aunty iP Cb { t ' ? S ( ¥ i + i bi M t ipa : y ' f int re 1 i) * a i Volume 57 Number 1 9 May 2003 ISSN 0024-0966 SEP 2 7 2003 LIBRARIES ociety Published quarterly by The Lepidopterists’ Society THE LEPIDOPTERISTS’ SOCIETY EXECUTIVE CounciL Lawrence F. Gary, President Scotr E. Mituer, Vice President J. Donatp Larontaine, Immediate Past President Konrap Frepier, Vice President Joun H. Accorn, Vice President Ketry Ricners, Treasurer Ernest H. Wiiiiams, Secretary Members at large: e J. Bolling Sullivan James K. Adams William E. Conner Philip DeVries Wayne F. Wheling Rebecca Simmons David Ahrenholz Jeffrey R. Slotten Charles V. Covell Jr. EprroriaL BoarD Roserr K. Rossrs (Chairman) Carta M. Penz (Journal) WitiiaM E. Mitter (Memoirs) Puitur J. Scuappert (News) Joun A. SnyDER (Web) Honorary Lire MEMBERS OF THE SOCIETY ‘Cuarzes L. Remincton (1966), E. G. Munroe (1973), lan F. B. Common (1987), Joun G. Franctemonrt (1988), Lincoin P. Brower (1990), Douctas C. Fercuson (1990), | Hon. Miriam Rotruscuip (1991), CLraupe 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 pro- _ mote the science of lepidopterology in all its branches, . . . to issue a periodical and other publications on Lepidoptera, to facilitate the exchange of specimens and ideas by both the professional worker and the amateur in the field; to secure cooperation in all mea- be sures’ directed towards these aims. Membership in the Society is open to all persons interested in the study of Lepidoptera. All Hemtees receive the Journal isd the ida News of The Lepidopterists’ Society Prospective members should send to the Assistant Treasurer full dues for the current year, to- gether with their full name, address, and special lepidopterological interests. In alternate years a list of members of the Society i is is- es sued, with addresses and special interests. 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Treasurer, 9417 Carvalho Court, Bakers- field, CA 93311, The additional cost for members outside the U.S. is to cover mailing costs. Journal of The Lepidopterists’ Society (ISSN 0024-0966) is published quarterly by The Lepidopterists’ Society, % Los Angeles County Museum of Natural History, 900 Exposition Blvd., Los Angeles, CA 90007-4057. Periodicals postage paid at Los Angeles, CA — and at additional mailing offices. POSTMASTER: Send address changes to The Lepidopterists’ Society, % Natural History Museum, 900 Exposition Blvd., Los Angeles, CA 90007-4057. Cover illustration: Detail of the wings of Cyrestis thyodamas (Nymphalidae) female, dry season form; Puli, Formosa. JOURNAL OF THE DuerrmortrRis1s: SOC LE IY Volume 57 2003 Number 1 Journal of the Lepidopterists’ Society 57(1), 2003, 1-16 ANT-ASSOCIATION AMONG SOUTHERN AFRICAN LYCAENIDAE ALAN HEATH! Department of Zoology, University of Cape Town, Rondebosch 7700, South Africa AND ANDRE J. M. CLAASSENS 203, High Level Road, Sea Point 8005, South Africa ABSTRACT. Known life history data, ant-associations and larval feeding habits for southern African lycaenids are summarized and dis- cussed with a focus on recently acquired knowledge. Of the 392 lycaenid species represented in southern Africa, over three quarters are ant- associated, two thirds of which are obligate. The Poritiinae is represented by two tribes of algae/lichen feeders that are not ant-associated. Of the Miletinae, three quarters are obligately ant-associated, and all are believed to prey on Homoptera or their secretions during their larval stage. Within the Lycaeninae, the tribe Theclini are all believed to be facultative. The tribe Aphnaeini constitutes one third of all lycaenids, almost all obligately ant-associated. The tribe Polyommatini accounts for a third of all lycaenids and is 95% ant-associated, containing ‘similar proportions of facultative and obligate associations. The presence, absence and function of myrmecophilous organs at various larval stages is discussed. Ovipositing below the soil or on sand surface is recorded for the first time. The various trophic strategies of larvae are discussed. Crematogaster (Myrmicinae) and Anoplolepis (Formicinae) ants together account for almost 80% of obligate relationships. It is suggested that many synonymies exist among obligately ant-associated taxa. Additional key words: evolution, myrmecophily, trophallaxis, aphytophagy, entomophagy, detritus, acoustics, South Africa. Of 3607 Afrotropical butterfly species, 42% are in head protected beneath a tough carapace. Many such the family Lycaenidae (Ackery et al. 1995), a similar larvae have organs that serve specific functions in their figure (47%) is given for southern Africa” by Pringle et association with ants. A dorsal nectary organ (DNO), al. (1994). Of the 397 Australian butterflies, 36% are in present on the seventh abdominal segment of some the family Lycaenidae (Braby 2000). These figures im- larvae provides honeydew for ants to imbibe. A pair of ply that a higher level of diversification has occurred in tentacle organs (TOs) located on the eighth abdominal the Lycaenidae than in other families, and Pierce segment aiid several minute perforated cupola organs (1984) suggested that this may have been caused by (PCOs) are present on the larva’s cuticle which allise- ant-association (myrmecophily). Many species of ly- crete substances that can influence ant behavior (Cot- caenids are ant-associated in the early stages (Fiedler trell 1984). Note that ant-organs found in ant-associated 1991), ants being among the leading predators of in- Riodininae differ from those in other lycaenids; see sects (Hoélldobler & Wilson 1990). Lycaenid larvae are DeVries (1991, 1997) for a comparison. By using these vulnerable to predatory ant species, hence various organs, free-living mutualistic caterpillars provide strategies have evolved to prevent or reduce attack. nutritious secretions and emit chemical signals (Hen- Dense and long hair serves as one effective strategy, ning 1983b) that can manipulate ant behavior to re- endophagy (feeding inside plant material) is another. duce aggression and obtain protection from predators Other lycaenids possess an extra-thick cuticle with the and parasitoids (Lenz 1917, Hinton 1951, Pierce & ' Address for correspondence: 1 The Close, Limekiln Lane Bal- Mead 1981, Pierce 1984, Pierce et al. 1987, DeVries dock, Herts, SG7 6PJ U.K. (Email: alan.heath3@virgin.net). 1988, 1991, Fiedler 1991). In some cases this manipu- * Southern Africa: Countries south of Angola, Zambia and the lation is extended to inducing regurgitations from Zambezi River in Mozambique. The region includes Namibia, dhol ante (et hhealllenes) bli 9 ea awe Botswana, Zimbabwe, South Africa, Swaziland, Lesotho and south- CXCHUINC GUMS) (CKO) OMe axis) or enabling the larva to prey ern Mozambique. directly on the ant brood (par Aon (Henning 1983 3b, Pierce 1995, Heath 1998, Elmes et al. 1991, Thomas 1983, Thomas & Wardlaw 1992). The DNO was first described by Guenée (1867), subsequently more knowledge was gathered about the myrmecophilous organs (Newcomer 1912, Ehrhardt 1914, Hinton 1951, Clark & Dickson 1956). Their structure and function was studied in greater detail by Malicky (1969, 1970). More recently, secretions from these organs were chemically analyzed (Maschwitz et al. 1975, Pierce 1983, DeVries & Baker 1989). Many, but not all ant-associated lycaenids possess these or- gans, whose presence and possibly function can vary throughout the larval stage (Cottrell 1984). Some ly- caenid tribes and genera are known to be more strongly ant-associated than others, but even within a single genus, the type of association can be quite var- ied (Pierce 1989). Ant-associated lycaenid larvae are known to pro- duce a substrate borne call to recruit ants (DeVries 1988, 1990, 1991, Travassos & Pierce 2000). Southern African aphnaeine larvae also produce sounds (Heath 1998), as do pupae (Schlosz 1991, Claassens 1991). Numerous field and laboratory studies have con- tributed to our knowledge of African ant-lycaenid as- sociations. One of the earliest accounts detailing ant- association in African lycaenids was Lamborn (1914) who recorded observations on early stages of 27 species (14 genera) from southern Nigeria. Nine of these genera are also represented in southern Africa. Observations in Kenya and Uganda by Jackson (1937) described obligate ant-associations in seven species and discussed 25 facultative associations. The first ma- jor study on South African lycaenid early stages was by Clark and Dickson (1971), who described and illus- trated 125 species at different stages of their develop- ment, including myrmecophilous organs. Behavioral studies under laboratory conditions were done subse- quently by Claassens (1972, 1976) and Claassens and Dickson (1977). On a global scale, lycaenid-ant associ- ations have been reviewed by Cottrell (1984), Fiedler (1991) and Pierce et al. (2002). This paper summarizes and discusses our under- standing of myrmecophilous lycaenids in southern Africa during the past ten years. Recent accounts in- clude: Schlosz and Brinkman (1991), Williams and Joannou (1996), Heath and Brinkman (1995a, b), Heath (1997a, b, 1998), Claassens and Heath (1997), Edge and Pringle (1996), Heath and Claassens (2000), Lu and Samways (2001). The number of species for each genus follows Pringle et al. (1994), Ackery et al. (1995), Williams (1999) and Heath (2001). A compre- hensive list of food plants and ant-associates for south- ern African Lepidoptera is found in Kroon (1999). JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Our classification here follows Scott (1985), Eliot (in Corbet et al. 1992), Ackery et al. (1999) and Pierce et al. (2002), who treat Riodininae® as a subfamily within Lycaenidae, along with Poritiinae (including Pentilini and Liptenini), Miletinae (Miletini, Liphyrini and Spalgini) and Lycaeninae (Theclini, Aphnaeini, Ly- caenini and Polyommatini). Examples of adult ly- caenids from the region are illustrated (Figs. 1-20). METHODS USED FOR STUDYING LYCAENIDS IN THE FIELD AND LABORATORY Inducing oviposition. Inducing oviposition facili- tates life history studies in the laboratory. The method for inducing oviposition among Chrysoritis Butler is to place not more than three known or ‘suspected’ host ants together with some stems preferably of a known larval foodplant, inside an open plastic container about 130 x 100 mm and 70 mm deep covered tightly with fine netting (e.g., ladies’ stockings) and kept warm, but not hot, or in the sun. A female Chrysoritis is added once the ants have settled down. More than three ants often results in an escape hole chewed in the netting. The female butterfly is fed daily with a small lump of tissue paper soaked in weak sugar water placed on the netting. Eggs are often laid on the netting. Attempts to induce oviposition among Aloeides Hiibner in this manner have been unsuccessful, but oviposition among captive Thestor Hiibner occurred on almost any surface without either ants or vegetation present (Heath & Claassens 2000). Obtaining early stages in the field. Methods for finding early stages in the field vary with life history characteristics. Many myrmecophilous species spend part or all of their juvenile phases in or close to subter- ranean ant nests which makes them difficult to dis- cover and study. Others are found under rocks and as- sociated with a particular plant or ant nest. The method used to find Chrysoritis larvae is to search the debris beneath potential food plants where ants are present. On occasions, the larvae are found in small fi- brous shelters built on the plant stems by the ants, pre- sumably intended for Homoptera. For Aloeides we search in the soil beneath potential food plants or be- neath nearby rocks. In Trimenia Tite & Dickson, the early stages are sought by digging in the gravel or by prizing open cracks in bedrock where nests of the host ants occur (Heath & Brinkman 1995b). For Phasis Hiibner, hollowed out stems of the foodplants are ex- amined where ants are present. For Thestor and Ty- lopaedia Tite & Dickson, we search beneath large boulders covering ant nests in areas where the adults > Riodininae is not represented in southern Africa. VOLUME 57, NUMBER 1 Fics. 1-20. Lycaenidae from southern Africa. 1, Iolaus (Io- laphilous) trimeni Wallengren. 2, Phasis thero thero (Linnaeus) fe- male. 3, Myrina silanus ficedula Trimen male. 4, Thestor protumnus protumnus (Linnaeus) female. 5, Aloeides aranda (Wallengren) male. 6, Aloeides pallida grandis Tite & Dickson male. 7, Erikssonia acraeina Trimen male. 8, Pentila tropicalis tropicalis (De Boisduval) male. 9, Tylopaedia sardonyx sardonyx (Trimen) female. 10, Cigari- tis natalensis (Westwood) male. 11, Lepidochrysops trimeni (Bethune-Baker) male. 12, Lepidochrysops oreas oreas Tite female. 13, Chrysoritis brooksi brooksi (Riley) female. 14, Chrysoritis thysbe thysbe (Linnaeus) female. 15, Chrysoritis nigricans nigricans (Aurivillius) male. 16, Chrysoritis palmus palmus (Stoll) female. 17, Lycaena orus Stoll male. 18, Lepidochrysops robertsoni Cottrell male. 19, Lachnocnema bibulus (Fabricius) female. 20, Axiocerces amanga (Westwood) male. are known to fly. Rocks under which juvenile Thestor species are found, are often turned over by baboons looking for Thestor larvae and pupae, and scorpions. Study of behavior in captivity. Due to the sub- terranean habits of many juveniles, close study of the interactions between ants and larvae in nature is not feasible. Hence, for taxa like Aloeides, Thestor, Lepi- dochrysops, Orachrysops and Trimenia, where a formicarium similar to that described by Claassens (1972, 1974) is required, a nest of host ants with brood and queen is installed within the formicarium. Before being introduced into the nest section of the formicar- ium, larvae are left among the ant brood for about two hours. This is to acquire the scent of the ant colony and avoid subsequent attack by ants, but also to see what interaction takes place between larvae and brood. Pieces of suspected food plant are kept in water within the arena, as described by Claassens and Dick- son (1977). Chrysoritis species do not generally enter ant nests but are constantly attended while feeding on the foodplant and resting in the debris beneath it; their ie) early stages can be studied in the natural environment. It is nevertheless possible to rear these in captivity with or without the host ant. CATEGORIES OF ANT-ASSOCIATION AND FEEDING STRATEGIES There have been many attempts to categorize feed- ing strategies and relationships between lycaenids and ants; e.g., Fiedler (1991), DeVries (1991), Pierce (1995), Eastwood and Fraser (1999), Heath and Claassens (2000) and Pierce et al. (2002). The follow- ing three broad categories of ant-association are used here: (1) ‘Not ant-associated’ (direct and close interac- tion with ants is absent or rare, even if ants are pres- ent). (2) ‘Facultative’ (intermittently attended by ants but not wholly dependent on them for survival under field conditions). (3) ‘Obligate’ (dependent on one species of ant for survival under field conditions). Three categories of larval feeding strategy used here are ‘algae/lichen-feeding’, ‘herbivorous’ and ‘ento- mophagous’.. The latter term being re-defined by Pierce et al. (2002) to mean dependant upon any in- sect-derived resource, and may include homopteran secretions, ant regurgitations and/or the insects them- selves (carnivory). More than one of these categories may be employed during the larval phase. GENERA, LIFE HISTORY AND ANT-ASSOCIATIONS For completeness, all 64 lycaenid genera occurring in southern Africa are included here, whatever their ant-association. More specific details are shown in Table 1 and summarized in Table 2. PORITIINAE: Larvae of Pentilini and Liptenini (together including 13 genera) are clothed with long hairs and not directly ant-associated; they possess no DNO or TOs. Larval food is mostly algae or lichen (Bampton 1995), but Delonewra were recorded taking honeydew from Homoptera in the company of ants (Pringle et al. 1994), although ants ignore the larvae (Jackson 1937). Pentilini (three genera): Alaena De Boisduval, 1847; Pentila Westwood, 1852; Ornipholidotos Bethune-Baker, 1914. Liptenini (ten genera): Durbania Trimen, 1862: Durbaniella van Son, 1959; Durbaniopsis van Son, 1959: Cooksonia H. H. Druce, 1905; Mimacraea Butler, 1872; Euthecta Bennett, 1954: Teriomima Kirby, 1887; Baliochila Stempffer & Bennett, 1953: Cnodontes Stempffer & Bennett, 1953; Deloneura Trimen, 1868. MILETINAE: Larval food can be regurgitations from ants but is mostly Homoptera or their secretions. Homoptera are almost always attended by ants but di- 4 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TABLE 1. Life history details for all known Southem African lycaenids, showing presence of DNO or TOs, larval feeding category* and ant-as- sociation. Abbreviations: * = probable but unconfirmed. DNO, TOs: + = yes; — = no; Feeding category: A = Algae or Lichen feeder; H = Her- bivorous; E = Entomophagous. Ant-association: N = none; F = facultative; O = obligate; ? = unknown. Principal sources: 1 = Clark and Dickson (1971); 2 = Pringle et al. (1994); 3 = Jackson (1937); 4 = Heath and Claassens (2000); 5 = Heath (1997); 6 = Bampton and Congdon (pers. com.); 7 = Stemptter (1967); 8 = Henning (1983a, b); 9 = Claassens (1976); 10 = Edge and Pringle (1996); 11 = Schlosz and Brinkmann (1991); 12 = Heath (pers. obs.); 13 = Kroon (1999); 14 = Lamborn (1914); 15 = Heath and Brinkmann (1995b); 16 = Williams and Joannou (1996); 17 = Lu and Samways (2001); 18 = Larsen (1991); 19 = Pennington (1956); 20 = Atsatt (1981); 21 = Braby and Woodger (1994); 22 = Henning and Hen- ning (1982); 23 = Henning (1984a); 24 = Ackery and Rajan (1990); 25 = van Someren (1974); 26 = Claassens and Dickson (1977); 27 = Claassens and Dickson (1980); 28 = Fiedler and Hagemann (1992); 29 Clark and Dickson (1956); 30 = Congdon and Bampton (1995); 31 = Edge (1990). Feeding Ant- Principal Genus Species DNO TOs category* association Ant sources PORITIINAE Pentilini (8) Alaena amazoula = = A N 1, 29 brainei = = A N 2,6 nyassa - - A N 2,6 maregaritacea = - A N 2,6 Pentila pauli = = A N 12 swynnertoni = = A N 2,6 ti opicalis — = A N 2,6 Ornipholidotos peucetia = = A N 19, 133 Liptenini (22) Durbania amakosa = = A N 1, 12, 29 limbata = — A N 9. 