17) Description. Male. Head: Frons with sparse, smooth scaling below mid eye, copper brown; vertex roughened above, yellowish white. Labial palpus pale red brown mesally, light copper laterally. Antennal scaling pale red brown; scape copper orange, with pale red-brown frosting. Thorax: Copper orange. Forewing (Fig. 6): Length 9.0—-10.0 mm (x = 9.4; n = 10). Upper side rust to light cop- per, pale rust brown apically; narrow, pale yellow-brown streak through lower half of discal cell; fine silver-white dot at end of dis- cal cell; thin, silver-white longitudinal line from base to beyond middle of DC. Fringe pale red brown. Under side light gold. Hind- wing: Upper side white. Fringe yellowish white. Under side yellow- ish white. Genitalia: As in Fig. 12 (photograph of JWB slide 1057, Mexico, Mexico; 5 preparations examined). Uncus moderately broad, slightly tapered distally, with truncate tip. Socius, gnathos, and transtilla as in unda. Valva short, costa slightly arched dorsally, with a lightly sclerotized subrectangular lobe near base; sacculus moderately long, curved, attenuate distally, ending in a short free pointed tip; base of sacculus with an ill-defined, semicircular lobe. Aedeagus moderately long, not as strongly bent as in ponera and spinacallis; distal end with 2-4 small spinelike teeth; cornuti 12-15 in a dense fascicle. Female. FW length 8.0 mm (n = 1). Essentially as described for male, except smaller forewing length and hindwing with dense gray- brown overscaling. Genitalia: As in Fig. 17 (drawn from JWB 1137 slide, Mexico, Veracruz; 1 preparation examined). Papillae anales unmodified, broad. Sterigma moderately heavily sclerotized; ostium bursae moderate in size; antrum a deep, slightly tapered, strongly sclerotized cup. Ductus bursae moderately long, with distinct junc- tion between corpus bursae and ductus bursae. Corpus bursae ovoid; signum an irregular sclerotized patch bearing the capitulum and long spine; posterior- and anterior-directed sclerites absent. Holotype 3, Mexico, Puebla, 10 km E Esperanza, 15 August 1987 (J. Brown & J. Powell, UCB). , Paratypes. MEXICO: Puebla: 10 km E Esperanza, 27 6, 15 Au- gust 1987 (J. Brown & J. Powell, UCB); 10 rd km E Escalante [prob- ably a lapsus of Esperanza], 4d, 15 August 1987 (J. Brown & J. Pow- ell, UCB). Veracruz: 22 rd km W Ciudad Mendoza, 2150 m, 12 4, 1 ?, 13 August 1987 (J. Brown & J. Powell, UCB). Diagnosis. Argyrotaenia octavana is most similar to A. unda, from which it can be distinguished by the characters described in the diagnosis of A. unda above. 121 Etymology. The species name refers to the eighth month of the year—all specimens were collected in August. Argyrotaenia coconinana Brown & Cramer, new species (Figs. 7, 14, 19) Description. Male. Head: Frons with sparse, smooth scaling be- low mid eye, pale red brown; vertex roughened above, pale red brown. Labial palpus light pale red brown mesally, slightly darker pale red brown laterally. Antennal scaling brown; scape pale red brown, with yellowish white scales. Thorax: Pale brown gold. Forewing (Fig. 7): Length 11.0-13.0 mm (x= 12.2; n = 10). Upper side pale red brown; a whitish to yellowish-white longitudinal streak extending from near base toward termen through DC; a faint, ill-de- fined, pale red-brown line immediately above yellowish-white streak, and a well-defined dark copper-brown line immediately be- low; a well-defined, short line in apical region concolorous with line below yellowish-white streak. Fringe mostly yellowish white, with some pale red brown distally. Under side pale gray. Hindwing: Up- per side white, with pale gray overscaling. Fringe concolorous with hindwing. Under side pale gray. Genitalia: As in Fig. 14 (drawn from USNM slide 89386, Arizona, USA; 4 preparations examined). Uncus extremely broad, uniform in width or slightly expanded distally, with truncate tip. Socius small, fused to tegumen. Arms of gnathos mod- erately long, united distally into pointed tip. Transtilla a simple, sclerotized, narrow band. Valva long, subrectangular, costa weakly arched downward at apex; sacculus moderately long, curved, ex- tending ca. 0.8 distance from base to apex, attenuate distally, con- spicuously dentate in distal 0.25. Aedeagus comparatively short, evenly curved near middle; weakly notched distally, with mesal, sclerotized, thornlike process and weak, short, subdistal, accessory lobe; vesica with fascicle of 10-15 long, slender cornuti, curved at base and in distal 0.1. Female. FW length 11.0-12.0 mm (x = 11.4; n = 10). Essentially as described for male, except slightly greater forewing length. Gen- italia: As in Fig. 19 (drawn from USNM slide 89387, Arizona, USA; 5 preparations examined). Papillae anales moderately large, flat. Sterigma a slender, strongly sclerotized, rounded band, with a lightly sclerotized disc mesally; ostium bursae slightly recessed; antrum small, in the form of a short, sclerotized, incomplete band at junc- tion of ostium bursae and ductus bursae (=colliculum?). Ductus bursae moderately long, with distinct junction between corpus bur- sae and ductus bursae. Corpus bursae ovoid; signum with large, sub- triangular, sclerotized flanges projecting anterad and posterad from capitulum. Holotype ¢, USA, Arizona, Coconino Co., Hochderffer Hill, 12.5 mi [20 km] NNW Flagstaff, 8500’ [2615 ml], 16 July 1964 (J. G. Franclemont, USNM). Paratypes. USA: ARIZONA: Coconino Co.: Same locality as for holotype, 14d, 16 July 1964, 4 d, 19, 17 July 1964, 1 4, 18 July 1964, 14,29, 19 July 1964 (all J. G. Franclemont, USNM, UCB, BMNH); Fort Valley, 7.5 mi [12 km] NW Flagstaff, 7350-7500’ [2262-2308 ml], 1 6, 26 June 1961, 1 6, 6 July 1961, 4, 9 July 1961, 1°, 11 July 1961, 2 4, 12 July 1961 (all R. W. Hodges, USNM), 14, 4 July 1964, 5 d, 8 July 1964, 2d, 11 July 1964, 5d, 13 July 1964, 6d, 14 July 1964, 1d, 15 July 1964, 5 d, 16 July 1964, 1 4, 17 July 1964 (all J. G. Franclemont, USNM); West Fork, 16 mi [25.5 km] SW Flagstaff, 6500’ [2000 m], 2 4, 4 July 1961, 1 4, 8 July 1961, 2 4, 19, 13 July 1961, 1 9, 19 July 1961 (all R. W. Hodges, USNM), 2.4, 3 July 1964 (J .G. Franclemont, USNM); West Fork Oak Creek, 19 road mi [30.4 km] SW Flagstaff, 6500’ [2000 m], 1 4, 13 July 1995, 2 4, 16 July 1995 (J. Powell & F. Sperling, UCB); Walnut Canyon, 6.3 mi [10 km] ESE Flagstaff, 6500’ [2000 m], 5 d, 5 July 1965 (J. G. Fran- clemont, USNM), 1 d, 14 July 1995, bl. (J. Powell & F. Sperling, UCB). Cochise Co.: Rustler Park, Chiricahua Mtns, 8500’ [2615 m], 1 9, 3 July 1972, 1 d, 12 July 1972, 1 2, 14 July 1972, 1 2, 27 July 1972, at light (J. Powell, UCB); East Turkey Creek, Chiricahua Mts., 6400’ [1970 ml], 1 2, 16 June 1966 (J. G. Franclemont, USNM). 122 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 9-14. Male genitalia of Argyrotaenia. 9. A. ponera; 10. A. unda; 11. A. spinacallis; 12. A. octavana; 13. A. bialbistriana; 14. A. coconinana. VOLUME 53, NUMBER 3 123 hey b BAe anes 2 se Fics. 15-19. Female genitalia of Argyrotaenia. 15. A. spinacallis; 16. A. unda; 17. A. octavana; 18. A. bialbistriata; 19. A. coconinana. 124 Pima Co.: Summerhaven, Mount Lemon, | 8, 21 July 1995, 1 2, 9 July 1996 (R. Nagle, RNC). NEW MEXICO: Grant Co.: Cherry Creek Camp, 13 mi [20.8 km] N Silver City, 6900’ [2123 m], 1 4, 1 9, 10 July 1964 (F,, P, & M. Rindge, AMNH). Diagnosis. Argyrotaenia coconinana is most similar to A. bial- bistriata, and their specific distinctness is not entirely certain. The two can be separated by the following: (1) in females of A. bialbis- triata the longitudinal streak through the discal cell is always silver- white rather than yellowish white or pale cream, and there is a sec- ond silver-white longitudinal streak along the lower edge of the discal cell; (2) males of A. bialbistriata have considerably darker scaling bordering the lower edge of the pale forewing longitudinal streak and in the apical region; (3) males and females of A. coconi- nana have a greater forewing length; and (4) the signum of A. co- coninana has a well developed, triangular sclerite projecting posterad from the capitulum that is lacking in A. bialbistriata. Etymology. The species name is derived from the county of Co- conino in Arizona. Argyrotaenia bialbistriata Brown & Cramer, new species (Figs. 8, 13, 18) Description. Male. Head: Frons with sparse, smooth scaling be- low mid eye, pale red brown; vertex roughened above, pale red brown. Labial palpus pale red brown mesally, slightly darker red brown laterally. Antennal scaling brown; scape pale red brown, with yellowish white scales. Thorax: Pale brown-gold. Forewing (Fig. 8): Length 9.8-9.9 mm (xX = 9.8; n = 2). Upper side pale red brown; a whitish to yellowish-white longitudinal streak extending from near base toward termen through DC; a faint, ill-defined, pale red-brown line immediately above yellowish-white streak, and a well-defined dark copper-brown line immediately below; a well-defined, short line in apical region concolorous with line below yellowish-white streak. Fringe mostly yellowish white, with some pale red brown dis- tally. Under side pale gray. Hindwing: Upper side white, with pale gray overscaling. Fringe concolorous with hindwing. Under side pale gray. Genitalia: As in Fig. 13 (photograph of JWB slide 1091; Du- rango, Mexico; 2 preparations examined). Uncus extremely broad, uniform in width, with truncate tip. Socius small, fused to tegumen. Arms of gnathos moderately long, united distally into pointed tip. Transtilla a simple, sclerotized, narrow band. Valva long, subrectan- gular, costa weakly arched downward at apex; sacculus moderately long, curved, extending ca. 0.8 distance from base to apex, attenuate distally, conspicuously dentate in distal 0.25. Aedeagus compara- tively short, curved near middle; weakly notched distally, with mesal, sclerotized, thornlike process and weak, short, subdistal, accessory lobe; vesica with fascicle of 10-15 long, slender cornuti, curved at base and in distal 0.1. Female. FW length 9.5-10.0 mm (x = 9.7; n = 3). Essentially as described for male, except forewing maculation much more pale, with a second whitish longitudinal streak along the lower edge of the discal cell. Genitalia: As in Fig. 18 (drawn from JWB slide 1110, Du- rango, Mexico; 3 preparations examined). Papillae anales moder- ately large, flat. Sterigma a slender, strongly sclerotized, rounded band, with a lightly sclerotized disc mesally; ostium bursae slightly recessed; antrum small, in the form of a short, sclerotized, incom- plete band at junction of ostium bursae and ductus bursae (=collicu- lum?). Ductus bursae moderately long, with distinct junction be- tween corpus bursae and ductus bursae. Corpus bursae ovoid; signum without sclerotized projection posterad of capitulum. Holotype 4, Mexico, Durango, 10 mi (16 km) W E] Salto, 9000’ [2769 m], 6 June 1964 (J. E. H. Martin, CNC). Paratypes. MEXICO: Durango: 10 mi (16 km) W EI Salto, 9000’ [2769 m], 1 5, 19,6 June 1964 (J. E. H. Martin, CNC). Additional specimens examined. USA: Arizona: Cochise Co.: Barfoot Park, Chiricahua Mts., 8500’ [2615 m], 1 9, 14 June 1985 (P. Jump, UCB); Rustler Park, Chiricahua Mtns, 8500’ [2615 m], 12,3 July 1972, at light (J. Powell, UCB). JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Diagnosis. As mentioned in the diagnosis of A. coconinana, A. bialbistriata is extremely similar to the latter. The most conspicuous character separating the two is the shape of the signum in the fe- male. In A. bialbistriata the signum always lacks the large, distally attenuate projection posterad of the capitulum that is always well developed in A. coconinana. Comments. The two female specimens cited above from the Chiricahua Mountains of southeastern Arizona present somewhat of an enigma, suggesting the sympatric and synchronic occurrence of A. bialbistriata and A. coconinana in the Chiricahua Mountains. There are at least two different interpretations of this information. (1) The two species are sympatric and synchronic in the Chiricahua Mountains of southeastern Arizona, and can be separated by differ- ences in the shape of the signum. This interpretation is supported further by a number of qualitative characters listed in the diagnosis of A. cocninana. (2) The two are conspecific, and the apparently dis- tinct shape of the signum and other qualitative differences merely represent infraspecific variation. Although this explanation may be correct, there are no intermediate expressions of the signum shape; i.e., all females throughout the range of A. coconinana (except for the two females from the Chiricahua Mountains) are extremely con- sistent in this character state. The apparent disjunct occurrence of A. bialbistriata in the Chir- icahua Mountains of southeastern Arizona and Durango, Mexico, is illustrated by several other Lepidoptera, e.g., Sparganopseustis mar- tinana Powell (Tortricidae) (Powell 1986), Dorithia trigonana Brown & Obraztsov (Tortricidae) (Brown & Powell 1991), and Hy- paurotis crysalus (Edwards) (Lycaenidae) (Brown 1989). Etymology. The species name refers to the two white longitudi- nal streaks of the forewing of the female. ACKNOWLEDGMENTS We thank the following for allowing us to examine material in their care: Kevin Tuck (BMNH), P.T. Dang (CNC), Jerry Powell (UCB), and Ray Nagle, Tucson, Arizona. We thank the following for helpful comments on the manuscript: Jerry Powell, University of California, Berkeley; Steve Nakahara, USDA, Systematic Entomology Labora- tory, Beltsville, Maryland; David Smith, USDA, Systematic Entomol- ogy Laboratory, National Museum of Natural History, Washington, D.C.; William Miller, University of Minnesota, St. Paul; and Ronald Hodges, Eugene, Oregon. Linda Lawrence, USDA, Systematic Ento- mology Laboratory, provided the enhanced digital images of the male genitalia; and David Adamski, USDA, Systematic Entomology Labo- ratory, provided the drawings of the female genitalia. National Science Foundation grant BSR84-11459 to J. Powell funded field work in Mexico in 1987, during which much of the material examined was col- lected. The taxonomic study was supported in part by a National Sci- ence Foundation grant to A. Cramer in the context of the Smithsonian Institution Research Training Program (1998). LITERATURE CITED Brown, J. W. 1989 (1988). Records of Hypaurotis crysalus (Ed- wards) (Lycaenidae) from western Mexico. J. Res. Lepid. 27:135. Brown, J. W. & J. A. POWELL. 1991. Systematics of the Chrysoxena group of genera (Lepidoptera: Tortricidae: Euliini). Univ. Calif. Publ. Entomol. 111:1—87. Horak, M. 1984. Assessment of taxonomically significant struc- tures in Tortricinae (Lep., Tortricidae). Mitt. Schweiz. Entomol. Gesel. 57:3-64. LANDRY, B., J. POWELL & F. SPERLING. 1999. Systematics of the Ar- gyrotaenia franciscana (Lepidoptera: Tortricidae) species group: evidence from mitochondrial DNA. Ann. Entomol. Soc. Am. 92:40-46. Lipscoms, D. 1994. Cladistic analysis using Hennig86, version 1.5. Documentation for Hennig86 computer program. George Washington University. 122 pp. Opraztsov, N. S. 1961. Descriptions of and notes on North and Central American species of Argyrotaenia, with descriptions of VOLUME 53, NUMBER 3 a new genus (Lepidoptera, Tortricidae). Am. Mus. Novit. 2048:1—42. POWELL, J. A. 1986. Synopsis of the classification of Neotropical Tortricinae, with descriptions of new genera and species (Lepi- doptera: Tortricidae). Pan-Pacif. Entomol. 62:372-398. POWELL, J. A., J. RAZOWSKI & J. W. BRowN. 1995. Tortricidae: Tor- tricinae, pp. 138-150. In: Heppner, J. B. (ed.), Atlas of Neo- tropical Lepidoptera, checklist: part 2, Hyblaeoidea-Pyraloidea- Tortricoidea. Association for Tropical Lepidoptera. Scientific Publishers, Gainesville, Florida. 125 RAZOWSKI, J. 1997. Generic composition of the New World Archip- ini (Lepidoptera, Tortricidae) with description of two new gen- era and two new species. Misc. Zool. 20:125-130. WALSINGHAM, LorD T. DE Grey. 1909-1915. Biologia Centrali- Americana. Insecta, Lepidoptera Heterocera, Vol. 4. British Museum (Natural History), London. 482 pp. + figs. Received for publication 15 December 1998; revised and accepted 15 January 2000. GENERAL NOTES Journal of the Lepidopterists’ Society 53(3), 1999, 126-127 FIELD OBSERVATIONS ON LARVAL DIAPAUSE IN THE FLORIDA VICEROY, LIMENITIS ARCHIPPUS FLORIDENSIS Additional key words: hibernacula, photoperiod, subspecies, thermoperiod. Recent publications treating nymphaline butterflies of Florida do not mention the occurrence of diapause in the Florida viceroy, Limenitis archippus floridensis (Stkr.) (Gerberg & Arnett 1989, Minno & Emmel 1993, Smith et al. 1994, Emmel 1997). In addition, Williams and Platt (1987) could not induce diapause in laboratory strains of L. a. floridensis derived from wild stock from Dade Co. No diapause occurred among 220 larvae in three broods reared under laboratory controlled conditions (25 + 2°C) with photoperiods ranging from 8L:16D to 18L:6D, and they speculated (p. 351) that “.,. a high proportion of the Florida subspecies may have lost the ability to construct hibermacula and enter diapause in the third instar.” Here we report that some individuals in natural populations of L. a. floridensis do, in fact, begin to diapause in northcentral and southwestern Florida, whereas, others apparently remain “on the wing” throughout the year. Three separate field observations of apparent diapause were made by us during 1995 and 1996 (Table 1). Each involved the collecting of either hibernacula and (or) larvae from foliage of willow, Salix caroliniana L. (Salicaceae). All of the hibernacula were found between 1.5-2.5 m above the substrate. The five occupied hibermacula each contained a third instar larva (in apparent diapause). Only one of the hibernacula we found had had its larva eaten by a small insectivorous bird, as evidenced by a small hole having been pecked into the basal portion of the empty hiber- naculum. Since there was evidence of larval feeding on the branches supporting these hibernacula, we suspect that the remaining larvae had completed development and had left their foodplants to pupate. At the Everglades locality, one egg, one second instar larva on its perch, and two fifth instar larvae also were observed alongside the three occupied hibernacula during late January. Adult L. a. floridensis also were seen on the wing at the Corkscrew Park and Everglades Localities. At the Gainesville locality, the 25 unoccupied hibernacula apparently had recently been vacated by larvae, since their silken girdles covered the newly emerging spring buds, and their color was a copper-brown rather than gray (both useful characters for estimating hibernaculum age). Our observations indicate that some L. a. floridensis larvae apparently possess diapause capabilities. The observations at Everglades N. Pk. suggest differential responses to diapause inducing variables, since all life stages were found contemporaneously at the sites during the latter part of January, a time during the “dead of winter’, when the ambient photophase is of shortest duration. Some individuals of L. a. floridensis within a deme may begin to diapause by responding equally well to environmental combinations of photoperiod, temperature, and levels of relative humidity. [The induction of diapause is controlled mainly by photoperiod in all northern populations of Limenitis spp. (Clark & Platt 1969, Hong & Platt 1975)]. Such differences in diapause control mechanisms among insects have been described by Beck (1980) and Saunders (1982). Our field observations suggest that some L. a. floridensis fly all year long (contradicting Gerberg & Arnett 1989, who report that these insects only fly between April-Sept.). Induction of diapause in second instar larvae of the temperate subspecies, L. a. archippus correlates with the appearance of the whitish, dorsal mid-abdominal saddlepatch on the young larvae. This saddlepatch overlies both the primordial gonads and paired ventro- lateral neurosecretory ganglia. Platt (1989) presented evidence that this region of the larvae may be photosensitive, and that it influences both normal and abnormal development in northem populations of L. a. archippus. Southern populations of L. a. floridensis also possess larval saddlepatches, but the role of these saddlepatches remains unstudied. TABLE 1. Records of 30 hibernacula of L. a. floridensis collected from northcentral and southwestern Florida during 1995-1996. Hibernacula Date Location County Occupied Empty Observer 18-20/1/95 Corkscrew Collier 3 0 D. Flaim N. Park 18-20/1/95 Everglades Dade y) 0 D. Flaim N. Park Newnan Lake Alachua 0 25 Gainesville 23/111 /96 A. Platt A well-documented intergrade zone exists between L. a. archippus and L. a. floridensis in northern Florida and southern Georgia (Remington 1958, 1968, Williams & Platt 1987, Ritland 1990, Ritland & Brower in review). Gene flow exists between these two subspecies throughout the Florida penninsula, and northward to Athens, Georgia, and the great Dismal Swamp in southeastern Virginia (36°40’N; Clark & Clark 1951). Alleles controlling facultative diapause may pass from these northern populations into more southern populations, and vice-versa. It might be argued that the empty hibermacula we collected near Gainesville were those of L. arthemis astyanax, rather than belonging to L. archippus floridensis, since L. arthemis astyanax also occurs in Florida. However, five lines of evidence point strongly in favor of these hibermacula belonging to L. a. floridensis: 1) several authors (Kimball 1965, Gerberg & Amett 1989, Platt & Maudsley 1994) report that L. arthemis astyanax is rare especially in southern Florida; 2) we identified all of the maturing larvae as being those of L. a. floridensis (although all North American Limenitis larvae are difficult to tell apart, especially in their early instars); 3) during the time of our observations, no adult or immature specimens of L. arthemis astyanax were sighted by us; 4) all of our observations were made in open marshy habitats typical for L. a. floridensis, but not for L. arthemis astyanax; and, finally 5) The empty hibernacula found near Gainesville all were of relatively large size (between 1.75-2.5 mm long), a further indication that they belonged to L. archippus floridensis. ACKNOWLEDGEMENTS We thank L. F. Gall for comments and revisions suggested on an earlier version of this note. P. C. Platt and G. D. Flaim assisted us with our field observations in Florida. We thank Colleen Wilkens of UMBC Department of Biological Sciences for typing this manuscript. LITERATURE CITED BECK, S. D. 1980. Insect photoperiodism. 2nd ed. Academic Press, New York, New York. x + 387 pp. Cuark, A. H. & L. F. CLarK. 1951. The butterflies of Virginia. Smiths. Misc. Colls. 116(7):1—195. Cuark, S. H. & A. P. Pratt. 1969. Influence of photoperiod on development and larval diapause in the viceroy butterfly, Limenitis archippus. J. Insect Physiol. 15:1951—1957. EMMEL, T. C. 1997. Florida’s fabulous butterflies and moths. Vol. II. World Publs., Tampa, Florida. 128 pp. VOLUME 53, NUMBER 3 GERBERG, E. J. & R. H. ARNETT JR. 1989. Florida butterflies. Natl. Sci. Publs., Inc. Baltimore, Maryland. 90 pp. Hone, J. W. & A. P. Piatt. 1975. Critical photoperiod and day length threshold differences between northern and southern populations of the butterfly Limenitis archippus. J. Insect Physiol. 21:1159-1165. KIMBALL, C. P. 1965. Arthropods of Florida and neighboring land areas. Lepidoptera of Florida. Vol. 1. Florida Dept. Agric. Div. Plant Industry. Gainesville, Florida. 168 pp. MINNO, M. C. & T. C. EMMEL. 1993. Butterflies of the Florida Keys. Active Record Scientific Publishers, Inc., Gainesville, Florida. 168 pp. Pxatt, A. P. 1989. Possible photosensitivity of the abdominal saddlepatch region of Limenitis archippus larvae (Lepidoptera: Nymphalidae). Mol. Entomol. 3:57—-66. Puatr, A. P. & J. R. MaupsLey. 1994. Continued interspecific hybridization between Limenitis (Basilarchia) arthemis astyanax and L. (B.) archippus in the southeastern U.S. (Nymphalidae). J. Lepid. Soc. 48(3):190-198. REMINGTON, C. T. 1958. Genetics of populations of Lepidoptera. Proc. Tenth, Intl. Congr. Entomol. 2:787—-805. REMINGTON, C. L. 1968. Suture-zones of hybrid interaction between recently joined biotas. Evol. Biol. 2:321428. Journal of the Lepidopterists’ Society 53(3), 1999, 127-129 127 RITLAND, D. B. 1990. Localized interspecific hybridization between mimetic Limenitis butterflies (Nymphalidae) in Florida. J. Lepid. Soc. 44:163-173. RITLAND, D. B. & L. P. BROWER. In review. A mimicry related cline in viceroy butterfly (Limenitis archippus) wing coloration: model-switching from monarchs (Danaus plexippus) to queens (Danaus gilippus) in the southeastern United States. Holarctic Lepid. SAUNDERS, D. S. 1982. Insect clocks. 2nd ed. Pergamon Press., Ltd. New York, New York. 409 pp.; xvii. SmitH, D. S., L. D. MILLER, & J. Y. MILLER. 1994. The butterflies of the West Indies and south Florida. Oxford Univ. Press, Oxford, England. 