12) Durbaniella clarki A N QD W2) Durbaniopsis saga - - A N D1 Cooksonia neavei = = A N 9 Mimacraea marshalli = = A N 2) neokoton = = A N 2 Euthecta cooksoni P ? A* N* 2 Teriomima puellaris P >? A N 2 puella P > A N 2 zuluana i > A N 2 Baliochila aslanga i ? A N 2 Baresi P > A N 2 neavei P ? A N 2 lipara ? a A N 2 singlaris P i A N 2 Cnodontes 3 species ? ? A* N* 19 Deloneura sheppardi = = A N 1 coe = - A N 2,3, 19 subfusca - = A N 2,3), 6, 19) MILETINAE Spalgini (1) Spalgis lemolea ? P E N 3, 8,14 ates neni (32) Lachnocnema bibulus = = E N 1, 8, 14, 29 durbani = = E N 1, 2, 29 brimo = = E N* 9 Thestor basutus = = E O A. custodiens 1, 4, 16, 29 brachycerus - = E O A. custodiens 12 pictus - - E O A. custodiens 12 protumnus - - E O A. custodiens 1,12 rileyi = = E O A. custodiens 12 rossouwi = = E O A. custodiens 12 strutti = = E O A. custodiens 4 yildizae - = E O A. custodiens 4 dicksoni = = E O A. custodiens 1, 2, 29 +20 species 8 ie E* O* A. custodiens* 2 Liphyrini (3) Aslauga 3 species ae +* h* N* 3, 6, 7, 14 LYCAENINAE Aphnaeini (131) Aphnaeus erikssoni ae 4% H* O Crematogaster sp. 9) hutchinsonii + + H O Crematogaster sp. IL, B, LY), Bil marshalli 4% Ae H* O* Crematogaster sp.* 2 Cigaritis natalensis + + H O C. castanea 1, 2, 6, 29 ella + + H O C. castanea 1, 2, 6, 18 namaqua + + H O Crematogaster sp. 8,6 phanes + + H O C. castanea 8,6 apelles +% + H O Crematogaster spy 2,6 +5 species pe +* H* O* Crematogaster sp.* 2,6 Lipaphnaeus aderna +* +* H O* Crematogaster sp.* 6, 30 Chloroselas pseudozeritis + + H O C. gerstaeckeri 2,6,3 argentea +% +* H O Crematogaster sp. 2,6 mazoensis +* +* H O Crematogaster sp. 2,6, 19 VOLUME 57, NUMBER 1 TABLE 1. Continued. Feeding Ant- Principal Genus Species DNO TOs category* association Ant sources Zeritis sorhagenii ? ? H > 2 Axiocerses tjoane + + H F 1, 6, 29° amanga + + H F* 3,6 unicea +* cp H F 2,6 Phasis praia +% - H O C. peringueyi 1,4 clavum +* +* H O* ¢ rematogaster sp.* 2 pringlei + + H O C. peringueyi 125 Ik thero +4 + H O C. peringueyi 1, 4, Tylopaedia sardonyx + + H O C. melanogaster ital, Argyraspodes argyraspis > ? H ? ? 5 Trimenia argyroplaga ? + H O A. custodiens 12 malagrida ? + ? O A. custodiens 15 +3 species - ? O* A. custodiens* 2 Aloeides apicalis + + H O Monomorium fridae 4 aranda +f + H O Pheidole capensis IZ 9s) clarki + + H O* Unidentified sp.* iL, 2 damarensis + + H O* Unidentified sp.* il, 2 dentatis Pg + H O L. capensis 2,48 depicta + + H O* Unidentified sp.* 1,2 gowani + + H O* Unidentified sp.* 1 pallida + + H, ES O L. capensis 4 pierus + + H O L. capensis 4, 29 thyra + + H O L. capensis 2, 4, 26 trimeni + + H O L. capensis i ® rossouwt +* +* H O Lepisiota sp. 22 taikosama + +* H O* Unidentified sp.* 2, 29 almeida + +* H O* Unidentified sp.* 2, 29 +39 species +% +* H Unidentified sp.* 2 Erikssonia acraeina + + H Lepisiota sp. 2523 Chrysoritis adonis + + H O C. liengmei 5 aethon + + H O C. liengmei 5 aridus + + H O Crematogaster sp. 5 aureus + + H O C. liengmei 5 azurius + + H O C. liengmei 5 beaufortia + + H O C. peringueyi 5 beulah + + H O Crematogaster sp. 5 blencathra + + H O Crematogaster sp. 5 braueri + + H O C. liengmei 5 brooksi + + H O C. peringueyi 5 chrysantas + + H O C. melanogaster 5 chrysaor + + H O C. liengmei 5, 29 daphne + + H O C. liengmei 5 dicksoni + + E O C. peringuey 5 endymion + + H O C. peringuey 5 felthami + + H O C. peringuey 5, 29 irene + + H O Crematogaster sp. 5) lycegenes + + H O G. liengmei 5 lyncurium + + H O C. liengmei 5 midas + + H O C. peringueyi 5 natalensis + + H O* Crematogaster sp.* 5 nigricans + + H O Crematogaster sp. 5 oreas + + H O Myrmicaria nigra 5 orientalis + + H O C. liengmei 5 palmus + + H O C. liengmei 5, 29 pan + + H O C. liengmei 5 pelion + + H O* Crematogaster sp.* 5 penningtoni + + H O Crematogaster sp. 5 perseus + + H O C. melanogaster 5 phosphor +* +* H* OF Crematogaster sp.* 5 plutus + + H O C. peringueyi 5 pyramus + + H O C. peringueyi 5 pyroeis + + H O Myrmicaria nigra 5, 29 rileyi - . H O C. peringueyi 5 swanepoeli + + H O C. liengmei 5 thysbe + + H O C. peringueyi 5, 29 trimeni + + H O C. peringueyi 5 turneri + + H O C. liengmei 5 uranus + + H O C. liengmei 5 violescens + + H O C. peringueyi 5 ZeuxO + + H O C. liengmei 5 zonarius - + H O C. peringueyi 5 Crudaria leroma + + H O A. custodiens IL By P98) wykehami + + H O A. custodiens It capensis x ie H* O* A. custodiens* ls Lycaenini (2) Lycaena clarki H N 1, 12 orus H N 1, JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TABLE 1. Continued. Feeding Ant- Principal Genus Species DNO TOs category* association Ant sources Theclini (47) Amblypodiiti [2] Myrina silenus + + H F 1, 3,14, 29 dermaptera + + H F 1, 29 Tolaiti [21] TIolaus aemulus + + H F* 1, 2, 29 alienus + + H F 1, 2, 29 aphnaeoides 4% +% H F* 2 australis +* 4% H F* 2 bakeri +* 4% H F* 2 bowkeri + + H F 1, 2, 12, 29 diametra +% +* H F* 2 lalos iii 4% H F* 2 lulua ahs +% H F* 2 mimosae + + H F 1, 2, 12, 29 nasisti +% +* H F* 2,12 obscurus +% +% H F* y) pallene +% +% H F* X12 penningtoni +* Et H F* 2 poultoni 4% 4% H F* 2 sidus + + H F* 1, 2, 12, 29 silarus + + H F 2 19) silas + + H F I, 2, 12, QS) subinfuscata +* 38 H F 2, WP trimeni +* 76 H F* 2,8, 12 violacea +% +* H F* 9. 19) Hypolycaeniti (9) Hypolycaena peeDeus + = H F 1, 3, 14, 29 ochmophila 4% a H F* 2 caeculus >? ? H F 2,6 +2 species +* —* H* F* 2 Leptomyrina hirundo + = H F iL, 2 lara + — H F QE e29) gorgias + = H F IL, 2 henningi +% a H F* ) Deudorigiti [15] 15 Deudorix antalus + = H F Il, 2, XY) dariaves +% =+ H F* 2 dinochares + = H F IL, 2, D4 dinomenes +* ma H F* 2 diocles + = H F 1, 2, 29 lorisona + a H F* 2 magda at = H F* 2 penningtont +* =e H ge 2; vansoni +* —* H F* 2 caerulea +% a H* F* 2 zeloides ae ats H* F* 2 Capys alphacus + = H F 1, 27, 29 penningtoni + - H F PD, IS disjunctus + = H F Is 2. 1S) connexivus + - H F 2,12 Polyommatini (146) Lycaenesthiti [26] Anthene amarah + + H F I, B, 2 butleri + + H F* 1, 29 contrastata +* +* H F* 2 crawshayi + + H F* 3 definita + + H F 1, 27, 29 kersteni + + H F : lemnos + + H F* 1, 29 liodes + + H F* 2,14 lunulata + + H F 3 wilsoni = = ? O Unidentified sp. 3 otacilia + + H F 1, 6, 25, 29 talboti + + H F* IL nigeriae + + H O Unidentified sp. 3 +13 species ? P H* F* 2 Polyommatiti [120] Cupidopsis cissus + + H F* 1S jobates + + H F* it Pseudonacaduba _ sichela P 2D H P 2 Lampides boeticus + + H F 1, 3, 27, 29 Uranothauma antinorii = = H N* 2 poggei = = H N 2,6 VOLUME 57, NUMBER 1 TABLE 1. Continued. Feeding Ant- Genus Species DNO TOs category* association Ant nubifer - - H N vansomereni —* —* H N* Cacyreus dicksoni = = H N lingeus - - H N marshalli = = H N tespis — - H N virilis + = H N Harpendyreus notoba + - H h* tsomo +* = H F* noquasa +* - H F* Leptotes pirithous - - H h* brevidentatus + + H F* jeanneli + + H Bs babaulti 4% +% H 1p pulcher +% 4% H F* Tuxentius calice + + H F* melaena + + H F* hesperis +* +* H 1a Tarucus has + + H F* thespis + + H F bowkeri + + H F* Zintha hintza + + H F Zizina antanossa + + H F* Zizeeria knysna + + H B Actizera lucida + + H F* stellata + + H F* Zizula hylax + + H ae Brephidium metophis + + H ee Oraidium barberae ? P H F* Azanus ubaldus + + H F* jesous + + H F* natalensis + + H F moriqua + + H F* mirza +* +* H F* Eicochrysops messapus + + H F* eicotrochilus +* ale H F* hippocrates + + H F* Euchrysops osiris + + H F barkeri + + H F malathana + + H F dolorosa + + H F subpallida +* +* H F* Lepidochrysops _asteris + - H, E*™ O} Camponotus sp.* bacchus + = H, E* O* Camponotus sp.* ignota + ~ H,E O C. nwweosetus ketsi + — H, E* O* Camponotus sp.* methymna + H,E O C. maculatus oreas + = H,E O C. maculatus patricia + - H, E O C. maculatus puncticilia + = H, E* O* Camponotus sp.* trimeni + = H,E O C. niveosetus variablis + = H,E O C. niveosetus +49 species +* —* lal, 1a O* Camponotus sp.* Orachrysops lacrimosa + + Jal Jigs niobe + + H* se ariadne + + H* O C. natalensis +7 species +* +* H* F* Oberonia bueronica P > H F Chilades trochylus + + H * Thermoniphas weal P > H N “For Feeding category, and Ant-association, see section “Categories of ant-association and feeding strategies” above. b As A. bambana (misidentified). ‘A. amanga is attended only by Pheidole species but often found unattended (Bampton pers. com.). dP thero has DNO but not in final instar *Has DNO but absent in final instar (also A. depicta, A. pallida and A. thyra). ‘DNO present in final instar (also A. pierus). §DNO absent in final inmstar (Henning 1983) but earlier instars unknown. Herbivorous but in final instar, feeds solely on ant eggs. ‘As A. larydas (misidentified). JAsC. palerion (misidentified). KAs Syntarucus telicanus (misidentified). 'As Castalius melaena (misidentified). ™ Lepidochrysops are herbivorous for first two instars, thereafter mainly carnivorous on ant brood, supplemented by trophallaxis. "As Lepidochrysops caffrariae (misidentified). °Currently regarded as facultative—it can be reared without ants. BO Re BO BSD BO BDO DN DD DD DD BDO DO DD BO FF DD DD RS Re Eb LP Principal sources nso 3 top bo go 19 P9 fo G9 bo NNN NNNNoNTD to D bo bo 29 , 29 5 PAY) , 29) , 29} JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TABLE 2. Summary of ant-association: Conf. = based on published and confirmed observations. Predict. = confirmed + predicted but un- confirmed associations. Obligate Taxon Total species Conf. Predic. PORITIINAE 30 Pentilini 8 0 0 Liptenini 22; 0 0 MILETINAE 36 Spalgini 1 0 0 Lachnocnemini 32 9 29 Liphyrini 3 0 0) LYCAENINAE 326 Aphnaeini 131 66 126 Lycaenini 2 0 0 Theclini 47 0 0 Polyommatini 146 9 62 TOTALS 392 84 217 rect interaction between ant and larva may not neces- sarily occur. Liphyrini (one genus): Aslauga Kirby, 1890 (14 Afrotropical species, three in southern Africa). Scarce, arboreal butterflies with a distinctive wingshape. Vir- tually nothing is known of the early stages of southern African Aslauga species but suspected to be the same as those observed by Jackson (1937) and Lamborn (1914). Larval skin is leathery, the carapace being ex- traordinarily heavy; head small and carried on an ex- tendable neck which can be retracted under a cara- pace; TOs present, but DNO absent. Although the presence of TOs is unique in this subfamily, they are small and do not evert. Larvae feed on Coccidae (Ho- moptera) tended by Crematogaster ants that do not in- teract directly with the larvae. Spalgini (one genus): Spalgis Moore, 1879 (four Afrotropical species, one in southern Africa). Larvae feed on Coccidae (Homoptera), do not associate with the ants directly, but ants are always present tending Homoptera. Larvae cover themselves with a waxy se- cretion produced by the Homoptera and lack both DNO and TOs (see Lamborn 1914, Jackson 1937). Lachnocnemini (two genera): In these genera, lar- vae lack DNO and TOs. Lachnocnema Trimen & Bowker, 1887 (38 Afrotropical species, 3 in southern Africa). Feeding habits of L. bibulus Fab. have been described by South African authors, but by a different account of Lachnocnema from Kenya (Cripps & Jack- son 1940), larvae were carried by ants down to their nest, and trophallaxis occured. The genus was revised by Libert (1996a-c) who described 25 new species, hence the life history accounts from Kenya most prob- ably apply to what are now considered different species. Accounts are given by Clark and Dickson (1971) of L. bibulus and L. durbani Trimen from Facultative None Unknown Conf. Predic. Conf. Predic. Conf. Predic. 0 0 8 8 0) 0 0) 0 18 22, 4 0 0) 0) 1 a 0) 0 0) 0) 2 3 alk 0 0) 0 0 3 3 0 3 3 0 0) 62 2 0 0) 2 2 0 0) 20 47 0 0 ON 0 15 73 8 10 114 1 38 123 39 49 231 4 South Africa. Early instar larvae of Lachnocnema bibu- lus feed on young psyllids (Homoptera) and their droppings. Mature larvae creep up behind adult psyl- lids, seize their wings, and devour them. Larvae do not associate with ants directly but ants are always present tending the Homoptera. Thestor Hiibner, 1819 (29 species, all endemic to southern Africa). Medium-size, moth-like, with stout bodies, and either blackish or yellow and brown; they always settle on the ground or rocks. Adults possess a vestigial proboscis and do not nectar. Most Thestor species are univoltine. All species appear to associate with Anoplolepis (Formicinae) ants (Claassens & Dickson 1980, Cottrell 1984, Claassens & Heath 1997). During the first three instars of T. protumnus aridus Van Son and T. basutus (Wallengren), larvae feed on Homoptera (Clark & Dickson 1960, 1971, Williams & Joannou 1996). In T. yildizae Kogak, T. pic- tus Van Son and T. basutus food of the final two instars is regurgitated food from ants (Fig. 21). We suspect that all Thestor are entomophagous but only in T. ba- sutus is the life history fully known. The remarkable Thestor larva (Fig. 22) lacks both DNO and TOs, and possesses an extremely small head with an extendable fleshy “neck”. Larval antennae are elongate and pro- ject forward; each with a long terminal seta. In T. yildizae these antennae remain in contact with the ant’s mandibles during exchange of food. It is possible these antennae simulate the ant’s mandibles, facilitat- ing the larva’s acceptance as a nestmate and/or to stim- ulate the ant to regurgitate. Williams and Joannou (1996) observed females of Thestor basutus capeneri (Dickson) ovipositing on blades of grass infested by grass-feeding coccids Pulvi- naria iceryi (Signoret) (Homoptera: Coccidae) which in turn were tended by Anoplolepis custodiens Smith VOLUME 57, NUMBER 1 Fic. 21. Final instar larva of Thestor yildizae Kogak being fed re- gurgitations from a “Pugnacious ant” Anoplolepis custodiens F. Smith. ants. They were observed to oviposit on a wide variety of vegetation, often without ants or Homoptera being present. In the wild and in captivity the first three in- stars were predaceous on coccids, the ants taking no interest in the larvae. After moulting, fourth instar lar- vae in captivity refused to feed on coccids and were left in the formicary where they entered the artificial ant nest. While in the ant nest, they were ignored by the ants but subsequently died, presumably of starva- tion (Williams & Joannou 1996). Similar observations were recorded by Clark and Dickson (1960, 1972) for T. basutus basutus. In November 2002 four final instar larva of T. basu- tus basutus (Wallengren) were taken from an Anoplolepis custodiens ant nest in KwaZulu-Natal and studied in captivity together with Anoplolepis ants from a nearby locality (AH, AJMC, S. P. Quek). A larva was often seen to approach two ants engaged in trophallaxis and insert its head between theirs. It pro- ceeded to imbibe regurgitations passed between them and continued to accept regurgitations from the donor ant after the other ant had departed. The larva was oc- casionally seen to ‘pull’ on ant eggs and larvae and drag them beneath its carapace; it is assumed these were eaten. The larva also appeared to scavenge for detritus on the nest substrate. Although T. basutus appears to utilize four food sources in its larval stage, during the week it was studied (prior to pupation) the major food was obtained by trophallaxis. An earlier study in cap- tivity of T. basutus larvae from KwaZulu-Natal with an ant species from Cape Town resulted in the larva feed- ing on detritus only and then dying after almost four weeks without any trophallaxis having been observed (Heath & Claassens 2000). This would indicate larvae are highly specific to certain Anoplolepis ants. The larva occasionally ‘groomed’ an ant, the latter remaining motionless while the former may have de- rived some form of detritus from the process (AH, AJMC unpublished obs.). This grooming behavior was also recorded in the case of Lepidochrysops larvae (Polyommatini) and its ant-associate Camponotus maculatus Fab. (Formicidae) (Claassens 1976). 9 1cm Fic. 22. Final instar larva of Thestor yildizae, lateral view (up- per), dorsal view with head extended (lower). Scale bar = 1 cm. The behavior of T. basutus differed from the ob- served behavior of T. yildizae and T. pictus larvae which solicited regurgitations from individual passing ants. The ant species associated with Thestor have so far been recorded as A. custodiens but this is currently believed to constitute a species complex. A prelimi- nary molecular study of A. custodiens at Harvard Uni- versity suggests that at least three distinct clades exist (S.P. Quek unpublished). LYCAENINAE: Some ant-association occurs in all tribes except Lycaenini. Almost all species have a DNO and most have TOs. Theclini (six genera): The number of larval instars is normally four and the DNO first appears in the sec- ond or third instar, the TOs appearing in the second instar except in endophytic feeders, where they are ab- sent in all instars. Myrina Fabricius, 1807 (five Afrotropical species, two in southern Africa); Iolaus Hiibner, [1819] (116 Afrotropical species, 21 in southern Africa); Hypoly- caena Felder, 1862 (extralimital*: 28 Afrotropical species, five in southern Africa); Leptomyrina Butler, 1898 (nine Afrotropical species, four in southern Africa); Capys Hewitson, 1965 (14 Afrotropical species, four in southern Africa); Deudorix Hewitson, 1863 (extralimital: 89 Afrotropical species, nine in southern Africa). All six genera are regarded as having a facultative association with ants. Ants (often Cre- matogaster species) may be present but do not attend the larvae permanently, and larvae do not depend on any species of ant for survival. Larvae of the first three genera possess DNO and TOs but those of Leptomy- 4 Extralimital: Genus also represented outside Africa. 10 rina, Capys and Deudorix are endophytic and lack TOs (Clark & Dickson 1971). Leptomyrina larvae feed inside the fleshy leaves of Crassuleaceae (Jackson, 1937). Capys larvae feed inside Protea flower buds (Proteaceae) (Murray 1935, Clark & Dickson 1971); the hollowed out buds are later often occupied by Crematogaster ants (AH pers. obs.). Deudorix larvae feed inside pods and immature fruit of a variety of trees (Pinhey 1965, Clark & Dickson 1971). Aphnaeini (14 genera): All known life histories of the Aphnaeini indicate an obligate ant-association. The number of larval instars is normally six, the DNO first appearing in second or third instars, but TOs occur in all instars. Chrysoritis Butler, 1898 (42 species, all endemic to southern Africa). A revision and molecular phylogeny of Chrysoritis can be found in Heath (2001) and Rand et al. (2000). The genus comprises small to medium- sized robust butterflies, associated with open veld, coastal and montane fynbos’. Most Chrysoritis are multivoltine, except in montane areas, but C. dicksoni (Gabriel) which flies in non-montane localities, is en- tomophagous and univoltine. With this one exception, Chrysoritis are all herbivorous feeding on a wide vari- ety of plants (see Heath 1997a) and can be reared in captivity without ants. They are nevertheless consid- ered obligate, as larvae are continuously attended by ants and oviposition only occurs in the presence of the correct ant species (Heath 1997a). Crematogaster ants are the most common associates for Chrysoritis, but one species of Myrmicaria (Myrmicinae) is known to associate in two species of this genus (Heath 1997a). Excepting C. dicksoni, larvae in nature rest in a corral (or byre) beneath the food plant and are tended by three or more ants. Larvae were often found singly or in pairs but as many as five or six of varying sizes could be found together within a corral (Heath 1997a). In Chrysoritis the DNO appears in the second in- star, a feature shared only with Crudaria Wallengren. The larva’s DNO is frequently visited and stimulated by the ant’s antennae and the secretion is eagerly taken by the ant. The TOs are active, everted quickly and as quickly withrawn; this happens whenever the larva is disturbed. When ants are over-eager to access honey- dew, the larva will evert its TOs causing them to jump in alarm. Among this large and otherwise herbivorous genus, C. dicksoni larvae are entomophagous and were observed in the field and laboratory by Heath (1998). The first two instars fed on ant regurgitations but stayed on the plant close to scale insects, also tended ° Fynbos: Characteristic treeless shrubland vegetation of the southern and south-western Cape of South Africa. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY by the host ant. The DNO was present in the second instar, but secreted no honeydew, however, when everted it attracted, but stupified nearby ants. When offered a red Homopteran juvenile, the second instar larva ate it and half a second one, but died the follow- ing day. The final instar also fed on ant regurgitations living and pupating within the carton nest of the host ant Crematogaster peringueyi Emery. It is assumed that the intervening three instars had the same feeding behavior as first and last instars. In areas where C. dicksoni flies, armored scale insects (Homoptera) are always nearby and may somehow be necessary for the survival of the larva (Heath 1998). The DNO was still present in the sixth (final) instar, although it was infre- quently visited by ants. The final instar also had spe- cialized setae on each segment that frequently at- tracted the host ant who seemed to nibble at them. These setae appeared as a curved structure resembling a bottle-brush, and are illustrated by Heath (1998). Heath and Brinkman (1997b) inferred that in the wild, females are only stimulated to oviposit by ants attend- ing this unidentified species of scale insect. Colonies of C. dicksoni adults occupy small areas, seldom larger than a tennis court, and where females oviposit on a wide variety of plants near the host ant nest. Aloeides Butler, 1819 (58 Afrotropical species, 53 in southern Africa). A large genus of small to medium- size brown or orange butterflies obligately associated with ants. All species are found in open or grassland habitat. The genus is in need of taxonomic revision, as we believe it contains many taxa of dubious status. Un- til recently, the only ant genus known to be associated with Aloeides larvae was Lepisiota (=Acantholepis) (Formicinae). However, two additional ant hosts have recently been discovered (Heath & Claassens 2000)— Monomorium fridae Forel (Myrmicinae) and Pheidole capensis Mayr (Myrmicinae). Larval food plants are species of Aspalathus (Fabaceae), Hermannia (Stercu- liaceae), Sida (Malvaceae) (Kroon 1999), and Gnidia (Thymelaeaceae) (A. Gardiner pers. com.). Aloeides larvae have TOs in all instars but the DNO first ap- pears in the third. In some species, e.g., A. apicalis Tite & Dickson, A. pallida grandis Tite & Dickson, A. thyra (Linnaeus) and A. dentatis (Swierstra), the DNO is absent in the final instar (Heath & Claassens 2000, Henning 1983a, b). However, in at least four species, A. pierus (Cramer) A. gowani Tite & Dickson, A. trimeni southeyae Tite & Dickson and A. aranda (Wallengren), larvae retain their DNO until pupation (Clark & Dickson 1971, Heath & Claassens 2000). Early instar larvae of A. pallida grandis are assumed to feed on species of Aspalathus always found close to where the larvae are found. Heath and Claassens (2000) VOLUME 57, NUMBER 1 observed that, in captivity final instar larvae remained inside the nest of Lepisiota capensis Mayr for four months, and grew without foraging outside. Despite ample ant brood of all stages present in the nest, the larvae fed only on ant eggs. Despite the DNO of A. thyra being absent in its final instar, it is herbivorous on Aspalathus species and rests in an L. capensis ant nest (Claassens & Dickson 1977) but it is not known if trophallaxis or ant eggs form a supplementary part of its diet. In contrast, A. apicalis and A. aranda which associate with Monomorium and Pheidole ants respectively, were not inside the ant nest but were generally tended by four or five ants in a cor- ral just below the soil surface close to the food plant, often two or three meters from the ant nest (AH, AJMC unpublished). Ovipositing females of A. molomo coalescens Tite & Dickson were observed inserting their abdomens deep into the sand beneath a species of Gnidia (Thymelaeaceae) (A. Gardiner pers. com.). Eggs of A. aranda (Wallengren) were found buried ca. 1 cm in the sand beneath its food plant Aspalathus sp. (C. Penz, P. De Vries, AH pers. obs.). Erikssonia Trimen, 1891 (three Afrotropical species, one in southern Africa). E. acraeina Trimen is scarce and local, its larval food plant is Gnidia kraussiana Meisner and it has an obligate ant association with Lepisiota sp. (Henning 1984a). The final instar have both DNO and TOs. This orange-red butterfly appar- ently mimics unpalatable species of Acraea Fabricius (Nymphalidae). Erikssonia is closely related to Aloe- ides and could be synonymized with Aloeides based upon genitalia (Heath 1997a), but other small differ- ences occur in the adult (Henning & Henning 2001). Eggs are laid among soil particles beneath the food- plant (Pringle et al. 1994). Phasis Hiibner, 1819 (four species, all endemic to southern Africa). Large brown lycaenids restricted to the southern and south-western Cape; their larval food plants include Rhus (Anacardiaceae) and Melianthus (Melianthaceae) and they have an obligate ant associa- tion with C rematogaster peringueyi. The DNO on Phasis thero (Linnnaeus) larvae appears on third and subsequent instars (Clark & Dickson 1971) but is ab- sent in the final instar (Heath & Claassens 2000). TOs are present in all instars (Clark & Dickson 1971). Lar- vae and pupae are found inside hollow stems and al- though associated with their host ants, interaction has not been studied closely. Trimenia Tite & Dickson, 1973 (five species, all en- demic to southern Africa). Medium-to-large orange and brown butterflies with silvery spots on the ventral surface of all wings. The genitalia are all very similar. They are restricted to arid habitats in the southern and south-western region of South Africa; they are univol- tine and all are presumed to be aphytophagous (Heath 1997a). A final instar larva and pupa of Trimenia mala- grida maryae (Dickson) were found in small fissures about 5 cm deep inside the bedrock tended by Anoplolepis custodiens ants (Heath & Brinkman 1995b). In captivity it could not be determined what the larval food was since the larva shunned any light. When disturbed, it was tended by many ants, with a concentration around the head of the larva (AH pers. obs.). There was no vegetation within a meter of the site where the larva was found, but after collection it survived among ants without vegetation for two weeks before pupating, supporting the notion that it is aphy- tophagous, at least in the final instar. We suspect it was feeding on ant eggs or ant regurgitations. Despite the presence of TOs, the final instar larva had no DNO and was very similar to that of Trimenia argyroplaga (Pringle in Pringle et al. 1994). A T. argyroplaga larva in its penultimate instar (presumed) had an active DNO, but in the final instar it was absent. The larva was seen to accept ant regurgitations, presumably its sole diet as it did not feed on ant brood or eggs. The larva often sought a dark place between the nest and arena to rest, and sometimes at night, it would go into the arena but the ants would manoever it back to the nest entrance again (Heath & Claassens 2000). Argyraspodes Tite & Dickson, 1973 (one species, endemic to southern Africa); Zeritis De Boisduval, 1836 (extralimital; six Afrotropical species, one in southern Africa). Life histories of these two genera are unknown, although the former is closely related to Tri- menia and may also be aphytophagous. Tylopaedia Tite & Dickson, 1973 (one species, en- demic to southern Africa). This large, robust lycaenid is orange and black and univoltine. Tylopaedia sar- donyx peringueyi (Aurivillius) is recorded as using a species of Aspalathus for its larval food plant (Schlosz & Brinkman 1991). The same authors observed the ant-associate to be Crematogaster melanogaster (Emery) and noted that the fernale would not oviposit without the presence of ants. Lipaphnaeus Aurivillius, 1916 (four Afrotropical species, one in southern Africa); Chloroselas Butler, 1885 (=Desmolycaena, Trimen) (13 Afrotropical species, three in southern Africa); Crudaria Wallen- gren, 1875 (three Afrotropical species, all in southern Africa); Aphnaeus Hiibner, 1819 (20 Afrotropical species, three in southern Africa); Axiocerses Hiibner, 1819 (25 Afrotropical species, five in southern Africa); Cigaritis Donzel, 1847 (=Spindasis Wallengren; Jeet Riley) (extralimital; 37 Afrotropical species, ten in souther Africa). The life history of these six genera have not been studied in recent years. Their larvae all possess DNO and TOs (Clark & Dickson 1971, Edge 1990), and are all believed to be herbivo- rous and obligately ant-associated, excepting Axio- cerses, which are facultative. TOs occur in all instars, the DNO first appears in the third instar, excepting Crudaria where it appears in the second. Clark and Dickson (1971) described additional saucer-like glands referred to as “dewpatches” on the dorsum of the A2—-A4 segments of Cigaritis and Crudaria final instar larvae. These glands secrete a fluid the ants consume. These six genera are mostly associated with Crematogaster ants, except for Crudaria which asso- ciate with Anoplolepis custodiens (Heath 1997a). Lar- vae and pupae of Crudaria wykehami Dickson taken from an ant nest beneath a large flat stone were heav- ily parasitized (ca. 80%) by wasps and a tachinid fly (AH unpublished obs.) Lycaenini (one genus): Lycaena Fabricius, 1807 (extralimital; three Afrotropical species, two in south- er Africa). Small coppery-red butterflies. Larvae naked, lacking DNO and TOs, and not ant-associated. Polyommatini (25 genera): Normally having four larval instars, the DNO appears in the second or third instar, the presence of TOs variable. Euchrysops Butler, 1900 (five in southern Africa). Medium-sized blue or brown lycaenids. The phy- tophagous larvae possess DNO and TOs, the former appearing in the second instar, the latter in the third and have a facultative ant-association (Clark & Dick- son 1971). The genitalia are of the same type as Lepi- dochrysops from which Euchrysops are not easily sep- arated on morphological grounds (see Gardiner 1998). Lepidochrysops Hedicke, 1923 (127 Afrotropical species, 59 in southern Africa). Medium to large, blue or brown with spotted undersides frequenting open and sparsely wooded grassland. Morphologically simi- lar to Euchrysops with remarkably uniform genitalia. All are assumed to be phytopredacious (herbivorous in early instars, later becoming predacious on other in- sects), associated with Camponotus ants, and in many respects, similar in appearance and behavior to the palaearctic Maculinea (see Frohawk 1916, Elmes et al. 1991, Thomas 1983, 1995). The larva feeds on flower buds for the first two instars, then it is carried by a species of Camponotus ant to its brood chamber in the third instar where it is predacious on ant brood. De- spite the large size of this genus, few life histories have been studied in any depth. Our current knowledge is based upon the life history of eleven species by Clark and Dickson (1971), Claassens (1972, 1974, 1976), Henning (1983a, b) and Williams (1990), some being incomplete. However, Claassens (1972, 1974, 1976) JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY studied the interaction between larvae and ants in the laboratory, and observed that larvae of L. trimeni (Bethune-Baker) and L. methymna (Trimen) did not possess TOs at any stage, but the DNO appeared in the second instar and remained until pupation. Claassens also observed that trophallaxis supple- mented the diet of ant brood, also that the larva some- times groomed an ant. Henning (1983b) studied L. ig- nota (Trimen & Bowker), and made similar feeding observations. Henning (1983b) demonstrated that lar- vae of L. ignota and Aloeides dentatis chemically mimic the brood of their attending ants Camponotus niveosetosus and Lepisiota capensis. Corn grits were soaked in epidermal extracts of ant brood or larvae and then offered to the appropriate species of ant. Treated grits were carried by ants to their brood chamber, un- treated grits were ignored. Gas chromatograms of epi- dermal extracts confirmed that chemicals found on the larva were similar to those on the ant brood. In captiv- ity, a fourth instar larva of L. p. plebeia (Butler) was observed feeding on ant eggs as soon as they were laid, when ant brood supply was exhausted (Williams 1990). Orachrysops Vari, 1986 (11 species, all endemic to southern Africa). Formerly included in Lepi- dochrysops, these medium-sized dull blue and gray lycaenids are superficially similar to Lepidochrysops, but their genitalia differ. Little is known about ant- association in their early stages. Formerly thought to be phytopredacious, as in Lepidochrysops (Henning & Henning 1994) but recent work suggests otherwise (Edge & Pringle 1996). The larvae of O. niobe (Tri- men) feed on Indigofera (Fabaceae), and in captivity, were bred to adult without ant presence. However it is unknown if ant-association is obligate or facultative under natural conditions and whether other sources supplement its diet. The DNO and TOs appear in the second and subsequent instars, the TOs are not well developed. The ant suspected of being an associate is Camponotus niveosetosus (Mayr) (Edge & Pringle 1996), recorded among the roots of the food plant where the larvae shelter. Recently, Lu and Samways (2001) showed that O. ariadne (Butler) has an oblig- ate ant-association. Larvae were found in soil beneath the foodplant at depths up to 10 cm, always attended by C. natalensis (Smith). Anthene Doubleday (25 species in southern Africa), Zintha Eliot (one species in southern Africa), Tuxentius Larsen (three species in southern Africa), Leptotes Scud- der (five species in southern Africa), Lampides Hiibner (one species in southern Africa), Tarucus Moore (three species in southern Africa), Harpendyreus Heron (three species in southern Africa), Pseudonacaduba Stempffer (one species in southem Africa), Eicochrysops Bethune- VOLUME 57, NUMBER 1 Baker (three species in southern Africa), Cupidopsis Karsch (two species in southern Africa), Thermoniphas Karsch (one species in southern Africa), Oboronia Karsch (one species in southern Africa), Actizera Chap- man (two species in southern Africa), Zizeeria Chapman (one species in southern Africa), Zizina Chapman (one species in southern Africa), Brephidium Scudder (one species in southern Africa), Oraidium Bethune-Baker (one species endemic to southern Africa), Azanus Moore (five species in southern Africa), Chilades Moore (one species in southern Africa), Zizula Chapman (one species in southern Africa). All 20 of these genera (71 species) are facultatively ant-associated, most having DNO and TOs but often without attendant ants (see Clark & Dickson 1971). Uranothauma Butler (four species in southern Africa), Cacyreus Butler (five species in southern Africa). Not ant-associated, TOs and DNO are absent, except for two species of Cacyreus which have a DNO. Vibrational communication. Using a cardboard poster tube with a paper membrane at one end, final instars of Chrysoritis thysbe (Linnaeus), C. dicksoni, Aloeides pierus and A. pallida grandis were heard to produce ticking or drumming sounds, but C. thysbe also made an intermittent high-pitched buzzing sound (Heath 1998). Sounds have also been noted in the pu- pae of Chrysoritis brooksi Riley and P. irene Penning- ton (Schlosz 1991). These sounds likely play an addi- tional role in communication with ants (De Vries 1990, 1991). Ant-associations and feeding strategies. More than three quarters of the 392 southern African lycaenids are ant-associated, over two thirds of these have obligate relationships. The Poritiinae constituting 8% of lycaenids (30 species) have no direct ant- association® and the Miletinae (36 species) represent- ing 9%, three quarters of which are obligately ant- associated although the remainder feed in the presence of ants. The subfamily Lycaeninae (326 species) accounts for 83% of species, of which the Aphnaeini (131 species) contains a third of all southern African lycaenids, almost all being obligate. Theclini (47 species) and Polyommatini (146 species) account for 12% and 37% respectively; the former being fac- ultative, and latter split between facultative and oblig- ate. The Lycaenini has only two species, neither ant- associated. ° In Deloneura and certain other Liptenini further north in Africa e.g., Epitola, larvae are found in the company of ants but no interac- tion has been observed. It is possible that ant-derived detritus sup- plements the algae on which the larvae feed, or it may enrich the al- gae, making it more attractive, but these hypotheses have yet to be confirmed. 