346 pp. WILLIAMS, T. F. & A. P, Parr. 1987. Absence of diapause in a laboratory strain of the Florida viceroy butterfly. Am. Mid]. Nat. 117(2):346-352. David FLAIM AND AusTIN P. PLATT, Department of Biological Sciences, University of Maryland, Baltimore County, 1000 Hilltop Circle, Baltimore, Maryland 21250. Received for publication 6 February 1997; revised and accepted 8 September 1999. NOTES ON THE GENUS STHENOPIS (HEPIALIDAE) IN ALBERTA, CANADA Additional key words: semivoltine, biennialism. The nearctic genus Sthenopis Packard (Hepialidae) currently contains five species (Davis 1983), of which S. argenteomaculatus (Harris), S. purpurascens (Packard) and S. quadriguttatus (Grote) purportedly occur in Alberta (Bowman 1951). Despite their large size and peculiar habits, little is known about their biology and specimens are rare in collections. The purpose of this note is to report on the adult biology and distribution of the genus in Alberta. S. argenteomaculatus does not occur in Alberta, and S. quadrigut- tatus was placed into synonomy with S. purpurascens by Nielsen et al. (1999) based on morphological characteristics. Our observations of sympatric populations of S. purpurascens and S. quadriguttatus color morphs support this view. Specimens examined and study sites. A total of 96 Sthenopis specimens from Alberta and Saskatchewan were examined, from the following sources: Northern Forestry Centre (NFC) (Canadian Forest Service, Edmonton), University of Alberta Strickland Museum (UASM) and the private collections of the authors. Behavior observations and habitat notes were based on the following Alberta localities: Finnegan Ferry (51°8’N, 112°5’W), 15- July-1985 (DDL); Didsbury (51°40’N, 114°8’W), 23-July-1987 (BCS); 23 km N. of Lac La Biche (54°55’N, 112°05’W), 22-July- 1997 (BCS); Rock Island Lake (55°35’N, 113°25’W), 26-July-1997 (BCS); Gregoire Lake Provincial Park (56°35/N, 111°10’W), 24- July-1997 (BCS); 10 km S Cooking Lake (53°21’N, 113°05’W), 28- 31-July-1997 (DDL, BCS); Palisades Research Centre, Jasper National Park (52°58’N, 118°04’W) 1030 m, 8-July-1998 (BCS); Redwater Natural Area (53°55’N, 112°57’W), 28-July-1999 (BCS). Sthenopis argenteomaculatus occurs from Québec to New England, and westward to Minnesota and Ontario (Strecker 1893, Forbes 1923, Riotte 1992, Handfield 1999). It is also reported from Alberta (Bowman 1951) and Saskatchewan (Hooper 1981), and Ives and Wong (1988) state this species occurs throughout the prairie provinces. However, this species has often been confused with Sthenopis purpurascens (Forbes 1923), and specimens labeled as S. argenteomaculatus in the Bowman collection (UASM) and the NFC are variants of S. purpurascens. Hooper (1981) and Ives and Wong (1988) provide a figure of a specimen identified as S. argenteo- maculatus. Comparisons with illustrations of S. argenteomaculatus from eastern North America (Solomon 1995, Handfield 1999) and specimens from Nova Scotia (BCS) reveal that the figures in Hooper (1981) and Ives and Wong (1988) are actually S. purpurascens. Furthermore, the peak flight period of S. argenteomaculatus is in mid- to late June, whereas S. purpurascens has a much later peak, from mid-July to August (Handfield 1999). Hooper (1981) states that in Saskatchewan, “adults [of the Hepialidae] emerge from mid- July to September”. Based on this, previous reports of S. argenteo- maculatus for Alberta and Saskatchewan should be referred to S. purpurascens. Sthenopis purpurascens ranges from British Columbia and the Northwest Territories east to Labrador and New York (Grote 1864, Forbes 1923, Prentice 1965, Handfield 1999), and as far south as the White Mountains of Arizona in the west (D. Wagner pers. comm.). In Alberta, this species is most common throughout the boreal mixed wood and aspen parkland ecoregions, and occurs locally in the mountain and prairie regions. The boreal forest localities include a range of habitats; the Cooking Lake site consists of mature trembling aspen (Populus tremuloides) woods, with an understory of beaked hazelnut, Corylus cornuta, and wild red raspberry, Rubus idaeus. The Redwater site is sandy, open jack pine (Pinus banksiana) forest, interspersed with stands of trembling aspen and paper birch (Betula papyrifera). Green alder (Alnus crispa) is the most common understory shrub. The Palisades Research Centre locality is within the montane ecoregion (Strong & Leggat 1992), and consists of dry, open meadows with stands of trembling aspen and lodgepole pine (Pinus contorta). S. pupurascens also occurs in riparian balsam poplar (Populus balsamifera) groves in the mixed grass prairie ecoregion (Finnegan Ferry site); populations here are likely restricted to riparian areas, since the larvae bore in the roots of poplar and aspen, Populus spp. (Prentice 1965, Gross & Syme 1981). It appears that S. purpurascens occurs throughout most of the province where suitable host plants occur. The light color form (formerly Sthenopis quadriguttatus) occured together with typical S. purpurascens at all 1997 localities, with the exception of Gregoire Lake PP. The fact that both phenotypes were collected together at several sites suggests that the habitat requirements and phenology of the two phenotypes are very 128 similar, supporting the synonomy of quadriguttatus into purpurascens proposed by Nielsen et al. (1999). Furthermore, the two phenotypes are almost identical in wing pattern, shape and size; only the ground color varies. Similar morphs (with salmon or brown ground color) occur in Hepialus behrensi (Stretch) (D. Wagner pers. comm.) and Gazorycta noviganna (Barnes & Benjamin) (C. Schmidt unpubl. data). The relative frequency of phenotypes in the specimens examined is unbiased in males (17 “quadriguttatus” : 15 “purpurascens”, X? = 0.125, 0.50 < p < 0.75) but a significantly higher proportion of females exhibited the quadriguttatus phenotype (46:18; X? = 12.25, p < 0.001). Assuming both phenotypes are equally likely to be attracted to light, it appears that the mechanisms determining phenotype may be sex-linked. Flight observations. Male and female Sthenopis purpurascens were observed flying at dusk; nightly flight activity was very brief, occurring between 2210 h and 2300 h (Cooking Lake and Red- water), with sunset at 2135 h (MST). Crepuscular flight activity is characteristic of hepialids, including other members of the genus Sthenopis (Winn 1909, McCabe & Wagner 1989, Wagner & Rosovsky 1991). Females displayed a nearly stationary, hovering flight, usually less than one meter above the shrub understory. One female was observed ovipositing between 2245 h and 2300 h, while exhibiting this type of behavior (Redwater site). Eggs were broadcast over vegetation consisting of low, herbaceous plants and scattered aspen saplings and alder shrubs. Although it was difficult to determine the rate at which eggs were dropped, captive females can lay approximately 1.5 eggs per second. Males were only rarely observed flying, and flight became rapid and erratic when disturbed. Members of the genus Sthenopis are unusual in that males possess long-range sex attractants, whereas this strategy is usually characteristic of female Lepidoptera (Mallet 1984, Wagner & Rosovsky 1991). Two types of courtship behaviour have been observed in Sthenopis: Males of S. thule and S. argenteomaculatus form mating swarms (leks) which females enter to mate (Winn 1909, Covell 1981), and male S. auratus are sessile and call for females, fanning their wings over the scent tufts (McCabe & Wagner 1989). These two mating strategies may be density-dependent, with male lekking behaviour occurring at higher densities (Wagner & Rosovksy 1991). Since our observations of S. purpurascens failed to turn up males exhibiting courtship behavior, it remains unclear which courtship type this species exhibits. Female S. purpurascens are more likely to be collected, since there is a high female bias in examined specimens (64 2: 32 d) and sex ratios are unbiased in the Hepialidae (Wagner & Rosovsky 1991). This is likely due to the fact that females have a longer nightly flight period during which oviposition occurs, while males are only active for a brief time period (D. Wagner pers. comm.). In hepialids where females release sex attractants, collections are often male- biased (Wagner & Rosovsky 1991). Phenology. Data labels of examined specimens range from July 5th to August 14th, and indicate the peak flight occurs during the last two weeks of July. Larvae of S. purpurascens likely take two years to develop (Vallée & Béique 1979); this can result in adults being much more common in alternating years. Out of the 16 years represented by the specimens examined (between 1931 and 1999), only five are even-numbered years; this phenomenon has also been observed in other hepialids (Wagner et al. 1989). It is thought that biennial species remain synchronous through complex interactions with predators and parasitoids, abiotic catastrophes, and plant defense mechanisms (Mikkola & Kononenko 1989, Mikkola 1976, Wipking & Mengelkoch 1994). The odd-year biennialism in S. purpurascens is synchronous with other biennial Lepidoptera species in the west; many of the Yukon species of Boloria (Nymphalidae), Erebia, Oeneis (Satyridae) and Xestia (Noctuidae) have a greater adult abundance in odd-numbered years (Lafontaine & Wood 1997). This odd-year zone extends from Hudson Bay westward to Fennoscandia (Lafontaine & Wood 1997), but it is not known what causes synchrony between such a wide range of taxa. Further research is needed to determine the courtship and JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY mating behavior of S. purpurascens, whether or not density- dependent lekking occurs, if there is a pre-dawn flight period, and to verify if both phenotypes interbreed. We thank David Wagner for useful suggestions and additional information. Dr. George Ball and Danny Shpeley kindly made material in the Strickland Museum collection available, and Greg Pohl provided access to the collections at the Northern Forestry Centre. LITERATURE CITED Bowman, K. 1951. Annotated list of the Lepidoptera of Alberta. Can. J. Zool. 29:121—165. CoVvELL, C. V. JR. 1981. A field guide to the moths of eastern North America. Houghton Mifflin. Boston. 496 pp. Davis, D. R. 1983. Hepialidae. In Hodges, R.W. et al. (eds.), 1983. Check list of the Lepidoptera of America north of Mexico. E. W. Classey Ltd., London. 284 pp. Forbes, W. T. M. 1923. The Lepidoptera of New York and neighboring states. Pt. 1. Cornell Univ. Agric. Expt. Sta. Mem. 68, 729 pp. Gross, H. L. & P. D. SyME. 1981. Damage to aspen regeneration in northern Ontario by the ghost moth, Sthenopis quadriguttatus (Grote). Canadian Forestry Service, Research Notes 1:30-31. GroTE, A. R. 1864. Descriptions of North American Lepidoptera. Proc. Entomol. Soc. Philadelphia 3:73-92. HANDFIELD, L. 1999. Le guide des papillons du Québec. Version populaire. Broquet. Ottawa. 536 pp. + 123 plates. Hooper, R. R. 1981. Saskatchewan ghost moths. Blue Jay 39:141-142. Ives, W. G. H. & H. R. Wonc. 1988. Tree and shrub insects of the prairie provinces. Canadian Forestry Service, Northern Forestry Centre. Edmonton, Alberta. Information Report NOR-X-292. LAFONTAINE, J. D. & D. M. Woop. 1997. Butterflies and moths of the Yukon. In H. V. Danks & J. A. Downes (eds.), Insects of the Yukon. Biological Survey of Canada (Terrestrial Arthropods), Ottawa. 1034 pp. MALLET, J. 1984. Sex roles in the ghost moth Hepialus humuli (L.) and a review of mating in the Hepialidae (Lepidoptera). Zool. J. Linn. Soc. 79:67-82. McCasg, T. L. & D. L. WAGNER. 1989. The biology of Sthenopis auratus (Grote) (Lepidoptera: Hepialidae). J. New York Entomol. Soc. 97:1—10. MIKKOLA, K. 1976. Alternate-year flight of northern Xestia species (Lepidoptera, Noctuidae) and its adaptive significance. Annales entomologici Fennici 42:191-199. MikkoLa, K. & V. S. KONONENKO. 1989. Flight year of the alternate-year Xestia moths (Lepidoptera, Noctuidae) in north- eastern Siberia—a character from the Ice Ages? Nota Lepid. 12:144-152. NIELSEN, E. S., G. S. RoBINSON & D. L. WAGNER. 1999. Ghost- moths of the world: a global inventory and bibliography of the Exoporia (Mnesarchaeoidea and Hepialoidea) (Lepidoptera). J. Nat. History. In press. PRENTICE, R. M. 1965. Forest Lepidoptera of Canada recorded by the Forest Insect Survey. Vol. 4: Microlepidoptera. Canada Dept. of Forestry, Forest Entomology and Pathology Branch. Ottawa. 296 pp. RIoTTE, J. C. E. 1992. Annotated list of Ontario Lepidoptera. Royal Ontaria Museum, Publications in Life Sciences, Misc. Publ. Toronto. 208 pp. SOLOMON, J. D. 1995. Guide to insect borers of North American broadleaf trees and shrubs. U.S. Dept. Agric., Forest Service. Agric. Handbk. 706. Washington, DC. 735 pp. STRECKER, H. 1893. Descriptions of two new species of North American Bombycidae. Proc. Acad. Nat. Sci. Philadelphia 45:282-284. STRONG, W. & K. R. Leccat. 1992. Ecoregions of Alberta. Alberta Forestry Lands and Wildlife, Edmonton, Alberta. 59 pp. VOLUME 53, NUMBER 3 VALLEE, G. & R. BEIQUE. 1979. Dégats d'une hépiale et susceptibilité des peupliers. Phytoprotection 60:23-30, WAGNER, D. L. & J. Rosovsky. 1991. Mating systems in primitive Lepidoptera, with emphasis on the reproductive behaviour of Korscheltellus gracilis (Hepialidae). Zool. J. Linn. Soc. 102:277-303. Wacner, D. L., D. R. Tost, B. L. PARKER, W. E. WALLNER & J. G. LEONARD. 1989. Immature stages and natural enemies of Korscheltellus gracilis (Lepidoptera: Hepialidae). Ann. Entomol. Soc. Am. 82:717-724. Winn, A. F. 1909. The Hepialidae, or Ghost-moths. Can. Entomol. 41:189-193. WIPKING, W. & C. MENGELKOCH. 1994. Control of alternate-year flight activities in high-alpine Ringlet butterflies (Erebia, Journal of the Lepidopterists’ Society 53(3), 1999, 129-130 129 Satyridae) and Burmet moths (Zygaena, Zygaenidae) from temperate environments. In H.V. Danks (ed.), Insect life-cycle polymorphism. Theory, evolution and ecological consequences for seasonality and diapause control. Kluwer Academic Publishers, Dordrecht, Netherlands. 378 pp. B. CHRISTIAN SCHMIDT, 208 - 10422 78 Ave., Edmonton, Alberta, CANADA T6E 1P2, Davip D. Lawrik, 301 - 10556 84 Ave., Edmonton, Alberta, CANADA T6E 2H4. Received for publication 5 April 1999; revised and accepted 9 December 1999. TWO LARGE TROPICAL MOTHS (THYSANIA ZENOBIA (NOCTUIDAE) AND COCYTIUS ANTAEUS (SPHINGIDAE)) COLONIZE THE GALAPAGOS ISLANDS Additional key words: _ light traps, island colonization. The arrival and establishment of a species on an isolated oceanic island is a relatively rare event. The likelihood of colonization depends on a variety of features of the species, including dispersal ability, availability of food (hostplants or prey) and ability to reproduce. In this note, I discuss two recent Galapagos records of tropical moths in the context of island colonization. Thysania zenobia (Cramer) is a tropical migratory species which has been occasionally collected in the Holarctic region (Ferguson et al. 1991). Its life history is unknown, but legumes are considered probable larval foodplants (Covell 1984). Between 20 and 25 April 1996, three fresh males were collected in a Mercury vapor light trap near Asilo de la Paz, Floreana Island, at 338 m elevation. The trap was located at the border of the agricultural zone and native forest. In March 1997, I collected another specimen in a forest of the endemic composite, Scalesia pedunculata Hook at Los Gemelos, Santa Cruz island, at 580 m elevation, feeding in a bait trap (mixture of rotting fruit). The fresh condition of these specimens suggested that they were from a population extant on the island, rather than migrant. These Galapagos specimens are identical in wing pattern and size to series from continental United States reported by Covell (1984), Cocytius antaeus (Drury) is one of the larger hawk-moths of the Neotropical region. Members of the Annonaceae have been reported as hostplants (Kimball 1965). Dyar (1901) and Matteson (1933) described its life cycle. I collected two specimens on Santa Cruz Island. On 26 May 1996, I captured a fresh female in a mercury vapor lamp trap at Media Luna (580 m elevation), the fresh condition again suggesting an existing population. This habitat is a mature forest of the endemic Miconia robinsoniana Cong. (Melastomataceae), native ferns and the introduced tree Cinchona succirubra Klotzsch (Rubiaceae). One month later, one worn male was collected by Godfrey Merlen at an outdoor fluorescent light at the Charles Darwin Research Station (sea level). Although I have never collected larvae of this species, farmers in Santa Cruz and San Cristobal Islands have reported the presence of “voracious green hornworms” feeding on leaves and branches of the introduced custard apple (Annona cherimola Mill). It is likely that these reports refer to C. antaeus, because no other Galapagos sphingids feed on members of the Annonaceae. The lack of specimens of these two moth species in previous lepidopteran surveys of the islands suggests that these are relatively recent additions to the fauna. Hayes (1975) did not report their presence in the archipelago but his species list was based on specimens collected by early expeditions with less efficient light traps (kerosene lamps) and collections made by amateur entomologists. Recently (1989 and 1992), Bernard Landry carried out an intensive Lepidoptera survey on the islands but he never collected the species (Landry pers. comm.). However, it is also possible that the absence of these species from earlier collections is due to flight time. I trapped both species late at night (2300 h to 2400 h) and few collections have been made during these hours by earlier collectors. ; Several features of the biology of these two species may have contributed to their ability to reach the Galapagos. Both, C. antaeus and T. zenobia have a history of long dispersal events by active flying to new areas, including oceanic islands (Ferguson et al. 1991, Schreiber 1978). The occurrence of many species of Annonaceae, all of which were introduced by humans in the present century (Lawesson et al. 1987), has probably favored the establishment of C. antaeus. Although the hostplant of T. zenobia is unknown legumes are a likely candidate (Covell 1984). There are many species of legumes on the Galapagos islands, including native and endemic species, and one of these could provide a suitable hostplant. I suggest that many of the macrolepidoptera that have colonized the Galapagos arrived by natural means and not as a direct result of human activity. However, their establishment has been facilitated by the increase in the number of introduced plant species, deforestation and other human-related activities. Voucher specimens have been deposited in the entomological collection of the Charles Darwin Research Station Museum on Isla Santa Cruz, Galapagos. ACKNOWLEDGMENTS I thank Bernard Landry, Charles V. Covell, Charlotte Causton, Alan Tye and Robert Bensted-Smith for their helpful comments. Valentina Cruz offered her valuable companionship and help in the field. Field work was supported by UNESCO in the project “Ecological monitoring in the Galapagos”. LITERATURE CITED COVELL, C. V. 1984. A field guide to moths of Eastern North America. Hougton Mifflin Co., Boston. 496 pp. Dyar, H. G. 1901. Life histories of some North American moths. Proc. U.S. Natl. Mus. 23:255-284. FERGUSON, D. C., D. J. Hitpurn & B. Wricut. 1991. The Lepidoptera of Bermuda: their food plants, biogeography, and means of dispersal. Mem. Ent. Soc. Canada 158. 105 pp. 130 Hayes, A. H. 1975. The larger moths of the Galapagos islands (Geometroidea: Sphingoidea & Noctuoidea). Proc. Calif. Acad. Sci., ser. 4, 40:145-208. KIMBALL, C. P. 1965. The Lepidoptera of Florida. Arthropods of Florida and neighboring land areas. State of Florida Dept. of Agriculture, Gainesville, Florida. 361 pp. LAWESSON, J. E., H. ADSERSEN & P. BENTLEY. 1987. An updated and annotated check list of the vascular plants of the Galapagos islands. Rept. Bot. Inst. Univ. Aarhus. 16:1—74. MATTESON, J. H. 1933. America’s largest hawkmoth. Lep. News 1 (2):3-5. Journal of the Lepidopterists’ Society 53(3), 1999, 130-131 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY SCHREIBER, H. 1978. Dispersal centres of Sphingidae (Lepidoptera) in the Neotropical region. The Hague, Boston, Biogeographica 10:1—195. LAZARO ROQUE-ALBELO, Charles Darwin Research Station, Galapagos Islands, P. O. 17-01-3891, Quito-Ecuador. Received for publication 19 January 1999; revised and accepted 10 December 1999. AN ANTI-PREDATOR BEHAVIOR IN LARVAE OF LIBYTHEANA CARINENTA (NYMPHALIDAE, LIBYTHEINAE) Additional key words: ant-predation, Celtis, frass chains. Strategies of avoiding predators are well documented in larvae of Lepidoptera, and include a variety of morphological, chemical and behavioral traits (Malicky 1970, Brower 1984, Heads & Lawton 1985, Stamp & Casey 1993, Freitas & Oliveira 1992, 1996, Loeffler 1996). One rather unusual behavioral defense found in larvae of species of Charaxinae and Limenitidinae is the construction of frass chains (DeVries 1987, Freitas & Oliveira 1996). Frass chains are stick-like structures formed by fecula and silk where the larvae rest when not feeding, and provide an efficient refuge against predation by “walking” arthropod predators by isolating the larvae from the leaf blade when not feeding (Freitas & Oliveira 1996). In this note, a similar behavior is reported in larvae of the Libytheinae Libytheana carinenta (Cr.) at two sites in southeastern Brazil. Field observations were conducted in two fragments of deciduous forests in S40 Paulo state; the “Fazenda Trés Barras”, in Castilho, in February 1995 and in the “Reserva da Mata Santa Genebra”, Campinas, in March 1997, March-April 1998 and March-April 1999. The larvae were observed feeding on Celtis iguanae (Ulmaceae) in Castilho, and on C. spinosa in Campinas. Larvae of Libytheana carinenta were observed resting on the midvein of partially eaten leaves of Celtis spp. (Fig. 1). This behavior was observed in all instars, being present even in fully grown fifth instar larvae. Most of the observed eggs (n > 50) were laid individually on very young leaves (with five observations of eggs on spines on the shoot tips), and larvae start to eat alongside the central vein just after hatching, resulting in the formation of the stick-like structure on the expanding leaf. Larvae rest on the tip of this structure, returning to the leaf blade only for eating. No larvae of L. carinenta were observing feeding asymmetricaly on the leaf tip, or on only one side of the leaf. Although Celtis spinosa does not bear extrafloral nectaries, ants were frequently seen on branches and leaves of Celtis spp. in forest edges. In this habitat ants commonly associate with Homoptera, especially on shoot tips, and were observed preying on small moth caterpillars, suggesting that they could be effective predators of butterfly larvae as well. As also recorded by Freitas and Oliveira (1996) for another ant-butterfly interaction, these and other ants were never observed climbing onto L. carinenta’s stick-like Fig. 1. Third instar larva of Libytheana carinenta resting in the remaining central vein of a Celtis spinosa leaf. VOLUME 53, NUMBER 3 structures. Therefore the behavior of constructing and resting on these structures may provide a refuge against ant predation on the host plant, and in this way could be analogous to the behavior of resting on frass chains, observed in the Eurytelinae, Charaxinae and Heliconiinae. Considering Libytheinae as the most basal lineage of Nymphalidae (Harvey 1991), the results presented here could be important to the understanding of the anti-predator strategies present in other Nymphalidae. For example, in the Limenitidinae different degrees of complexity of similar refuges are known, ranging from the simple remaining midvein (as described in the present paper) to true frass chains, with several known variations (Morrell 1954, Fukuda et al. 1972). Data on larval biology of Libytheinae are scarce, and no defensive strategies have been reported, making this the first report of a defensive behavior in Libytheinae larvae. Similar simple structures are present in larvae of several moth species in different areas of tropical forests in Brazil, as well as in the pierid genus Dismorphia (Brown 1992), and in some Heliconiini, that construct or leave island-like structures on leaf edges (Benson et al. 1975) which were considered to be analogous to the frass-chains (see discussion in Freitas & Oliveira 1996). All these behaviors represent different ways for the larvae to maintain themselves isolated from the leaf blade; their occurrence in several different lineages of Lepidoptera shows that the fixation of this trait in many cases could be explained by predation pressure. ACKNOWLEDGEMENTS This study was conducted during the data gathered for a Ph.D. thesis on Nymphalidae systematics and evolution. K. S. Brown, G. Machado, P. S. Oliveira, I. Sazima, D. Bowers and two anonymous reviewers made useful criticisms and comments on the manuscript. K. S. Brown and F. Vanini helped in field work. C. E. G. Patto helped with the pictures. This research was supported by a fellowship from the Brazilian CNPq. LITERATURE CITED BENSON, W. W., K. S. BRowN JR. & L. E. GILBERT. 1975. Coevolution of plants and herbivores: passion flower butterflies. Evolution, 29:659-680. BROWER, L. P. 1984. Chemical defense in butterflies, pp. 109-134. In RB. I. Vane-Wright and P. R. Ackery (eds.), The biology of butterflies. Academic Press. 131 Brown, K. S. Jr. 1992. Borboletas da Serra do Japi: Diversidade, habitats, recursos alimentares e variacao temporal, pp. 142-187, 18 figs. In Morellato, L. P. C. (ed.), Hist6ria natural da Serra do Japi. Ecologia e preservacao de uma 4rea florestal no sudeste do Brasil. Campinas, Editora da Unicamp/Fapesp. DEVRIES, P. J. 1987. The butterflies of Costa Rica and their natural history, Princeton University Press, Princeton, New Jersey. Freimas, A. V. L. & P. S. OLIvEIRA. 1992. Biology and behavior of the Neotropical butterfly Eunica bechina (Nymphalidae) with special reference to larval defense against ant predation. J. Res. Lepid. 31:1-11. . 1996. Ants as selective agents on herbivore biology: effects on the behavior of a non-myrmecophilous butterfly. J. Anim. Ecol. 65:205—210. FukubaA, H., K. Kuso, T. Kuzuya, A. TAKAHASHI, B. TANAKA, M. WakabaYASHI & T. SHIROZU. 1972. Insects’ life in Japan. Hoikusha Publishing Co., Ltd. Harvey, D. J. 1991. Higher classification of the Nymphalidae (Appendix B), pp. 255-273. In H. F. Nijhout, The development and evolution of butterfly wing patterns. Smithsonian Press. Heaps, P. A., & J. H. Lawron. 1985. Bracken, ants and extrafloral nectaries. III. How insect herbivores avoid ant predation. Ecol. Entomol. 10:29-42. LOEFFLER, C. C. 1996. Caterpillar leaf folding as a defense against predation and dislodgement: staged encounters using Di- chomeris (Gelechiidae) larvae on goldenrods. J. Lepid. Soc. 50:245-260. Maticky, H. 1970. New aspects of the association between lycaenid larvae (Lycaenidae) and ants (Formicidae, Hymenoptera). J. Lepid. Soc. 24:109-202. MorRELL, R. 1954. Notes on the larval habits of a group of nymphalid butterflies. Malay. Nat. Journ. 8:157—-164. STAMP, N. & T. M. Casey. (EDs.). 1993. Caterpillars: ecological and evolutionary constraints on foraging. Chapman & Hall, London. ANDRE VICTOR LUCCI FREITAS, Museu de Historia Natural, Instituto de Biologia, Universidade Estadual de Campinas, CP 6109, 13083- 970, Campinas, SP, Brazil. Received for publication 19 January 1999; revised and accepted 10 December 1999. Date of Issue (Vol. 53, No. 3): 16 June 2000 4 ; —— valet pit adi ee ere ee ea Pe, “le ah a , ; tthe Sei THG ar 4 ’ 1 Ww i mr ih es Lig t ' r ' \ é - cates) * Lay olf ~ a Oe » Breeden Nis = She ite et: winet tbe we lee ree hearer Bal wip val! b gina aM? wae at, | ids nope Ws nw Oty tae aor ghee oy que direps hgh a 4 4 , i i ie ° ee ' é Syed alii nica Gh As on . epee yi Thea beeen Ane £ svleagben Rb y/ 2 ; + ke rete air EP seaman ten en yours they, ~; ? is J 5 + ‘ a” re, ee EDITORIAL STAFF OF THE JOURNAL M. Deane Bowers, Editor Entomology Section University of Colorado Museum Campus Box 218 University of Colorado Boulder, Colorado 80309, U.S.A. email: bowers@spot.colorado.edu Associate Editors: Gerarpo Lamas (Peru), Kenetm W. Pumir (USA), Rosert K. Rossins (USA), Feux A. H. Speruinc (USA), Davin L. Wacner (USA), Curister WikLuND (Sweden) NOTICE TO CONTRIBUTORS Contributions to the Journal may deal with any aspect of Lepidoptera study. 