13 TABLE 3. Lycaenids obligately associated with ant genera. Number of obligately ant-associated lycaenids (%) Confirmed Confirmed Ant genus only and predicted Myrmicinae Pheidole 1 (1) 1 (0.5) Monomorium 1 (1) 1 (0.5) Crematogaster 51 (61) 62 (28) Myrmicaria 2 (2) 2D (il) Formicinae Anoplolepis 13 (15) 37 (17) Lepisiota 7 (9) 7 (3) Camponotus 7 (9) 60 (28) Unidentified 2 (2) 47 (22) Totals 84 217 It can be seen from Table 1 that algae or lichen feeders are unique to the two tribes of Poritiinae. The Miletinae feed mostly on Homoptera but also on ho- mopteran secretions, ant regurgitations and possibly detritus and ant early stages at times. Among the Ly- caeninae, almost all the entomophagous records are from the Polyommatiti, with only two confirmed cases from the Afrotropical Aphnaeini, although there may well be more still undiscovered’. Symbiont ant genera. The two main ant genera confirmed in obligate association with lycaenids are Crematogaster (Myrmicinae) and Anoploplepis (Formicinae), together 76%, whereas Camponotus and Lepisiota (Formicinae), represent 9% each (Table 3). Three accounts of ant distribution (Samways 1983, Donnelly & Giliomee 1985, H. Robertson pers. com.) indicate the two dominant ant genera in Southern Africa are Pheidole (Myrmicinae) and Anoplolepis, the latter more so in open habitats. C rematogaster species are also numerous in denser vegetation while Lepi- siota and Camponotus although not always numerous, are to be found in most habitat types in southern Africa (H. Robertson pers. com.). Despite the com- parative dominance of Pheidole ants, only one species has been identified as an obligate symbiont (to a wide- spread species of Aloeides). DISCUSSION Reliability of data. Considerable disparity exists in the Polyommatini between confirmed and predicted ant-association (Table 2) mainly because Lepi- dochrysops (Polyommatiti) life histories are widely be- lieved to be uniform, but few have been confirmed. * Cottrell (1984), Elmes et al. (2001) list Cigaritis takanonis Mat- sumura from Japan as accepting ant regurgitations. Jackson (1937) infers insect feces as additional food for Chloroselas pseudozeritis (Trimen). Fiedler (1991) infers possible ant regurgitations for several Aphnaeini. 14 Obligately ant-associated Afrotropical lycaenid species show little morphological differences among closely related species (Heath 1997b), and recently Heath (2001) synonymized many Chrysoritis taxa, reducing the species by 28%, inferring that similar syn- onymies probably exist in Aloeides, and we believe this is the case for Lepidochrysops. We suspect that southern Africa has an oversplit taxonomy among the ant-associated lycaenids, particularly those with an ob- ligate relationship. Changes in feeding strategy. Although most lep- idopterous larvae are herbivores, some lycaenid species are known to switch from one trophic strategy to another midway through their larval phase; e.g., the Maculinea (Thomas 1983, 1995). In southern Africa this occurs in Lepidochrysops, Thestor, and at least one Aloeides species (Cottrell 1984, Heath & Claassens 2000). More than one food source can also be exploited at the same time e.g., trophallaxis and car- nivory in Cigaritis (Sanetra & Fiedler 1996), and in some Maculinea (Thomas & Wardlaw 1992), and in Lepidochrysops (Claassens 1976, Henning 1983) and Thestor (AH, AJMC) in southern Africa. Dorsal nectary organ and tentacle organs. The DNO and TOs are completely absent in the Poritiinae and Miletinae except in Aslauga (Liphyrini) but one or both are usually present among the Lycaeninae. In some Aphnaeini (Phasis, some Aloeides) the DNO is present in earlier instars but absent in the final instar. Second instar Chrysoritis dicksoni had a DNO that did not secrete honeydew but rather it gave off a pheromone, causing the ants to remain close and stupified (Heath & Brinkmann 1995a). Although the TOs are absent in all instars of Lepidochrysops, the DNO appears in the second instar (Clark & Dickson 1971) and is retained until pupation, as in Palaearctic Maculinea. Fiedler (1998) suggested DNO secretions supplement the adoption procedure. However, Hen- ning’s (1983b) experiments using corn cob grits do not support this for Lepidochrysops. Fiedler (1998) attributed the lack of TOs in Macu- linea to the endophytic life-habit of the early larval in- stars rather than its life-habit within the ant nest, but in Afrotropical Polyommatini (including Euchrysops) the TOs generally appear in the third or fourth instar (Clark & Dickson 1971). Lepidochrysops larvae cease to be herbivorous after the second instar, and hence the TOs would not have been present during any early instar. An alternative hypothesis for Lepidochrysops is that within the nest, the TOs became redundant as a means of recruiting ants and may also have been a hin- derance within the confined space. JOURNAL OF THE LEPIDOPTERISTS SOCIETY ACKNOWLEDGMENTS We thank Adrian Armstrong and the Natal Parks Board (South Africa) and the Chief Executive Officer of the Western Cape Nature Conservation Board for permission to study and collect material in areas under their control. Hamish Robertson, South African Mu- seum, Cape Town is acknowledged for identifying ant species pro- vided by the authors and for giving related input on this paper. Ivan Bampton and Colin Congdon are acknowledged for some recent life history data. Simon Joubert and Swee Peck Quek are thanked for helping to locate the T. basutus larvae and ant associates in KwaZulu-Natal. Phil DeVries is especially thanked for his many helpful comments on the manuscript. LITERATURE CITED Ackery, P. R., R. DE Jonc & R. I. VANE-WricuT. 1999. The but- terflies: Hedyloidea, Hesperioidea and Papilionoidea. In Kris- tensen, N.P. (ed.), Handbook of Zoology 4 (35):263-300. Berlin, de Gruyter. AcKErY, P. R. & D. Rajan. 1990. 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Journal of the Lepidopterists’ Society 57(1), 2003, 17-24 MOLECULAR SYSTEMATICS OF BIRDWING BUTTERFLIES (PAPILIONIDAE) INFERRED FROM MITOCHONDRIAL ND5 GENE KIYOTARO KONDO, TSUTOMU SHINKAWA Department of Liberal Arts, The University of the Air, 2-11 Wakaba, Mihama-ku, Chiba, Japan 261-8586 AND HIROTAKA MATSUKA Nature photographer, 4-7-17 3F Tsurumaki, Setagaya-ku, Tokyo, Japan 154-0016 ABSTRACT. Birdwing butterflies including three genera, Trogonoptera, Troides and Ornithoptera, were subjected to molecular system- atic analysis using sequences of the mitochondrial gene ND5. All three genera descend from a common ancestor and were monophyletic. Tro- gonoptera might have emerged from an ancestral species perhaps in the Miocene, from which Troides and Ornithoptera were also originated. Ornithoptera was further split in two subclusters, one totally corresponding to the subgenus Schoenbergia which lacks male sex marks in the forewing. The other subcluster includes species having sex marks. Green O. priamus, orange O. croesus, and blue O. urvillianus are regarded as an example of intraspecific variety of O. priamus by some authors, but they were totally different phylogenetically. Trogonoptera is limited to the Sundaland, but Troides is distributed across the Wallace line. It may be that Troides arose in Sundaland, but Ornithoptera probably arose in old Wallacea and migrated eastwards producing the various species we see today. Additional key words: Numerous publications were made on birdwing butterflies. Zeuner (1943) united their taxonomy with the geohistory of the Indo-Australian archipelago, con- sidering continental drift. His work was called paleon- tology without fossils. Larvae of troidine butterflies feed on various Aris- tolochiaceae which contain toxins. Even adult butter- flies are toxic, therefore being protected from preda- tion. Troidini is highly varied and widely distributed. According to Hauser (http://www.insects-online.de/ gartfron.htm), a total of 10 genera are recognized in the tribe Troidini from all over the world except Africa, of which three genera are collectively referred to as “birdwing butterflies” for their beauty and birdlike size. These brilliant butterflies live in tropical rain- forests encompassing the Oriental and Australian fau- nal regions. Recent progress in DNA systematics opened a new era in lepidopterology, especially to trace evolutionary process in the light of geohistory, and to reevaluate tra- ditional classification (Brower 1994, Sperling 1993, Sperling & Harrison 1994, Yagi et al. 1999). With re- spect to birdwing butterflies, Morinaka et al. (1999) and Morinaka et al. (2000) reported DNA-based sys- tematic analyses for various troidine butterflies. In the latter study, one species of Trogonoptera, six of Troides and all of Ornithoptera were analyzed. In our study presented here, one Trogonoptera, three Troides and all Ornithoptera were analyzed. We therefore provide an independent test for Morinaka et al. (1999, 2000) studies of birdwing butterflies. Fur- birdwing butterflies, molecular systematics, ND5 gene, phylogenetic tree, Wallacea. thermore Ornithoptera croesus, urvillianus, and eu- phorion were sometimes treated as subspecies of O. priamus by some authors, but we tentatively re- garded them as separate, and evaluated whether this is true. MATERIALS AND METHODS Samples. Butterflies listed in Table 1 were pre- served in 100% alcohol except for four species. Flight muscles from one each of single adult individuals are used to extract DNA. Muscles were digested with AL buffer and proteinase K according to QIAGEN Dneasy Tissue Kit. In four species, O. alexandrae, O. victoriae, O. urvillianus and O. euphorion, legs were removed from old dried museum specimens. They were crushed in a 1.5 L tube and homogenized thoroughly with AL buffer. The DNA was washed ac- cording to the QIAGEN protocol. The DNA was dis- solved in 400 UL of PE buffer. DNA analyses. Primers V1,#A1 and KA2 for am- plification of a part of mitochondrial ND5 gene (873 bases) were designed (Yagi et al. 1999, Su et al. 1996). The most conserved region of ND5 nucleotide se- quences of Drosophila melanogaster, D. yakuba, Carabus japonicus and Anopheles gambiae, which are included in the EMBL data base were used. The poly- merase chain reaction (PCR) was carried out in 50 uL of solution comprised of 130 ng template DNA, 0.2 uM each primer, and 2.5 units of ExTag DNA polymerase and dNTPs and PCR buffer according to Takara protocol. The amplification protocol was 30 cycles of 18 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Table 1. Samples analyzed in this study and GenBank database. No. Species Sampling place DDBJ numbers 1 Papilio memnon Kagoshima, Japan ABO84426 2 Atrophaneura varuna Cameron Highland, Malaysia ABO84427 3 Trogonoptera brookiana Cameron Highland, Malaysia ABO84428 4 Troides hypolitus Sulawesi, Indonesia ABO84429 5 Troides helena Bali, Indonesia ABO84430 6 Troides amphrysus Sumatra, Indonesia ABO84431 i Ornithoptera tithonus Irian Jaya, Indonesia ABO84432 8 Ornithoptera goliath Irian Jaya, Indonesia ABO84433 9 Ornithoptera rothschildi Irian Jaya, Indonesia ABO84434 10 Ornithoptera paradisea Irian Jaya, Indonesia ABO84435 11 Ornithoptera chimaera Aseki, PNG** ABO84436 12 Ornithoptera meridionalis Aseki, PNG** ABO84437 13 Ornithoptera croesus Halmahera, Indonesia ABO84438 Ornithoptera aesacus* Obi, Indonesia 14 Ornithoptera victoriae Bougainville, PNG** ABO84439 15 Ornithoptera priamus Wau, PNG** ABO84440 16 Ornithoptera urvillianus Bougainville, PNG** ABO84441 17 Ornithoptera euphorion Cairns, Australia ABO84442 18 Ornithoptera alexandrae Popondetta, PNG** ABO84443 *DNA amplification was unsuccessful ** Papua New Guinea 94°C for 30 sec, 50°C for 30 sec and 72°C for 1 min in the PCR Thermal Cycler PK2400. The PCR product was separated by 1.0% agarose gel electrophoresis. The gel containing 873 base-DNA fragment was cut out, and the DNA fragment was extracted and purified by the QIAquick gel extraction Kit. In O. aesacus, DNA was not successfully amplified, and this species was eliminated from further analyses. For nucleotide sequencing of the ND5 DNA frag- ment, primers A3, C2 as well as V1, Al and KA2 were used. Nucleotide sequences of both strands of the DNA fragment were determined with the Big Dyeter- minator Cycle Sequencing Ready Reaction Kit. Nu- cleotide sequences of primers were as follows: V1: 5’-CCTGTTTCTGCTTTAGTICA-3" Al: 5’-AATADTAGGTATAAATCATAT-3’ A3: 5’-TTCGAATTTAGCTTTATGTGG-3" C2: 5’-ATCYTTWGAATAAAAYCCAGC-3° KA2: 5’-GTATAATATATTGTTAAACCTGTAG-3’ Sequencing and phylogenetic study. Nucleotide sequences were edited and aligned using Sequencher DNA Sequencing Software. A part of the ND5 nu- cleotide sequences (813 bases) accurately determined in all species was subjected to phylogenetic analysis and registered in the Genbank, as listed in Table 1. Phylogenetic trees were constructed with the Neighbor-Joining (NJ) method. NJ method with the Bootstrap test was performed using the CLUSTAL X program (Felsenstein 1985, Thompson et al. 1997). Evolutionary distances were computed by the Kimura's two-parameter method (Kimura 1980). Max- imum Parsimony method (MP) and UPGMA were also applied using standard default procedures in PAUP 4.0 (Swofford 1993). Scanning electron-microscopy of sex marks. Sex marks were dissected with the ordinary wing por- tion in the male forewings involving the Cu veins, kept in 99.5% alcohol, ultrasonically cleaned for 30 min (OMRON HU-10,46KHz), and dried for scanning electron-microscopy attached to a carbonized sticky tape. The samples were gold-spattered (200 nm) and a Hitachi T300 scanning electron microscope was used for observation. RESULTS DNA phylogeny. Fig. 1A, B and C show phyloge- netic trees for 16 species studied plus three outgroup species. Besides the NJ method, MP and UPGMA produced basically similar trees with some differences. O. aesacus was not included in which DNA sequenc- ing was not successful. Our analyses show that: (1) The three birdwing butterfly genera Tro- gonoptera, Troides and Ornithoptera combined, were monophyletic. (2) An ancestral species gave rise to Trogonoptera, and the ancestor of Troides plus Ornithoptera. (3) Ornithoptera evolved in two subclusters. One totally corresponded to the subgenus Schoenbergia, lacking sex marks in the male forewings like Tro- gonoptera and Troides, and is almost completely en- demic to main island of New Guinea. Schoenbergia appeared to split into two species groups; the rothchildi group and the paradisea group in NJ. VOLUME 57, NUMBER 1 19 P.memnon P.memnon cal dnos63n0 A. varuna T. brookiana Trogonoptera 58 T. hypolitus Saplosy T. helena T. amphrysus 0. tithonus 0. goliath 0. rothchildi 56 o1Bsaquaoyrs 0. paradisea 0. chimaera .meridionalis 94 0. croesus 0. victoriae 0. priamus 52 0.urvillianus yUDW-XAS YIM OUazdoYyz1UUO 0. euphorion 0. alexandrae ®.01 substitutions/site 1A NDS NJ-tree T.brookiana lf O.chimaera 99 O.meridionalis O.croesus O.priamus 10 changes 1B NDS MP-tree A.varuna T.brookiana T.hypolitus T.hypolitus T.helena T.helena T.amphrysus T.amphrysus O.tithonus O.tithonus O.goliath O.goliath O.rothschildi O.rothschildi O.paradisea O.paradisea O.chimaera O.meridionalis O.croesus O.victoriae O.victoriae O.priamus O.urvillianus O.urvillianus O.euphorion O.euphorion 0.01 substitutions/site 1C NDS UPGMA-tree O.alexandrae O.alexandrae Fic. 1. Genealogical trees based on the base sequences of the ND5 gene, 813 base pairs, birdwing butterflies. A, B and C are based on NJ, MP and UPGMA methods, respectively. Numbers below branches are bootstrap values. Trogonoptera, Troides and Ornithoptera were mono- phyletic, Trogonoptera being most primordial, which yielded the other two. Of three Troides species for which permissions were obtained and analyzed, T. hypolithus lives west of the Wallace line. Ornithoptera was split in two subclusters; Schoenbergia, and all other species with sex mark in male forewing. The other subcluster included all species having strik- ing velvet-black male sex marks, and residing in islands from the Moluccas to the Solomons except for three species that live in mainland New Guinea and the north- east coast of Australia; O. priamus, O. alexandrae and O. euphorion. It also split in two species groups each repre- sented by O. croesus and O. urvillianus. Unfortunately, O. aesacus was not included in the analyses, but it ap- parently belongs to this subcluster, because of the pres- ence of a sex mark of the same structure. O. priamus, croesus, urvillianus, euphorion were totally paraphyletic. Table 2 shows genetic distance analyzed using Kimura’s 2 parameter method. Species numbers in Tables 1 and 2 coincide. Sex marks. The basal scales found in the sex marks were uniformly black, while they had refraction lattice in their surface that express iridescent color in the other areas of the wing. The cover scales which are spe- cific to the sex marks were cylindrical with falcate tips, and their radical sockets were enlarged and arranged in tandem with those of the basal scales (Fig. 2). DISCUSSION Previous studies. So-called birdwing butterflies in- volve three genera; Trogonoptera, Troides and Or- nithoptera. The following three subgenera are recog- nized in the genus Ornithoptera based on morphological evidence (D’Abrera 1975, Haugum & Low 1978, Scriver et al. 1995): Schoenbergia (Pagenstecher 1893), including O. go- liath, O. rothschildi, O. tithonus, O. paradisea and O. meridionalis. Aetheoptera (Rippon, 1894), including O. victoriae and O. alexandrae. Ornithoptera (Boisuduval, 1832), including O. croe- sus, O. aesacus, O. urvillianus and O. priamus, in- volving its subspecies which live in various areas from Seram, entire New Guinea and adjacent is- lands to York peninsula, and O. ewphorion which lives in the northern Queensland. There are two successive reports (Morinaka et al. 1990, Morinaka et al. 2000) which are the only pub- lished studies on DNA phylogeny of birdwing butter- flies. Our results are different from theirs in two im- portant ways: (1) In their study Trogonoptera shares common an- cestor with all other Troidine butterflies but is para- phyletic with Troides plus Ornithoptera. Ours suggested monophyly of all three genera; i.e., Trognoptera shares a common ancestor with Troides plus Ornithoptera; (2) Their results are not parsimonious with respect to the sex mark. According to Morinaka et al. (2000), O. alexandrae with a sex mark is monophyletic with JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TABLE 2. Kimura 2-parameter distance matrix (%) of the birdwing butterflies. 