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Authors of Book Reviews and Obituaries are exempt from page charges. Correspondence: Address all matters relating to the Journal to the editor. PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A. CONTENTS Pegs Dreruatys: Division OF THIS SHOWY NEOTROPICAL GENUS, PLUS A NEW SPECIES AND THE IMMATURES AND FOO TWO SPECIES FROM COSTA RICAN DRY FOREST (Hesperupae: Pyrcinaz) John M. Burns and Daniel H. je NEW PRONOPHILINE BUTTERFLIES FROM THE VENEZUELAN TEPUYES (NYMPHALIDAE: SATYRINAE) Angel L. Viloria and Tomasz W. Pyrez eae Freie ae and May? BerenBauny 8 2) Seis Se wea eee 8 FrEpING PREFERENCE or Hezicontus Erato (Lev.: NYMPHALIDAR) IN RELATION TO LEAF AGE AND CONSE LARVAL PERFORMANCE Daniela Rodrigues and Gilson Rudinei Pires Moreira ae, GineraL Notes Field observations on larval diapause in the Florida viceroy, Limenitis archippus floridensis David Flaim and Austin P. Platt - a Notes on the genus Sthenopis (Hepialidae) in Alberta, Canada B. Christian Schmidt and David D. Lawrie... Two large tropical moths (Thysania zenobia (Noctuidae) and Cocytius antaeus (Sphingidae)) colonize the Galapag TERE Sa ORCL NOIDA a eee ci eee ee ee ee ea : An anti-predator behavior in larvae of Libytheana carinenta (Nymphalidae, Libytheinae) André Victor Lucci Freitas This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanance of Paper). (ee 54 | fies ENT- Volume 53 2000 JOURNAL of the LEPIDOPTERISTS’ SOCIETY Published quarterly by THE LEPIDOPTERISTS’ SOCIETY Publié par LA SOCIETE DES LEPIDOPTERISTES Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN Publicado por LA SOCIEDAD DE LOS LEPIDOPTEROLOGOS JUL 07 2000 22 June 2000 Number 4 ISSN 0024-0966 THE LEPIDOPTERISTS’ SOCIETY EXECUTIVE CoUNCIL Joun W. Brown, President Susan S. Borxin, Vice President Micuaet J. Situ, Immediate Past President Mirna M. Casacranbe, Vice President Manuet A. Baucazar-Lara, Vice President Davin C. Irtrner, Treasurer Ernest H. WiiiaMs, Secretary Members at large: Ronald L. Rutowski M. Deane Bowers George L. Balogh Felix A. H. Sperling Ron Leuschner Andrew V. Z. Brower Andrew D. Warren Michael Toliver Brian Scholtens EpiroriAL Boarpb Rosert K. Rossins (Chairman), Joun W. Brown (Member at large) M. Deane Bowers ( Journal) WiuiaM E. Mixier (Memoirs) Puiu J. Scuarrert (News) Honorary Lir—t MEMBERS OF THE SOCIETY Cuares L, Remincton (1966), E. G. Munroe (1973), Ian F. B. Common (1987), Joun G. Franciemont (1988), Lincotn P. Brower (1990), Douctas C. FEercuson (1990), Hon. Miriam RotuscuiLp (1991), CLraupEe Lemaire (1992), Freperick H. Rinpce (1997) The object of The Lepidopterists’ Society, which was formed in May 1947 and formally constituted in December 1950, is “to promote the science of lepidopterology in all its branches, . . . to issue a periodical and other publications on Lepidoptera, to facilitate the exchange of specimens and ideas by both the professional worker and the amateur in the field; to secure cooperation in all measures” directed towards these aims. Membership in the Society is open to all persons interested in the study of Lepidoptera. All members receive the Journal and the News of The Lep- idopterists’ Society. Prospective members should send to the Assistant Treasurer full dues for the current year, together with their full name, address, and special lepidopterological interests. In alternate years a list of members of the Society is issued, with addresses and special interests. 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The additional cost for members outside the U.S. is to cover mailing costs. Journal of The Lepidopterists’ Society (ISSN 0024-0966) is published quarterly by The Lepidopterists’ Society, %o Los Angeles County Museum of Natural History, 900 Exposition Blvd., Los Angeles, CA 90007-4057. Periodicals postage paid at Los Angeles, CA and at additional mailing offices. POSTMASTER: Send address changes to The Lepidopterists’ Society, o Natural History Museum, 900 Exposition Blvd., Los Angeles, CA 90007-4057. Cover illustration: Adult of the tiger swallowtail, Papilio glaucus (Papilionidae). Pen and ink drawing by Deane Bowers, University of Colorado. JOURNAL OF Toe LerprporvreERIstTs’ SOCIETY Volume 53 Journal of the Lepidopterists’ Society 53(4), 1999, 133-137 2000 Number 4 DISTRIBUTION AND HOSTPLANT RECORDS FOR EUPACKARDIA CALLETA FROM SOUTHEASTERN TEXAS WITH NOTES ON MANDIBULAR MORPHOLOGY OF ATTACINI (SATURNIIDAE) VALERIE A. PASSOA 10603 Brettridge Drive, Powell, Ohio 43065, USA AND STEVEN Passoa! USDA/APHIS/PPQ, The Ohio State University, Museum of Biological Diversity, 1315 Kinnear Road, Columbus, Ohio 43212, USA ABSTRACT. Eupackardia calleta (Westwood) is recorded on privet (Ligustrum) from Kingwood, Texas. This new county record represents a northeastern range extension of 100 miles and confirms privet as a hostplant under natural conditions. The role of the mandible in digestion of the hostplant is discussed. The last instar larval mandible of most Attacini either lacks teeth or has the cutting edge bluntly serrated with reduced teeth. In contrast, the distinctive mandible of E. calleta contains three large teeth and deep molar ridges, an autapomorphy for the genus. Well-developed mandibu- lar teeth are present on the last instar of a few unrelated Saturniinae and Ceratocampinae; therefore this character is homoplastic in Saturni- idae. A structurally complex mandible of the “sphingid-type” occurs in several saturniids such as Antheraea pernyi (Guér.- Mén.) (Saturniidae). This mandible type is illustrated. There are two ontogenetic patterns of mandibular development in Saturniidae. In one case, teeth are present in the first instar, then lost in later molts. The mandibular development of E. calleta represents an alternative scenario where teeth are present throughout the larval stage. Additional key words: Ligustrum, ontogenetic development, Sphingidae, Notodontidae. Eupackardia calleta (Westwood) is a member of the Attacini, usually considered the sister group to Roth- schildia (Ferguson 1972, Peigler 1989, Friedlander et al. 1998). This species is distributed from Central America (Honduras, Guatemala) north to Arizona and southern Texas (Victoria Co., Calhoun Co.) (Ferguson 1972, Wolfe 1995, Tuskes et al. 1996). The systematics and biology of E. calleta have been reviewed by several authors including Ferguson (1972), Weast (1989), Miller (1976), Lemaire (1978), and Tuskes et al. (1996). Weast (1989) suggested that systematic studies of E. calleta are needed. Tuskes et al. (1996) illustrated three larval forms of the United States populations, but did not study their mandibular morphology. In this paper we present biological data on E. calleta from southeastern Texas, then compare the mandibular ‘Address correspondence to this author morphology to other Attacini, with emphasis on re- lated nearctic taxa. Ontogenetic and phylogenetic ob- servations are also included. MATERIALS AND METHODS In order to survey saturniid mandibles, preserved larval specimens or exuviae associated with cocoons and emerged adults were taken from the authors’ col- lection and dissected. The mandible was either slide mounted in Euperal or preserved in alcohol. For analysis with a J.O.E.L. JSM 820 scanning electron mi- croscope, each sample was attached to a stub with car- bon paint and coated with a thin conductive layer of gold-palladium. Mandibular terminology is based on Godfrey (1972). Material examined: Eupackardia calleta: TEXAS (various localities, ex ova from female moths in lab cul- ture): 10 eggs, 10 first instars, 27 second to fourth in- 134 TABLE 1. Mandibular morphology of selected late instar Attacini larvae occurring in the Western Hemisphere. Locality data designate source of the study specimens. Species examined Source Cutting edge of mandible Callosamia angulifera (Wlk.) USA Teeth reduced, margin bluntly serrate Teeth reduced, margin bluntly serrate Teeth reduced, margin bluntly serrate Teeth present, well- Callosamia promethea (Drury) USA Callosamia securifera (Maas.) USA Eupackardia calleta (Ww.) Texas, Mexico developed Hyalophora cecropia (L.) USA Teeth absent Hyalophora columbia USA Teeth absent (S. I. Smith) Hyalophora euryalus (Bdv.) | USA Teeth reduced, margin bluntly serrate Teeth reduced, margin bluntly serrate Prob. Rothschildia cincta Prob. Baja Teeth reduced, margin (Tepp.) Cali- bluntly serrate fornia Rothschildia erycina (Shaw) USA Hyalophora gloveri (Stkz.) USA Teeth reduced, margin bluntly serrate Rothschildia forbesi Benj. Texas Teeth absent Rothschildia lebeau Honduras Teeth absent (Guér.-Mén.) Rothschildia orizaba (Ww.) Ecuador Teeth reduced, margin bluntly serrate Samia cynthia (Drury) USA Teeth reduced, margin bluntly serrate stars, 6 pupal cases (with larval exuviae), det. V. A. Pas- soa, mandible slides #581, 582 S. Passoa collection; Harris Co., Kingwood, XII-1995, on Ligustrum, V. A. Passoa, 6 mature larvae, det. V. A. and S. Passoa, mandible slide # 616 S. Passoa coll. MEXICO: various localities, collection data un- known, 2 first instars, 18 second to fourth instars, 3 pu- pal cases (with larval exuviae), det. V. A. Passoa; Mich. [Michoacan], Urupan (sic) [Uruapan], 2-X-1941, coll. [D. M.] Delong, det. V. A. and S. Passoa (1 last instar larva with mandibles slide mounted (The Ohio State University collection). One to ten mandibular pairs of each species (Table 1) were examined, depending on material available. RESULTS AND DISCUSSION During November 1995, the senior author collected seven third instar larvae of E. calleta on privet (Ligus- trum) from her backyard in Kingwood, Harris Co., Texas. This is a northeastern range extension of ap- proximately 100 miles (161 km) and a new county record. It remains to be determined if E. calleta is now established in Harris County. Tuskes et al. (1996) considered Leucophyllum frutescens (ceniza, purple sage) to be the main host- plant for E. calleta in Texas. Privet is normally treated JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY as an artificial laboratory host (Ferguson 1972,Tuskes et al. 1996). Ours is the second published record for E. calleta eating ornamental privet under natural condi- tions, supporting the suggestion of Weast (1989) that privet is an important hostplant of this species near ur- ban areas of Texas. In fact, E. calleta appears to be oligophagous on several genera of Oleaceae, including privet in Texas (present study) and Mexico (Peigler pers. comm.), Fraxinus gregeti A. Gray in western Texas (Peigler pers. comm.), and Forestiera angustifo- lia Torrey (Stone 1991). A mature larva of E . calleta on privet from Kingwood, Texas, is illustrated in Fig. 1. At maturity, the larva lacks long and dark scoli which are characteristic of earlier instars (Fig. 2). In spite of the availability of ceniza nearby, E. calleta was found only on privet in the senior author's backyard. Examination of E. calleta mandibles from Texas showed that these structures are atypical compared to other saturniids described in the literature. Bernays and Janzen (1988) characterized late instar saturniid mandibles as “short, with a broad base, and without obvious teeth”. This definition does not fit E. calleta which has large teeth in both the early (Fig. 3) and last instars (Figs. 4, 5, and 6). Except for E. calleta, members of the Attacini typi- cally lack well-developed mandibular teeth in the last instar or have the cutting edge of the mandible ser- rated with blunt lobes (Table 1). Mandibular teeth are also absent in mature larvae of two Indo-Australian Attacini, Attacus atlas (L.) (Heppner et al. 1989) and Coscinocera hercules (Miskin) (specimens in Passoa collection). Our data on saturniid mandibles agree with results of a survey of notodontid mandibles published by Godfrey et al. (1989) in several respects. Last instar mandibles of the Notodontidae usually lack teeth, except for a few apparently unrelated exceptions. This is also true in Saturniidae. Outside of the Attacini, mature larvae of Actias selene (Hbn.), A. luna (L.), Argema mittrei (Guér.- Mén.) (Saturni- inae) and Citheronia regalis (F.) (Ceratocampinae) (specimens in Passoa collection) have well-devel- oped mandibular teeth, so the presence of teeth on saturniid mandibles is homoplastic across several subfamilies. Large teeth also occur in the Antheraea genus com- plex, for example, Antheraea pernyi (Guér.- Mén.) and its purported synonym A. hartii (Moore), A. polyphe- mus (Cram.), and Opodiphthera eucalypti (Scott) (spms. in Passoa coll.). In addition, the mandible of A. pernyi is unusual because it matches the “sphingid- type” as defined by Bernays and Janzen (1988). Sphin- gid mandibles were characterized as being “ridged in a VOLUME 53, NUMBER 4 135 3 Fics. 4-6. Mandible of last instar Eupackardia calleta (West- wood), scale line = .10 mm in all figures. 4, oral surface, ventral view, Fics. 1-3. Eupackardia calleta (Westwood) from Texas. 1, late 70x. 5, teeth and deep molar ridges, ventral view, 150x. 6, teeth on instar larva on privet (lateral view). 2, mid-instar larva on privet (dor- cutting margin, dorsal view, 95x. sal view). 3, early instar mandible, oral surface, ventral view, pho- tographs under polarized light, 100x. 136 Fics. 7-9. Mandible of last instar Antheraea pernyi (Guér.- Mén.). 7, complex pattern of teeth on dorsal and oral surfaces, ven- tral view, 27x, scale line = 1 mm. 8, overlapping teeth of dorsal sur- face, ventral view, 33x, scale line = 1 mm. 9, tooth of dorsal surface, ventral view, 70x, scale line = .1 mm. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY variety of complex ways”. The mandible of A. pernyi (Figs. 7, 8, and 9) has a series of complex ridges on the dorsal surface that appear to form overlapping teeth, in addition to several smaller toothlike projections on the oral surface. The mandibles of E. calleta and A. pernyi suggest it is wiser to characterize saturniid mouthparts with ecological or physiological criteria (cutting versus grinding), instead of using a phylogenetic approach at the family level as did Bernays and Janzen (1988) in their study of saturniid and sphingid mandibles. There appear to be two ontogenetic pattems for sat- urniid mandibles. In one pattern, first instar larval mandibles have teeth; subsequently these teeth are lost in later molts. This pattern is typical for Costa Ri- can saturniid species in four subfamilies (Bernays & Janzen 1988). A second pattern is found in E. calleta where early instars have mandibular teeth (Fig. 3) that are retained throughout larval life (Figs. 4, 5, and 6). This dichotomy was also noted by Godfrey et al. (1989) in notodontid genera. Godfrey et al. (1989) stated that characterization of certain taxa by mandibular morphology was “problem- atical” because in some species the mandibular margin was intermediate between toothed and smooth. Inter- mediate conditions are common with Saturniidae, as shown in Table 1 by the term “margin bluntly serrate”. A bluntly serrated mandible has round conical projec- tions that resemble small teeth, or an irregular man- dibular margin with indentations. Callosamia is an ex- ample of the first condition where from 10-15 small teeth cover the cutting margin of the mandible. Irregular mandibular margins are found in Hyalo- phora and Rothschildia where it is difficult to discern teeth, but the mandibular margin is not straight. Dock- ter (1993) warmed that mandibular wear must be taken into account when describing mandibles. It is unclear whether bluntly serrated teeth in Saturniidae are a transition state between toothed and smooth man- dibles, as noted by Godfrey et al. (1989) in notodontids, or instead represent worn mandibular teeth as de- scribed by Dockter (1993) in Heterocampa and Soura- kov (1996) in larvae of satyrid butterflies. Examination of unworn mandibles on freshly molted larvae are needed to resolve this question. Thus, it may be prema- ture to characterize saturniid mandibles as usually toothless (Bernays & Janzen 1988) until more ontoge- netic studies are published. In summary, the mandibular shape of Macrolepi- doptera depends on several factors including head size and food toughness (Bernays 1986), food particle size and chemistry (Bernays & Janzen 1988), the need to pierce the chorion of the egg at eclosion, and a need to VOLUME 53, NUMBER 4 seal the oral cavity during ingestion (Godfrey et al. 1989). Unlike notodontids which use toothed man- dibles to pierce the leaf epidermis (Dockter 1993), E. calleta does not skeletonize leaves during any instar (V. A. Passoa pers. obs.). Therefore, it seems likely that a toothed mandible is required for other reasons. Bernays and Janzen (1988) noted that larvae which feed on toxic plants tend to be associated with mandibles that contain teeth. They (Bernays & Janzen 1988) suggested that complete mastication of the leaf tissue allows more complete digestion, a strategy not possible with hostplants containing tannins that inhibit digestive efficiency. However, the digestive physiology of E. calleta may be more complicated than other Sat- urniidae. Caterpillars of E. calleta secrete a pungent liquid containing phenolic compounds and biogenic amines from their scoli when disturbed, but it is un- known if plant toxins are sequestered for this secretion (Deml & Dettner 1993). Until the metabolism and de- fensive chemistry of E. calleta are clarified, it may be premature to assume mandibular teeth are an aid to coping with plant toxins. Nevertheless, E. calleta lar- vae do utilize toxic plants (Weast 1989). Besides the need to understand mandibular mor- phology in an ecological context, it should be noted that a survey of saturniid mandibles would aid identifi- cation of the immatures. Minet (1994:70) mentioned mandibular secondary setae as an apomorphy of the Lasiocampidae, although these setae are also indepen- dently evolved in other macrolepidopteran families. Mandibular secondary setae are present (Agapema) or absent (E. calleta) in Saturniidae, thus this character has potential for identification purposes. Mandibular teeth associated with the cast larval exuvium can be used to separate cocoons of E. calleta (teeth present) and Rothschildia (teeth absent). Both genera are easily confused due to their similar pupal morphology. A fi- nal example is A. luna and A. polyphemus. Although these genera are often confused as larvae (Ferguson 1972:207), their mandibles are completely different, be- ing either simple and toothed (A. luna) or complex with ridges (A. polyphemus). More descriptive studies of sat- urniid mandibles will no doubt yield further examples. ACKNOWLEDGMENTS Reviews by G. L. Godfrey (Haskell Indian Nations University, Lawrence, Kansas), R. S. Peigler (University of the Incamate Word, San Antonio, Texas), J. P. Tuttle (Tucson, Arizona), and R. D. Weast (Johnston, Iowa) significantly improved the manuscript. W. D. Ol- hausen (Jesse Jones Park and Nature Center, Humble, Texas) pro- vided keen observations concerning E. calleta in Harris County, 137 Texas. We are grateful to K. S. Johnson (Ohio University, Athens, Ohio) for Callosamia larvae and pupae, K. L. Wolfe (Escondido, California) for a R. erycina larval exuvium, C. A. Conlan (San Diego, California) for E. calleta pupal cases, M. D. Schmidt (Springboro, Ohio) for a H. euryalus pupa, and R. S. Peigler for use of his Oleaceae hostplant records. At the The Ohio State University (Columbus, Ohio), we thank J. C. Mitchell for the scanning electron microscope photographs, D. J. Horn and G. D. Keeney for greenhouse space used to grow host- plants, and N. F. Johnson for use of the insect collection. Voucher specimens of E. calleta and its hostplant are deposited in The Ohio State University’s Insect Collection and Herbarium. LITERATURE CITED BErRNAys, E. A. 1986. Diet-induced head allometry among foliage- chewing insects and its importance for graminivores. Science 231:495—496. Bernays, E. A. & D. H. JANZEN. 1988. Saturniid and sphingid caterpillars: two ways to eat leaves. Ecology 69:1153-1160. DEML, R. & K. DETTNER. 1993. Biogenic amines and phenolics characterize the defensive secretion of saturniid caterpillars (Lepidoptera: Saturniidae): a comparative study. J. Comp. Phys. B 163:123-132. DocxTER, D. E. 1993. Developmental changes and wear of larval mandibles in Heterocampa guttivitta and H. subrotata (Notodontidae). J. Lep. Soc. 47:32-48. FERGUSON, D.C. 1972. In Dominick, R. B. et al. (eds.), The moths of America north of Mexico. Bombycoidea. Saturniidae. Fasci- cle 20.2. 275 pp., 22 colored pls. FRIEDLANDER, T. P., K. R. Horst, J. C. REGIER, C. MITTER, R. S. PEIGLER & Q. Q. FANG. 1998. Two nuclear genes yield concor- dant relationships within Attacini (Lepidoptera: Saturniidae). Mol. Phyl. Evol. 9:131-140. GopFkEy, G. L. 1972. A review and reclassification of larvae of the subfamily Hadeninae (Lepidoptera, Noctuidae) of America north of Mexico. U. S. D. A. Tech. Bull. No. 1450. GopFRrEy, G. L., J. S. MILLER & D. J. CARTER. 1989. Two mouth- part modifications in larval Notodontidae (Lepidoptera): their taxonomic distributions and putative functions. J. New York En- tomol. Soc. 97:455-470. HEPPNER, J. B., H. Y. WANG & Y. C. SEN. 1989. Larval morphology of Taiwan Saturnilidae [sic]: Attacus atlas (Linnaeus). J. Taiwan Museum 42:89-97. LEMAIRE, C. 1978. Les Attacidae Americains. 1. Attacinae. Neuilly-sur-Seine, France. Published by the author. 238 pp. MILLER, T. A. 1976. Observations of Eupackardia calleta in south- em Texas (Saturniidae). J. Lep. Soe. 30:127-130. MinET, J. 1994. The Bombycoidea: phylogeny and higher classifi- cation (Lepidoptera: Glossata). Entomol. Scand. 25:63-88. PEIGLER, R. S. 1989. A revision of the Indo-Australian genus Atta- cus. The Lepidoptera Research Foundation, Inc. Beverly Hills, California. 167 pp. Sourakoy, A. 1996. Notes on the genus Calisto, with descriptions of the immature stages (part 1) (Lepidoptera: Nymphalidae: Satyrinae). Trop. Lepid. 7:91—112. TuSKES, P. M., J. P. TurrLe, & M. M. Cou.ins. 1996. The wild silk moths of North America. Cornell University Press, Ithaca, New York. 250 pp. WeasT, R. D. 1989. Saturniidae. Ecological and behavioral obser- vations of select Attacini. Self-published. Johnston, Iowa. 53 pp. Wo LrFE, K. L. 1995. Photo Contest-1995. Eupackardia calleta. Trop. Lepid. 6: iv (fig. 8). Received for publication 11 May 1997; revised and accepted 10 De- cember 1999. Journal of the Lepidopterists’ Society 53(4), 1999, 138-141 METHOD OF HANDLING AFFECTS POST-CAPTURE ENCOUNTER PROBABILITIES IN MALE HYPOLIMNAS BOLINA (L.) (NYMPHALIDAE) DARRELL J. KEMP School of Tropical Biology, James Cook University, P.O. Box 6811 Cairns, Queensland, 4870, AUSTRALIA AND MYRON P. ZALUCKI Department of Zoology and Entomology, The University of Queensland, St Lucia, Queensland, 4072, AUSTRALIA ABSTRACT. Mark-recapture studies of butterfly populations are often plagued by low recapture rates, which make population estimation problematic. One reason for low recaptures is that the handling process of capture, marking and release contributes to low and unequal catch- ability of marked individuals. Here we report the results of an experiment conducted to evaluate the hypothesis that cooling individuals prior to release minimizes handling effects. The post-capture difference in site fidelity of territorial male Hypolimnas bolina (L.) (Lepidoptera: Nymphalidae) was compared among three groups: (1) males handled normally, (2) males chilled prior to release, and (3) uncaught controls. Un- chilled males showed significantly reduced site fidelity compared to both control and chilled butterflies. Furthermore, chilled butterflies re- sumed activity after capture in a manner similar to uncaught controls. These results indicate that chilling has the potential to minimize the ad- verse effects of handling on subsequent butterfly catchability. Since ‘equal catchability of caught and uncaught individuals is a critical assumption of mark-release-recapture programs, this method has the potential to greatly increase the accuracy of subsequent population esti- mates. On this basis, in population studies on butterflies, the precise method of handling may prove a more meaningful consideration than the question of whether or not to handle. Additional key words: _catchability, mark-release-recapture, censusing, population estimation. Mark-recapture methods represent a powerful tool In recent times, many workers have taken to cooling for the estimation of animal population parameters, individuals before their release (e.g., Bull et al. 1985, especially those of invertebrates (Seber 1973, South- Zalucki & Kitching 1985, Suzuki & Zalucki 1986, Ru- wood 1978, Begon 1979). These methodologies have towski 1992, Zalucki 1993, Wickman & Jansson 1997). often been applied to the study of butterfly popula- This technique stems from the idea that increased im- tions; however, on many occasions, such programs are mobilization of individuals immediately prior to their plagued by low recapture rates, sometimes in the or- release helps to reduce their subsequent degree of der of 1 to 10% (Brussard & Ehrlich 1970, Urquhart et ‘panic’ (Wickman & Jansson 1997). In this way, indi- al. 1970, Cook et al. 1971, Urquhart & Urquhart 1976, viduals are supposedly more likely to remain in the im- Watt et al. 1977, Cullenward et al. 1979). This makes the mediate study area and behave normally, rather than estimation of population size and related measures prob- disperse and avoid further capture (as found by Leder- lematic (Rosenberg et al. 1995), and researchers must house 1982). The use of this technique, however, is often resort to less powerful methods, such as transect- based on mostly anecdotal information, and little, if based surveys to census their populations (e.g., Pollard any, published research has been conducted to evalu- 1977, Eberhardt 1978, Thomas 1983, Dent 1997). ate its merit. One of the many potential causes of low recapture The aim of this paper is to evaluate the hypothesis rates in the Lepidoptera is that the actual process of that cooling individuals prior to release may increase capture and handling may affect the behaviour of their subsequent catchability relative to individuals marked individuals, reducing their chance of being re- handled in the normal manner. This hypothesis is in- captured (Gall 1984). This effect would contribute to vestigated using territorial mate-locating males of the low catchability in marked individuals, and lead to sig- nymphalid species Hypolimnas bolina (L.). nificant bias in subsequent population estimates (Gall 1984, Rosenberg et al. 1995). Although crucial to the MATERIALS AND METHODS application of mark-recapture methods, few authors Male H. bolina in resident territories were censused attempt to validate the assumption of ‘equal catchabil- along a 1020 m transect (Rutowski 1992) located on ity (Gall 1984). Several attempts to assay the effects of campus at James Cook University in Townsville, Aus- capture in butterflies have indicated that the capture tralia (19°15’S, 146°45’E). Sampling was conducted in and handling process may have strongly negative ef- two rounds in 1998; from 9-12 February and 16-19 fects on subsequent catchability (Singer & Wedlake April, and once in 1999; from 22-25 February 1999. 