4 5 6 7 1 2 3 1 P. memnon = 2 A. varuna 12.861 = 3 T. brookiana 14.921 9.424 = 4 T. hypolitus 15.831 10.841 11.126 5 T. helena 13.884 9.844 10.975 6 T. amphrysus 16.134 10.690 11.978 7 O. tithonus 15.827 13.301 13.155 8 O. goliath 14.921 11.696 11.403 9 O. rothschildi 14.323 11.121 WLW 10 O. paradisea 15.221 11.691 11.710 11 O. chimaera 13.731 10.409 10.267 12 O. meridionalis 14.621 11.263 10.985 13 O. croesus 14.175 9.853 9.148 14 O. victoriae 15.082 9.998 9.012 15 O. priamus 14.476 9.714 8.872 16 O. urvillianus 15.678 11.413 12.000 17 O. euphorion 16.141 12.155 11.748 18 O. alexandrae 15.676 10.985 11.448 8 9 10 8 O. goliath = 9 O. rothschildi 7.586 = 10 O. paradisea 7.276 7.014 = 11 O. chimaera 7.674 7.549 5.904 12 O. meridionalis 6.742 6.899 5.259 13 O. croesus 6.852 7.854 6.298 14 O. victoriae 9.082 8.117 7.661 15 O. priamus 8.229 7.976 7.800 16 O. urvillianus 9.896 9.914 8.906 17 O. euphorion 9.246 6.748 6.723 18 O. alexandrae 8.809 6.339 7.549 15 16 17 15 O. priamus = 16 O. urvillianus 5.109 = 17 O. euphorion 6.483 5.503 = 18 O. alexandrae 7.299 5.500 1.620 Outgroup status changed: 2 taxa transferred to outgroup Total number of taxa now in outgroup = 2 Number of ingroup taxa= 16 Schoenbergia having no sex mark. In many of their trees, sex-marked O. victoria shares a common ances- tor with all others including mixed species with and without sex mark. The sex mark is a conspicuous inherited synapomor- phy, and may be used to validate any attempt of sys- tematics of birdwing butterflies. Namely, trees based on DNA should be consistent with the dichotomy of the species by presence/absence of the mark. The stated inconsistencies suggest some confusion with their results, and we do not quote their reports except their data on O. aesacus, which we do not have. The new phylogenetic classification of the tribe Troi- dini using immature characteristics is fundamentally different from those based on adult morphology (Par- sons 1996). In Parsons’ study, origin of Ornithoptera was distinct from Troides, and the author assumed that the former has evolved in northward-drifting Australia, 7.400 - 8.916 5.640 - 12.176 10.434 11.718 = 9.876 9.162 11.576 7.982 11.879 10.879 11.426 8.543 10.427 9.716 10.133 8.111 10.841 9.565 10.125 8.360 10.434 9.152 10.129 8.111 9.172 8.339 9.016 8.370 9.881 9.742 9.856 10.364 9.595 9.457 9.856 9.493 11.916 10.155 10.708 10.749 11.924 10.169 11.743 10.372 12.201 10.155 11.727 106.58 11 12 13 14 1.995 = 6.564 6.320 = 7.515 7.549 3.293 - 7.240 etal 3.293 0.994 8.476 7.949 6.461 5.919 8.246 WT 2 6.627 7.038 8.099 7.445 7.445 7.579 18 while the latter evolved allopatrically on landmasses on the Eurasian plate. Two successive reports by Morinaka et al. (1999, 2000) indicated monophyly of Troides and Ornithoptera, and totally rejected Parsons’ (1996) ideas, but the position of Trogonoptera in their study is obscure. We also rejected Parsons’ (1996) views and demonstrated the monophyly of all three genera, un- like Morinaka et al. (1999, 2000). Origin of birdwing butterflies based on our study. A bar of 0.01 in Fig. 1 corresponds to one mil- lion years required for 1% of base replacements. Based on the studies of ground beetles (Carabus, Carabidae, Coleoptera) in Japan and European Alps, Su et al. (1998) indicated a value of 4 + 0.5 million years necessary for this magnitude of base replace- ments. They proposed a new figure of 3.6 million years more recently (Su pers. com. 1999). It is therefore possible that ancestral Trogonoptera gave rise to the VOLUME 57, NUMBER 1 ae, % ., Eee eee Fic. 2. Scanning electronmicrograph of the male sex mark, Ornithoptera priamus. The basal (ordinary) scales and the cover scales which are specific to the sex mark area are shown. They are arranged alternatively and in tandem. The latter lost a fan-like appearance as the basal scale, club-like with a falcate tip, and have enlarged radical sockets which probably emit scent. Most scales were removed to show sockets clearly. The scale bar is 10 microns. ancestor of Troides plus Ornithoptera in the early or middle Miocene, if linearity and identical rate of nu- cleotide evolution are assumed as the stated beetles. More studies are necessary to solve this problem more accurately, however. In the late Mesozoic Era, angiosperm trees started to form rainforests. In Sundaland, they stably existed for 130 million years and represented a cradle for bio- diversity. Sundaland is the only place where Tro- gonoptera lives today, and it may be that Trogonoptera arose from an ancestral troidine butterfly in Sunda- land. Troides is most varied there, decreasing in num- bers of species towards surrounding areas north to Tai- wan, west to India and east to Papua New Guinea. Only one each species is found in these extremes of the territory of the genus Troides. Possibly, Troides also arose in rainforests of the Sundaland. The diversification of Ornithoptera and Troides pos- sibly took place in a landmass corresponding to today’s Wallacea. We think this happened east of the Wallace line in any way, because Fig. 1 showed that Troides hy- politus, which lives east of the stated line, is basal within Troides species analyzed and therefore more closely related to the ancestor of Ornithoptera. Many more species of Troides are necessary of course, to evaluate this important hypothesis. Diversification of Ornithoptera. There are two discrete monophyletic subclusters in the genus Or- nithoptera. One group, corresponding to the subgenus Schoenbergia, diversified in two species groups repre- sented by O. rothschildi and O. paradisea as indicated in Fig. 1 based on NJ, but other methods gave some- what different results, although the monophyly of Schoenbergia was supported in all three trees. Schoenbergia was thus natural but Aetheoptera ap- peared unnatural according to our results. Therefore, Aetheoptera is a synonym of Ornithoptera. Subgenus Ornithoptera plus Aetheopteran species appeared monophyletic, however, and all species share a striking synapomorphy, a male sex mark, which characterizes this subcluster. NJ and UPGMA gave the same result, but MP gave some doubts that this subcluster consists of croesus group and urvillianus group. Origin of such diversification of Ornithoptera is un- known, but we propose a hypothesis based on the geo- formation of the area where Ornithoptera lives today. Owing to a complex tectonophysics of the oceanic plates, old Wallacea is much different from today’s land masses. Australia was still far south during the Miocene. New Guinea was not yet formed (Van Bem- melen 1949), but it later rose as the Indo-Australian plate collided with the Pacific plate and initiated oro- genic movements. The Bird’s head peninsula (BH) of the western end of today’s New Guinea was still an iso- lated island and located far west, just east of Halma- hera which were being formed out of a group of is- lands (Hall & Nichols 1990, Burrett et al. 1991). Volcanic island arcs extended south to the line where the Pacific plate disappeared beneath of Indo- Australian plate. Owing to elevation and northward 1000 km LL @eeoeseeoee8d @ee0202000008 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY EARLY MIOCENE 18 Ma VOLUME 57, NUMBER 1 drift of New Guinea, these island arcs were later roughly separated in eastern and western groups of is- lands (Fig. 3). We hypothesize that the ancestral Ornithoptera arose somewhere in old Wallacea and reached an area corresponding to old BH which was still an island, and produced ancestral Schoenbergian species before or after it fused with New Guinea main island which was being formed by the northward drift of Australia. Mt. Arfak of today’s BH is a home of many rare birdwing butterflies, including O. rothschildi, which is unique to this mountain. This species may represent the most primordial patterns of Schoenbergia. On the other hand, we further assume that a sepa- rate group reached volcanic island arcs, evolved there and migrated towards east via island arc, and produced Ornithoptera plus Aetheoptera. Sex mark was proba- bly produced at an early stage of this evolutionary process, perhaps around the time it departed from Wallacea to migrate eastwards and eventually became a hallmark of all species descendant to an ancestral species which entered to the island arc. Distribution. While Schoenbergia is almost con- fined to mainland New Guinea, sex-marked species are found in islands from the Malucca to the Solomons with three exceptions; O. euphorion, O. alexandrae and O. priamus which live in Australia and New Guinea. The south-west corner of main island of Papua is regarded as a part of Australia, unlike the rest of New Guinea (Ollier & Bain 1994). We showed that O. euphorion is not a subspecies of O. priamus. It be- longs to sex-marked species group and probably, ar- rived at an ancient landmass which corresponds to to- day’s Queensland and South-east New Guinea. O. alexandrae which is closely akin to O. euphorion ac- cording to Fig. 1 is endemic to main island New Guinea, being confined to a small area near Popon- detta. We assume that O. alexandrae was also a species originally evolved in the volcanic island arc. It has a velvet-black sex mark and probably evolved in isolated islands that once existed off the northeast coast of old Papua, which later became a part of todays New Guinea due to the gross elevation of land. Eastern Papua, especially Huon Peninsula area, is known for a large-scale land elevation, proven by coastal shelves containing corals and sea shells found even in the alti- tudes of 200 m (Bloom et al. 1974). << The origin of O. priamus is obscure. It is interesting to note that Morinaka et al. (2000) showed a tree indi- cating that O. aesacus and O. priamus share a close common ancestor. This is good circumstantial evi- dence that O. priamus arose in old Wallacea and mi- grated eastwards and finally invaded entire Papua New Guinea with neighboring island and northern Aus- tralia. O. aesacus occurs only in a small island of Obi, closely south to Halmahera, and its bluish-green col- oration suggests a kinship with O. priamus. Possibly O. aesacus is a surviving relic of the ancestral species of O. priamus. This question can be solved when various subspecies of O. priamus were analyzed along with O. aesacus and O. croesus. Further studies. Many puzzles remained unsolved. (1) A complete study of Troides is necessary. Troides is the only birdwing butterfly genus containing species which live on the both sides of the Wallace line. The eastern margin of Sundaland is marked by a deep ocean ditch formed by disappearance of the Pacific plate beneath the Eurasian plate, thus stably existed since Mesozoic Era, forming a strong barrier against migration of animals; i.e., the Wallace line. It may be that Troides arose in Sundaland and perhaps migrated across the Wallace line with trade wind. Which particular Troides species is most basal re- mains a puzzle. We suspect if Troides rhadamantus do- hertyi is a candidate of the relic because of its simplis- tic yellow patterns, small size compared with other subspecies of T. rhadamantus, strong tendency to pro- duce a melanic form and its delimited distribution in the Talaud Islands of Indonesia. Yellow pigment for- mation may be still weak in this species. We still do not know whether Ornithoptera arose west of the Wallace line, but a thorough study of Troides species may give a clue to this interesting question by evaluating how Ornithoptera is related with various Troides species re- siding west and east of the line. (2) A complete study of various subspecies of O. pri- amus is necessary. This species is most widely spread and probably tells about routes of expanding of Or- nithopteran distribution, not only O priamus. (3) Questions in the species level are many. For ex- ample, Troides magellanus and T. prattorum are simi- larly patterned sharing pearly shine in the upper side of female hindwing. O. magellanus in common in the Phillipines, but O. prattorum is restricted to elevated Fic. 3. Southern Pacific landmasses in the Miocene (Burrett et al. 1991). Heavy-dotted landmasses are on the Asian plate, but thin-dotted landmasses are on the Indo-Australian plate. A thick arrow indicates drifting direction of Australia, and a thin arrow drifting of Asian islands. Plates submerged at the barbed lines towards the direction of the barbs. Ma, million years; K, Kalimantan; S, Sulawesi; B, Bachan; BS, Bang- gai-Sula; O, Obi; Bu, Buru; Se, Seram; H, Halmahera; BH, later Bird’s head peninsula of Irian Jaya; IMA, Inner Melanesian are. 24 altitudes in small island of Buru far south in the Banda sea. Whether both are close in term of DNA is very in- teresting and perhaps a complex tectonophysical movements of the area may shed some lights on this strange distribution of two sister species. These are few examples of puzzles. To solve them, we attempted to obtain fresh alcohol specimens in vain. Birdwing butterflies are protected fauna. Proba- bly, a comprehensive international collaborative proj- ect is necessary to persuade Governments of the coun- tries where these lovely butterflies live. ACKNOWLEDGMENTS We are obliged to Dr. Aoki, Invertebrate Laboratory 1, The Re- search Institute of Evolutionary Biology, The Tokyo University of Agriculture, who supplied specimens of four species, O. alexandrae, O. victoriae, O. urvillianus and O. euphorion, from which he re- moved legs for us. Time of collection of these species were all earlier than 1967. For all species other than the 4 species stated above, copies of certificates were submitted to the editorial board which are dated, stamped, signed, and issued by each respective Government authorities, according to the rules of the Convention on Interna- tional Trade in Endangered Species of Wild Fauna and Flora. We are indebted to Mr. Hiromichi Makita in various ways. Also, we are obliged to Ms. Yoko Sugita for preparation of the manuscript. Although S. Morinaka was an undergraduate student of the Uni- versity of the Air, his work was carried out elsewhere without our awareness prior to publication. LITERATURE CITED D’ABRERA, B. 1975. Birdwing butterflies of the world. Lands- downe, Melbourne, Australia. 42 pp. VAN BEMMELEN, R. W. 1949. The geology of Indonesia, Vol 1A, General Geology of Indonesia and Adjacent Archipolagoes. Martinus Nijhoff, The Hague, Holland. 719 pp. BLoom, A. L., W. S. BROECKER, J. M. A. CHAPPELL, R. K. MATTHEWS & K. J. MESOLELLA. 1974. Quaternary sea level fluctuations on a tectonic coast; new 230Th/234U date from the Huon Peninsula, New Guinea. Quaternary Res. 4:85-205. BroweER, A. V. Z. 1994. Phylogeny of Heliconius butterflies in- ferred from mitochondrial DNA sequences (Lepidoptera; Nymphalidae). Mol. Phylogenet. Evol. 3:159-174. BurRRETT, C. D., R. BERRY & R. VERNE. 1991. Asian and South- western Pacific continental terranes derived from Gondwanna, and their biogeographic significance. Aust. Syost. Bot. 4:13-24. FELSENSTEIN, J. 1985. Confidence limits of phylogenies: an ap- proach using bootstrap. Evolution 39:783-791. HALL, R. & G. J. NICHOLS. 1990. Terrane amalgamation in the Philippine Sea margin. Tectonophysics 181:207-222. Haucu, J. & A. M. Low. 1978-1985. A monograph of the birdwing butterflies, 1-5. Scandinavian Science Press Ltd., Klampenborg. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY KimurA, M. 1980. A simple method for estimating evolutionary rate of base substitutions through comparative studies of nu- cleotide sequences. J. Mol. Evol. 32:128-144 . MoRrInaka, S., T. MAEYAMA, K. MAEKAWA, D. ERNIWATI, S. N. PRI- Jono, I. K. Ginarsa, T. NAKAZAWA & T. HIDAKA. 1999. Molec- ular phylogeny of birdwing butterflies based on the representa- tives in most genera of the tribe Troidini. Entomol. Sci. 2:374-358. MorInaka, S., N. MINAKA, M. SEKIGUCHI, ERNIWATI, S. N. PRIJONO, I. K. Grnarsa & T. Miyata. 2000. Molecular phylogeny of Birdwing butterflies of the Tribe Troidini (Lepidoptera: Papil- ionidae) using all species of the genus Ornithoptera. Biogeog- raphy 2:103-111. OLLIER, C. D. & J. H. C. Barn. 1994. Geology. In Ryan, P. (ed.), Encyclopedia of Papua and New Guinea. Vol 1. Melbourne University Press, Melbourne, Australia. 478 pp. Parsons, M. J. 1996. Gondwannan evolution of the Troidine swal- lowtails (Lepidoptera: Papilionidae): cladistic reappraisals using mainly immature stage characters, with focus on the birdwing Ornithoptera Boisduval. Bull. Kitakyushu Mus. Nat. Hist. 15:43-118. SCRIBER, J. M., Y. TSUBAKI & R. C. LEDERHOUSE (EDS). 1995. Swal- lowtail butterflies; their ecology and evolutionary biology. Sci- entific Publishers, Gainesville. 