1981, Lederhouse 1982, Morton 1984). On each sample round, transects were censused VOLUME 53, NUMBER 4 So ° NN Difference in count probability oS fo) Bh NN) o rer) Control Chilled Unchilled Fic. 1. The average (+ SE) difference in the probability of en- countering a male H. bolina between pre- and post-capture cen- suses, as a function of which treatment group he was assigned to. Control males were not caught, chilled males were caught and cooled before release, and unchilled males were caught but not chilled. hourly for the first two days to identify individuals of high territory site fidelity (for census techniques see Pollard 1977). Notes on size and wing wear were taken to allow reliable identification of these individuals in subsequent censuses (Rutowski 1992). At 1000 h on the third day (termed the marking round), each resi- dent was randomly assigned to one of three treatment groups. Unchilled’ males were captured at their site, marked with an identification number on both hind- wing undersides, and released immediately (total han- dling time of 40-50 seconds). ‘Chilled’ males were similarly captured and marked, but were held for a further 210 seconds in a paper envelope placed on a block of ice wrapped in newspaper (Wickman & Jans- son 1997), then released directly back to their perch. Notes were taken on the immediate behavior of indi- viduals in these two treatments upon release. Butter- flies of the third group, designated as control, were not caught. Hourly transects were then conducted for the remainder of the third day and the next day to deter- mine the behavior of males in all three groups. Cen- suses were conducted no sooner than 0900 h and no later than 1500 h, and only in the presence of sunshine before and during the entire transect. This was done to ensure that individuals were censused only at times of maximum site fidelity. During the course of each tran- sect, individual males were deemed re-counted if they were within 20 meters of their designated territory, and actively engaged in defense of that site. No effort was made to search for individuals that were not im- mediately obvious at their sites following marking, hence the observation technique was kept similar be- fore and after marking. An index of site fidelity was calculated for each male by dividing his total counts by his total number of 139 count opportunities (i.e. the number of times his terri- tory was passed during transect censuses). Since the ratio of before: after-marking counts in each treatment group was homogenous between sampling rounds (Chi-squared homogeneity test on before- and after- counts across the three rounds: x”, < 2.79, p > 0.24 for each group), data were pooled across all rounds. Ini- tially, pre- and post-marking round count probabilities were compared for control group butterflies only. This was done to check that the activity of untreated but- terflies was homogenous and that no other factor, for instance weather, affected the probability of counting butterflies between censuses. Mean differences (pre- and post marking round) in count probabilities between butterflies of each treatment group were then com- pared using a one-way analysis of variance (ANOVA). Three comparisons were planned prior to analysis; these were (1) between chilled and unchilled butter- flies, (2) between chilled and control group butterflies, and (3) between unchilled and control group butter- flies. These contrasts were evaluated using least signifi- cant difference (LSD) test for planned, non-orthogonal comparisons (Sokal & Rohlf 1995). Prior to conducting analyses, data sets were transformed using the angular transformation, and Kolmogorov-Smirnov goodness of fit tests were used to confirm that the transformed data were normally distributed (Kolmogorov-Smirnov d < 0.24, p > 0.20 for all groups). Levene tests were used to check homoscedasticity among ANOVA groups; these were non-significant in all cases (p > 0.175), which confirmed that data were homoscedastic. Two- tailed probabilities were used, and sample means throughout this paper are given + | standard error. RESULTS A total of 30 primary territory residents were identi- fied during the three sampling occasions, and these were randomly distributed amongst treatments as fol- lows: 11 chilled, 11 unchilled, and eight control. Of the total 226 count opportunities before the marking round, 191 counts were registered, and primary resi- dents exhibited a mean site fidelity of 0.87 + 0.02 (each being present on approximately 87% of all count opportunities). Mean pre-marking site fidelity did not differ significantly among the three butterfly groups (control, chilled, unchilled; ANOVA on angular-trans- formed data, F,,. = 0.15, p = 0.86). In addition, the mean site fidelity of control group butterflies was not significantly different between pre- and post-marking censuses (paired t-test on angular-transformed data, t = 0.63, df = 7, p = 0.55). Post-marking censuses revealed that four marked individuals (one chilled and three unchilled) had de- 140 serted their territories. All other males were counted at least once defending their pre-designated territory area. The mean difference between pre- and post- marking round fidelity varied significantly among treatment groups (ANOVA on angular-transformed data, F, ,, = 3.97, p < 0.05; Fig. 1). Fidelity was re- duced to a varying degree in all groups following the marking round. The reduction in site fidelity of un- chilled butterflies was significantly greater than that of either control or chilled group butterflies (LSD test for planned comparisons, p < 0.05). The significant differ- ence between the two captured treatment groups demonstrated an effect due to the method of handling individual males before their release. Furthermore, the change in fidelity among chilled butterflies did not differ significantly from that shown by control group butterflies (LSD test, p = 0.85). Hence, relative to un- caught controls, the process of capture and marking had no appreciable effect on the subsequent territorial fidelity of butterflies that were chilled prior to their release. Captured and marked butterflies in each treatment group showed clear behavioral differences following their release. While all chilled males remained perched on the substrate where they were placed, non-chilled males either resumed active territory defense (2 males), roosted on the underside of shaded foliage near the site (2 males), or flew quickly out of the area (7 males). Hence, while all chilled butterflies remained in the vicinity of their territory (i.e., less than 20 meters away) in the first instance, only 4 (36.4%) of the 11 non-chilled butterflies did so. DISCUSSION Although based on relatively small sample sizes, the results of this experiment clearly support the hypothe- sis that cooling may reduce capture effects and in- crease post-handling catchability of territorial male H. bolina. Not only did the process of chilling affect the probability of resighting captured males, but chilled males also resumed activity in a manner similar to that of butterflies that were never caught. This result is sig- nificant, since it shows that chilling may not simply re- duce or ‘manage’ the adverse effect of handling, but that this process has the potential to actually nullify short-term effects of capture. To our knowledge, this has not been demonstrated previously for any butter- fly species. These results contrast with the generally negative effects of handling of butterflies obtained in previous studies, for example Singer and Wedlake (1981), Leder- house (1982), and Morton (1984). In these studies, however, no reference is given to any method of chill- JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY ing butterflies prior to their release, and it must be as- sumed that butterflies were simply released immedi- ately following marking. The experiment conducted here on H. bolina corroborates the finding that butter- flies handled in this manner may be less likely to be re- captured, but that this adverse handling effect may be mediated by a chilling treatment prior to release. One concer with capturing and marking butterflies is that individuals will suffer from an increased level of predation owing to the wing mark and subsequent loss of crypsis or aposematism (Gall 1984, Reynolds et al. 1997). If present, such an effect is often difficult to dis- tinguish from any potential effects arising from cap- ture and handling. Fortunately, in this study, the con- trasting results between treatment groups allowed the separation of specific marking effects. Since both treatment groups were given similar marks, chilled males provided an adequate control for the effects of wing marks in the unchilled group. That only the un- chilled group of butterflies were less likely to be re- counted than controls clearly suggests that, at least in the short term, these were affected by the method of handling and not by the marks placed upon their wings. A similar conclusion was reached by Morton (1984), who found that colored wing marks did not sig- nificantly affect recapture of the highly cryptic wood- land butterfly Melanargia galathea (L.). The limitation with the current experiment, however, is that post- marking site fidelity was only measured for a period of two days. On this basis, the presence of a longer term effect due to wing marks in H. bolina cannot be com- pletely discounted. The question of why captured butterflies may be less likely to be resighted is not specifically addressed by this study, but some insight is afforded by casual be- havioral observations made on released individuals. Clear differences in behavior were evident between treatment groups, and this resulted in more unchilled butterflies leaving their immediate area of capture in the short term. This difference alone may be responsi- ble for differences in encounter probabilities of un- chilled and chilled individuals, especially in the case of a site-tenacious butterfly such as male H. bolina (Rutowski 1992). In unchilled H. bolina, the decision to leave the territory is more likely to be made as a consequence of immediate ‘panic’ upon release. This behavior appears similar to the defensive or evasive re- sponse of butterflies to failed predatory attacks, or failed capture attempts. Chilled butterflies, however, remain in their territory area for quite some time whilst warming up, and therefore any active decision to abandon the site must be made at some later stage. On average, the difference between groups after VOLUME 53, NUMBER 4 marking is that unchilled butterflies must decide whether to return to their former territory, and the chilled butterflies must decide whether to leave. If both these decisions are sufficiently unlikely, then dis- crepancies will exist between the recounts of chilled and unchilled group individuals, leading to the results observed in this study. In theory (depending on specific post-capture be- havioural responses), this principle may hold for a wide variety of butterfly species being sampled in a wide variety of circumstances. Indeed, both Leder- house (1982) and Singer and Wedlake (1981) suggested that initial dispersal or displacement from the site of capture may have accounted for their observed cap- ture effects in butterflies handled without chilling. The advantages of chilling may therefore lie in the preven- tion of early dispersal immediately following capture, as implied by Wickman and Jansson (1997). Since ‘equal catchability of caught and uncaught individuals is a critical assumption of mark-release-recapture pro- grams, this method has the potential to greatly in- crease the accuracy of subsequent population esti- mates. On this basis, in studies that intend to employ mark-release-recapture techniques, the actual method of handling may prove a more meaningful considera- tion than the question of whether or not to handle. ACKNOWLEDGMENTS Thanks are due to Dr. C. J. Hill and Prof. F .S. Chew for first sug- gesting the potential benefits of cooling butterflies before release to D. J. K. Prof. R. E. Jones kindly offered suggestions and criticized an earlier manuscript. This research was supported by an Australian Postgraduate Research Award to D. J. K. LITERATURE CITED BEcon, M. 1979. Investigating animal abundance: capture-recap- ture for biologists. University Park Press, Baltimore. BRUSSARD, P. F. & P. R. EHRLICH. 1970. 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Museum of Natural History Con- tributions to Science, Los Angeles. ZALUCKI, M. P. & R. L. KitcHinc. 1985. The dynamics of adult Danaus plexippus L. around patches of its host plant Asclepias spp. J. Lepid. Soc. 38:209-19. Received for publication 5 April 1999; revised and accepted 16 December 1999. Journal of the Lepidopterists’ Society 53(4), 1999, 142-152 EARLY STAGES OF CALIGO ILLIONEUS AND C. IDOMENEUS (NYMPHALIDAE, BRASSOLINAE) FROM PANAMA, WITH REMARKS ON LARVAL FOOD PLANTS FOR THE SUBFAMILY. CARLA M. PENZ Department of Invertebrate Zoology, Milwaukee Public Museum, 800 West Wells Street, Milwaukee, Wisconsin 53233, USA, and Curso de Pés-Graduagao em Biociéncias, Pontificia Universidade Catélica do Rio Grande do Sul, Ay. Ipiranga 6681, Porto Alegre, RS 90619-900, BRAZIL ANNETTE AIELLO Smithsonian Tropical Research Institute, Apdo. 2072, Balboa, Ancon, REPUBLIC OF PANAMA AND ROBERT B. SRYGLEY Smithsonian Tropical Research Institute, Apdo. 2072, Balboa, Ancon, REPUBLIC OF PANAMA, and Department of Zoology, University of Oxford, South Parks Road, Oxford, OX] 3PS, ENGLAND ABSTRACT. Here we describe the complete life cycle of Caligo illioneus oberon Butler and the mature larva and pupa of C. idomeneus (L.). The mature larva and pupa of each species are illustrated. We also provide a compilation of host records for members of the Brassolinae and briefly address the interaction between these butterflies and their larval food plants. Additional key words: The nymphalid subfamily Brassolinae includes Neotropical species of large body size and crepuscular habits, both as caterpillars and adults (Harrison 1963, Casagrande 1979, DeVries 1987, Srygley 1994). Larvae generally consume large quantities of plant material to reach maturity, a behavior that may be related as much to the low nutrient content of their larval food plants (Auerbach & Strong 1981) as to their large body size (e.g., 15 g for a living, mature larva of Caligo memnon (Felder), DeVries 1983). Several species of brassolines lay eggs in clusters, and their larval feeding activity may produce remarkable damage to their food plants. For ex- ample, two Caligo memnon females were reported to lay, jointly, 165 eggs on banana plants in approximately three weeks in an outdoor enclosure (Young & Muyshondt 1985). Caligo caterpillars were reported to cause severe damage to banana plantations (Malo & Willis 1961, Har- rison 1962, 1963, 1964), and larvae of Brassolis isthmia Bates are known to defoliate coconut palms (Dunn 1917; R. B. Srygley, C. M. Penz pers. obs.). Apart from studies of population control (Malo & Willis 1961, Harrison 1962, 1964), few investigations have focused on the early stage biology of Caligo (see Young & Muyshondt 1985, and references therein). Here we describe the complete life cycle of C. il- lioneus oberon Butler, describe a mature larva of C. idomeneus (L.), and review larval food plant records for 39 brassoline species. Central America, host records, monocotyledonous plants, larval food plants. METHODS Between 25 May and 31 December, 1994 we searched for ovipositing female butterflies along Pipeline Road, Soberania National Park, Panama, mo- tivated by a study on Caligo mating behavior (Srygley & Penz 1999). The study area was a mosaic of old sec- ondary and primary forests and pasture grasses with ap- proximately 2.2 m annual precipitation (Ridgely 1976) and a wet season extending from late April to mid De- cember. Our observations showed that C. illioneus oberon oviposit mostly at dusk (approx. 1700-1900 h) and only occasionally at dawn (0530-0630 h), and we therefore concentrated our observations in the twilight hours. Field collected early stages were reared in plastic containers at ambient temperature (25—29°C). Wild female Caligo illioneus oberon were captured in two traps at the edge of the forest along Pipeline Road. To induce oviposition, captured females were re- leased into an outdoor insectary (3 x 3 x 3 m) inside of which grew Musa sapientum L. and Heliconia latis- patha Benth.(Musaceae), Calathea latifolia (Link) K. (Marantaceae), and three species of unidentified palms (Arecaceae). Females were supplied also with fresh cut leaves of Saccharum spontaneum L. (Poaceae) and Cyrtostachys sp. (Arecaceae), both exotics on which we had observed oviposition by female Caligo illioneus and Opsiphanes sp. respectively (R. B. Srygley pers. obs.). VOLUME 53, NUMBER 4 Preserved larvae and pupal skins of C. illioneus oberon are in the collection of the Milwaukee Public Museum. The head capsule and pupal skin of C. idomeneus are currently in the Smithsonian Tropical Research Institute, to be relocated to the National Museum of Natural History in the future. RESULTS Caligo illioneus oberon Butler Oviposition behavior and food plants. In the field, females laid clusters of 9-13 eggs (n = 4 clusters) in a row along the midvein on the underside of medium-aged to old leaf blades of Saccharum sponta- neum, an introduced Asian grass that invaded natural grasslands of Panama during the 1970's. Following its introduction, S. spontaneum gradually replaced the pasture grasses Hyparrhenia rufa (Nees) Stapf and Panicum maximum Jacq. (both introduced from Africa) on Pipeline Road (N. Smith pers. comm.). The native larval food plant for C. illioneus is not known in Panama, and our captive females did not oviposit on any of the plants available in the insectary. Egg (developmental time = 6 days, n = 13). White, spherical, approximately 1.5 mm wide, adorned with vertical ribs; description refers to a cluster of 13 eggs laid 27 August 1994. First instar (duration = 8 days, n = 4). Head: brown with simple black setae; two dark brown verti- cal stripes flank epicranial and frontal sutures from apex of head, terminating at approximately halfway the length of the front. Body: translucent green; broad, lemon-yellow middorsal stripe bordered by an irregu- lar reddish-brown stripe that is prominent on the tho- rax and divided by a thin white discontinuous midline stripe that is more prominent on the thorax than on the abdomen; two thin, lateral, lemon-yellow longitu- dinal stripes; thoracic and abdominal legs grayish- white; ventral side grayish-white; caudae held sepa- rated, reddish to dark brown, each with a black sub-terminal seta arising at one-third to one-half the length of the caudae, and a terminal seta which is white at base and black at tip. The larvae molted syn- chronously. Second instar (duration = 5 days, n = 4). Head: dark brown anteriorly, lateral and post-genal regions translucent white; three pairs of scoli: dorsal scoli light brown (approximately half the height of the head), subdorsal scoli whitish (two-thirds the height of the dorsal scoli), lateral scoli whitish (approximately one- third the height of the subdorsal scoli); front dark brown; two thin whitish lines arise from base of dorsal scoli, converge toward and flank epicranial suture, ter- 143 minating at upper end of front; frontal suture whitish. Body: predominantly green; brown middorsal stripe divided by discontinuous white midline stripe; thoracic and abdominal legs grayish white; ventral side grayish white; caudae held separated, pink with black tips and numerous short white setae. The larvae molted syn- chronously. Third instar (duration = 5-6 days, n = 4). Head: as in second instar. Body: as in second instar, except for a broad, red lateral line divided by a thin, white spiracu- lar stripe; single dark brown triangular middorsal pro- jection at posterior end of abdominal segment A3. Shed caudae were not eaten after molt to fourth instar. Larvae molted asynchronously. Fourth instar (duration = 6—7 days, n = 4). Head: patterned in creamy-white and dark brown; dorsal scoli light brown anteriorly and reddish-brown poste- riorly at the base (approximately same height as the head); subdorsal scoli creamy-white (approximately two-thirds the length of dorsal scoli); lateral scoli creamy-white (approximately one-half the length of the subdorsal scoli); one pair of creamy-white tuber- cles below lateral scoli; epicranial suture darkened; front creamy-white with vertical, medial brown stripe; two brown stripes arise from the base of dorsal scoli converge toward and flank epicranial and frontal su- tures, terminating halfway down the length of the front; adfrontal region dark brown above stemmatal region; post-genae reddish-brown; base of head red- dish-brown from occiput to mandibles; mandibles creamy-white, darkened at the cutting edge. Body: color varied from light mustard to greenish; thoracic segments Tl and T2 with middorsal white midline stripe, flanked by reddish stripes; remaining segments with thin grayish middorsal stripe; large reddish- brown, triangular middorsal projection located at poste- rior end of abdominal segment A3; very small middorsal projection at posterior end of A5; supra-spiracular white stripe along the entire length of the body; white sub- spiracular stripe on a continuous longitudinal swelling: thoracic and abdominal legs reddish; ventral side red; caudae pale pinkish-brown patterned with reddish- brown dorso-laterally, where pattern develops into thin broken lines. The larvae molted asynchronously and three out of four aggregated at rest. Fifth instar (duration = 6-7 days, n = 4, Fig. 1a). Head: as in fourth instar, except for a brown stripe arising from dorsal scoli that flanks the epicranial and frontal sutures, and terminates above antennal socket: head densely covered with short creamy-white setae. Body: predominantly beige; dark brown middorsal stripe runs along entire length of body; on segments Tl, T2, and anterior end of T3, the dark brown mid- 144 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fic. 1. Caligo illioneus oberon, photographs by R. Srygley; (a) fifth instar (October, 1994); (b) pupa (24 October, 1994). Fic.2. Caligo idomeneus (Aiello Lot 81-77), photographs by A. Aiello; (a) final instar (2 December, 1981); (b) final instar, head (2 December, 1981); (c) final instar, body detail (2 December, 1981); (d) pupa (23 December, 1981). dorsal stripe is divided by a thin creamy-white stripe; triangular middorsal projections same as in fourth in- star; body patterned with brown from dorsal midline towards the sides, where pattern develops into thin, broken, longitudinal brown lines; dark brown supra- spiracular line bordered by creamy-white lines; creamy- white subspiracular stripe on a continuous longitudinal swelling; ventral side pink; caudae held separated, light brown at base, becoming dark brown at tip, approxi- mately two-thirds the length of the head. Larvae molted asynchronously and did not aggregate at rest. Sixth instar (duration = 14-16 days, n = 3). Head: as in fifth instar, except for four pairs of scoli; addi- tional stripe from dorsal scoli terminating above stem- matal region; dorsal scoli dark brown with white tips and basal creamy-white spots posteriorly; subdorsal and lateral scoli posteriorly brown at base; sub-lateral scoli dark brown posteriorly (one-half of the length of VOLUME 53, NUMBER 4 the lateral scoli). Body: as in fifth instar, except for small dark brown triangular middorsal projection at posterior end of abdominal segment A2; large triangu- lar middorsal projection at posterior end of A3; and small triangular middorsal projections at posterior ends of A4 and A5; ventral side brown; caudae held separated, slightly longer than head height. Larval development and behavior. Egg develop- ment took 6 days (n = 13) and larval development (from hatching to pupation) took 4449 days (n = 3). First through fourth instar larvae fed gregariously and generally rested together on the leaf blade. Fifth and sixth instar larvae rested away from each other on the stem of the plant. Late instar larvae found in the field were solitary (R. B. Srygley pers. obs.). Pupa (duration = 13-15 days, n = 3, Fig. 1b). Beige with fine, dark brown cryptic markings, giving the gen- eral appearance of a dried, curled and sun-bleached leaf; head with transverse keel at apex; long, black se- tae located immediately above and on the surface of the eyes; antennae with a black longitudinal stripe, and a black transverse line marking each antennal seg- ment; thoracic segment T2 with a prominent keel along dorsal midline, more developed in female than in male pupae; posterior edge of wing pad forming a crest; prominent hump at the base of wing pad; wing surface with two small silver spots located near base of wing; abdomen with conspicuous long black setae along dorsal midline; abdominal segments A5—10 with dark brown lateral line, A4—10 with brown ventral line; A6 humped; brown middorsal stripe arising at head and terminating at cremaster; abdominal segments with transverse oblique markings that resemble leaf venation. Pupal mass: 2.3 g (n = 1, male). Caligo idomeneus (L.) A wandering final instar larva of C. idomeneus was found off the food plant by R. Kimsey at Fort Clayton (Canal Area, Panama) on 1 December, 1981 and reared to adult (Aiello Lot 81-77). The larva had about 15 white fly eggs cemented to the underside of the thorax and head. The eggs were removed with forceps and preserved in 80% ethanol. The oviposition behay- ior and larval food plants of C. idomeneus are un- known, but the captive mature larva readily accepted Heliconia latispatha Benth. (Musaceae) and Calathea latifolia (Link.) K. (Marantaceae) which it ate for 20 days prior to pupation. Final instar (n = 1, Figs. 2a—c). Head: beige with brown stripes; three pairs of beige scoli: largest scoli dorsal, clothed in long setae, enlarged towards the pointed apex and abruptly curved outward toward the sides of the head; subdorsal scoli about two thirds the 145 length of the dorsal scoli, clothed in long setae, gently curved upwards to pointed apex; lateral scoli smallest, about one half the length of the subdorsals, conical; front with dark vertical dash; adfrontal area dark brown; upper section of epicranial suture dark brown; broad stripe lateral to adfrontal area, terminating at stemmatal level with a darker vertical dash; broad stripe from base of dorsal scoli, terminating on stem- mata with a darker vertical dash; dark brown stripe from mid point of inside of each dorsal scolus, termi- nating at epicranial suture; dark brown stripe along curve from mid point of outside of each dorsal scolus, to sides of head just in front of subdorsal scolus; base of head dark brown from occiput to mandibles. Body: brown, except paler dorsally on abdominal segments A2—A6; broad subspiracular white stripe on Al—A7 with oblique brown intrusions from above, just poste- rior to each spiracle on A2—A7; four soft laterally flat- tened triangular middorsal projections, one on each of A3—A6; large oval middorsal spot, lying between the projections of A3 and A4, dark brown with a beige pos- terior-pointing arrow; caudae held separated, brown, broad. Pupa (17 days, Fig. 2d). About 4.5 cm long, and 2 cm wide at widest point; beige with fine brown cryptic markings, giving the general appearance of a dried, curled and sun-bleached leaf; head ridged from center of eye to vertex; eye area adorned with stout dark brown upright setae; antennae with a median black stripe for their entire length, and with cross lines set approximately 0.5 mm apart; mesothorax mid-dorsally humped and keeled; a lateral keel begins near the tho- racic spiracle, passes along the forewing, parallel to the inner margin, and, at a level with abdominal segment Al, smoothes to become a raised area following the forewing inner margin to the tornus; each forewing bears two white enameled triangles toward the base of the inner margin and just ventral to the wing keel; clear patches and a small dark triangle are found at the midpoint of each mesothoracic leg; on the abdomen a dark brown midventral line terminates at the tip of the cremaster, as does an oblique dark brown line that be- gins at the spiracle on abdominal segment A6; dorsum adomed with dark brown, upright setae from the mesothoracic hump through abdominal segment A8, on which the setae are somewhat appressed; spiracles narrowly elliptical, and that of AS is obscure. Diagnostic characters of early stage morphol- ogy. The mature larvae of Central American species of Caligo can be easily diagnosed by head and body color, and by the number and morphology of the head scoli. The mature larva of Caligo eurilochus sulanus Fruh- storfer has a dark tan head adorned with four pairs of 146 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TABLE 1. Larval food plants of the butterfly subfamily Brassolinae (Nymphalidae). Abbreviations: ARE = Arecales, BRO = Bromeliales, CYC = Cyclanthaceae, GEN = Gentianales, gym = gymnosperm, POA = Poales, ZIN = Zingiberales, ovp = oviposition record. References: Aiello, unpbl’; Aiello & Silberglied, 1978”; Barcant, 1970°; Biezanko et al., 1974*; Burmeister, 1873°; Casagrande, 1979°; Condie, 1976’; Cubero, 1985°, d Almeida, 1922°; d’Aratijo e Silva et al., 1968"; DeVries, 1985", 1987"; Fontaine, 1913", Harrison, 1963"; Hayward, 1969’; Moss, unpbl (in Ackery, 1988)!°; Miiller, 1886"; Rothschild, 1916'*; Small, unpbl!