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 Papilio machaon groups of swallowtail butterflies. Evolution 48:408—422. Su, Z-H., T. OHAMA, T. S. OKADA, K. NAKAMURA, R. ISHIKAWA & S. Osawa. 1996. Phylogenetic relationships and evolution of the Japanese Carabinae ground beetles based on mitochondrial ND5 gene sequences. J. Mol. Evol. 42:124-129. Su, Z-H., O. Tominaca, M. Okamoto & S, Osawa. 1998. Origin and diversification of hindwingsless Damaster ground beetles with the Japanese Islands as deduced from mitochondrial ND5 gene sequences (Coleoptera, Carabidae). Mol. Biol. Evol. 15:1026-1039. SWOFFORD, D. L. 1993. PAUP (phylogenetic analysis using parsi- mony) version 3.1.1. Smithsonian Institution, Washington. THOMPSON, J. D., T. J. Gipson, F. PLEWNIAK, F. JEANMOUGIN & D. G. Hiccins. 1997. The Clustal X Windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tool. Nucleic Acids Research 24:4876—4882. Yact, T., G. SASAKI & H. TAKEBE. 1999. Phylogeny of Japanese pa- pilionid butterflies inferred from nucleotide sequences of the mitochondrial ND5 gene. J. Mol. Evol. 48:42-48. ZEUNER, F. E. 1943. Studies in the systematics of Troides Huebner (Lepidoptera, Papilionidae) and its allies; distribution and phy- logeny in relation to the geological history of the Australian Archipelago. Trans. Zool. Soc. Lond. 25:107-184. Received for publication 7 June 2000; revised and accepted 14 August 2002. Journal of the Lepidopterists’ Society 57(1), 2003, 25-35 WHY NATURAL HYBRIDS ARE HARD TO DETECT AND VERIFY: EXEMPLIFIED BY A RARE PRIMARY HYBRIDIZATION EVENT BETWEEN TWO TIGER SWALLOWTAIL BUTTERFLY SPECIES IN NORTHERN MICHIGAN JENNIFER DONOVAN AND J. MARK SCRIBER Department of Entomology, Michigan State University, East Lansing, Michigan 48824 USA ABSTRACT. Phenotypically intermediate specimens are often attributed to interspecific hybridization. Such observations of putative in- terspecific hybridizations are sufficiently rare to warrant literature records. However, it is seldom that additional evidence is available to docu- ment such an event. To illustrate this process, we have examined several lines of evidence documenting such a rare natural interspecific (“Pri- mary,” F-1) hybridization event in northern Michigan (Charlevoix Co.) between a P. canadensis female and a P. glaucus male. We describe analyses using a historical framework of extensive spatial and temporal sampling of multiple traits to illustrate the complexity of approaches re- quired for identifying hybrids. We emphasize the importance of thorough ecological as well as morphological trait analysis relative to parental types for documenting true natural hybrids. Additional key words: Papilio glaucus, Papilio canadensis, diapause, morphological analysis, host plants. Although rare, interspecific hybridization in natural populations has been reported from several Lepidoptera families (Remington 1968, Sperling 1990). However, ex- tensive study is required to convincingly demonstrate or document such events. Use of multivariate morpho- metric analyses are often required even in large and distinct species such as the giant silkmoths of the Hyalophora cecropia group (Collins 1984) or butterflies such as the Limenitis species group (Platt 1983, Boyd et al. 1999). We used a multi-trait approach combined with an extensive historical data base for these diagnos- tic traits between Papilio canadensis and P. glaucus. The swallowtail butterflies of the Papilionidae family are also a large, well-known showy group of more than 560 species with generally well-studied nat- ural histories and host plant relationships (Scriber et al. 1995). The genus Papilio has historically included nearly half (>200) of the species in the family world- wide, although the precise phylogenetic relationships are still being clarified (Munroe 1960, Hancock 1983, Miller 1987, Sperling 1987, Scriber 1995, Reed & Sperling 1999). Hand-pairing of these Papilio has pro- duced extensive interspecific hybridization with the production of viable offspring in laboratory research programs (see reviews in: Ae 1995, Brown et al. 1995, Clarke 1995, Scriber et al. 1990, 1995, 2003). Natural hybrid zones have also produced specimens “interme- diate” in appearance that have been assumed to be in- terspecific hybrids. Several examples have been re- ported from the Papilio machaon species group in North America (between the P. machaon, P. polyxenes, or P. zelicaon; Sperling 1987) and in other areas be- tween P. machaon and P. hospiton (Clarke & Larson 1986, Clarke 1995). Additional putative interspecific hybrids of Papilio have also been reported on other continents (Hancock 1983, Collins & Morris 1985, Johnson & Matusik 1987, Tyler et al. 1994). We have also conducted extensive interspecific hy- bridization with tiger swallowtails of the North Ameri- can Papilio glaucus group (reviewed in Scriber et al. 1995) and have used multivariate morphometric analy- ses of adult wing traits (Luebke et al. 1988, Scriber 1990, Scriber 2002a) and diagnostic larval characters (Hagen et al. 1991, Scriber 1998) and electrophoreti- cally-detectable species-diagnostic allozymes to iden- tify natural populations of introgressed interspecific hybrids in the field (Hagen & Scriber 1991, Scriber 1996a). Morphological traits from known lab-hybrids have been used to identify suspected intermediates between P. glaucus and P. rutulus (Scott & Shepard 1976, Clarke & Clarke 1983, Scriber et al. 1990): P. ru- tulus and P. multicaudatus (Brower 1959, Garth and Tilden 1986); PR eurymedon and P. rutulus (Wagner 1978, West & Clarke 1988, Scriber et al. 1995); P. glau- cus and P. multicaudatus (Scriber et al. 1995, Rahn 2001); and P. glaucus and P. canadensis (Scriber 1982, Luebke et al. 1988, Scriber et al. 1996, 2002a). Our studies of the natural hybrid zone between Pa- pilio glaucus and P. canadensis that exists from Min- nesota and Wisconsin through Michigan and central New York State to southern New England have demon- strated the existence of several historically-stable, geo- graphically-defined, and ecologically-significant trait step clines that differ interspecifically in these two tiger swallowtail species (Scriber 2002b; Table 1). These trait differences include differential abilities to detoxify tulip tree leaves (Liriodendron tulipifera, and other species of Magnoliaceae) and quaking aspen leaves (Populus tremuloides, and other species of Sali- caceae). These differences (Scriber 1986b, Lindroth et al. 1988, Scriber et al. 1991) have a genetic basis, and hybrids are able to detoxify and grow on plants in both families due to intermediate levels of autosomally con- trolled detoxification enzymes (Scriber 1986b, Scriber et al. 1989, 1999). In 3-choice oviposition bioassays (quaking aspen, black cherry, and tulip tree leaves) this female (#15116) laid a little more than 40% of her eggs on aspen, which closely fits the typical profile for a P. canadensis female (Scriber et al. 1991, Scriber 1994). Females of P. glaucus typically place fewer than 5% of their eggs on quaking aspen in such an arena as “mis- takes” (Scriber 1993). Another major interspecific ecological trait differ- ence is that P. glaucus individuals have an environ- mentally-determined pupal diapause induction (they directly develop into adults at long photoperiods: Rockey et al. 1987a, Vallella & Scriber 2002) whereas P. canadensis has an “obligate diapause” that is pho- toperiod insensitive (Scriber 1988) and sex-linked on the X-chromosome (Rockey et al. 1987b). Hybrids are variable in this respect, depending on the direction of the cross (obligate diapause tendencies are inherited from the father’s X-chromosome in female offspring since the females are the heterogametic sex in Lepi- doptera). Therefore, a male P. canadensis parent in an interspecific hybrid will produce hybrid daughters that all diapause, even under long day photoperiods. Below, we describe results of the first verification of a natural “primary” interspecific hybridization event for tiger swallowtail butterflies. Despite the occur- rence of hybrid “morphotypes” in Wisconsin (Luebke et al. 1988) and elsewhere (Scriber 2002b) and evi- dence of extensive genetic introgression of species- diagnostic allozymes and Magnoliaceae host detoxifi- cation abilities in the last few years (Hagen 1990, Hagen et al. 1991, Hagen & Scriber 1991, Scriber 1996, 2002a, Ording 2001), we have never found a pri- mary F-1 hybrid individual in the field. Of thousands of individuals collected and examined, none have been heterozygous for all of the diagnostic allozyme traits (PGD, LDH, and HK), despite wing traits, larval detoxification and oviposition behavior that would be considered diagnostically “intermediate” between parental species types (Ording 2001, Scriber 2002b). The LDH-100 allele is apparently quickly (and totally) selected out of the hybrid populations in Wisconsin, Michigan, eastern New York, and southern Vermont (Scriber 2002a, b), and other than in this family from our Charlevoix female we have never seen LDH-100 north of Clinton Co. in southern Michigan P. glaucus territory (Fig. 1; Nielsen 1999). METHODS Female butterflies collected in the field were brought to the lab for 3-choice oviposition preference assays (tulip tree, which is toxic to P. canadensis; quak- ing aspen, which is toxic to P glaucus; and black cherry, which is mutually and naturally acceptable to both species for larval survival and growth). These JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY leaves were arranged along the sides of round clear plastic arenas that rotate in front of lights 10 times per hour (see methodology details in Scriber 1993). Eggs were counted and collected daily while females are fed 20% honey water solution. Neonate larval eclosion from the eggs occurred at approximately 5-7 days in controlled environment chambers at 25°C and 16:8 photoperiods. These fresh neonate (first instar) larvae were distributed evenly across the three host plants and reared in controlled environment chambers with conditions set as described below. Leaves of tulip tree, quaking aspen and black cherry from Ingham County were changed each 48 hours and survival noted for all individual larvae in all treatments. The results of our odd brood #15116 were discov- ered in the process of conducting a larger study of the impacts on larval/pupal offspring fitness of interspe- cific hybrids relative to parental types (Donovan 2001). The pure parental genotypes (Papilio glaucus and P. canadensis) were reared simultaneously with hybrid larvae of reciprocal pairing types (canadensis females mated to glaucus males; and glaucus females mated to canadensis males) on three host plants (tulip tree, quaking aspen, and black cherry) at three different temperatures (15°C, 23°C, and 31°C). For each family of all 4 genotypes during 1999 and 2000, 36 larvae were randomly distributed to the 9 treatments (two larvae per treatment) with 2 growth chamber repli- cates of each. For the purposes of comparison here, the temperature treatments and chamber replications were lumped to highlight the 3 host plant effects (Table 2). In 1999, 5 canadensis families (180 larvae), 2 glaucus families (72 larvae), 6 Pc x Pg families (216 larvae) and 4 families of Pg x Pc (144 larvae) were bioassayed. In 2000, there were 5 canadensis families (180 larvae), 4 (144 larvae) of Pc x Pe, 4 families (144 larvae) of Pg x Pc, with 5 glaucus families (180 larvae). Pupae were weighed and set up in small cylindrical screen cages for eclosion as adults and wing expansion. After 6 weeks, remaining pupae were presumed to be in diapause and were moved from chambers to storage in dark coolers maintained at 4-5°C until the following Spring when they were again set up in screen cylinders for adult eclosion. Adult specimens were scored for morphological wing traits as in Luebke et al. (1988), sometimes mated for livestock rearing, and then frozen alive at -80°C for subsequent electrophoresis. Electrophoresis techniques using cellulose acetate plates for the diagnostic allozymes (LDH = lactate de- hydrogenase; PGD = 6-phosphoglucose dehydroge- nase; and HK = hexokinase) were basically conducted as in Hagen and Scriber (1991) modified slightly as in Stump (2000). VOLUME 57, NUMBER 1 nS) | TasBLeE 1. Summary of physiological, biochemical, and behavioral differences between P. glaucus and P. canadensis, and their modes of in- heritance, if known. See text for additional explanation. Character P. glaucus Environmental YES determination of pupal diapause Oviposition preference tuliptree Larval survival (Aspen) very low Larval survival (Tuliptree) high Hexokinase (Hk) alleles 100 Lactate dehydrogenase 100 (Ldh) alleles 6-Phosphogluconate 100, 50 dehydrogenase (Pgd) Adult hindwing width 10-40% black on anal cell P. canadensis Inheritance Reference NO X-linked 19; aspen X-linked 3 high polygenic 45,6 very low polygenic 4,5,16 110 autosomal 8 80, 40 X-linked 17h, 3 125, 80, and X-linked 1. 73 150 55-90% autosomal 9, 10 1. Hagen and Scriber 1989; 2. Rockey et al. 1987a; 3. Scriber 1994; 4. Scriber 1986b; 5. Scriber 1988; 6. Scriber et al. 1989; 7. Hagen 1990; 8. Hagen et al. 1991; 9. Luebke et al. 1988; 10. Scriber 1982; 16. Scriber 2002a. RESULTS Detection and verification of hybridization. Among female butterflies of Papilio canadensis col- lected in northern Michigan (Charlevoix County 1999) we obtained offspring (from a single family derived from a field captured female) that were clearly primary hybrids. Neonate larval survival, and even survival through the final instar, of some offspring on tulip tree leaves (Liriodendron tulipifera of the Magnoliaceae) was our first clue. In addition, morphological traits of larvae were intermediate, and pupae exhibited direct development (non-diapause) resulting in eclosion of adults within a 1-3 weeks, including females. This lack of diapause was puzzling since basically all pupae of P. canadensis usually enter an obligate diapause, con- trolled by an X-linked trait. Adult offspring that emerged also had wing patterns that clearly looked to be similar to our lab-paired interspecific (“reference”) hybrid specimens. Electrophoresis using allozymes “diagnostic” (with nearly fixed differences) for the 2 tiger swallowtail species (P. canadensis and P. glaucus) confirmed the identity of the field-captured female as “canadensis” and confirmed all of the sons and daugh- ters as primary F-1 hybrid offspring (indicating that the unseen father was a P. glaucus). We have examined this Charlevoix population and others near the hybrid zone for many years, yet have never found any evidence of a primary F-1 hybrid (heterozygous for all diagnostic allozymes). The results we report here (having occurred at a distance consid- erably north of the center of the Michigan hybrid zone; >150 km), may be partly explained due to sev- eral warmer than normal years and recently docu- mented general northward movement of several typi- cal P. glaucus traits, from the south (Scriber 2002a, b). Extensive interspecific genetic introgression from the southern species (“glaucus”-type traits) was known to have occurred northward along seasonal isoclines of to- tal degree day accumulations of 2600-2300 (above a base 50°F), especially along the warm Lake Michigan shoreline and since 1998 (Ording 2001, Scriber 2002a). Our observations began with the comparative study of multiple families of larvae from lab-paired hybrids (both reciprocal types) for comparison of fitness with parental species as part of another study (Donovan 2001). One of the first “odd” characteristics of family #15116 that we observed in offspring of this female P. canadensis collected in Charlevoix County, Michigan in 1999 was that many (92%; Table 2) of the neonate larvae survived the entire first instar feeding on tulip tree (Liriodendron tulipifera). Exceedingly few neonate larvae of P. canadensis have ever survived the first instar on tulip tree (less than 1% of 446 individu- als from dozens of families; Scriber et al. 1995), but these offspring from family #15116 were also surviving into the later instars and pupae as well (18 of 36; Table 2). When in the final instar, it was clear that the super- anal (dorsal) stripe was only weakly (faintly colored) yellow, instead or sharply yellow with pointed protu- berances as observed typically with “canadensis.” These larval “tail” patterns looked much more like those seen in the hybrids or “glaucus” larvae (JMS, pers. obs.). After pupation, many individuals of this family de- veloped directly into adults within 10-16 days (non- diapausing), including females. This was also a very atypical character for P. canadensis, which have an en- vironmentally non-sensitive (obligate) diapause (see Table 1). These results led us to suspect the possibility of interspecific hybridization as a possible explanation. Careful checks of our rearing records and data charts JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TABLE 2. The 10-day and full larval survival of Papilio canadensis, P. glaucus, and their primary (both reciprocal) hybrids as a function of host plant (TT = tulip tree; BC = black cherry; QA = quaking aspen). The number of larvae for each plant is indicated. The overall percent of direct developing (i.e., non-diapausing) pupae is also presented for each genotype. This study was part of a larger hybrid vigor study in 1999 and 2000 (Donovan 2001). The odd brood we discovered and report here (#15116 from Charlevoix Co. Michigan) is included for comparison. 