*; Srygley, unpbl”®; Srygley & Penz, unpbl*!; Stauffer et al., 1993”; Urich & Boos, 1981”; Urich & Emmel, 19914; Yepez et al., 1985”; Young, 1977°°, 198677; Young & Muyshondt, 1975°°, 1985”. Butterfly species Plant: order family species Reference 1. Blepolenis (as Opsiphanes) batea (Hiibner) ARE Arecaceae “palm” 18 POA Poaceae Panicum lanatum Sw. (as capim amargoso) 10 (no. 2388) Blepolenis batea (Hiibner) ARE Arecaceae Arecastrum (as Syagrus) romanzoffianum 4 (Cham.) Becc. ARE Arecaceae Butia (as Syagrus) capitata (Mart.) Becc. 4 2. Brassolis astyra Godart ARE Arecaceae “diversas especies de Palmae” 15 ARE Arecaceae “palmen” 17 Brassolis astyra astyra Godart ARE Arecaceae Arecastrum (as Cocos) romanzoffianum 10 (no. 2362 (Cham.) Bece. (as geriv) ARE Arecaceae Astrocaryum ayri Mart. (as brejauva) 10 (no. 2362 ARE Arecaceae Bactris sp. 10 (no. 2362 ARE Arecaceae Butia (as Cocos) eriospatha (C. Mart. ex Drude) _ 10 (no. 2362 Becc. (as butidzeiro) ARE Arecaceae Cocos nucifera L. (as baba de boi) 10 (no. 2362 ARE Arecaceae Cocos nucifera L. (as coqueiro da Bahia) 10 (no. 2362 ARE Arecaceae Cocos nucifera L. (as coqueiro anao) 10 (no. 2362 ARE Arecaceae Copernicia cerifera Mart. (as carnaubeira) 10 (no. 2362 ARE Arecaceae Livistona chinensis (Jacq.) R. Br. 10 (no. 2362 (as pent-séo da China) ARE Arecaceae Livistona rotundifolia (Lamarck) Mart. 10 (no. 2362 ARE Arecaceae Phoenix dactylifera L. (as tamareira) 10 (no. 2362 ARE Arecaceae Roystonea (as Oreodoxa) oleracea (Jacq.) 10 (no. 2362 O.F. Cook (as palmeira imperial) ARE Arecaceae Roystonea (as Oreodoxa) regia (Kunth) 10 (no. 2362 O.F. Cook (as palmeira real) POA Poaceae Saccharum officinarum L. (as cana de ag¢icar) 10 (no. 2362 ZIN Musaceae Musa sapientum L. (as bananeira) 10 (no. 2362 3. Brassolis isthmia Bates ARE Arecaceae Chaemodora sp. 27 ARE Arecaceae Cocos nucifera L. 12 4. Brassolis sophorae (L.) ARE Arecaceae Acrocomia aculeata (Jacq.) Lodd. ex Mart. 29, ARE Arecaceae Archontophoenix alexandrae (Muell.) 22 H. Wendl. & Drude ARE Arecaceae Arecastrum romanzoffianum (Cham.) Becc. 22, ARE Arecaceae Astrocaryum spp. 10 (no. 2365) ARE Arecaceae Attalea sp. 10 (no. 2365) ARE Arecaceae Bactris major Jacq. 22 ARE Arecaceae Bactris spp. 10 (no. 2365) ARE Arecaceae Butia (as Cocos) eriospatha (C. Mart. 10 (no. 2365) ex Drude) Becc. (as butidzeiro) ARE Arecaceae Caryota mitis Lour. (as C. plumosa horticola) 22 ARE Arecaceae Caryota urens L.. ae) ARE Arecaceae Chrysalidocarpus lutescens H. Wendl. 22 ARE Arecaceae Chrysalidocarpus lutescens H. Wendl. 10 (no. 2364) (as areca bambi) ARE Arecaceae Cocos nucifera L. 8h, WS, 22 ARE Arecaceae Cocos nucifera L. (as coqueiro da Bahia) 10 (nos 2363— 2365) ARE Arecaceae Cocos nucifera L. (as coqueiro anao) m (no. 2365) ARE Arecaceae Copernicia cerifera Mart. (as carnatiba) 10 (no. 2365) ARE Arecaceae Desmoncus spp. 10 (no. 2365) ARE Arecaceae Euterpe spp. 0 (no. 2365) ARE Arecaceae Hyophorbe lagenicaulis (L.H. Bailey) H.E. Moore 2 ARE Arecaceae Livistona chinensis (Jacq.) R. Br. (as pent- 10 (nos 2364, sdo da China) 2365) ARE Arecaceae Livistona sp. 2) ARE Arecaceae Mauritia flexuosa L. f. 22, ARE Arecaceae Neodypsis decaryi Jumelle 22, ARE Arecaceae Orbignya spp. 10 (no. 2365) ARE Arecaceae “palms” 18 ARE Arecaceae Phoenix canariensis hort. ex Chabaud 22, ARE Arecaceae Phoenix dactylifera L. 22 ARE Arecaceae Phoenix dactylifera L. (as tamareira) 10 (no. 2365) VOLUME 53, NUMBER 4 Butterfly species 5. 6. Ts 8. 10. Ul. 12. Caligo arisbe Hiibner Caligo atreus Kollar Caligo atreus dionysos Fruhstorfer Caligo beltrao (Illiger) Caligo eurilochus (Cramer) Caligo eurilochus brasiliensis (Felder) Caligo eurilochus sulanus Fruhstorfer . Caligo idomeneus (L.) Caligo illioneus (Cramer) Caligo illioneus oberon Butler Caligo illioneus pampeiro Fruhstorfer Caligo martia (Godart) Caligo memnon (Felder & Felder) Plant: order ARE ARE ARE ARE ARE ARE ARE ARE ARE ARE ARE ARE POA ZIN ZIN ZIN ZIN ARE ZIN ZIN CYC ZIN ZIN ZIN ZIN ZIN ZIN ZIN ZIN ZIN ZIN ZIN ZIN ZIN ZIN ARE ZIN ZIN ZIN ZIN ZIN ZIN ZIN ZIN ZIN ZIN ZIN ZIN POA ZIN ZIN ZIN POA POA ZIN ZIN ZIN ZIN ZIN ZIN ZIN TABLE 1. family Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Poaceae Musaceae Musaceae Musaceae Marantaceae Arecaceae Marantaceae Musaceae Cyclanthaceae Musaceae Musaceae Cannaceae Marantaceae Marantaceae Musaceae Musaceae Zingiberaceae Marantaceae Musaceae Musaceae Musaceae Musaceae Musaceae Arecaceae Musaceae Musaceae Zingiberaceae Marantaceae Musaceae Musaceae Marantaceae Musaceae Musaceae Marantaceae Musaceae Musaceae Poaceae Musaceae Musaceae Musaceae Poaceae Poaceae Cannaceae Cannaceae Marantaceae Musaceae Musaceae Musaceae Musaceae Continued. 147 species Phoenix reclinata Jacq. Pritchardia pacifica Seemann and Wendland Ptychosperma macarthurii (A.A. Wendl.) G. Nicholson Roystonea (as Oreodoxa) oleracea (Jacq. ) O.F. Cook (as palmeira imperial) Roystonea (as Oreodoxa) regia (Kunth) O.F. Cook (as palmeira real) Roystonea oleracea (Jacq.) O.F. Cook Roystonea regia (Kunth) O.F. Cook Roystonea venezuelana L.H. Bailey Sabal mauritiiformis (H. Karst.) Griseb. and H. Wendl. Sabal umbraculiferus Mart. Scheelea macrocarpa Karsten Washingtonia filifera (Linden ex André) H. Wendl. Saccharum officinarum L. (as cana de agtcar) Musa sapientum L. (as bananeira) Ravenala madagascariensis Sonn. Strelitzia nicolai Regel & Kérn. Caeté Asterogyne martiana H. Wendl. (H. Wendl.) ex Hemsl. Calathea sp. Heliconia spp. Cyclanths Heliconia sp. Musa sp. Canna indica L. Caeté Calathea zebrina (Sims) Lindl. Musa sapientum L. (as bananeira) “plusieurs Musaceae” Hedychium coronarium J. Konig (as lirio do brejo) Calathea latifolia (Willd. ex Link.) Klotzsch Hedychium sp. Heliconia latispatha Benth. Heliconia latispatha Benth. Musa sapientum L. “plusieurs Masaceas (sic) Euterpe edulis Mart. (as palmito) banana Musa sapientum L. (as bananeira) » Hedychium coronarium J. Konig (as lirio do brejo) Calathea sp. Heliconia sp. Musa sp. unidentified Heliconia latispatha Benth. Musa sp. Hedychium coronarium J. Kénig (as lirio do brejo) Heliconia spp. Musa sapientum L. (as bananeira) Saccharum spontaneum L. Heliconia sp. Musa sp. banana Echinochloa crus-galli (L.) P. Beauv. (as capim canivao) Pennisetum purpureum Schumach. (as capim elefante) Canna indica L. Canna sp. Calathea latifolia (Willd. ex Link.) Klotzsch Heliconia latispatha Benth. Heliconia spp. Musa sapientum L. Musa sp. Reference wp bw Nw wb 10 (nos 2364, 2365) 10 (nos 2364, 2365) NWwWW Wb Ww bh Ol 10 (no. 2365) 10 (no. 2365) 10 (no. 2369) 10 (no. 2367) 6 10 (no. 2367) 9 10 (no. 2367) 20 17 1 (lot 80-26) 20 (ovp) 20 (ovp) 9 10 (no. 2368) 18 10 (no. 2368) 10 (no. 2368) 12 12 12 15 1 (lot 81-77) 16 10 (no. 2371) ll 10 (no. 2371) 21 12 12 18 10 (no. 2372) 10 (no. 2372) (lot 85-57) PNWANWN AH IS bo oe) 4,29 148 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TABLE 1. Continued. Butterfly species Plant; order family species Reference Caligo memnon memnon (Felder & Felder) ZIN Musaceae Heliconia sp. 12 ZIN Musaceae Musa sp. 12 Caligo memnon telamonius (Felder & Felder) GEN Rubiaceae Coffea sp. (as cafeeiro) [dubious record] 10 (no. 2373) 13. Caligo oberthurii oberthurii (Deyrolle) ARE Arecaceae “low, evergreen palm” 18 14. Caligo oileus (Felder & Felder) ZIN Musaceae Musa sp. 16 Caligo oileus scamander (Boisduval) ZIN Musaceae Heliconia sp. 12 15. Caligo placidianus Staudinger ZIN Musaceae Musa sp. 16 16. Caligo praxsiodus Fruhstorfer POA Poaceae Saccharum officinarum L. (as cana de a¢icar) 10 (no. 2374) 17. Caligo prometheus epimetheus ZIN Musaceae banana 18 (Felder & Felder) 18. Caligo teucer (L.) ZIN Musaceae Heliconia sp. 16 ZIN Musaceae Musa sp. 8} Caligo sp. ARE Arecaceae Cyrtostachys sp. 20 ZIN Musaceae Musa sapientum L. (as bananeira) 10 (no. 2366) ZIN Zingiberaceae Hedychium coronarium J. Konig (as lirio do brejo) 10 (no. 2366) 19. Catoblepia amphirhoe (Hiibner) ARE Arecaceae Arecastrum (as Cocos) romanzoffianum (Cham.) 10 (no. 2375) Becc. (as geriva) ARE Arecaceae “palmeras” 15 20. Catoblepia orgetorix championi Bristow ARE Arecaceae palms 12 21. Dasyophthalma rusina (as geraensis) (Godart) ARE Arecaceae Bactris tomentosa Mart. (as uricana) 10 (no. 2376) Dasyophthalma rusina (Godart) ARE Arecaceae Euterpe edulis Mart. (as palmito) 10 (no. 2377) POA Poaceae Bambusa sp. (as bambi) 10 (no. 2378) 22. Dynastor darius (F.) BRO Bromeliaceae Aechmea fasciata (Lindl.) Baker 10 (no. 2381) BRO Bromeliaceae Aechmea nudicaulis (L.) Griseb. 24 BRO Bromeliaceae Ananas comosus (as sativus)(L.) Merr. (as abacaxi) 10 (nos 2379— 2381) BRO Bromeliaceae Ananas sp. (as anands selvagem) 10 (no. 2379 BRO Bromeliaceae Ananas sp. (as anands) 10 (no. BRO Bromeliaceae Billbergia nutans H. Wend. ex Regel 10 (no. BRO Bromeliaceae Billbergia speciosa Thunb. 10 (no. 2381 BRO Bromeliaceae Billbergia spp. 10 (no. BRO Bromeliaceae Bromelia fastuosa Lindl. (as bananilhado mato) _10 (no. 10 ( BRO Bromeliaceae Bromelia fastuosa Lindl. (as caraguaté) nos 2379, 2380) BRO Bromeliaceae Bromelia fastuosa Lindl. (as banana do mato) 10 (no. 2379) BRO Bromeliaceae Ortgiesia (as Aechmea) gamosepala (Wittm.) 10 (no. 2381) L.B. Sm. & WJ. Kress BRO Bromeliaceae Tillandsia zebrina hort. ex Baker 10 (no. 2379) BRO Bromeliaceae unidentifed 17 Dynastor darius mardonius Fruhstorfer BRO Bromeliaceae unidentified 15 Dynastor darius stygianus Butler BRO Bromeliaceae Aechmea magdalenae (André) André ex Baker Il BRO Bromeliaceae Aechmea sp. 12 BRO Bromeliaceae Agallostachys pinguin (L.) Beer (as Bromelia 11 pinguin L.) BRO Bromeliaceae Ananas comosus (L.) Merr. [accepted by larvae 2, 1 (lot 78-84) in captivity] BRO Bromeliaceae Ananas sp. 12 BRO Bromeliaceae Bromelia plumieri (E. Morren) L.B. Sm. 1 BRO Bromeliaceae Bromelia sp. 12 BRO Bromeliaceae “pineapple and other bromeliads, gravata” 18 23. Dynastor macrosiris (Doubleday) BRO Bromeliaceae Aechmea nudicaulis (L.) Griseb. 23, 9A 24. Dynastor napoleon (Doubleday) BRO Bromeliaceae Aechmea nudicaulis (L.) Griseb. 24 BRO Bromeliaceae Aechmea sp. 10 (no. 2382) BRO Bromeliaceae Ananas comosus (as sativus) (L.) Merr. (as abacaxi) 10 (no. 2382) BRO Bromeliaceae gravata 18 25. Eryphanis aesacus bubocula (Butler) ARE Arecaceae palms 12 POA Poaceae Bambusa vulgaris Schrad. ex J.C. Wendl. 8 POA Poaceae Chusquea scabra Soderstr. & C.E. Calderén 8 POA Poaceae Olyra caudata Trin. 8 26. Eryphanis automedon (Cramer) POA Poaceae Bambusa sp. 3 27. Eryphanis polyxena lycomedon POA Poaceae bamboo Ww (Felder & Felder) POA Poaceae Bambusa arundinacea (Retz.) Willd. 1 (lot 82-9) POA Poaceae Saccharum spontaneum L. iL, Bil 28. Eryphanis reevesii (Doubleday) POA Poaceae Bambusa (as Guadua) sp. 10 (no. 2383) POA Poaceae Bambusa vulgaris Schrad. ex J.C. Wendl. 10 (no. 2383) (as bambti comum) VOLUME 53, NUMBER 4 Butterfly species Eryphanis reevesii (Doubleday) (as Caligo rivesit) 29. Narope cyllastros Doubleday Narope cyllastros cyllastros Doubleday Narope cyllastros testacea Godman & Salvin 30. Opoptera aorsa (Godart) (as Opsiphanes aorosa) 31. Opoptera staudingeri (Godman & Salvin) Opoptera (as Opsiphanes) staudingeri (Godman & Salvin) 32. Opoptera (as Opsiphanes) syme (Hiibner) 33. Opsiphanes bogotanus Distant Opsiphanes bogotanus bogotanus Distant 34, Opsiphanes cassiae (L.) Opsiphanes cassiae cassiculus Stichel Opsiphanes cassiae lucullus Fruhstorfer 35. Opsiphanes cassina aiellae Bristow Opsiphanes cassina fabricii (Boisduval) 36. Opsiphanes invirae (Hiibner) Plant: order POA POA POA POA POA POA POA POA POA POA POA POA POA POA ARE ZIN ZIN ZIN ARE ZIN ZIN ZIN ZIN ZIN ZIN ARE ARE ARE ARE ARE ARE ARE ARE ARE ARE ARE ARE ARE ARE ARE ARE ARE ARE ARE ARE ARE ARE ARE ARE ARE TABLE 1. family Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Arecaceae Marantaceae Marantaceae Musaceae Arecaceae Musaceae Musaceae Musaceae Musaceae Musaceae Musaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Continued. species Bambusa vulgaris Schrad. ex J.C. Wendl. (as bambi: comum) Olyra latifolia L. (as taquarinha) Pennisetum purpureum Schumach. (as capim elefante) Bambusa sp. Olyra latifolia L. Bambusa spp. Bambusa (as Guadua) sp. Bambusa sp. (as bambi) Bambusa sp. Bambusa vulgaris Schrad. ex J.C. Wendl. (as bambti comum) Chusquea longifolia Swallen Chusquea sp. Chusquea sp. Bambusa (as Guadua) sp. palms Calathea inocephala (Kuntze) H.A. Kenn. & Nicolson Calathea latifolia (Willd. ex Link.) Klotzsch banana unidentified Heliconia sp. Heliconia sp. and “différentes plantes musacées” Musa sapientum L. Musa sp. banana Musa sapientum L. (as bananeira) Cocos nucifera L. Livistona sp. palm Acrocomia vinifera Oerst. Bactris guineensis (L.) H.E. Moore (as Bactris minor) Bactris sp. Cocos nucifera L. Erythea salvuadorensis (H.Wendl. ex Becc.) H.E.Moore (as Brahea saldorensis) Roystonea regia (Kunth) O.F. Cook Arecastrum (as Cocos) romanzoffianum (Cham.) Bece. (as geriva) Arecastrum (as Syagrus) romanzoffianum (Cham.) Becc. Butia (as Cocos) eriospatha (C. Mart. ex Drude) Bece. (as butidzeiro) Butia (as Syagrus) capitata (Mart.) Becc. Cocos nucifera L. (as coqueiro da Bahia) Copernicia cerifera Mart. (as carnatiba) Livistona australis (R. Br.) C. Mart. Livistona australis (R. Br.) C. Mart. (as pent-sdo austral) Livistona chinensis (Jacq.) R. Br. (as pent-sdo chinés) Livistona chinensis (Jacq.) R. Br. Livistona rotundifolia (Lamarck) Mart. Palmeira de leque Phoenix canariensis hort. ex Chabaud Prestoea sp. Raphia sp. (as palmeira omamental) Roystonea (as Oreodoxa) oleracea (Jacq.) O.F. Cook (as palmeira imperial) 149 Reference 10 (no. 2383) 10 (no. 2383) 10 (no. 2383) 17 15,17 10 (no. 2384) 10 (no. 2384) 12 10 (no. 2387) 8 12 11 10 (no. 2395) 11, 12 1 (lot 81-41) 20 18 13 16 5 15 3 18 10 (no. 2389) 1 (lots 77-76, 82-1, 83-17, 87-3) 1 (lot 91-25) 1 (lot 95-8) 11, 12 28 11, 12 11, 12, 28 28 28 10 (nos 2390, 2392) 4 10 (nos 2390, 2392) 4 10 (no. 2390) 10 (no. 2390) 4 10 (nos 2390, 2392) 10 (no. 2392) 4 10 (no. 2390) 10 (no. 2392) 4 8 10 (no. 2391) 10 (no. 2390) 150 Butterfly species Plant; order Opsiphanes invirae amplificatus Stichel (as O. i. remoliatus) Opsiphanes invirae amplificatus (Stichel) Opsiphanes invirae amplificatus Stichel (as ampliplacita) Opsiphanes invirae cuspidatus Stichel 37. Opsiphanes merianae Stichel 38. Opsiphanes quiteria (Cramer) Opsiphanes quiteria badius Stichel Opsiphanes quiteria meridionalis Staudinger Opsiphanes quiteria meridionalis (as philon) Staudinger Opsiphanes quiteria quirinus Godman & Salvin 39. Opsiphanes tamarindi Felder & Felder Opsiphanes tamarindi sikyon Fruhstorfer Opsiphanes tamarindi tamarindi Felder & Felder Opsiphanes sp. gym ZIN ARE ARE ZIN ARE ARE ARE ARE ARE ARE ARE ARE ARE ARE ARE ARE ARE ZIN ZIN ZIN ZIN ZIN ZIN ZIN ZIN ZIN ARE ARE ZIN ZIN TABLE 1. family Cycadaceae Musaceae Arecaceae Arecaceae Musaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Arecaceae Musaceae Musaceae Musaceae Musaceae Musaceae Cannaceae Musaceae Musaceae Musaceae Arecaceae Arecaceae Musaceae Musaceae scoli, dark brown body with six middorsal projections (see figs. 6 and 7 in Malo & Willis 1961 p. 532). That of C. atreus dionysos Fruhstorfer has a tan colored head with fine vertical striations and three pairs of scoli, TABLE 2. Plant: order family Butterfly genera Blepolenis Brassolis Caligo Catoblepia Dasyophthalma Dynastor Eryphanis Narope Opoptera Opsiphanes JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Continued. species Cycas circinalis L. (as palmeira de jardim) [dubious record] Musa sapientum L. (as bananeira) “giriva and palms” Phoenix sp. Musa sapientum L. (as bananeira) Bactris major Jacq. palms ornamental palm Arecastrum romanzoffianum (Cham.) Becc. Bactris sp. Chrysalidocarpus (as Areca) lutescens H. Wendl. Astrocaryum ayri Mart. (as brejauva) Euterpe edulis Mart. (as palmito) Chrysalidocarpus lutescens H. Wendl. Cocos nucifera L. Geonoma sp. palms Prestoea allenii H.E. Moore Musa sapientum L. (as bananeira) Musa sp. Heliconia collinsiana Griggs Heliconia latispatha Benth. Musa sp. Canna indica L. Heliconia latispatha Benth. Heliconia sp. Musa sp. Cocos nucifera L. (as coqueiro ano) Cyrtostachys sp. Heliconia latispatha Benth. Musa sapientum L. (as bananeira) Reference 10 (no. 2390) 10 (nos 2390, 2392) 18 15 10 (no. 2386) 1 (lot 82-44) 12 3 15 1 (lot 81-74) 10 (no. 2393) 10 (no. 2394) 10 (no. 2394) 8 26 8 12 8 10 (no. 2396) 14,17 28 28 28 1 (lot 84-9) 1 (lots 80-40, 80-43) 12 12 10 (no. 2385) 20 1 (lot 82-18) 10 (no. 2385) with the dorsal pair enlarged at tip and curved outward (see fig. 32, El in DeVries 1987 p. 248), a tan colored body with many fine striations on dorsum, and five middorsal projections (DeVries 1987). The head cap- Summary of larval food plant records for brassoline butterflies. Numbers represent species as listed in Table 1. ARE Arecaceae 33, 34, 35, 36, 37, 38 POA Poaceae 21 25, 26, 27, 28 29 30, 31, 32 ZIN Musaceae 33, 34, 36, 39 QE Marantaceae Zingiberaceae Canaceae Cyclanthaceae 5, GO, 7, 3, © Le 7, 12 6 10, 12 33 39 BRO Bromeli- aceae 22, 23, 24 VOLUME 53, NUMBER 4 sule of the mature larva of C. memnon is banded with tan and dark brown with four pairs of scoli (see fig. 5 in Young & Muyshondt 1985 p.162; fig. 32, E2 in De- Vries 1987 p. 248), with the dorsal pair enlarged at tip. The body is light brown with a dark brown middorsal stripe, dark brown striations, and six middorsal projec- tions (note that fig. 5 in Young & Muyshondt 1985 p.162, and fig. 31 F in DeVries 1987 p. 248, do not portray the same body color pattern). The mature larva of C. illioneus has a head patterned in brown and creamy white adorned with three pairs of scoli plus a lateral tubercle, and the body is beige with a dark brown middorsal stripe and four middorsal projections (Fig. la). That of C. idomeneus has a beige head pat- terned with brown, three pairs of scoli with the dorsal pair enlarged at tip and curved outward (Fig. 2b). The body is brown with a lighter colored area dorsally and a large oval middorsal spot between A3 and A4, a broad white subspiracular stripe, and four middorsal projections (Fig. 2a). The pupae of all species are very similar, and those of C. illioneus and C. idomeneus seem to differ only in the size of the white triangular marking at the base of the wing (more prominent in C. idomeneus, Fig. 2d). Early stages of C. oileus have never been formally described. Larval food plants of the Brassolinae. It is well known that brassoline immatures are restricted to monocotyledonous plants (Ehrlich & Raven 1965, Ackery 1988, Table 1), but little correlation has been found between plant use and brassoline classification (Ackery 1988). We found that larval food plants in- clude four of the eight monocot superorders (Table 1), a distribution suggesting that brassolines are generalist monocot feeders. However, all food plant records to- gether indicate that the majority of species feed on plants in the families Arecaceae, Musaceae, and Poaceae (Tables 1 and 2). Therefore, the apparent lack of correspondence between plant use and brassoline classification should be reexamined. Available records are sufficient to show that brasso- line genera vary both in diet breadth and their associa- tion with monocot families (Table 2; see Stichel 1909 and Bristow 1981, 1982, 1991 for taxonomic classifica- tion of the butterflies). For instance, although individ- ual species of Caligo have been reported to feed on 1-4 plant genera in 14 families, collectively Caligo has a larval food plant range that includes 11 genera in 7 families (Table 2, see also Ackery 1988) suggesting multiple events of host colonization during its evolu- tionary history. Similar patterns occur in Eryphanes and Opsiphanes: species of Eryphanes typically feed on Poaceae, except for E. aesacus which has also been found on Arecaeae; and Opsiphanes tend to associate 151 with Arecaceae and Musaceae, except for O. bo- gotanus and O. tamarindi, whose food plant range also includes Marantaceae and Canaceae respectively. Available information suggests that other brassolines are restricted to a single plant family (Blepolenis and Catoblepia on Arecaceae, Narope and Opoptera on Poaceae, Dynastor on Bromeliaceae). Noteworthy is that the food plant range of the putative basal genus Brassolis includes Arecaeae, Poaceae and Musaceae; the plant families upon which most brassolines feed as immatures. Although patterns of host association can be recognized at the generic level, their examination in an evolutionary context awaits a well supported phy- logeny for this group of butterflies. We hope that the summary presented here encour- ages research aimed at furthering our understanding of the patterns of food plant utilization and evolution in brassoline butterflies. ACKNOWLEDGMENTS We are grateful to: R. Kimsey, who brought in the C. idomeneus larva; C. Galdames, for identifying plants collected by R.B. Srygley and C.M. Penz; W. Hahn for guidance on palm nomenclature; P.J. DeVries, G. Lamas and A. Neild for their help and suggestions; and P. Ackery, P. J. DeVries and D. Jenkins for comments on the manu- script. We also acknowledge InReNaRe (now A.N.A.M.) for grant- ing permission to conduct research and collect butterflies in Panama, and the Smithsonian Tropical Research Institute (STRI) for providing a vehicle to conduct research. Financial support was provided by STRI (to C. Penz), National Geographic Society (to R.B. Srygley), and National Science Foundation DEB98-06779 (to C.M. Penz). LITERATURE CITED ACKERY, P. R. 1988. Hostplants and classification: a review of nymphalid butterflies. Biol. J. Linn. Soc. 33:95-203. AIELLO, A. & R. E. SILBERGLIED. 1978. Life history of Dynastor darius (Lepidoptera: Nymphalidae: Brassolinae) in Panama. Psyche 85:331-345. AUERBACH, M. J. & D. R. STRONG. 1981. Nutritional ecology of He- liconia herbivores: plant fertilization and alternate hosts. Ecol. Monogr. 51:63-83. BarcanT, M. 1970. The butterflies of Trinidad and Tobago. Lon- don, Collins. BIEZANKO, C. M., A. RUFFINELLI & D. LINK. 1974. 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Venez. N. S. 8(1):95-103. STICHEL, H. 1909. Brassolidae. Das Tierreich 25:1-244. Uricu, F. C. & J. O. Boos. 1981. Metamorphosis of Dynastor macrosiris Westw. (Lepidoptera Brassolidae). Living World, 1981:34. Uricu, F.C. & T. C. EMMEL. 1991. Life histories of Neotropical butterflies from Trinidad: 5. Dynastor darius darius (Lepidoptera: Nymphalidae:Brassolinae). Trop. Lepidoptera 2(2):145-149. YEPEZ, G., F. FERNADEZ Y. & J. CLavyo. 1985. Presencia de Brasso- lis sophorae (L.) (Lepidoptera: Brassolidae) causando danos en palmas de chaguaramos, Roysotea oleracceae (Jacq.), en el estado Carabobo, Venezuela. Bol. Entomol. Venez. N. S. 4(3):23-24. Younc, A. M. 1977. Notes on the defoliation of coconut palm (Co- cos nucifera) by the butterfly Opsiphanes quiteria quirinus in north-eastern Costa Rica. Deutsche Entomologische Zeit- schrift, 24:353-365. Younc, A. M. 1986. Natural history notes on Brassolis isthmia Bates (Lepidoptera: Nymphalidae: Brassolinae) in northeastern Costa Rica. J. Res. Lep, 24:385-392. YouNG, A. M. & A. MuysHONDT. 1975. Studies on the natural his- tory of Central America butterflies in the family cluster Satyri- dae-Brassolidae-Morphidae (Lepidoptera: Nymphaloidea). IIT. Opsiphanes tamarindi and Opsiphanes cassina in Costa Rica and E] Salvador. Studies on the Neotropical Fauna 10:19-56. YOuNG, A. M. & A. MuysHONDT. 1985. Notes on Caligo memnon Felder and Caligo atreus Kollar (Lepidoptera: Nymphalidae: Brassolinae) in Costa Rica and El Salvador. J. Res. Lepid. 24:154-175. Received for publication 23 April 1999; revised and accepted 10 October 1999 Journal of the Lepidopterists’ Society 53(4), 1999, 153-158 THE SPHINGIDAE (HETEROCERA) OF THE “EL OCOTE” RESERVE, CHIAPAS, MEXICO Oxca Lipia GOMEZ-NUCAMENDI, ROBERT W. JONES! AND ALEJANDRO MorOn-Rios * El Colegio de la Frontera Sur, Apdo. Postal 63, C.P. 29290 San Cristébal de las Casas, Chiapas, MEXICO ABSTRACT. A study of the family Sphingidae was conducted in the Reserve, “El Ocote,” located in the northeast portion of the state of Chiapas, Mexico. Collections were made principally using a blacklight between 1994 and 1997. A total of 60 species were collected, from 20 gen- era in five tribes and three subfamilies. Xylophanes, Manduca, Eumorpha and Erinnyjis had the greatest number of species. Nyceryx mulleri Clark is a new record for Chiapas. Eighty percent of the species were collected in the first two months of the rainy season. It was estimated that the present collection accounted for 75% of the species of Sphinigidae in the reserve, based on the accumulation of species per collection ef- fort. The fauna of the Sphingidae collected from “E] Ocote” was compared with that reported from two other reserves of southern México, “Los Tuxtlas”, Veracruz and “Chajul”, Chiapas. “El Ocote” and “Chajul’” were the most similar with 87% affinity. Additional key words: tropical forests, biodiversity, neotropical region, invertebrate inventory. RESUMEN. ‘Se presenta un estudio faunistico de la familia Sphingidae de la selva “El Ocote”, ubicada al noreste del estado de Chiapas, México. Se efectuaron colectas con trampa de luz tipo pantalla entre 1994 y 1997, que aportaron 60 especies de 20 géneros comprendidos en cinco tribus y tres subfamilias. E] mayor numero de especies se distribuye en los géneros Xylophanes, Manduca, Eumorpha y Erinnyis. Nyceryx mulleri Clark es un nuevo registro para Chiapas. E] 80% de las especies se colecté en los dos primeros meses de la temporada lluviosa. La esti- maci6n de la riqueza de especies por esfuerzo de colecta indica que esta investigacién aporta el 75% de las especies de esta regién. Se incluye una comparacion con la fauna de Sphingidae de Los Tuxtlas, Veracruz y Chajul, Chiapas, resultando que El Ocote y Chajul presentan una afinidad de 87%. There has been tremendous international publicity and concern surrounding the unprecedented rate at which tropical forests are being lost. The concern is well warranted because although tropical forests cover only about 7% of the terrestrial surface of the planet, they probably support around 50% of the flora and fauna (Myers 1986). Despite this diversity, these forests are increasingly threatened and destroyed. Although much of the international attention has been directed at the larger tracts of tropical forest such as the Amazon and the Congo basin of Africa, Mexico has important tracts of tropical forests that merit seri- ous conservation efforts. However, it is estimated that 80% of the tropical forests of Mexico have already been destroyed and those that remain are seriously threatened (Estrada et al. 1995). This is the case with the tropical forest reserve, “El Ocote” in the north- eastern Chiapas. Of the 48,140 hectares designated as reserve lands in 1982 (Diario Oficial de la Federacién 1982), only 57%, (27,437 ha), can now be considered as forested and not directly affected by agricultural ac- tivities (Garcia et al. 1996). As for other tropical forests, a high diversity of ver- tebrates have been reported from “El Ocote” Reserve (Dominguez et al. 1996, Mufioz et al. 1996, Navarrete- Gutiérrez et al. 1996), but there is a lack of inventories _of the invertebrate fauna. Knowledge of the inverte- brate diversity could complement the findings con- " Present address: Licenciatura en Biologia, Universidad Aut6- noma de Querétaro, Apdo. Postal 184, C.P. 76010, Querétaro, Qro., MEXICO * Address correspondence to this author cerning the vertebrate fauna, and give further infor- mation on the species most threatened by habitat de- struction, and, in general, give an indication of the bi- ological significance of the reserve based on species richness (Toledo 1988). The present study is a contribution to the knowl- edge of the Lepidoptera fauna of southeastern Mexico, and in particular, of the family Sphingidae of the “El Ocote” Reserve of northwestern Chiapas. An inven- tory was conducted of the Sphingidae of the Reserve, from which a comparison was made of the species richness of this family reported from two other tropical forests of southern Mexico. “Los Tuxtlas” of the Gulf Coast of Veracruz, and “Chajul” of the Lacandon region of eastern Chiapas. These forests, together with “El Ocote”, presumably formed a single tract of tropical for- est stretching from the Gulf coast of Veracruz to what is now the border with Guatemala (Challenger 1998). MATERIALS AND METHODS Description of the study area. The protected for- est and fauna reserve known as the “Selva El] Ocote’” is located in the northwest portion of the state of Chia- pas (16°53’-17°05’N and 93°30’—93°47’W) in the mu- nicipality of Ocozocoautla de Espinoza (Fig. 1). The reserve has an area of 48,140 ha (Diario Oficial de la Federacién 1982) and ranges in altitude from 180 to 1500 m above sea level. The mean annual temperature and precipitation is 25.2°C and 2387 mm, respectively (INEGI 1984). The soils of the reserve are thin and fragile, of lime- stone origin, with large numbers of exposed rocks and boulders (Garcia et al. 1996). The topography is highly 154 93° 30’ i OCOZOCOAUILA SN JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY limOoe sR Md, 1Oeta3% Fic. 1. A. Geographic position of the “E] Ocote” Reserve in Southeastern México. B. Detail of the reserve and location of sampling areas: 1= El Encajonado, 2 = irregular and has contributed to the existence of very diverse floral assemblages, of which the high tropical semi-evergreen tropical forest is the dominant vegeta- tion. This forest type coexists with fragments of medium height evergreen tropical forest, low semi- evergreen tropical forest, low deciduous tropical for- est, and associated successional stages: “acahuales,” sa- vanna, and pastures (Ochoa-Gaona 1996). 9 km SW of Hjido Cuauhtémoc 3 = Finca Nueva Providencia. Sampling. Night sampling was conducted at three locations that mostly comprised well preserved medium height evergreen tropical forest: E] Encajon- ado, Ejido Cuauhtémoc and Finca Nueva Providen- cia (Fig. 1). These three locations provide a good rep- resentation of the spatial complexity of this type of tropical rain forest in the Reserve. Collections were mostly made during 1994. Supplementing field work VOLUME 53, NUMBER 4 155 TABLE 1. Characteristics of the collection localities compared in this study. Vegetation categories are: TRF = Tropical Rain Forest, ESF = Evergreen Seasonal Forest, TDF = Tropical Deciduous Forest , and SS = Short-tree Savanna (Breedlove, 1981). Locality Elevation (m) Latitude (N) Longitude (w) “El Ocote”, Chiapas 750 16°53’ 90°30’ “Chajul”, Chiapas 140 16°06’ 90°55’ “Los Tuxtlas”, Veracruz 675 18°25’ 90°13” was conducted in 1995, 1996 and 1997. Each collec- tion coincided with the new moon and had an average duration of five days. A twelve volt ultraviolet light and sheet were placed from 1800 to 0500 h each night to attract moths. At the time of capture, each captured specimen was injected in the thorax with 95% ethyl al- cohol and placed in individually labeled glassine en- velopes. Species were identified using Hodges (1971), D’Abrera (1986) and by comparison with reference material from the Entomology Collection of the Insti- tuto de Biologia, Universidad Nacional Auténoma de México (IBUNAM), in Mexico City. Collected speci- mens were deposited in the Entomology Collection of the El Colegio de la Frontera Sur, Unidad San Cristdbal, Chiapas (ECOSC-E) and the entomology collection of IBUNAM. Analysis of data. Estimations of the species rich- ness of the Sphingidae of “El Ocote” Reserve were based on the Clench equation of species accumulation: S(t) = at/ (1 + bt) where S (t) is the expected number of species at time (t), ais the list increase rate, b is the species accumula- tion parameter, and the asymptote is given as a/b (Sober6n & Llorente 1993). The model estimates total number of species present in the study area based on the characteristics of the decrease in new species col- lected as more time is spent in the field, this process will eventually generate an asymptote as an estimated total number of species (Sober6én & Llorente 1993, Leén-Cortés 1995). The model was fitted by the non linear regression module provided by the package SPSS (v.6.1) using Levenberg-Marquardt algorithm. The fauna of Sphingidae from “E] Ocote” Reserve was compared with that reported for the Chajul Bio- logical Station, Chiapas (Leén-Cortés & Pescador 1998) and Los Tuxtlas, Veracruz (Beutelspacher 1989) (Table 1), using the Simpson’s similarity Index. This in- dex is appropriate when compared faunas are dispro- portionate in size and number of shared taxa (Sanchez & Lopez 1988). We applied cluster analyses using the unweighted arithmetic average clustering method Mean annual temp.(°C) Mean annual precipitation(mm) Vegetation 25.2 2387 TRF, EST, TDF SS 25.0 3000 TRE, ESF 24.4 2900 TRF, TDF, ESF (UPGMA) to show the total relationships among these faunas (Crisci & Lopez 1983). RESULTS AND DISCUSSION Species richness and seasonal abundance. A to- tal of 60 species of the family Sphingidae were col- lected from the “El Ocote” Reserve, belonging to three subfamilies, five tribes, and 20 genera (Table 2). The genus with the largest number of species was Xy- lophanes with 15 species, followed by Manduca with 10 species, and both Ewmorpha and Erinnyis had five species each. These four genera represented 58% of the species collected in “El Ocote”. Leén-Cortés and Pescador (1998) reported the same pattern in the abundance of species per genera from the Chajul Bio- logical Station, in eastern Chiapas, and, in general, this also appears to be common to other tropical forests of America (Le6n-Cortés & Pescador 1998). The species collected from “El Ocote” Reserve rep- resented 64% of the sphingids collected in the state of Chiapas and are all new records for the locality. Of these, Nyceryx mulleri Clark is a new state record. With our records, Chiapas has 49% of the Sphingidae reported from Mexico by White et al. (1991). We estimated a species accumulation curve using Clench’s equation. Our collection represents 75% of the Sphingidae of “El Ocote” Reserve predicted over 100 nights. Additional collection effort may provide only 10 tol5 more sphingid species (Fig. 2). Three abundance categories were established using the criteria of Rabinowitz et al. (1986) and based on the numbers of specimens of each of the species col- lected. These categories were: “rare,” (1 to 2 speci- mens); “common,” (3 to 19 specimens); and “abun- dant,” (20 to 50 specimens). Using these groupings, 16 of the collected sphingids were rare, 40 were common, and 4 were abundant (Table 2). The number of species collected varied greatly be- tween seasons. Forty-six species (77%) were only col- lected during the rainy season (May to October), whereas only one species was found exclusively during the dry season, as compared to 13 species (21%) which were found in both wet and dry seasons. Of the species 156 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TABLE 2. List of species of Sphingidae (Insecta: Lepidoptera) collected in the “Selva El Ocote” Reserve in Chiapas, México during 1994-1997. F = February, A = April, M = May, J = June, Jl = July, Au = August, O = October and N = November, Total = Total number of specimens captured. Species Subfamily Sphinginae Tribe Sphingini Cocytius lucifer (Rothschild and Jordan,1903) Neococytius cluentius Cramer,1775 Manduca dilucida Edwards,1887 Manduca occulta Rothschild and Jordan, 1903 Manduca lefeburei (Giierin, 1844) Manduca ochus (Klug, 1836) Manduca rustica (Fabricius,1775) Manduca albiplaga (Walker,1856) Manduca muscosa (Rothschild and Jordan, 1903) Manduca corallina (Druce,1883) Manduca lichenea (Burmeister,1856) Manduca florestan Cramer,1782 Sphinx leucophaeta Clemens,1870 Sphinx merops (Boisduval,1870) Subfamily Ambulicinae Tribe Smerinthini Protambulyx strigilis (Linnaeus,1771) Adhemarius gannascus (Stoll,1790) Adhemarius ypsilon Rothschild and Jordan,1903 Subfamily Macroglossinae Tribe Dilophonotini Pseudosphinx tetrio (Linnaeus,1771) Isognathus rimosus Grote,1865 Erinnys alope (Drury,1770) Erinnyjis lassauxi (Boisduval,1859) Erinnys ello (Linnaeus,1758) Erinnys oenotrus (Cramer,1782) Erinnyis obscura (Fabricius,1775) Pachylia ficus (Linnaeus, 1758) Pachyliodes resumens (Walker,1856) Hemeroplanes ornatus (Rothschild and Jordan,1894) Hemeroplanes triptolemus (Cramer,1779) Madoryx oiclus (Cramer,1779) Madoryx pluto Cramer,1779 Callionima innus (Rothschild and Jordan,1903) Callionima parce Fabricius,1775 Callionima falcifera (Gehler,1943) Enyo lugubris (Linnaeus, 1777) Enyo ocypete (Linnaeus, 1758) Enyo gorgon (Cramer,1777) Perigonia lusca Fabricius,1777 Subfamily Macroglossinae Tribe Philampelini Eumorpha anchemola (Cramer,1780) Eumorpha triangulum Rothschild and Jordan,1903 Eumorpha elisa (Smyth,1901) Eumorpha satellita Linnaeus,1771 Eumorpha vitis (Linnaeus,1758) Subfamily Macroglossinae Tribe Macroglossini Cauthetia spuria Boisduval,1875 Nyceryx mulleri Clark,1917 Nyceryx riscus Schaus,1890 Xylophanes pluto (Fabricius,1777) Xylophanes tyndarus (Boisduval,1875) Xylophanes pistacina (Boisduval,1877) Xylophanes porcus (Hiibner,1829) Xylophanes ceratomiodes (Grote and Robinson,1867) Xylophanes anubus (Cramer,1877) Xylophanes amadis amadis Stoll Xylophanes amadis cyrene (Druce,1777) Xylophanes belti (Druce,1878) Xylophanes eumedon (Edwards, 1887) Xylophanes turbata Edwards, 1887 Xylophanes chiron nechus Drury,1770 Xylophanes libya (Druce,1878) Xylophanes neoptolemus (Stoll,1782) Xylophanes thyelia Linnaeus,1758 Month M, J FJ, Jl, Au, 0, N J.J FJ] M, J Sis a = Se = Sees eat ° os ZP o> Total Re RAP PR CONE De. St WANrPPR Ree Re wokRorP erie 8He AN — 1 RRENRNWON Rawr g® eee VOLUME 53, NUMBER 4 Number of species 0 20 40 60 80 100 Number of nights Fic. 2. Species accumulation curve of Sphingidae as a function of collection effort from “E] Ocote”, Reserve, Mexico. (OC = Cumula- tive number of expected species, (© = Cumulative number of ob- served species. of sphingids collected during the rainy season, 48 of these (80%) were collected during the first two months of the season in May and June. Towards the end of the wet season, the abundance of each species declined and with fluctuations at low densities, in a manner similar to that observed during the dry season. This seasonal pattern in abundance and activity is sim- ilar to that reported in other studies (Haber & Frankie 1989, Janzen 1984, 1986, Powell & Brown 1990, Pescador 1994, Gregg et al. 1993). Haber and Frankie (1989) and Janzen (1984) state that precipitation and vegetative productivity are closely linked to the life cy- cle of the sphingids. During the dry season, the major- ity of the sphingids are in the pupal stage, while the larval and adult stages are found during the wet sea- son. This pattern is reflected in reduced photosyn- thetic activity and productivity in host plants during the dry season, including the complete loss of leaves in some species, with the resumption of productivity dur- ing the wet season. Similarity among fauna of the Sphingidae in Southern Mexico. The similarity among the sphingid 0 10 20 30 40 50 60 70 80 90 100 “Los Tuxtlas”, Veracruz “El Ocote”, Chiapas “Chajul", Chiapas Fic. 3. Dendrogram of Sphingidae from three sites in southern Mexico. The unweighted arithmetic average (UPGMA) was used to cluster related groups. Los Tuxtlas Chajul El Ocote. Chajul-Los f Tuxtlas El Ocote-Los Tuxtlas El Ocote-Chajul 7 | | | | Ve oa Le B El Ocote-Chajul- F777. | eal Los Tuxtlas Se - | 0 5 10 15 20 25 30 35 40 45 50 55 60 | OSpecies Genera | Fic. 4. Number of genera and species from the three localities compared from southern Mexico: A. number of genera and species not shared between sites, and B. number of genera and species shared between sites. fauna at the three sites analyzed are presented in Fig. 3. Within the three localities, 29 genera and 100 species have been collected, of which only 15 genera and 44 species were shared. The dendrogram derived from the cluster analysis indicates that there is greater similarity between “El Ocote” and the “Chajul” region of the Lacandon Forest of eastern Chiapas, than with “Los Tuxtlas” located on the Gulf coast (Fig. 3). These two latter localities shared 18 genera and 58 species, whereas “E] Ocote” shared 17 genera and 52 species with “Chajul” and 16 genera and 46 species with “Los Tuxtlas”(Fig. 4). In 1998 forest fires devastated approximately 7.4% of “El Ocote” reserve during the National severe drought associated with the El Nifio phenomenon of 1997 (Aguilera 1998). Since this study represents the only non-vertebrate faunal data available of the species richness of a specific group from the reserve before the fires, the data presented here may serve as a base- line for estimating the speed and nature of the recovery of the reserve following this major disturbance event. In addition, Sphingidae, being highly vagile species, easily monitored, and with relatively high diversity may be an especially appropriate group of organisms in the study of what is hopefully a process of recovery. ACKNOWLEDGMENTS The authors wish to thank Jean Haxaire of the Muséo National d Histoire Naturelle de Paris and Carlos R. Beutelspacher-Baigs of the Instituto de Biologia, Universidad Nacional Aut6noma de Méx- ico for their comments and help in confirming the identification of some of the species. We are in debt with different colleagues of El Colegio de la Frontera Sur: Pedro F. Quintana-Ascencio read and gently commented on a preliminary version of the manuscript and helped with the calculations of the Clench model, Trinidad Aleman- Santillan made valuable comments at the beginning of the project, Miguel A. Vazquez-Sdnchez provided important logistic support during field work, and Manuel Girén-Intzin for his help in the field 158 and with the preparation of the specimens. El Colegio de la Fron- tera Sur (ECOSUR) provided logistical and financial support to the authors. LITERATURE CITED AGUILERA, M. 1998. Programa de reforestacion de areas forestales afectadas por incendios 1998. 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Concentrations of lowland sphingid and noctuid moths at high mountain passes in Eastern Mexico. Biotropica 22(3): 316-319. RABINOWITZ, D., S. CAIRNS, & 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 diversity. University of Michigan, Sunderland, Massachu- setts. 584 pp. SANCHEZ, O. & G. LOPEZ. 1988. A theoretical analysis of some in- dices of similarity as applied to biogeography. Fol. Entomol. Mex. 75: 119-145. SOBERON, J. & J. LLORENTE. 1993. The use of species accumulation functions for the prediction of species richness. Conserv. Biol. 7(3):480-488. TOLEDO, V. 1988. La diversidad biol6gica de México. Ciencia y Desarrollo 14(81):17-30. WuitE, A., J. WHITE, & J. DE LA Maza. 1991. La fauna de mari- posas de México. Part III. Sphingidae (Lepidoptera). Rev. Soc. Mex. Lepidop. 14(2):1-19. Received for publication 29 December 1998; revised and accepted 2 February, 2000. Journal of the Lepidopterists’ Society 53(4), 1999, 159-168 POLYPHENISM AND POPULATION BIOLOGY OF EUREMA ELATHEA (PIERIDAE) IN A DISTURBED ENVIRONMENT IN TROPICAL BRAZIL FABIO VANINI, VINiCIUS BONATO AND ANDRE VICTOR LUCCI FREITAS! Museu de Historia Natural, Instituto de Biologia, Universidade Estadual de Campinas, CP 6109, 13083-970, Campinas, Sao Paulo, Brazil ABSTRACT. A population of E. elathea was studied for 13 months, from May 1996 to May 1997 in a disturbed environment in suburban Campinas, southeastern Brazil. The population showed fluctuations in numbers throughout the study period, with well-marked peaks of abun- dance in June-July, November—December and February. Sex ratio was male biased in four months and the time of residence was higher in the dry season. Both sexes were polyphenic; paler phenotypes occurred in the dry season and darker phenotypes in the wet season. Paler pheno- types were more frequently recaptured and had higher residence values than darker ones. Differences in behavior were attributed to adapta- tion to seasonally different environments. Additional key words: Coliadinae, mark-recapture, urban butterflies. The recent surge of interest in the conservation of tropical environments has led to an increase in studies of the natural history and ecology of organisms resid- ing in the tropics (Noss 1996). These have included some long-term studies on population biology of neotropical butterflies, focused mainly on aposematic groups such as Heliconiini, Ithomiinae and Troidini (Turner 1971, Ehrlich & Gilbert 1973, Brown & Ben- son 1974, Drummond 1976, Young & Moffett 1979, Brown et al. 1981, Saalfeld & Araujo 1981, Vasconcel- los-Neto 1980, 1986, 1991, Freitas 1993, 1996, Haber 1978, Rogner & Freitas 1999). Population studies with the Pieridae, however, have focused on non-tropical species, especially agricultural pests (Watt et al. 1977, 1979, Tabashnik 1980) common in this family (Chew 1995). Seasonal polyphenism (Shapiro 1976, 1984) is an in- teresting feature of some pierid butterflies (the “whites” and “small yellows”). Numerous insect groups, especially butterflies, are polyphenic in response to sea- sonal abiotic factors (Shapiro 1976). Degree of melanization, number of wing spots (especially eye- like markings on the wings in Satyrinae), and color variations have been attributed to adaptation to sea- sonally different environments and thermoregulation (Shapiro 1976, Brakefield & Larsen 1984, Kingsolver & Wiernasz 1987, Braby 1994, Van Dyck et al. 1997, Windig et al. 1994). In general, seasonal morphs are related to dry vs. wet season in the tropics, and spring vs. summer or fall in temperate regions (Nylin 1989). Many different environmental factors induce poly- phenism (see Shapiro 1984 and Jones 1992). Different behaviors linked to different color patterns also have been reported (Shreeve 1987, Nakasuji & Nakano 1990, Van Dyck et al. 1997). The pierid Eurema elathea (Cramer, 1777) is a small Neotropical butterfly common in lawns, pastures, and other disturbed environments (DeVries 1987, Brown ‘address correspondence to this author 1992). The polyphenism in this species was reported by Brown (1992), who noted that dry season forms were dorsally paler than wet season forms. The species is common on the Campus of the Universidade Estad- ual de Campinas (Unicamp), where it can be observed flying on the lawns and visiting several species of wild flowers (Oliveira 1996). This paper provides a detailed account of the popu- lation parameters of Eurema elathea, and our objec- tives are to: (1) examine the age-structure, size and sex ratio and size of the population; (2) describe the varia- tion in proportions of the different polyphenic types throughout the year; and (3) provide subsidies to the management of this species in urban habitats. MATERIALS AND METHODS A mark-release-recapture (MRR) study was carried out on the Campus of the Universidade Estadual de Campinas (Unicamp), Sao Paulo state, southeastern Brazil, as part of a project of study of urban ecology on the Campus of the University. Annual rainfall is about 1360 mm and mean temperature 20.6°C (data from the Instituto Agron6mico de Campinas). The regional climate is markedly seasonal, with a warm wet season from September to April and a cold dry season from May to August. During the research, the mean tem- perature of the coldest month was 17.6°C and of the warmest month was 24.8°C, with climate typical of the region (Fig. 1), except for May 1997, which was a rainy month and thus was included in the wet season. The vegetation on the campus where the study was conducted consists of large lawns with sparse trees and scattered small flowering shrubs. The lawn was mowed five times during the period of study, as part of normal procedures of maintenance of the campus. The study area was divided into nine plots corresponding to lawns separated by walking trails connecting the build- ings on the Campus (Fig. 2). Butterflies were marked and recaptured between 28 May 1996 and 27 May 1997 (1—3 times per week), in 160 RAINFALL (mm) $8 8 ss) o TEMPERATURE (°C) A M J J A S O N DOD J FM AM 1996 (MONTHS) | 1997 Fic. 1. Climagram for the Campinas region during the research (based on Santos 1965 and Walter 1985). Hatched = humid periods, black = superhumid periods, and dotted = dry periods. sessions with 2—3 persons lasting 2—3 hours near mid- day, totaling 84 field days. The sessions ended when more than 80% of individuals captured represented re- captures of marks already given or recorded on that day. Butterflies were net-captured, numbered on the un- derside of the forewings (felt-tipped pen), and re- leased. Wing wear (seven “age” classes), point of cap- ture, sex, forewing length and food sources were recorded (all following Freitas 1993, 1996, and Rogner & Freitas 1999). Forewing length was measured with a ruler, to the nearest mm. The age of individual but- terflies was estimated in seven categories based on wing wear (Ehrlich & Davidson 1960, Brussard & Ehrlich 1970, Ehrlich & Gilbert 1973), posteriorly grouped into three: new, intermediate and old (as in Freitas 1993, 1996 and Rogner & Freitas 1999). Values from one to seven, attributed to each wing wear class (where 1 = teneral and 7 = tattered) were used to com- pare the mean “age” of first captures in the different SAN Fic. 2. Study area on the Campus of the Universidade Estadual de Campinas, showing the eight grassy areas (closed figures) sepa- rated by open trails. The small circular area, covered by dense shrubs (Calliandra sp., Fabaceae) was not sampled. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY fa | = ee Fic. 3. Color types for males (top and middle rows) and females (bottom row) of Eurema elathea (background pale yellow, solid = black, dotted = orange). color types. The amount of black on the dorsal forewing inner margin was recorded for males and fe- males (details in next section). The mark-release-recapture (MRR) data were ana- lyzed by the Jolly-Seber (Southwood 1971) method for estimating population parameters (CMLR software developed by Dr. R. B. Francini, Unisantos) for the obtainment of estimated population numbers and standard errors. Daily results were tabulated as “num- ber of individuals captured per day” (NICD), and “number of individuals present per day” (NIPD). To estimate the NIPD, recaptured individuals were con- sidered to be present in the population on all previous days since the day of first capture (=marked animals at risk) (following Rogner & Freitas 1999). The data were grouped for analysis into “dry season” (May—August 1996) and “wet season” (September 1996—May 1997), in accord with the rainfall (climagram in Fig. 1). Phenotype. Males were classified in six color types (I to VI) based on the amount of black on the dorsal forewing inner margin (Fig. 3), following a classifica- tion used previously by Ruszcsyk and collaborators (unpublished data) for E. elathea in central Brazil. The extremes were “marginal black bar absent” (type I) Vo) SS AIST aa | (0) D F N Y . \ \ SS Wléaa WL, 50 % A J 1996 (MONTHS ) | 1997 M Fic. 4. Monthly percent of different male color types of E. elathea from May 1996 to May 1997. From top to bottom in first col- umn, types I to VI. VOLUME 53, NUMBER 4 50 Vo N D J F M A i996 | 41997 Fic. 5. Monthly percent of different female color types of E. elathea from May 1996 to May 1997. White = type I (pale), black = type II (dark). Because the type II was absent from May to October 1996, the first six months were combined in a single bar.. and “broad and complete marginal black bar” (type VI). In females only two color types were defined: “marginal black bar absent” (type I) and “short mar- ginal black bar” (type I). Intermediates, as defined by the length and width of the black stripe, were not ob- served in females. The variation appeared to be con- tinuous in males, causing some problems in defining the classes of intermediates especially in the first month (the first 40 individuals marked were dis- carded), resulting in a total number of assigned poly- phenic types different from the total by sex. RESULTS Phenotype. Males of the color types I to IV were more common in the dry season, and V and VI were more common in wet season. The latter two pheno- types (dark forms) represented 90-100% of the total individuals seen in seven months (all in the wet sea- son), and less than 50% in the other months (two in the wet season and all dry season months) (Fig. 4). Fe- males of the color type II (the dark wet season form) appeared first in November 1996, disappearing after April 1997 (Fig. 5). The proportions should be com- pared with the climate of the previous month, when the individual completed larval growth and pupated. ‘Thus, although September was a wet month, the pro- portion of lighter forms was very high, reflecting the dry weather of August; and even though April was rel- atively dry, the darker forms still predominated (Fig. 1). Population biology. In all, 1468 individuals of Eu- rema elathea were captured during the 12 months of 161 INDIVIDUALS PRESENT/DAY JJ A S O N D JF M A M 1996 (MONTHS) | 1997 Fic. 6. Number of individuals present per day (NIPD) of males (below) and females (above) of E. elathea from May 1996 to May 1997. Arrows indicate mechanized grasscuttings. the study, with 320 of these recaptured at least once. In the dry season (25 days), the number of individuals captured per day varied from 0 to 53 individuals (mean = 21.3: SD = 14.3) for males and 0 to 32 (mean = 13.6; SD = 8.2) for females. In the wet season (59 days), the number of individuals captured per day varied from 0 to 32 individuals (mean = 9.8; SD = 8.4) for males and 0 to 27 (mean = 7.9; SD = 7.8) for females. The num- ber of individuals present per day in the dry season varied from 0 to 64 individuals (mean = 27.9; SD = 17.2) for males and 0 to 39 (mean = 17.6; SD = 11.0) for females. In the wet season, the same varied from 0 to 35 individuals (mean = 11.0; SD = 9.6) for males and 0 to 29 (mean = 8.7; SD = 8.7) for females. The population (based on NIPD) presented three peaks of abundance for both sexes: June-July 1996, Novem- ber-December 1996 and February 1997 (Fig. 6). Mechanized grasscuttings (17 Aug. 1996, 13 Dec. 1996, 27 Feb. 1997, 23 Mar. 1997 and 17 May 1997, arrows in Fig. 6), led to decrease in adult abundance especially in 1997 after three grasscuttings at 24 and 55-day intervals. Males were recaptured up to six times in the dry season and up to four times in the wet season; females were recaptured up to four times in dry season and up to three times in wet season. In all, 200 males and 120 females were recaptured at least once (Table 1). The highest recapture rate was observed in male color type I (dry season), and the lowest in male color type VI (wet season) and in both female color types (Table 2). Multiple recaptures were significantly more frequent in males (Table 2). The results of Jolly-Seber analysis for males and females separately (Figs. 7 and 8) show patterns very similar to those of NIPD (Fig. 6). The total sex ratio (809 males and 659 females) was male biased (x? = 15.3, df = 1, p < 0.001), but when 162 TABLE 1. Recapture rates of adults of Ewrema elathea in the study. cap = total captured, recap = individuals captured at least once. Asterisks on the Chi square values indicate that the difference is significant (P < 0.05, df = 1). Chi squares were calculated considering “cap” versus “individuals never recaptured” (“cap” minus “recap’). Dry Wet Total % lo cap/recap % cap/recap % Sex cap/recap Males 370/104 28.1 Females 265/59 223 Total 635/163 25.7 439/96 21.9 394/61 15.5 833/157 18.8 Chi squares males vs females y? = 2.78 x2 = 5.39% dry vs wet 57 = O27" 809/200 24.7 659/120 18.2 1468/320 21.8 I = SO the months were analyzed separately, significant bias was observed in only four months (June, July and De- cember 1996, and May 1997), with females dominat- ing only in one month and sex ratio nearly 1:1 in four months (Fig. 9). Although the proportion of recap- tures of males was higher than females, differences were significant only in the wet season, with total re- capture rate significantly less than in the dry season (Table 1). Age structure and residence time. Most of the first captures of both sexes were individuals of “inter- mediate” age (49% of males and 52% of females). Based on the categories of wing wear, the “age” of first recapture of males (mean class = 2.65, SD = 0.86, n = 744) was lower than females (mean = 2.83, SD = 0.87, n = 660) (¢ = 4.0, df = 1402, p < 0.0001). Color type VI was captured at a lower “age” than all other types ex- cept I (Table 2). The population showed peaks in abundance of “new” individuals in late June, Au- TABLE 2. Population parameters of the different color types of Eurema elathea. Different superscript letters show significant differ- ences among the different phenotypes. AFC = mean age at first cap- ture (based on age categories), MRT = mean residence time (days), MUR = proportion of individuals with multiple recaptures (%), MOV = proportion of individuals that moved to another subarea (%). Males Females Marked 64 62 47 70 1446 355 529 131 Recaptured 24 14 12 19 40 76 95 25 % of recapture* 37.5% 22.6%» 95.5% 97.19» 97.49» 91.4 17.9% 19.1» 9.772» 9.862 2.892 2.787 2.899 2.43> 9.778 9.858 AFC** MRT (days)*** Ine Oa HOS AO OOP lO qWase le MUR 45.8 35.7 25.0 52.6 350 329 210 160 MOV 83.3 57.1 75.0 73.7 80.0 75.0 61.0 76.0 *p < 0.05, Chi-square tests ** D < 0.05, one way ANOVA *E* > 0.05, one way ANOVA JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Trl 500 400 300 200 100 1560 500 400 300 200 100 J J A Fic. 7. Estimated population size (Jolly-Seber) for E. elathea males (above) and females (below), from May to August 1996. The maximum number of individuals is given as the estimate plus the er- ror (superior part of vertical bars), and the minimum number is given as the number of individuals present per day (inferior part of vertical bars), assuming that the population could not be lower than this number. gust-September, late November and February, sug- gesting high rates of recruitment in these periods (Fig. 10), with some indication of protandry. Combining both seasons, residence time of males (mean = 9.8 days, SD = 7.7, n = 201) was not signifi- cantly higher than that of females (mean = 8.3 days, SD = 7.6, n = 119) (¢ = 1.710, df = 318, p= 0109) However, differences were significant in the wet sea- son (t = 2.127, df = 155, p = 0.03), and not in the dry season (¢ = 0.749, df = 161, p = 0.45). Residence times were marginally higher in the dry season for both males (t = 1.95, df = 199, p = 0.052) and females (¢ = VOLUME 53, NUMBER 4 200 100 O N D 519 ee 300 200 100 O N D 163 J F Fic. 8. Estimated population size (Jolly-Seber) for E. elathea males (above) and females (below), September 1996 to February 1997 (see Fig. 7 legend). 