10-day survival (%) Genotype & Year (N) TT BC QA P. canadensis 1999* (60) 45.0 93.3 83.3 2000* (60) 36.7 78.3 75.0 2-yr. mean 40.8 85.8 79.2 Pe x Pg 1999 (72) 75.0 84.7 84.7 2000 (72) rea 83.3 79.2 2-yr. mean 76.1 84.0 82.3 Pg x Pe 1999 (48) 83.3 91.7 83.3 2000 (48) 89.6 85.4 81.3 2-yr. mean 86.5 88.6 82.3 P. glaucus 1999 (24) 58.3 50.0 0.0 2000 (60) 85.0 80.0 15.0 2-yr. mean 71.7 65.0 Veo Family #15116 (12) 91.7 83.3 75.0 Survival to pupa (%) Direct development TT BC QA % (n = pupae) 3.3 45.0 36.7 0.0 (52) 15.0 53.3 36.7 0.0 (64) 9.1 49.2 36.7 0.0 59.7 58.3 26.4 33.3 (156) 45.8 52.1 30.9 34.0 (100) 52.8 55.2 28.6 33.6 56.3 54.2 29.2 32.6 (92) 35.4 29.2, 8.3 18.6 (43) 45.9 41.7 18.8 25.6 37.5 45.8 0.0 33.3 (30) 25.0 37.5 0.0 17.2 (29) 31.3 41.7 0.0 25.3 50.0 66.7 16.7 48.8 (16) * Populations selected for this hybrid fitness study (Donovan 2001) as P. canadensis (from Emmet, Cheboygan, Charlevoix, and Isabella Counties in Michie and Clark Co. Wisconsin) exhibited some introgression from P. glaucus, especially with regard to tulip tree detoxification abilities since the regional climatic warming that began in 1998. In the years from 1980-1997, larval survival on tulip tree from the same populations in Clark Co. Wisconsin and those in Michigan north of Clare was essentially non-existent (Scriber 1982, 2002a). convinced us that we were dealing with the offspring of a field-collected P. canadensis female that had somehow naturally mated with a P. glaucus male be- fore we captured her. We conducted electrophoresis on the offspring and the mother to confirm this hy- pothesis. Our electrophoresis analyses produced allozyme profiles that basically confirmed the assessment that this was a “pure” P. canadensis female that had mated to a P. glaucus male before we captured her in Charle- voix County. The mother was clearly a typical P. canadensis (hemizygous as LDH-80 & PGD-125; ho- mozygous HK-110; Hagen & Scriber 1989). The hind- wing black band width of the anal cell was 70% of the distance to the origin of the Cu2 vein, and also was clearly “canadensis” (“glaucus” bands are generally less than 40%: see Scriber 1982, Luebke et al. 1988). In addition, the submarginal yellow forewing band on the ventral side was solid as in “canadensis” (not a series of yellow spots as in “glaucus:” Luebke et al. 1988). The 2 male offspring were both heterozygous at the diagnostic PGD locus (100/125) and also heterozygous at the LDH locus (100/80) as would be expected for primary hybrids (bottom of table 3). Their HK alleles were not resolved clearly. The 6 daughters tested were also all as expected for a primary hybridization event (as we hypothesized), exhibiting the hemizygous sex- linked PGD 100 and the LDH 100 alleles which had to have been inherited from their putative P. glaucus father. The autosomal HK were heterozygous, as would be expected, for the 4 daughters that had clearly visualizable bands on the gel. Individual specimens collected from this Charlevoix County population from 1998-2001 were scored and analyzed for trends of differences in the individual black band widths of hindwings for females and males (Figs. 1, 2). The parental P. glaucus typically has band widths that are 40%-10%, while P. canadensis typically shows 55%-90%. Reference hybrids from lab pairings of 29 different families (with more than 500 lab-reared adult offspring) range from 35%-60% (Scriber 1982, 2002a). It seems from the scoring indices (Figs. 1, 2) that 1998 and 1999 populations were characteristically P. canadensis in nature. However, both 2000 and 2001 males and 2001 females have individuals with signifi- cantly narrower band widths, which are likely to repre- sent interspecifically intermediate traits. Analyses us- ing t-tests show significant mean differences of the 2001 females from both 1998 and 1999 females (p < 0.027 and p < 0.020, respectively; Fig. 1). Females from 2000 were intermediate and not statistically nar- rower than those from 1998 and 1999. Males from the VOLUME 57, NUMBER 1 TABLE 3. Summary of male allozyme frequencies (the most common alleles)for Papilio populations in Michigan compared to Charlevoix Co. and the odd family (#15116). Some data pre-1992 from Hagen et al. (1991), and some 1998-2000 (from Stump 2000; Ording 2001; Scriber et al. unpubl.). (* = diagnostic for the species P. glaucus) LDH Latitude & Counties (n) 100* 80 40 (n) 45.8-45.3N (Northern Lower Peninsula) Charlevoix 1992 50) 0 92 8 (50 1999 (8) 0 87 13 (8 2000 20) 0 90 10 (33 Cheboygan 47) 0 94 6 (47 Emmet 1992 28) 0 100 0 fo) 1999 24) 0 91 9 (24) Presque Isle 50) 0 100 0 (50 42.8-42.0N (Southern Michigan) Allegan 24 92 8 0 (24 Clinton (3 100 0 0 (3 Ingham Pre-1992 61 85 13 2 (61 1992 12) 100 0 (0) (12) Jackson (4 100 0 0 (4 Lenawee 34) 100 0 0 (35) St. Joseph 29 94 6 0 (29 Washtenaw (29) 93 a 0 (29) ** Offspring of Hybrid Family #15116 Males (2) 50 50 0 (2) Females (6) 100 0 0 (6) 2000 population have narrower bands than those from both 1998 (p < 0.001) and 1999 (p < 0.019). While the 2001 males exhibit hybrid-like (narrower) bands in some individuals, the mean 55.8% appeared slightly shifted back toward the “canadensis” type. Nonethe- less, 2001 males were still significantly narrower than both 1998 (p < 0.002) and 1999 (p < 0.012; Fig. 2. DISCUSSION Natural hybrids have always been exceedingly diffi- cult to document in the field for many reasons. Pri- mary among these reasons is that hybrids do not differ greatly in appearance from the parental types in vari- ous morphological characters (Platt 1983, Arnold 1997, Porter et al. 1997). Multivariate analyses with known parental types and known hybrids (lab-paired as “reference groups’) are usually needed to identify the relatively rare hybrids and introgressed individuals from all of the field-collected “unknowns” (Collins 1984, Sperling 1987, Luebke et al. 1988, Scriber 1990, Boyd et al. 1999). Physiological (e.g., diapause regula- tion or host plant use abilities) or biochemical differ- ences (allozymes or mitochondrial DNA) can help with the taxonomic diagnoses, but such analyses are seldom undertaken. We conclude from our laboratory documentation that the female P. canadensis collected from northern PGD HK 100* 50* 125 80 150 (a) 00x 0 0 0 94 4 2 (3) 0 100 0 0 100 0 0 (8) 13 87 0 95 5 0 (33) 0) 100 1 0) 89 2 ® (32) 0) 100 0 0 80 14 6 (26) 0) 100 0 0 88 4 (7) 14 86 13 0 82 4 1 (36) 0 100 96 0 4 0 0 (0) = = 100 0 0 0 0 (3 100 0 93 ] 6 0) 0 3} 95 5 84 8 8 0 0 12 100 0 100 0 0 0) 0 100 0 94 3 3 0 0 (28 93 W 97 0 3 0 0 (5 100 0 100 0 0 0 0 (28 73 27 50 0 50 0) 0 (2) 50 50 100 0 0 0) (3) 50 Michigan (Charlevoix County) in June 1999 must have mated to a male P. glaucus to produce “primary” inter- specific hybrids observed in family #15116. Popula- tions of tiger swallowtail butterflies collected at this same location (approximately 45 degrees North lati- tude) have always exhibited strict “canadensis” traits in the past, prior to 1999 (Scriber 2002a, b). The larvae have never survived to pupation, and extremely few have even survived the neonate (first instar) stage on tulip tree leaves. Other individuals collected at this Charlevoix County site but reared on acceptable natural hosts such as black cherry (Prunus serotina) and quak- ing aspen (Populus tremuloides) all exhibited the dia- pause trait, even under long day photoperiods (16-18 hours; Table 2). Individuals of this population have his- torically had “canadensis” type allozymes (PDG, LDH, and HK; Donovan 2001, Ording 2001; Table 1) and have possessed morphological (black bands in hind wing) traits that were typically “canadensis” in nature (>55% of the distance of anal cell to origin of the Cu-2 vein), not “glaucus-like” (<40%, Scriber 1982; Fig. 1). However, our postulated 1999 primary hybridiza- tion event would have been possible only with the presence of a male P. glaucus at considerably greater distance North than ever previously reported from the center of the hybrid zone in Michigan (Scriber 1996a). We do know that 1998 and 1999 were exceptionally 30 Charlevoix Co. MI 1998 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Mean 64.1 (6.9) n= 17 0 5 10 is AO 25 30 25 20 25 50 BS GO 05 70 75 80 8S OO OG Charlevoix/Emmet Co. Ml 1999 Charlevoix Co. Ml 2000 Number of Individuals Charlevoix Co. MI 2001 oN FF OD Mean 65.5 (7.9) n= 11 0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 85 90 95 Mean 60.5 (10.3) n= 33 0 5 10) 15) 210-25) 30) S554 0) 45S FNS Si 60 65/97/0875) 80) “Si5eigi0ne9)5 MEan 58.3 (9.5) n= 38 0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 85 90 95 Black Band Widths(%) Fic. 1. Population black band widths (hind wing) trait frequencies for individual females collected in 1998, 1999, 2000, 2001 in Charlevoix County, Michigan. Our 1999 odd brood (#15116) female scored as a 70%. Immediately adjacent (6-12 km) Emmet Co. females were included in 1999, since we only collected 3 females from the Charlevoix Co. population that year. A significant trend toward narrower band widths seems to have occurred during this period, perhaps reflecting more extensive interspecific hybridization than our single documented family might sug- gest alone. P. glaucus individuals typically have 10%—-40%, P. canadensis 55-90%, and hybrids 35-60% (see text). warm years in total seasonal degree day accumulations and that there is clear evidence of extensive northward introgression of “glaucus” traits has been occurring in central Wisconsin, west central Michigan, New York, and Vermont since 1998 (Scriber 2002a, b). However, we have never seen a pure P. glaucus male as far north as Charlevoix County (Table 3) or anywhere from Min- nesota to Massachusetts north of the hybrid zone as delineated by seasonal degree day accumulations of less than 2700 F DD (based on wing characters and al- lozyme electrophoresis; Hagen et al. 1991, Scriber 1996, Stump 2000, Ording 2001). In 2000, intensive sampling of adults from this Thumb Lake site in Charlevoix County (n = 83 male captures) yielded about a dozen males with “glaucus- like” hindwing bands (20%-40%; Fig. 2). However, no adult primary hybrids were seen in those we were able to run for the 3 diagnostic allozymes (Table 3). We hoped to capture individuals of primary hybrids de- rived from offspring oviposited before the 1999 field- capture of this particular female (that produced brood #15116). It is also noteworthy that only one or two clearly “glaucus-type” males or females were detected in those captured in 1998 and 1999 based on band widths alone (Figs. 1, 2). Long distance dispersal of P. glaucus individuals may occur with strong storms, as was reported in 1997 for a dark female that was collected even further north in Dickinson County of the Upper Peninsula of Michi- gan (Scriber et al. 1998). Perhaps this wild male parent VOLUME 57, NUMBER 1 15.0 12.5 Charlevoix Co. MI 1998 10:0 MALES 5 4 Charlevoix Co. MI 1999 3 MALES Charlevoix Co. Ml 2000 MALES Number of Individuals 25 20 Charlevoix Co . MI 2001 15 MALES Mean 60.2 (7.5) n =50 (0) 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 85 90 95 Mean 62.5 (7.7) n= 18 0) 5 10,15 20 25 30 4.35 40 45 50 55 60 165 70 75) 810) 85| 90) 95 Mean 52.5 (9.9) n= 83 QO” 5) 10 i520) 52555310) 35) 4045 50 55° 60) 65 70 75 80) 85 90 95 Mean 55.8 (7.8) n= 83 10) 5. 10 1S 20 259 30 65) JO. 6650) 55 CO 85.70 76 BO 8G) 80) OS Black Band Widths(% ) Fic. 2. Population relative black band widths for individual males captured in Charlevoix County, Michigan during 1998, 1999, 2000, and 2001. A significant shift toward narrower bands was clear after 1999. of brood #15116 was an isolated 1999 “blow-in.” How- ever, we do know that considerable northward gene flow has been happening along the west coast of Michigan and the Islands off the Leelanau peninsula with the warm lake effects and “season extensions” northward beyond the expected latitudinal limits (Scriber & Gage 1995, Ording 2001, Scriber 2002a, b). Natural hybridization (e.g., of our female P. canadensis with a P. glaucus male postulated as an ex- planation for brood #15116) would not have been suprising based upon the results of experimental field mating preference studies with tethered, size-matched virgin females of each tiger swallowtail species. In Charlevoix County during 1997 in 2-choice interspe- cific field mating preference bioassays of free-flying P canadensis males, 82% of the 476 copulations ob- served were with the heterospecific P. glaucus female, rather than with the conspecific (P. canadensis) female (Deering 1998, Deering & Scriber 2002). In contrast, free-flying P. glaucus males in Florida had preferred their conspecific P. glaucus females in 98% of all copu- lations observed for the 1997 and 1998 field seasons (Deering & Scriber 2002). Without P. glaucus females in the area to select from, the mating with our P. canadensis female (#15116, or others) seems more probable for any P. glaucus males recently flown in or blown in (Scriber et al. 1998). We did observe directly developing adults, both 42 N latitude JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY P. glaucus Fic. 3. Michigan map of counties. Historically, the hybrid zone separation between the two tiger swallowtail species, Papilio canadensis and P. glaucus, centered approximately across the 43 degree North latitude band (Scriber 1996a, 2002a, Nielsen 1999). Charlevoix county is in up- per northwest of lower Michigan peninsula. males (n = 2) and females (n = 6) from offspring of this female (brood #15116) that survived to pupation. However, other offspring of this female that may have grown and pupated in the field at Charlevoix County (i.e., from eggs laid before her capture) would cer- tainly not have led to a successful second generation this far north, even if they had found mates. Insuffi- cient growing degree-day accumulations (DDs) ex- isted to permit larval growth and pupation of a second generation before the Fall freezes and leaf abscission (Scriber & Gage 1995, Scriber 1996b, Tesar & Scriber 2002). The 20-year average was less than 2200 F DD’s, and even in these very warmest years, 1998, and 1999 there were still less than 2500 F DD’s (Seriber & Lederhouse 1992, Scriber 2002b). It seems clear that selection against these or any similar non-diapausing hybrid genotypes (P. canadensis fe- males and P. glaucus males) would be severe, whereas those of reciprocal parental crosses (P glaucus fe- males and P. canadensis males) would be able to sur- vive Fall and Winter as dispausing pupae (Rockey et al. 1987a, b, Scriber 2002b). In fact, this brood mortality of direct developing adults in areas with insufficient thermal unit accumu- lations to complete the (second) generation, may ex- plain the very strong selection gradient for univoltin- ism at the hybrid zone. One sex-linked allozyme disjunction (step-cline) for lactate dehydrogenase is evident along a northward cline across the hybrid zone in New England and here in Michigan (LDH-100 is also “diagnostic” for the glaucus species whereas LDH-80 and LDH-40 are diagnostic for “canadensis”; Hagen 1990, Ording 2001, Scriber 2002b). Other sex- linked allozymes and autosomal traits such as tulip tree VOLUME 57, NUMBER 1 detoxification abilities have moved northward exten- sively, but not LDH-100 (Scriber 2002a). The “true” second generation capability also stops short of those latitudinal distances observed for the species-diagnostic PGD-100, HK-100, tulip tree detoxification abilities, and the narrow black hindwing bands (Ording 2001, Scriber 2002a, b). It is important to realize that not all hybrids are evo- lutionary “dead-ends” (Arnold & Hodges 1995, Fu- tuyma & Shapiro 1995, Arnold 1997). The interspe- cific hybridization of P. glaucus and P. canadensis that occurs across the Great Lakes region hybrid zone does not always result in maladapted offspring. In fact, the larval growth rates and pupal sizes of interspecific hy- brids are sometimes greater than either parental species type (Scriber et al. 2003). In no case were per- formances of reciprocal hybrid genotypes less than ei- ther parent when reared on combinations of three hosts (tulip tree, quaking aspen, and black cherry) at three different temperatures (15°C, 23°C, and 31°C; Donovan 2001). We suspect that such genetic intro- gression from interspecific hybridization may con- tribute significantly to the different genetic combina- tions that may be locally suited to islands in the Great Lakes islands such as South Manitou and North Man- itou Islands of The Sleeping Bears Dunes National Park (Ording 2001) or to local climatic “cold pockets” such as the area just east of this Charlevoix County site (Scriber 1996b), or in latitudinal/altitudinal zones where seasonal thermal unit resources for completing a generation are “constrained” (Collins 1984, Scriber 2002b). It will be interesting to follow the genetic changes in these populations and the extent of inter- specific hybridization if the Great Lakes regional cli- mate continues to warm as seen globally (Parmesan & Yohe 2003). Since some hybrids have been unique enough to have incorrectly been assigned species status (Tyler et al. 1994), and since morphological traits alone are of- ten insufficient to confirm hybrid status, we have pro- vided a multi-trait analysis as a mini-review. Spatially and temporally extensive trait analyses may be the only way to assess the extent genetic introgression across hybrid zones and for identifying parental versus hybrid status. We have recently (since 1998) seen extensive introgression of tulip tree detoxification abilities (from P. glaucus) to locations more than 200 miles North of the 1980-1997 hybrid zone center (Scriber 2002a). These most recent observations were based upon more than 3080 larvae of 136 families and for 800-900 field-captured adults (for morphometric introgression of diagnostic adult wing traits) in Michigan alone. Since 1999, in this Charlevoix population, neonate sur- 33 vival on tulip tree has increased from 10% in 2000, to 35% and 33% in 2001 and 2002. This hybrid family (#15116) from Charlevoix was apparently an early forerunner of additionally extensive hybridization and backcross introgression that has been documented in the Great Lakes and New England region since 1998. It also supports the general lack of pre-zygotic repro- ductive isolation observed between these species in the field (Deering & Scriber 2002). ACKNOWLEDGMENTS This research was supported in part by the College of Natural Sciences, NSF grants (DEB 9510044; and DEB 9981608), and the Michigan Agricultural Experiment Station (Project #1644), and the Michigan State University Foundation. We are grateful for help in the field and lab from Holly Hereau, Matt Lehnert, Jenny Muehlhaus, Michelle Oberlin, Gabe Ording, and Howard Romack and Aram Stump. We thank Guy Bush and Jim Smith for use of their laboratories. 