1.99, df = 117, p = 0.049) (see also Table 3). The max- imum residence time in the dry season was 52 days for a male and 54 days for a female, and in the wet season 28 days for a male and 21 days for a female. The resi- dence times were not different among the different phenotypes of males and females (F «sey = OLU3, 9 = 0.504) (Table 2). Survival of males and females in each 50 % SEX RATIO MJ JA S ON DJ FM A M 1996 (MONTHS ) | 1997 Fic. 9. Sex ratio for E. elathea from May 1996 to May 1997, as percent of males in each day’s captures. vi Wy / Tt " ~ woeimonTas) | tos? Fic. 10. Age structure of males (below) and females (above) of E. elathea (black = fresh individuals, hatched = intermediate, white = worn individuals) from May 1996 to May 1997. 164 TABLE 3. Residence time (in days) of Eurema elathea adults in the dry and wet seasons. Days elapsed between marking and last recap- ture represent the minimum permanence (MP) for each individual. MP (days) Males (%) Females (%) Total (%) Dry Season 1-5 36 (34.6) 29 (49.1) 65 (39.9) 6-10 23 (22.1) 12 (20.3) 35 (21.5) 11-15 WP (IL A)) 5 (8.5) 27 (16.5) 16-20 8 (7.7) 4 (6.8) 12 (7.4) 21-25 8 (7.7) 4 (6.8) 12 (7.4) 26-30 2 (1.9) 4 (6.8) 6 (3.7) 31-35 3 (2.9) — 3 (1.8) > 35 2 (1.9) 1 (1.7) 3 (1.8) Total 104 59 163 (100) Mean + sd 10.9 + 9.1 9.7+9.8 Max 52 54 Wet Season 1-5 34 (35.4) 32 (52.4) 66 (42.0) 6-10 33 (34.4) 15 (24.6) 48 (30.6) 11-15 16 (16.7) 11 (18.0) Q7 (17.2) 16-20 9 (9.4) 2 (3.3) 11 (7.0) 21-25 2 (2.1) 1 (1.6) 3 (1.9) 26-30 2 (2.1) — 2 (1.3) Total 96 61 157 (100) Mean + sd 8.8 +5.6 6.9+4.2 Max 28 21 season are somewhat different (Table 3), with esti- mated median residence time (“life expectancy” of Cook et al. 1967) of 5.53 days for males and 4.72 days for females in the dry season, and 4.95 days for males and 4.87 days for females in the wet season. Forewing length. For the entire sample, the aver- age forewing length of females (mean = 17.9 mm, SD = 1.19, n = 661) was statistically greater than males (mean = 17.4 mm, SD = 1.05, n = 782) (¢ = 7.51, df = 1441, p < 0.001), and this difference was significant in seven months (Table 4). No significant differences in wing length were observed among the different color TaBLE 4. Mean forewing length (+ SD) of Eurema elathea in this study. Asterisks indicates that means are different between sexes in that month (¢-tests, p < 0.05). N = sample size. Month/year Males N Females N May/1996* 17.6 + 0.9 60 18.4 + 1.2 52 Jun/1996* 174+1.1 147 17.9'+ 1.1 108 Jul/1996* 17.3 + 1.1 Ill 17.9 + 1.3 72 Aug/1996 17.2 + 1.4 oy 17.8 + 1.3 32 Sep/1996 17.8 + 1.1 19 18.1+ 1.4 11 Oct/1996* 16.7 + 0.9 39 17.3 + 1.0 38 Nov/1996* 17.1+1.1 60 17.5 + 1.1 62 Dec/1996* 17.3 + 0.8 58 17.9 + 1.2 40 Jan/1997* 17.3 + 0:9 65 17.9 + 1.2 74 Feb/1997 17.7 + 0.9 133 17.8 + 1.2 120 Mar/1997 17.8 + 0.8 24 18.0 + 1.2 25 Apr/1997 17.8 + 0.9 20 17.9 + 1.1 23 May/1997 17.5 + 1.2 14 17.0 +14 4 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 18.5 18.0 17.5 17.0 AVERAGE WING LENGTH (mm) Fic. 11. Mean (+ SD) forewing length of males and females of E, elathea (based on monthly recruitment). Different letters above the bars indicate significant differences in the means (one way ANOVA, p < 0.005). types within sexes (Fig. 11). Average wing length of males was equal in the dry (mean = 17.4 mm, SD = 1.11, n = 350) and the wet season (mean = 17.4:mm, SD = 0.99, n = 432) (t = 0.942, SD = 780 p = 0.34). In females, average wing length in the dry season (mean = 18.0 mm, SD = 1.21, n = 264) was considered greater than in the wet season (mean = 17.8 mm, SD = 1.17, n = 397) (¢ = -2.437, df = 659, p = 0.015). Natural history of the adults. Adults started ac- tivity around 0800 h in summer and 1000 h in winter, varying greatly with the weather (on some cold days during winter, the activity only began after 1100 h). As a rule, the peak of activity of the adults was between 1200 h and 1400 h, especially in the wet season, when many individuals were in courtship behavior or mat- ing, and females usually were looking for plants for oviposition. After 1600 h, activity diminished, and es- pecially in the dry season, the butterflies congregated in some grass patches in the study area to roost in loose aggregations. Movements among the subareas were observed in all color types of both sexes. Males of color types I and V had the greatest number of individuals moving be- tween sub-areas (Table 2). In the dry season, butter- flies appeared to be more resident, engaging mainly in short flights, while in the wet season they were more active and frequently observed in long flights. Due to the fact that the different subareas were of unequal size, the proportions of individuals moving to other subareas do not reflect the flight distances. VOLUME 53, NUMBER 4 Adults were usually seen feeding on flowers. Thir- teen species of flowers were used as nectar sources. The most visited was Emilia sonchifolia (Asteraceae) (103 of a total of 150 records), but virtually any plant species in blossom was observed being used by the adults (see Oliveira 1996 for a list of flowers used in this same area). DISCUSSION Population biology. Male biased sex ratios have been observed in many natural populations of butter- flies, even when the sex ratio in laboratory was 1:1 (e.g., Brussard & Ehrlich 1970, Ehrlich & Gilbert 1973, Brussard et al. 1974, Watt et al. 1977, 1979, Brown & Ehrlich 1980, Ehrlich 1984, Ehrlich et al. 1984, Mat- sumoto 1984, 1985, Freitas 1993, 1996). Behavioral differences may contribute to this bias (Ehrlich 1984, Freitas 1996); Shapiro (1970) argues, however, that in several pierids this bias is related to the density of males. In this study, females accounted for 44.9% of the 1468 individuals collected in one year (male: female ratio 1.2:1), but the ratio was different from 1:1 in only four months. Residence time of E. elathea is not high if compared to tropical butterflies in the genus Heliconius (Turner 1971, Benson 1972, Ehrlich & Gilbert 1973), but is similar to that obtained for some Ithomiinae (Vascon- cellos-Neto 1980, Freitas 1993, 1996, Rogner & Fre- itas 1999). Dispersal of adults probably affects resi- dence time, explaining the lower time of residence, mainly in the wet season. Especially the lower resi- dence time of females in the wet season could be re- lated to higher dispersal rates in this sex (see also Shapiro 1970 and Freitas 1993, 1996). Increased dis- persal after the first rains has been reported in Ithomi- inae in tropical seasonal forests (Vasconcellos-Neto 1980). Also contrasting with populations of Heliconius but- terflies, that maintain relatively constant numbers throughout the year (Turner 1971, Ehrlich & Gilbert 1973, Araujo 1980, Rogner & Freitas 1999), E. elathea fluctuated markedly in abundance throughout the year, in a way similar to that observed in Ithomiinae (Nymphalidae) and Troidini (Papilionidae) (Brown & Benson 1974, Drummond 1976, Haber 1978, Young & Moffett 1979, Vasconcellos-Neto 1980, 1986, 1991, Brown et al. 1981, Freitas 1993, 1996). Such fluctua- tions may be common in populations of temperate- zone pierids (Watt et al. 1977, 1979, Tabashnik 1980). For E. elathea in this study, these fluctuations are in part related to local grasscuttings, which probably de- stroyed many immatures, and made the adults leave the area looking for food sources. In the present work, 165 grasscuttings occurred in periods of population de- cline, on all occasions (Fig. 6), affecting the population and leading to very low numbers of butterflies in May 1997. Recolonization of the area probably occurred by individuals arriving from nearby populations, and the increase in the frequency of intermediate and old indi- viduals after the first grasscutting (on 17 August 1996) supports this idea. The high proportion of first cap- tures of individuals classified as intermediate and old suggests that migration might be common among sub- populations, and could provide new stock to a recently cut site. These population features suggest that this butterfly species would persist in metapopulations (Hanski & Gilpin 1997) in the study area. Further in- vestigation on this subject could reveal important pat- terns in the ecology of butterflies in urban environ- ments, but the results suggest that grasscuttings should be done after the first rains, when the popula- tion starts to be more vagile. Also, the grasscuttings in the dry season destroyed the roosting places of the butterflies, that could affect the survival of the individ- uals in the dry environment. Even if reproductive diapause was not investigated in the present study, some comparisons can be made with other butterfly species in the seasonal tropics. Several species of tropical butterflies are known to ex- hibit reproductive seasonality (e.g., Jones & Rienks 1987, Braby 1995), and this pattern is supposed to be linked with rainfall, and consequently with the avail- ability of host plants (Braby 1995). The pattern ob- served in E. elathea in the present study is similar to that exhibited by E. hecabe in Australia (Jones & Rienks 1987), with the maximum populational density reached in the middle of the dry season (but the ef- fects of lawn mowing, that is affecting the population numbers, cannot be discarded). In E. hecabe, “gravid” females were present in all samples, but the propor- tion of them dropped when the population reached a peak (Jones & Rienks 1987). Thus, we could hypothe- size that if E. elathea follows the same pattern, the population of this species present continuous breed- ing, but the number of reproductive females in the dry season (also the population peak) might be low. This subject is now under investigation by Ruszczyk and collaborators in a population in central Brazil. Polyphenism. In most studies of polyphenic species of butterflies, dry-season phenotypes were lighter and less conspicuous than wet-season pheno- types (Brakefield & Larsen 1984, Shapiro 1984, Brake- field 1987, Brakefield & Reitsma 1991, Jones 1992, Braby 1994, Windig et al. 1994). These differences have been related to ventral surface camouflage in the more sedentary color morphs produced in the dry sea- 166 son (Shapiro 1976, Brakefield & Larsen 1984, Brake- field & Reitsma 1991, Jones 1992, Van Dyck et al. 1997). In the present study, paler phenotypes were also more frequent in the dry season, when a brown substrate of dead leaves predominated on the ground where the butterflies rest. This paler phenotype may enhance survival of E. elathea through crypsis as the environment dries out and changes color (Owen 1971, Jones 1987, 1992). Thus, the different phenotypes could represent responses to seasonal differences in the environment and selective pressures such as pre- dation (Brakefield & Larsen 1984, Brakefield 1987, Brakefield & Reitsma 1991). Braby (1994) proposed that in satyrs, dark wet-season forms rely on anti- predator devices (prominent eyespot patterns) which are displayed at rest and function to deflect attacks; while lighter dry-season forms with reduced eyespots probably rely on crypsis for survival. Windig et al. (1994) proposed that selection on males tends to favor wing patterns contributing positively to mate-seeking activity and thermal budgets (small and dark wings), while selection on females tends to favor paler ground colors and, in wet season forms, conspicuous markings (as female type II in this study). Allied with differences in color and pattern, differences in behavior in differ- ent seasons and phenotypes were observed on several occasions (Guppy 1986, Shreeve 1987, Nakasuji & Nakano 1990, Van Dyck et al. 1997). A possible hy- pothesis to be tested in tropical seasonal environments is that paler dry-season forms would be more resident than darker wet-season forms, that could enhance the value of their cryptic coloration (as reported in Van Dyck et al. 1997). The results of recapture rates of males of different color types of E. elathea in this study agree with this, since the paler color type (type I) had a higher recapture rate, and the darker color type (type VI) had a lower recapture rate (see Table 2). Un- fortunately, the present study does not provide real distances traveled by the individuals, due to the fact that the different subareas were of unequal size (Fig. 2), and the hypothesis that paler phenotypes are more resident could not be tested. Behavioral differences among the different color types could also be related to wing melanization (and, consequently, to thermoregulation) and body size (wing length). Van Dyck et al. (1997) hypothesize that small phenotypic wing differences could result in con- siderable variation in thermoregulation: darker butter- flies could heat up more rapidly (Wasserthal 1975), and could spend more time flying and searching for fe- males. In the study area, the wet season is also the warm season, and dark butterflies probably would be- came warm faster than pale forms, resulting in in- JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY creased flight activity. This could be true in many trop- ical seasonal habitats. If this holds, dark butterflies would benefit in all seasons (warm or cool), and their absence or low numbers in the dry season would be explained only by the cryptic advantage of the paler forms in dry season. Future research in this subject is needed to investigate this point. The ventral pattern of the wings (not analyzed in this study) varies continually from plain white (mostly in the wet season) to patterned yellow and orange (mostly in the dry season) in both sexes. This is the side normally displayed by the butterflies at rest; the variation probably enhances camouflage and thermal effects in the different color types. Ruszezyk (pers. comm.) proposed that when E. elathea males are fly- ing, the black bar of the darker types resembles the black bar of the hindwing outer margin of the distaste- ful moth Utetheisa ornatrix (Arctiidae). The white pat- tern of the underside of these morphs also matches the light coloration of this moth. The two species often fly together and use similar substrates when perched, and this could be a good hypothesis to be investigated in the future. Size is also important, since larger butterflies could more easily attain and maintain higher temperatures (Willmer & Unwin 1981). Van Dyck et al. (1997) pro- pose that larger darker individuals would be adapted for patrolling and dispersal. Dry-season forms are in general larger than wet-season forms (Brakefield 1987, Brakefield & Reitsma 1991, Jones 1992, Braby 1994). Brakefield and Reitsma (1991) argue that this differ- ence could be the result of behavioral and life history components: larger butterflies were the result of op- portunistic development at the end of the wet season, while dry season forms derived more benefit from fast development (resulting in smaller size) which enables their progeny to complete development before the vegetation dries out. In most pierids, larger adults arise when larvae are reared at lower temperatures, and in two Eurema species larger size was induced by short photoperiods (Jones 1992). In this study, no sig- nificant differences in size in E. elathea were observed within the different color types, but the dry season fe- males were larger than wet season females (not wet season forms). According to Braby (1994), several ad- vantages could be associated with this increase in size, like increased longevity and capacity of storage in the fat body, both important in the dry season, when food resources are scarce. ACKNOWLEDGMENTS We thank K. S. Brown Jr., A. Ruszczyk, A. Gomes-Filho, K. Facure, D. Bowers, Fatima M. Souza and two anonymous reviewers VOLUME 53, NUMBER 4 for suggestions on the manuscript and help in diverse phases of field work. K. Mancini, G. Chaves, R. Cogni and R. Raimundo also helped in field work. We thank R. B. Francini for the copy of the software for population analysis, and the “Instituto Agrondmico de Campinas” (IAC) for the climatological data. A. V. L. Freitas thanks the CNPq for a fellowship. LITERATURE CITED Araujo, A.M. 1980. Estudos genéticos e ecolégicos em Heliconius erato (Lepidoptera, Nymphalidae). Actas do IV Congr. Lati- noam. Genética, Vol 2:199-206. BENSON, W. W. 1972. 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The mate location behavior of the male speck- led wood butterfly, Pararge aegeria, and the effect of phenotypic differences on hind-wing spotting. Anim. Behav. 35: 682-690. 168 SouTHWooD, T. R. E. 1971. Ecological methods with particular reference to the study of insect populations. Chapman & Hall, London. 524 p. TABASHNIK, B. E. 1980. Population structure of pierid butterflies. IIL. Pest populations of Colias philodice eriphyle. Oecologia 47: 175-183. TURNER, J. R.G. 1971. Experiments on the demography of tropical butterflies. II. Longevity and home range behavior in Helico- nius erato. Biotropica 3(1): 21-31. VAN Dyck, H., E. MaTTHYSEN & A. A. DHONDT. 1997. The effect of wing color on male behavioral strategies in the speckled wood butterfly. Anim. Behav. 53: 39-51. VASCONCELLOS-NETO, J. 1980. Dinamica de populagées de Ithomi- inae (Lepidoptera: Nymphalidae) em Sumaré-SP. M.S. Thesis, Universidade Estadual de Campinas, Campinas, SP. vi + 206 pp. . 1986. Interactions between Ithomiinae (Lepidoptera: Nymphalidae) and Solanaceae, pp. 366-377. In W. G. D'Arcy, (ed.), Solanaceae, biology and systematics. Columbia University Press, New York. 603 pp. . 1991. Interactions between ithomiine butterflies and Solanaceae: feeding and reproductive strategies, pp. 291-313. In P. W. Price, T. M. Lewinsohn, G. W. Fernandes & W. W. Benson (eds.), plant-animal interactions. Evolutionary ecology in tropical and temperate regions. John Wiley & Sons, Inc., New York. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY WALTER, H. 1985. Vegetation of the Earth. Springer-Verlag, Berlin, Germany. WASSERTHAL, L. T. 1975, The role of butterfly wings in regulation of body temperature. J. Insect Physiol. 21; 1921-1930. Wart, W. B., F. S. CHEW, L. G. SNYDER, A. G. Watr & D. E. ROTH- SCHILD. 1977. Population structure of pierid butterflies. I. Numbers and movements of some montane Colias species. Oe- cologia 27: 1-22. Wart, W. B., D. HAN & B. TABASHNIK. 1979. Population structure of pierid butterflies. II. A “native” population of Colias philodice eriphyle in Colorado. Oecologia 44: 44-52. WILLMER, P. G. & D. M. Unwin. 1981. Field analyses of insect heat budgets: reflectance, size and heating rates. Oecologia 50: 250-255. WINDIG, J. J., P. M. BRAKEFIELD, N. REITSMA & J. G. M. WILSON. 1994. Seasonal polyphenism in the wild: survey of wing pattern in five species of Bicyclus butterflies in Malawi. Ecol. Entomol. 19: 285-298. Younc, A. M. & M. W. Morretr. 1979. Studies on the population biology of the tropical butterfly Mechanitis isthmia in Costa Rica. Amer. Midl. Nat. 101: 309-319. Received for publication 13 October 1998; revised and accepted 10 December 1999. GENERAL NOTES Journal of the Lepidopterists’ Society 53(4), 1999, 169-170 IS SPERMATOPHORE NUMBER A GOOD MEASURE OF MATING FREQUENCY IN FEMALE CALLOPHRYS XAMI (LYCAENIDAE)? Additional key words: copulation, female remating, mating system. Recent developments in sexual selection theory suggest that the role of females in shaping the evolution of mating systems has been underestimated (Eberhard 1996). In particular, female mating fre- quency is considered a variable that affects the potential for, and the strength of, sperm competition (Drummond 1984) and cryptic fe- male choice (Eberhard 1985, 1996). In Lepidoptera, spermatophore counts have been used to determine the number of times a female has mated (Burns 1968, Drummond 1984, Eberhard 1985). How- ever, the validity of spermatophore number as a measure of female mating frequency is based upon a number of assumptions which need to be verified before inferences about mating systems are made (Burns 1968, Lederhouse et al. 1989, Braby 1996). Here, I re- port the number of spermatophores found in a field sample of fe- males of the lycaenid butterfly Callophrys xami (Reakirt); and, based upon previous information on the mating behavior and sper- matophore production patterns in this butterfly (Cordero 1993, 1998), I discuss possible biases incurred when using such a measure as an estimate of female mating frequency. I sampled females during eight sunny days, between 28 Decem- ber 1989, and 23 January 1990 (this multivoltine species reaches its highest density between October and January (Soberén et al. 1988)) in the Pedregal de San Angel ecological reserve, located in the south of Mexico City (description of the area in Soberon et al. 1988). All females observed during these days were collected and frozen until dissection. I measured the length of the right forewing of each fe- male with a calliper. I used this length as a measure of body size, considering that there was a positive correlation between wing length and body weight in a laboratory-reared sample of females (r = 0.8, p < .001, n = 27). I also determined the degree of female wing wear using the following scale: (1) similar to a recently emerged adult (wings mostly green on the ventral side with intact margins), (3) very worn female (wings mostly brown on the ventral side with worm margins), and (2) all individuals intermediate between (1) and (3). I evaluated the frequency of “successful” copulations by females (ie., copulations that resulted in spermatophore transfer) by count- ing the number of spermatophores and spermatophore remains in the corpus bursae. 14 12- Wing wear class: G1 2 See Number of spermatophores Number of females Fic. 1. Distribution of females with different number of sper- matophores and the relationship between number of sper- matophores and wing wear category. The number of spermatophores in the 28 females I collected ranged from 0 to 3 (Fig. 1). The percentage of females without sper- matophores was 32.1%, with one spermatophore was 46.4%, and with more than one spermatophore was 21.4%. The mean + SD number of spermatophores found in females with at least one sper- matophore was 1.37 + 0.6. I looked for a relation between female wing length and spermatophore number with Spearman correlation because wing length was not normally distributed. This correlation was not significant (r, = —0.07, p > 0.05, n = 27). Since all females collected were in wing wear conditions 1 or 2, I compared the num- ber of spermatophores of females in each condition with a Mann- Whitney U test without finding significant differences (U = 54, p > 0.05: Fig. 1). Since the work of Drummond (1984) is the only extensive sum- mary presenting data from Lycaenidae, I use the data in that paper as a reference. Considering average number of spermatophores per mated female, maximum number of spermatophores, and propor- tion of females multiply mated, female Lycaenidae relative to other lepidopterans show the lowest degree of polyandry, comparable only with the Satyrinae (Drummond 1984). However, C. xami shows some differences when compared with the four lycaenid species in- cluded in Drummond (1984). The average copulation frequency es- timated for mated females (1.37 + 0.6), the maximum number of spermatophores (3), and the proportion of females multiply mated (21.4%) in C. xami is higher than in the other four lycaenids (ranges of average copulation frequency values: 1.05-1.17; maximum num- ber of spermatophores: 2 in the four species; and proportion of fe- males multiply mated: 3.7-12.7%). Furthermore, average number of copulations, maximum number of spermatophores, and propor- tion of females multiply mated could be underestimated in C. xami, since no females in the very worn (“old”) wing wear condition were collected (Fig. 1). A sampling bias could exist if “old” females were more difficult to detect or to capture. However, our research group has been studying this species in the field for more than 10 years, and we have no evidence of any greater difficulty in observing and catching “old” females. It is possible that most females do not live long enough to become very worn and, therefore, are rare; in this case our estimates of copulation frequency would be unbiased. On the other hand, since outside the sampling period we have observed very worn C. xami females in the field, it is also possible that the abundance of “old” females varies in time as a result of, for example, varying predation pressure or weather conditions. Under these con- ditions, average and maximum number of spermatophores could vary with time depending on the age structure of females. The method used to evaluate female copulation frequency in the field is based on three assumptions (modified from Drummond 1984): Copulation always results in spermatophore transfer. In C. xami this is not true because there are some copulations of very short duration that do not result in the transfer of a spermatophore (Cordero 1993, 1998). However, these “interrupted” copulations are not common in the field (0/18 copulations observed in 1983-1985 and 2/27 copulations observed in 1989-1990; Cordero 1993). On the other hand, although the existence of interrupted copulations pre- vented the estimation of the total number of copulations performed by females, the figures obtained could be good estimates of the number of copulations resulting in spermatophore transfer. Males transfer only one spermatophore per copulation. In C. xami this is not true since in laboratory experiments we observed three copulations in which different males transferred two sper- matophores during one copulation (Cordero 1998). Violation of this assumption results in an overestimation of copulation frequency. 170 However, if the frequency of copulations resulting in the transfer of two spermatophores in the laboratory is a good estimate of their fre- quency in the field (3/199 copulations observed in the laboratory), its quantitative effect should be small. Spermatophores always leave recognizable remains within the corpus bursae of the female. This is not true in C. xami since in the laboratory it was not always possible to observe clear sper- matophore remains in very old females that had laid most of their eggs (pers. obs.). However, judging from wing wear, no female in this condition was sampled (see paragraph four above). In conclusion, the possible violation of the first and the last as- sumptions, and the fact that some of the females may have mated again had they not been collected, results in an underestimation of the frequency of copulations in females; whereas the fact that some males transfer more than one spermatophore in one copulation re- sults in an overestimation of the number of copulations. However, judging from the low frequency of “interrupted” copulations (4.4%), very worn females in the field (at least during the sampling period), and copulations resulting in the transfer of two spermatophores (1.5%), I conclude that spermatophore counts are a reasonably good estimate of female copulation frequency in C. xami. ACKNOWLEDGMENTS I thank Gabriela Jiménez and Dr. Rogelio Macias for their valu- able technical help, and Dr. J. M. Burns and an anonymous reviewer for their comments. This research was supported by a Consejo Na- cional de Ciencia y Tecnologia (México) scholarship. LITERATURE CITED Braby, M. F. 1996. Mating frequency in bush-brown butterflies (Nymphalidae: Satyrinae). J. Lepid. Soc. 50:80-86. Journal of the Lepidopterists’ Society 53(4), 1999, 170-172 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Burns, J. M. 1968. Mating frequency in natural populations of skippers and butterflies as determined by spermatophore counts. Proc. Nat. Acad. Sci. U.S.A. 61:852—-859. CorDERO, C. 1993. The courtship behavior of Callophrys xami (Lycaenidae). J. Res. Lepid. 32:99-106. . 1998. Ecologia del Comportamiento Sexual de los Machos de la Mariposa Callophrys xami, con Algunas Consideraciones Acerca de la Evoluci6n del Semen de Insectos, Doctoral Thesis, UACPyP/CCH, UNAM, México. DruMMoND III, B. A. 1984. Multiple mating and sperm competi- tion in Lepidoptera, pp. 291-370. In R. L. Smith, (ed.), Sperm competition and the evolution of animal mating systems. Acad- emic Press, New York. EBERHARD, W. G, 1985. Sexual selection and animal genitalia. Har- vard University Press, Cambridge, U.S.A. . 1996. Female control. Sexual selection by cryptic female choice. Princeton University Press, Princeton, U.S.A. LEDERHOUSE, R. C., M. P. AyRES & J. M.ScrRIBER. 1989. Evaluation of spermatophore counts in studying mating systems of Lepi- doptera. J. Lepid. Soc. 43:93-101. SOBERON, J., C. CORDERO, B. BENREY, P. PARLANGE, C. GARCIA- SAEZ & G. BERGES. 1988. Patterns of oviposition by Sandia xami (Lepidoptera, Lycaenidae) in relation to food plant ap- parency. Ecol. Entomol. 13:71-79. CARLOS CORDERO. Instituto de Ecologia, Universidad Nacional Auténoma de México, Apdo. Post. 70-275, C.P. 04510 D.E., and Cen- tro de Investigaciones Fisiologicas, Universidad Auténoma de Tlax- cala, Apdo. Post. 262, C.P. 90070 Tlaxcala, Tlaxcala, MEXICO (Ad- dress for correspondence) Received for publication 5 April 1999; revised and accepted 16 De- cember 1999. ADDITIONAL NOTES ON PROSERPINUS CLARKIAE AND ARCTONOTUS LUCIDUS (SPHINGIDAE) LIFE HISTORIES FROM THE PACIFIC COAST OF NORTH AMERICA Additional key words: Onagraceae, Rubiaceae, Gayophytum, Galium, Clarkia breweri, Clarkia modesta, Camissonia. Host associations for Proserpinus clarkiae (Boisduval) and Arctonotus lucidus (Boisduval) have recently been documented. Proserpinus clarkiae was found using Clarkia unguiculata (Lindley) in nature (Osborne 1995). Here, I compare results of my life history work on P. clarkiae with other results (Hardy 1959) on this species. The life history of A. lucidus is also known (Comstock & Henne 1942). However, the first natural host associations for A. lucidus were made by photographs and collections from Clarkia species in California, and are presented here along with observations on cap- tive rearing of this moth. The immature stages of these related sphingid species have been confused in the field by some, possibly due to their sympatry, common use of Clarkia hosts, and superficial resemblance. Thus, I will also discuss morphological differences among these and other sympatric Clarkia feeding sphingids. In presenting the biology of P. clarkiae (Osborne 1995), I re- peated the assertion made by Hodges (1971) that its life history was unknown. Since that time, Dr. Frederick Rindge (American Mu- seum of Natural History) has drawn my attention to a life history of P. clarkiae that predates both works. Larvae and a pupa reared from Vancouver Island (Hardy 1959) were described by Hardy (1959), and match the immatures of P. clarkiae from California. Hardy ob- tained seven ova by confining females over potted Galium aparine (Lewis & Szweykowski) (Rubiaceae). He reared at least one individ- ual to pupation on that plant, but a field host was not given. The sin- gle fifth instar larva of P. clarkiae from Vancouver Island had the lat- eral dark blotches contiguous in an undulating line, a trait consistent with some (< 5%) of the California material I reared (most Califor- nia larvae had oblique blotches disjunct) (Osborne 1995). This dark form may be typical of cool, wet, north coastal localities, where darker maculation may impart local selective advantages, or may be an artifact of captive rearing. Dr. Robert Raguso, who studied sphingid pollination of Clarkia species in central California (see Raguso & Pichersky 1995, Raguso et al. 1996, Raguso & Light 1998), sent me several suspected Pros- erpinus larvae, a reared pupa, and a photograph (Fig. 1) of a fifth in- star larva in nature on Clarkia breweri (A. Gray) E. Greene. These specimens were all collected from C. breweri and Clarkia modesta (Jepson) at Del Puerto Canyon, Stanislaus Co., California in May, 1991. However, instead of P. clarkiae, all were determined (by KHO) to be Arctonotus lucidus, a closely related species from a monotypic genus. Early instar A. lucidus larvae may be separated from P. clarkiae by the presence of a black anal horn which is absent in P. clarkiae. Fifth instar A. lucidus lose the anal horn, but have dor- sal and lateral markings of olive green (but briefly black just after molt [Comstock & Henne 1942]), not black or gray as in P. clarkiae. In addition, A. lucidus can be distinguished from P. clarkiae on the basis of dorsal, transverse intersegmental lines of tan or cream breaking the olive green field, and ventral whitish or gray. The ground color in fifth instar A. lucidus larvae is variable (Comstock & Henne 1942), ranging from black to olivacious green to light green, to pink (Comstock & Henne 1942; D. Rubinoff pers. comm.; K. H. Osborme unpubl. obs.). VOLUME 53, NUMBER 4 171 Fic. 1. Fifth instar Arctonotus lucidus larva on Clarkia breweri at Del Puerto Canyon, Stanislaus Co. CA., May, 1991. Photograph by Robert Raguso. Raguso’s photograph of an A. lucidus larva on C. breweri, and collections of A. lucidus from C. breweri and C. modesta, represent the first natural host records for this moth. Raguso (pers. comm. 1995) has seen [these?] larvae on Clarkia gracilis sonomensis (Hitche.) near Lake Berryessa, Napa Co., California. (These records must be considered as likely A. lucidus but could possibly be P. clarkiae.). Additionally, one wandering fifth instar A. lucidus (de- termined by KHO) was found by M. Lynn (pers. comm.) in May, 1997 in the immediate vicinity of abundant Camissonia bistorta (Nutt.), Camissonia strigulosa (Fisck. & Meyer) Raven [=Oenothera contorta Munz], and Clarkia purpurea (Curt.) Nels. & Macbr. at Lake Skinner, Riverside Co., California, suggesting these plants as possible hosts. The Cammissonia species are used as larval hosts by related sphingids in southem California, Euproserpinus phaeton Grote & Robinson using C. bistorta (Osborne 1995), Euproserpinus euterpe Hy. Edwards using C. strigulosa (Tuskes & Emmel 1981, K. H. Osborne unpubl. obs.), and Hyles lineata (L.) using both (K. H. Osbome unpubl. obs.). Clarkia breweri is restricted to central Cali- fornia from Alameda Co. south to Fresno Co. (Munz 1959) and C. modesta ranges through California from Tehema Co. south to Santa Barbara Co. (Hickman 1993). The wide range of A. lucidus (Holland 1903, Hodges 1971) from British Columbia at least as far south as San Diego Co., California (Brown & Donahue 1989; Osborne un- publ. obs.) indicates A. lucidus must use other host plant species. Galium, suitable for P. clarkiae in captivity (Hardy 1959), was re- jected by A. Iucidus as were two unnamed Oenothera species (Com- stock & Henne 1942). In captivity, A. lucidus larvae would accept leaves of C. breweri, C. modesta and Clarkia affinis (H. Lewis & M. Lewis) (Raguso pers. comm.), but were hesitant to accept Clarkia unguiculata (Lindley) and Fuchsia (K. H. Osborne unpubl. obs.; Ra- guso pers. comm.). Most prepupal A. lucidus larvae wandered and died without pupating in dry, sandy soil, and the one that did pupate, about 3 cm below ground in loose gravel, was attacked by mold and never emerged (K. H. Osborne unpubl. obs.; Raguso pers. comm.). Dan Rubinoff (pers. comm.) reported success getting A. lucidus to pupate on moist potting soil and when reared by Comstock & Henne (1942) larvae pupated as deep in the soil as possible (in cages). Hyles lineata is common on Clarkia (C. unguiculata at Gates Canyon, Solano Co, CA [unpublished records] and C. breweri, at Del Puerto Canyon [Raguso]) when A. lucidus and P. clarkiae may be present. Hyles lineata is easily distinguished from larvae of A. lu- cidus and P. clarkiae by its prominent orange or yellow anal horn in all larval stages and by distinctive (but variable) longitudinal mark- ings (Hodges 1971). ACKNOWLEDGMENTS I thank Dr. Frederick Rindge for informing me of prior life his- tory work on P. clarkiae, Dr. Robert Raguso who sent me immatures, photographs and notes, and reviewed this manuscript. Daniel Rubi- noff also reviewed the manuscript and made helpful suggestions. Fi- nally, my deepest gratitude goes to an anonymous reviewer who scoured the manuscript, making many insightful comments and im- provements. LITERATURE CITED Brown, J. W. & J. P. DoNAHUE. 1989. The Sphingidae (Lepi- doptera) of Baja California, Mexico. J. Lepid. Soc. 43:184—-209. 72 CoMSTOCK, J. A. & C. HENNE. 1942. The early stages of Arctonotus lucidus Bdy. (Lepidopt.). Bull. South. Calif. Acad. Aci. 41:167-171. Harpy, G. E. 1959. Notes on the life histories of four moths from southern Vancouver Island. Entomol. Soc. British Columbia, Proc. 56(4):49-52. Hickman, J. C. (ed). 1993. The Jepson manual: higher plants of California. University of California Press, Berkeley, Ca. Hopces, R. W. 1971. Moths of North America, north of Mexico. Fascicle 21 (Sphingidae). 170 pp. HOLLAND, W. J. 1903. The moth book. Doubleday, Page and Co., New York. 479 pp. OsporNE, K. H. 1995. Biology of Proserpinus clarkiae (Sphingi- dae). J. Lepid. Soc. 49(1):72-79. Munz, P. A. 1959. A California flora. University of California Press, Berkeley. 1681 pp. Racuso, R. A. & E. PicHErRSKy. 1995. Floral volatiles from Clarkia breweri and C. concinna (Onagraceae)—recent evolution of flo- ral scent and moth pollination. Plant Systematics and Evol. 194(1—2):55-67. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Racuso, R. A., D. M. Licnut & E. Picuersky. 1996. Electroan- tennogram responses of Hyles lineata (Sphingidae, Lepi- doptera) to volatile compounds from Clarkia breweri (Ona- graceae) and other moth-pollinated flowers. J. Chem. Ecol. 22(10):1735-1766. Racuso, R. A. & D. M. Licut. 1998. Electroantennogram re- sponses of male Sphinx perelegans hawkmoths to floral and ‘green-leaf volatiles’. Entomologia Experimentalis et Applicata. 86(3):287-293. TuskEs, P. M. & J. F EMMEL. 1981. The life history and behavior of Euproserpinus euterpe (Sphingidae). J. Lepid. Soc. 35:27-33. KENDALL H. OsBorNeE. 24292 Lysanda Drive, Mission Viejo, Cali- fornia, USA 92691 Received for publication 3 June 1999; revised and accepted 6 Febru- ary 2000. Journal of the Lepidopterists’ Society 53(4), 1999, 173-176 Abies, 26 Accuminulia, 60 buscki, 60 longiphallus, 60 Achillea lanulosa, 74 millefolium, 74 Actias selene, 134 luna, 134 Adamski, D., 29 Aglossa, 8 caperalis, 2 Agraulis, 17 Aiello, 142 Alberta, Canada, 127 Albu, V., 45 allelochemicals, 104 alpine habitat, 32 Amathes glareosa, 99 Amazonia, 65 Anopheles, 106 Antheraea, 134 hartii, 134 pernyt, 134 polyphemus, 134 antioxidant, 104 anti-predation, 130 apricot, 60 Arbutus menziesii, 22 xalapensis, 27 Archipini, 114 Arctiidae, 72 Arctonotus lucidus, 171 Arctostaphylos patula, 22 Area de Conservation Guanacaste (ACG), ian Arecaceae, 142 Arescoptera, 7 Argema mittrei, 134 Argentulia, 61 Argyrotaenia, 114 bialbistriata,114 coconinana, 115 octawvana, 114 ponera, 115 spinacallis, 114 unda, 114 Arispe, 8 Asclepiadaceae, 37 Asclepias, 37 asperula, 38 curassivica, 15, 38 latifolia, 38 oenotheroides, 38 syriaca, 43 texana, 38 viridis, 38 INDEX FOR VOLUME 53 (new names in boldface) Asia, 7 Attacini, 50, 133 Attacus atlas, 134 Australia, 1 Balcazar-Lara, M. A., 45 Basilarchia archippus archippus, 104 arthemis astyanax, 104 arthemis arthemis, 104 Berenbaum, M., 104 Bicyclus, 26 Bidens pilosa, 15 biennialism, 127 biodiversity, 65, 153 bird dropping, 104 Biston betularia, 99 betularia cognataria, 101 black swallowtail, 104 Blastobasinae, 29 Blastobasini, 29 Blepolenis batea, 146 Bolivia, 65 Boloria acrocnema, 32 improba, 32 Brassolinae, 142 Brassolis astyra, 142 astyra astyra, 142 isthmia, 142 sophorae, 142 Brazil, 11, 65, 108 British Columbia, 22, 74 Brown, J. W., 60, 114 Burns, J. M., 77 Byrsonima crassifolia, 89 Cacotherapiini, 4 Calathea latifolia, 142 California, 22 Caligo amphirhoe, 148 arisbe, 147 atreus, 147 atreus dionysos, 147 beltrao, 147 eurilochus, 147 eurilochus brasiliensis, 147 eurilochus sulanus, 147 illoneus, 142 illioneus oberon, 147 illioneus pampeiro, 147 idomeneus, 142 martia, 147 memnon, 142 memnon memnon, 148 memnon telamonius, 148 oberthurii oberthurii, 148 oileus, 148 oileus scamander, 148 placidianus, 148 praxsiodus, 148 prometheus epimetheus, 148 teucer, 148 Callophrys xami, 170 Callosamia, 22, 49 angulifera, 26, 134 promethea, 50, 134 securifera, 26, 134 Calvert, W. H., 37 Cammisonia, 171 bistorta, 172 strigulosa, 172 Carcelia reclinata, 72 catchability, 138 caterpillar, 22 Catoblepia amphirhoe, 148 orgetorix championi, 148 Catocrosis, 8 Ceanothus integurrimus, 22 Ceanothus silk moth, 22 Celtis, 130 censusing, 138 Central America, 142 Cephise, 77 nuspesez, T7 Chao, 65 Chiapas, Mexico, 153, 155 Chile, 60 Chileulia, 64 Chrysanthemum leucanthemum, 74 Chrysauginae, 1 Chrysobalanaceae, 77 Citheronia regalis, 134 Clark, R. M., 49 Clarke, C. A., 99 Clarkia breweri, 171 gracilis, 172 modesta, 171 purpurea, 172 unguiculata, 171 Cleora repandata, 99 Clepsis consimilana, 74 climate change, 32 Cocytius antaeus, 129 cognataria, 99 Coliadinae, 159 collecting activity, 32 Collins, M. M., 22 Colorado, 32 174 Combretaceae, 77 Conium maculatum, 72 copulation, 170 Cordero, 171 Coscinocera hercules, 134 Costa Rica, 29, 65, 77 Cotesia, 24 Crambinae, 1 Crambini, 7 Cramer, A., 114 crypsis, 22 Cucullinae, 55 Cupressaceae, 55 Curatella americana, 89 Curena, 7 Cynanchum, 38 barbigerum, 40 Cyrtostachys, 142 Danaus plexippus, 37 gilippus, 38 Dasyophthalma rusina, 148 developmental plasticity, 22 developmental rate, 72 Dichrorampha guenceana, 74 sedatana, 74 vancouverana, 74 Dione, 17 Diptera, 72 distribution, 133 Dolichomia, 8 Douglas-fir, 22 Drephalys, 77 alcmon, 77 eous, 79 helixus, 78 heraclides, 78 kidonoi, 77 miersi, 79 mourei, 79 olva, 79 olvina, 79 opifex, 79 phoenice, 79 phoenicoides, 79 Drosophila melanogaster, 49, 106 Dryas, 17 Dynastor darius, 148 darius mardonius, 148 darius stygianus, 148 macrosiris, 148 napoleon, 148 Ecuador, 65 El Ocote Reserve, 153 Emilia sonchifolia, 15 Endotricha, 1 flammealis, 2 Endotrichinae, 2 Endotrichini, 1 Epidendrum fulgens, 15 epipaschiine, 1 Erynnides, 78 Erynnis, 78 Eryphanis aesacus bubocula, 148 automedon, 148 polyxena lycomedon, 148 reevesti, 148 Eudamus, 78 helixus, 78 Eueides, 16 Euliini, 60 Eumorpha, 26 Eupackardia calleta, 133, 134 Eupatorium laevigatum, 15 Euproserpinus euterpe, 172 phaeton, 172 Eurasia, 74 Eurema elathea, 159 hecabe, 165 Euryalus cedrosensis, 49 Fabricius, 108 f. carbonaria, 102 feeding preference, 108 female remating, 170 fire ant, 37 Flaim, D., 127 Florida viceroy, 126 Forestiera angustifolia, 134 frass chains, 130 Fraxinus greggti, 134 Freitas, A. V. L., 11, 131 f. swettaria, 102 furanocoumarin, 104 Galapagos, 129 Galium, 171 aparine, 171 Galleriinae, 2 Gauna , 7 Gayophytum, 171 genitalia, 1, 80, 114 morphology, 5 Geometridae, 99 G6mez- Nucamendi, O. L., 153 Gonodontis bidentata, 99 Grant, B. S., 99 grape, 60 Grapholitini, 74 Guanacaste, 29 Gurania, 15 Heliconia latispatha, 142 heliconian butterflies, 108 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Heliconiini, 11 Heliconius, 165 besckei, 20 erato, 108 erato phyllis, 11 ethilla nacaea, 17 melpomene nanna, 19 nattereri, 19 numata robigus, 17 sara apseudes, 17 Hemiceras, 65 Hemileuca, 50 Hepialidae, 127 Herculia, 8 psammioxantha, 2 Hesperiidae, 77 Heterocera, 153 Heyda salicella, 169 hibernacula, 126 Hinsdale County, Colorado, 32 Hirtella, 77 racemosa, 77 holarctic, 74 host records, 142 host-plant selection, 108 hostplant, 133 Hyalophora, 49 cecropia, 49, 134 columbia, 134 columbia columbia, 24, 49 columbia gloveri, 24, 49 euryalus, 22, 49, 134 gloveri, 134 Hypatopa interpunctella, 31 tapadulcea, 29 Hyles lineata, 172 Hypolimnas bolina, 138 Hypsopygia, 8 Ichneumon sp., 72 immigrant, 169 Impatiens walleriana, 15 Inape, 61 industrial melanism, 99 invertebrate inventory, 153 island colonization, 129 Ithomiinae, 16 jackknife, 65 Janzen, D. H., 77 Jones, R. W., 153 Karban, R., 73 Kemp, D. J., 138 Kruger, M., 47 Lantana camara, 15 Larix, 26 laricina, 50 larval diapause, 126 larval food plants, 142 VOLUME 53, NUMBER 4 larval integument, 104 leaf age effects, 108 Leschenaultia adusta, 72 Leucophyllum frutescens, 134 Libytheana carinenta, 130 Libytheinae, 130 life cycle, 22 life history, 171 light traps, 129 Ligustrum, 133 Limenitis archippus floridensis, 126 Lithophane leautieri, 59 lemmeri, 55 longior, 59 subtilis, 59 thujae, 55 Los Tuxtlas, Veracruz, 155 lupine, 72 Lupinus albicaulis, 72 albifrons, 72 arboreus, 72 Lycaenidae, 170 Macena, 1 madrone, 22 Maine, 74 Malphighiaceae, 77 mandibular morphology, 133 manzanita, 22 Mapeta, 8 Marantaceae, 142 mark-release-recapture, 11, 138, 159 Matelea, 38 reticulata, 38 mating frequency, 170 mating system, 170 Melanargia galathea, 140 Mexico, 37, 114 Michigan, 55 Micromasta, 1 isoldalis, 7 Micronix amsel, 1 nivalis, 7 migration, 37 Mikania lundiana, 15 milfoil, 74 milkweeds, 37 Miller, W. E., 75, 169 Monarch butterfly, 37 monocotyledonous plants, 142 Mor6n-Rios, 153 morphological characters, 1 morphology, 1, 114 Morro de Japui, 11 de Vorturua, 11 Mount Uncompahgre, 32 Musa saientum, 142 Musaceae, 142 Narope cyllastros, 149 cyllastros cyllastros, 149 cyllastros testacea, 149 natural regions, 37 Nemophila, 72 Nemoris, 26 Neodavwisiai, 8 neotropical region, 153 Neotropics, 1, 29, 60 New Brunswick, 55 New Hampshire, 74 New York, 60, 74 Noctuidae, 1, 55, 129 Nola pygmaea, | Nolinae, 1 North America, 37, 74, 171 northern white cedar, 55 Notodontidae, 65, 133 Nymphalidae, 108, 130, 138, 142, 165 Ocrasa, 8 Oenothera, 172 Olethreutinae, 74, 169 Onagraceae, 171 ontogenetic development, 133 Opodiphthera eucalypti, 134 Opotera aorsa, 149 staudingeri, 149 syme, 149 Opsiphanes, 142 bogotanus, 149 bogotanus bogotanus, 149 cassieae, 149 cassieae cassiculus, 149 cassieae lucullus, 149 cassina aiellae, 149 cassina fabricii, 149 invirae, 149 invirae amplificatus, 150 invirae cuspidatus, 150 merianae, 150 quiteria, 150 quiteria badius, 150 quiteria meridionalis, 150 quiteria quirinus, 150 tamarindi, 150 tamarindi sikyon, 150 tamarindi tamarindi, 150 Osborne, K. H., 173 oviposition, 37 ox-eye daisy, 74 Palaearctic species, 169 Panama, 142 Pantepui, 90 Papilio, 26 polyxenes, 104 xuthus, 104 Papilionidae, 16, 104, 165 Paradrephalys, 77 croceus, 79 dumeril, 79 oria, 79 oriander, 79 talboti, 79 tortus, 79 Paradros, 77 parasitism, 72 Passiflora alata, 15 capsularis, 15 edulis, 15 jileki, 15 suberosa, 108 Passifloraceae, 11 passion vines, 108 Passoa, S., 133 Passoa, V., 133 Pastinaca sativa, 105 peach, 60 Pedaliodes, 90 chaconi, 95 darmarmelsi, 95 dejecta, 96 manis, 95 napaea, 96 prytanis, 95 roraimae, 95 terramaris, 90 yutajeana, 90 Peigler, R. S., 47 Penz, C. M., 142 peppered moths, 99 Perforadix, 1 sacchari, 1 Perisseretma, 7 Peru, 65 Perula, 7 pest species, 60 Petroselinum crispum, 105 Phacelia, 72 phenotypic plasticity, 22 Phigalia titea, 99 photoperiod, 126 Phycitinae, 2 phylogeny, 49, 114 Pieridae, 16, 159 Pieris brassicae, 104 pigmentation, 104 Plagiobothrys, 72 Plantago, 72 Platt, A. P., 127 Platyprepia virginalis, 72 Pleistocene, 90 plum, 60 Poaceae, 142 Pococera complex, 4 Pogue, M. G., 65 Pollard transect, 32 175 176 polymorphism, 22 polyphenism, 11, 159 ponera group, 114 population biology, 159 population estimation, 138 Populus, 169 Praepronophila, 93 Proeulia, 60 Pronophiline, 90 Proserpinus clarkiae, 171 Proteaceae, 77 Protopedaliodes, 90 kukenani, 93 profauna, 90 ridouti, 90 Prunus, 50, 60 armeniaca, 60 domestica, 60 emarginata, 22 persica, 60 Pseudasopia, 8 Pseudodrephalys, 77 Psuedotsuga menziesii, 22 Puerto Rico, 7 Puntarenas, 29 Pyralidae, 1 Pyralinae, 1 Pyralini, 1 Pyralis, 8 farinalis, 1 Pyraloidea, 7 Pyraustinae, 1 Pyrez, T. W., 90 Pyrginae, 77 Quercus alba, 50 Ramos, R. R., 11 red raylets, 11 Redcloud Peak, 32 Rhodnius prolixus, 106 RFLP, 49 Rodrigues, D., 108 Roque-Albelo, L., 129 Rothschildia, 133 cincta, 134 erycina, 134 forbesi, 134 lebeau, 26, 134 orizaba, 134 Roupalla montana, 77, 89 Rubiaceae, 171 Rubus rosaefolius, 15 ursinus, 72 Rudinei Pires Moriera, 108 Rumex, 72 Sabourin, M., 169 Saccharum spontaneum, 142 Salicaceae, 32 Salix, 169 arctica, 32 caroliniana, 126 reticulata nivalis, 32 Samia cynthia, 134 Sao Vicente, 11 Sarcostemma, 38 crispum, 40 cynanchoides, 40 Saturnia mendocino, 27 Saturniidae, 22, 49, 133 Satyridae, 43 satyrine, 90 Scenedra, 7 Scenidiopsis, 7 Schmidt, B. C., 129 Seidl, A. L., 32 semivoltine, 127 Shaffer, M., 1 snow willow, 32 Solenopsis invicta, 37, 43 Solis, M. A., 1 southwest, 114 spermatophore number, 170 Sphingicampa, 26 Sphingidae, 129, 133, 153, 156, 171 Spodoptera exigua, 100 Srygley, R. B., 142 Stachytarpheta polyura, 15 Sthenopus argenteomaculatus, 127 purpurascens, 127 quadriguttatus, 127 storage excretion, 104 subspecies, 126 Subtranstillaspis, 61 Date of Issue (Vol. 53, No. 4): 22 June 2000 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Sufetula, 1 grumalis, 7 pygmaea, 1 systematics, 60 Taboga, 1 inis, 7 Tachinidae 72 Tanyethira, 7 taxonomy, 49 tepuyes, 90 Texas, 37, 133 Thelaira americana, 72 bryanti, 72 thermoperiod, 126 Thomas, A. W., 55 Thuja occidentalis, 55 Thysania zenobia, 129 Timmerman, S., 104 Tithonia speciosa, 15 Tortricidae, 60, 114, 169 Tortrix ponera, 117 tritrophic interactions, 72 Trixis antimenorrhoea, 15 Troidini, 16 tropical forests, 153 Uncompahgre fritillary, 32 urban butterflies, 159 uric acid, 104 Utetheisa ornatrix, 166 Vale do Rio Quilombo, 11 Vancouver, 74 Varifula, 61 Venezuela, 90 Viloria, A. L., 90 Vitis, 60 Washington, 74 Webster, R. P., 55 Western Hemisphere, 9 winter moths, 55 Wisconsin, 55 Zalucki, M. P., 138 ee a a EDITORIAL STAFF OF THE JOURNAL M. Deane Bowers, Editor Entomology Section University of Colorado Museum Campus Box 218 University of Colorado Boulder, Colorado 80309, U.S.A. email: bowers@spot.colorado.edu Associate Editors: Gerarbo Lamas (Peru), KeNnELM W. Puiir (USA), Rosert K. Rossins (USA), Feix A. H. Speriinc (USA), Davip L. Wacner (USA), Curister WikLUND (Sweden) NOTICE TO CONTRIBUTORS Contributions to the Journal may deal with any aspect of Lepidoptera study. Categories are Articles, Profiles, General Notes, Technical Comments, Book Reviews, Obituaries, Feature Photographs, and Cover Illustrations. Obituaries must be authorized by the President of the Society. Requirements for Feature Photographs and Cover Illustrations are stated on page 111] in Volume 44(2). Journal submissions should be sent to the editor at the above address. Short manuscripts concerning new state records, current events, and notices should be sent to the News, Phil Schappert, Dept. 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For authors without institutional support, page charges are $25 per Journal page. For authors who are not members of the Society, page charges are $75 per Journal page. Authors unable to pay page charges for any reason should apply to the editor at the time of submission for a reduced rate or free publication. Authors of Book Reviews and Obituaries are exempt from page charges. Correspondence: Address all matters relating to the Journal to the editor. PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A. CONTENTS DistTRIBUTION AND HOSTPLANT RECORDS FOR EUPACKARDIA CALLETA FROM SOUTHEASTERN TEXAS WITH NOTES ON MANDIBULAR MORPHOLOGY OF ATTACINI (SATURNUDAE) Valerie A. Passoa and Steven PASSO 2... ce eceee cece eeee cece eee METHOD OF HANDLING AFFECTS POST-CAPTURE ENCOUNTER PROBABILITIES IN MALE HyPoLiMNas BOLINA (L..) (NYMPHALIDAE) Darrell.J:\Kemp and Myron3P. Zaluckt 225 22) 22 ae st ee a, 2 EARLY STAGES OF CALIGO ILLIONEUS AND C. IDOMENEUS (NYMPHALIDAE, BRASSOLINAE) FROM PANAMA, WITH REMARKS ON LARVAL FOOD PLANTS FOR THE suBFAMILY Carla M. Penz, Annette Aiello and Robert B. Srygley 2.0.0.0. Tue SpHincmar (HETEROCERA) OF THE “EL Ocore” Reserve, Curaras, Mexico Olga Lidia Gémez-Nucamendi, Robert W. Jones and Alejandro’ Mor6n-Rios 2a POLYPHENISM AND POPULATION BIOLOGY OF EUREMA ELATHEA (PreRIDAE) IN A DISTURBED ENVIRONMENT IN TROPICAL Braz Fabio Vanini, Vinicius Bonato and André Victor Lucci Freitas cece General Notes Is spermatophore number a good measure of mating frequency in female Callophrys xami (Lycaenidae)? 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