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Latitudinal variation in pho- toperiodic induction of pupal diapause in the spicebush swal- lowtail butterfly, Papilio troilus. Holarctic. Lepidoptera 8(1):37-41 WAGNER, W. H., Jr. 1978. A possible natural hybrid of Papilio eu- rymedon and P. rutulus (Papilionidae) from Idaho. J. Lepid. Soc. 32:226—-228. Received for publication 12 November 2001; revised and ac- cepted 22 July 2002. Journal of the Lepidopterists’ Society 57(1), 2003, 36-42 A FIELD INVESTIGATION OF DEPRESSARIA (ELACHISTIDAE) HOST PLANTS AND ECOLOGY IN THE WESTERN UNITED STATES DUANE D. MCKENNA Department of Organismic and Evolutionary Biology, Harvard University, 26 Oxford St., Cambridge, Massachusetts 02138, USA, E-mail: dmckenna@oeb.harvard.edu AND May R. BERENBAUM Department of Entomology, University of Illinois, 320 Morrill Hall, 505 S. Goodwin, Urbana, Illinois 61801, USA ABSTRACT. Depressaria Haworth is a relatively species-rich group of moths with a Holarctic distribution. In western North America there has been a striking radiation of Apiaceae-feeding members of the genus. To understand patterns of Depressaria distribution, host usage, and natural history in the western United States, we surveyed select potential host plants for larvae. Particular emphasis was placed on surveying plants in the genus Lomatium Raf. because most published Depressaria host plant records are from this genus. Surveys took place throughout the western United States from Utah and Wyoming to the Pacific Ocean, and from the Canadian border in Washington State to California and northem Arizona. Approximately 32,000 km of roadway were covered. When larvae were encountered they were collected and reared to adult- hood. Ten species of Depressaria were reared. Two additional potentially undescribed species, represented only by female specimens, were also reared. Our data support previously published accounts of Depressaria biology, host usage, and distribution, consistent with the fact that known host plant genera were targeted in the surveys. Substantial changes in land use have occurred in some parts of the western United States since the work of J. F. G. Clarke, an early student of Depressaria. Several of his published collection localities, including the type locality for D. whit- mani Clarke and most collection localities near the Pacific Coast and in dry grasslands, have been destroyed or seriously degraded by agricul- ture, grazing, roadway improvements, and other forms of development. Additional key words: _host-plants, Lomatium. Depressaria Haworth (Elachistidae) is a relatively two morphologically distinct groups: the douglasella- species-rich group of small moths with a Holarctic dis- group and the pastinacella-group. Fourteen of the 24 tribution (Hodges 1998). There are approximately 100 described species of Depressaria reported from North species of Depressaria, of which 24 (and 2 potentially America belong to the douglasella-group (sensu Han- undescribed species discussed herein) are found only nemann 1953) and 5 species belong to the pastina- in the Nearctic region (Hodges 1974). Three species cella-group. The remaining 5 described species belong (D. artemisiae Nickerl, D. daucella Denis & Schiffer- to two groups: the artemisiae-group (2 species) and miiller, and D. pastinacella Duponchel) have been in- thomaniella-group (3 species) (Hannemann 1953, troduced from the Palearctic and have established Hodges 1974). Larvae of all species in the douglasella- Nearctic ranges. Twenty-five of the 27 described group feed on plants in the genus Lomatium and a few species that occur in North America have been re- other closely related genera of Apiaceae. The pastina- ported from the western United States. Adult Depres- cella-group also feeds only on Apiaceae, but their host saria are similar in appearance to adults of the genera plants belong to several distantly related genera (Plun- Agonopterix Hiibner, Apachea Clarke, Exaeretia kett & Downie 1999). The artemisiae-group and the Stainton, and Nites Hodges, and can be separated thomaniella-group feed only on Asteraceae (Clarke from Apachea by the absence of a strong anteriorly di- 1933, 1941, 1947, 1952, Hodges 1974). rected scale tuft on the second segment of the labial All North American Depressaria are univoltine. Af- palpus, from Nites by the presence of ocelli, and from ter overwintering as adults they emerge from pre- Agonopterix and Exaeretia by the presence of veins reproductive diapause, mate, and oviposit on the Cu, and Cu, separate basally in the forewing (Hodges emerging umbels (Apiaceae-feeders) and meristem- 1974). atic tissue of their host plants. Except for some peren- All Depressaria for which feeding habits are known nial Artemisia L. species, all Depressaria host plants in feed on Apiaceae or Asteraceae (Berenbaum & Passoa North America are herbaceous perennials. Generally, 1999). In western North America there has been a newly hatched first instars build small silk webs in the striking radiation of Apiaceae-feeding Depressaria. developing umbels and leaves of their host plants. Lar- Seventeen of the 24 endemic North American species vae tie together a small amount of umbel or leaf mate- are known to feed on plants in the family Apiaceae rial, forming a tube from which they reach to feed on (Hodges 1974). Hannemann (1953) first noted that the nearby plant parts (Clarke 1952, Hodges 1974). North American Apiaceae-feeding Depressaria form Species-specific variations on this general feeding pat- VOLUME 57, NUMBER 1 tern observed during this study are discussed in the re- sults section. Depressaria pastinacella, an introduced species in North America, has played an important role as a model system for the study of plant-insect coevolution. This is due in part to its host specificity on a few genera of Apiaceae with copious secondary defenses (Thomp- son & Price 1977, Berenbaum 1981, 1983, 1990, Hen- drix 1984, Zangerl & Berenbaum 1993). In contrast, there have been relatively few studies of the native North American Depressaria, or of introduced species other than D. pastinacella (Thompson 1983a, b, 1998, Thompson & Moody 1985). This study further elucidates the distribution, host usage, and natural history of Depressaria in the west- ern United States by surveying potential host plants for Depressaria larvae and rearing them to adulthood. Special emphasis was placed on the douglasella-group and its known host plant genera Lomatium, Pteryxia, and Angelica (all Apiaceae). MATERIALS AND METHODS To elucidate patterns of Depressaria distribution, host usage, and natural history in the western United States, we surveyed potential host plants of Depres- saria in the region for larvae. Potential hosts included all plant genera from which Depressaria species had been reported in the literature as well as additional genera reported to be closely allied to the primary host genus, Lomatium Raf. (Plunkett & Downie 1999). The most frequent and widespread of these al- lied genera included Aletes J. M. Coult. & Rose, An- gelica L., Cymopterus Raf., Pteryxia (Nuttall ex Tor- rey et A. Gray) J. M. Coult. & Rose, and Tauschia Schltdl. Particular emphasis was placed on the genus Lomatium in the surveys because most reared speci- mens of Depressaria from the western United States have been obtained from larvae found feeding on Lo- matium. Survey sites were identified by consulting annota- tions on herbarium specimens from the University of Illinois at Urbana-Champaign, University of Michigan at Ann Arbor, Michigan State University at East Lans- ing, the Rocky Mountain Herbarium at Laramie, and from published type locality and other data for Depres- saria species collected in Arizona, California, Idaho, Montana, Oregon, Washington, and Wyoming (Clarke 1933, 1941, 1947, 1952, Hodges 1974). Additional populations of Depressaria were located by searching for potential larval host plants in suitable habitat along roadsides. Clarke (1933, 1941, 1947, 1952) effectively employed the same general method of collecting De- pressaria in the western United States. Fic. 1. Approximate location of survey sites in the western United States. Filled black circles represent sites where potential host plants were encountered and searched for larval Depressaria. Open circles indicate location of survey sites from which Depressaria were reared. In four trips to the western United States, more than 300 sites were surveyed for Depressaria species over the course of two spring-fall cycles (Fig. 1). “Sites” were discrete locations of variable size where potential host plants were found to grow and were searched for Depressaria. At least 54 species of Apiaceae, including 43 species of Lomatium, were surveyed for larval De- pressaria (Table 1). We identified Lomatium species and other host plants using keys in Cronquist et al. (1997). We photographed plants of uncertain identity and made notes on morphology to facilitate later iden- tification using keys and herbarium specimens (herbar- ium specimens were identified by Professor R. Hart- man, University of Wyoming, Laramie, a specialist on Lomatium and allied genera). All insect specimens were identified by the first au- thor using keys and published descriptions (Clarke 1933, 1941, 1947, 1952, Hodges 1974). Surveys took place primarily in Arizona, California, Idaho, Montana, Nevada, Oregon, Utah, Washington, and Wyoming. More than 32,000 km of roadway were covered. Umbels, leaves, stems, and other above-ground structures of potential host plants encountered were searched for larval Depressaria. If the larval population at a site was greater than approximately 10 individuals, one or more larvae were collected and placed with ad- equate plant material to support their development into a self-sealing plastic bag lined on the bottom with damp, long-fiber sphagnum moss. The larvae in plastic 38 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TABLE 1. Apiaceae surveyed for Depressaria. Abbreviations for states: AZ, Arizona; CA, Califormia; ID, Idaho; MT, Montana; NV, Nevada; OR, Oregon; UT, Utah; WA, Washington; and WY, Wyoming. Apiaceae States Angelica arguta Nutt. CA, ID, MT, OR, NV, UT, WA, WY Angelica lucida L. WA Cicuta douglasii (DC.) Coult. & Rose CA, OR, WA Cicuta maculata L. ID, MT, OR, WA, WY Cymopterus acaulis (Pursh) Raf. ID Cymopterus corrugatus M. E. Jones UT Cymopterus duchesnensis M. E. Jones UT Cymopterus ibapensis M. E. Jones UT Daucus carota L. CA, ID, MT, OR, UT, WA, WY L. ambiguum (Nutt.) J. M. Coult. & Rose ID, OR, WA, WY L. bicolor (S. Wats.) J. M. Coult. & Rose CA, ID, OR, WA L. brandegei (J. M. Coult. & Rose) J. F. Macbr. WA L. californicum (Nutt.) Mathias & Constance CA, OR L. canbyji (J. M. Coult. & Rose) J. M. Coult. & Rose CA L. caruifolium (Hook. & Arn.) J. M. Coult. & Rose CA L. ciliolatum Jepson CA L. circumdatum (S. Wats.) J. M. Coult. & Rose ID L. columbianum Mathias & Constance OR, WA L. cous (S. Wats.) J. M. Coult. & Rose ID, WA L. dasycarpum (Torr. & Gray) J. M. Coult. & Rose CA L. dissectum (Nutt.) Mathias & Constance AZ, CA, ID, MT, NV, OR, UT, WA, WY L. engelmannii Mathias ID L. farinosum (Hook.) J. M. Coult. & Rose OR L. foeniculaceum (Nutt.) J. M. Coult. & Rose AZ, CA, ID, MT, NV, OR, WY L. geyeri (S. Wats.) J. M. Coult. & Rose ID L. gormanii (T. J. Howell) J. M. Coult. & Rose ID, OR L. graveolens (S. Wats.) Dorn & Hartman ID L. grayi (J. M. Coult. & Rose) J. M. Coult. & Rose ID, NV, OR, WA L. hallii (S. Wats.) J. M. Coult. & Rose WA L. hooveri (Mathias & Constance) Constance & Ertter CA L. idahoense Mathias & Constance ID L. laevigatum (Nutt.) J. M. Coult. & Rose OR L. latilobum (Rydb.) Mathias CO, UT L. macrocarpum (Nutt. ex Torr. & Gray) J. M. Coult. & Rose CA, ID, MT, NV L. marginatum (Benth.) J. M. Coult. & Rose CA L. martindalei (J. M. Coult. & Rose) J. M. Coult. & Rose WA L. nevadense (S. Wats.) J. M. Coult. & Rose CA L. nudicaule (Pursh) J. M. Coult. & Rose CA, ID, NV, OR, WA L. orientale J. M. Coult. & Rose ID L. parryi (S. Wats.) J. F. Macbr. UT L. peckianum Mathias & Constance CA L. piperi J. M. Coult. & Rose CA L. rollinsii Mathias & Constance ID L. salmoniflorum (J. M. Coult. & Rose) Mathias & Constance ID L. sandbergii (J. M. Coult. & Rose) J. M. Coult. & Rose ID L. scabrum (J. M. Coult. & Rose) Mathias NV L. serpentinum (M.E. Jones) Mathias ID L. simplex (Nutt.) J. F. Macbr. AZ, ID, UT L. stebbinsii Schlessman & Constance CA L. suksdorfii (S. Wats.) J. M. Coult. & Rose WA L. torreyi (J. M. Coult. & Rose) J. M. Coult. & Rose CA L. tracyi Mathias & Constance CA L. triternatum (Pursh) J. M. Coult. & Rose CA, ID, MT, WA, WY L. utriculatum (Nutt. ex Torr. & Gray) J. M. Coult. & Rose CA, OR L. vaginatum J. M. Coult. & Rose CA, NV L. vaseyi (J. M. Coult. & Rose) J. M. Coult. & Rose CA L. watsonii (J. M. Coult. & Rose) J. M. Coult. & Rose MT Oreoxis alpina (Gray) Coult. & Rose UT Perideridia bolanderi (Gray) A. Nels. & J. F. Macbr. CA, ID, NV, OR, UT, WA Perideridia gairdneri (Hook. & Am.) Mathias AZ, CA, ID, NV, OR, UT, WA, WY Pseudocymopterus montanus (Gray) Coult. & Rose WY Pteryxia petraea (M. E. Jones) Coult. & Rose CA Pteryxia terebinthina (Hook.) Coult. & Rose CA, ID, MT, OR, NV, UT, WA, WY Tauschia glauca (Coult. & Rose) Mathias and Constance CA VOLUME 57, NUMBER 1 39 TABLE 2. Depressaria reared during this study. State records are marked with a “*” preceding the name. County records are marked with a “#° preceding the name. Abbreviations for states as in Table 1. Species Collection site Host plant “)D), angelicivora Custer Co., ID; Hwy 26, 8 km west of Stanley Ange slica arguta 292 #D. betina Modoc Co., CA; Hwy 395, 1 km N of Davis Creek (town) L. triternatum 1d D. daucella Whatcom Co., WA; S of Bellingham Oenanthe sarmentosa 29 #D. juliella Whatcom Co., WA; 13 km S of Hart’s Pass Cicuta maculata yy) *D. leptotaeniae Elko Co., NV; Hwy 227, 16 km SE of Elko L. dissectum 25 1¢ #D. leptotaeniae Custer Co., ID; Hwy 75, 20 km S of Challis L. dissectum 1d #D. leptotaeniae Lemhi Co., ID; Hwy 93, S of Salmon L. dissectum 1d #D. leptotaeniae Jerome Co., ID; Hwy 93, 5 km N of Twin Falls L. dissectum 1d #D. leptotaeniae Lincoln Co., ID; Hwy 93, South of Shoshone L. dissectum 1d D. leptotaeniae Powell Co., MT; Hwy 90, 18 km NW of Deer Lodge L. dissectum ile D. multifidae Idaho Co., ID; Hwy 13, 3 km S of intersection with Hwy 12 L. grayi 82 *D. multifidae Alpine Co., CA; Hwy 88, near intersection with Hwy 89 P. terebinthina var. californica 2° #D. multifidae Wasco Co., OR; Rowena Plateau, near Tom McCall Nature Preserve ___L. grayi 4d #D. multifidae Hamey Co., OR; Hwy 20 at Drinkwater Pass L. grayi 1d #D. multifidae Idaho Co., ID; Hwy 12 at intersection with Hwy 13 L. grayji 16 19 *D. multifidae Alpine Co., CA; Hwy 89, W of intersection with Hwy 395 P. terebinthina var. californica 92 #D. multifidae Adams Co, ID; Kleinschmidt Grade, 800 m from Snake R. L. grayi 12 #D. multifidae Lemhi Co., ID; Hwy 93, S of Salmon P. terebinthina var. foeniculacea 12 #D. multifidae Wallowa Co., OR; Opposite Idaho Powers’ Snake River Campground __L. grayi 19 D. pastinacella Specific locality data not recorded (see “Methods”) Heracleumlanatum, Pastnacasatioa N/A D. meee Washakie Co., WY; Hwy 16, W of Ten Sleep P. terebinthina var. calcarea 19 D. sp. A Park Co., WY; Hwy 20/14/16, 16 km E of Yellowstone National Park P. terebinthina var. foeniculacea 19 *D. sp. A Lemhi Co., ID; Hwy 93, $ of Salmon P. terebinthina var. foeniculacea 19 *D. sp. A Alpine Co., CA; Hwy 4, W of intersection with Hwy 89 P. terebinthina var. californica 29 *D. sp. A Alpine Co., CA; Hwy 4, W of intersection with Hwy 89 P. terebinthina var. californica 22 *D. sp. B Modoc Co., CA; Hwy 395, S of Davis Creek (town) L. bicolor ite D. togata Whatcom Co., WA; Hart’s Pass L. ambiguum 1d D. togata Whatcom Co., WA; Slate Peak L. brandegei 12 bags were reared on plant material from the same RESULTS species and same site from which they were collected. Larvae were not collected if fewer than 10 were found at a site, or if they were found within the boundaries of a national or state park. Depressaria pastinacella was encountered frequently but, because its life-history and ecology are so well documented, it was not col- lected in the surveys. After emerging, the moth and a gelatin capsule con- taining the pupal exuviae were mounted on a pin. The abdomen was removed and frozen at —80°C for future DNA extraction and analysis, and the genitalia were cleared in KOH and mounted on a microscope slide (without staining) to facilitate identification. County and state records were determined by consulting the most recent published distribution data for the Oecophoridae (sensu Hodges 1983) of western North America (Powell & Opler 1996) and are listed in Table 2 Vouchers have been deposited at the University of Illinois at Urbana-Champaign. Vouchers of the two po- tentially undescribed species and county records will be retained at Harvard University in Cambridge, Massachusetts until the completion of ongoing studies, at which time they will be deposited at the National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM). Fifty-four Depressaria larvae were collected and successfully reared (Table 1). These specimens repre- sent eleven of the 22 described species of Depressaria known from western North America, and 2 potentially undescribed species. Additional species may have been encountered, but they were not collected due to small population sizes or their residence in national parks. Most reared specimens belong to the dou- glasella-group. Despite extensive surveys of their re- ported host plants (when known), no members of the artemisiae-group or the thomaniella-group were found. Douglasella-group An unusual Depressaria that is probably an un- described species was reared from Pteryxia tere- binthina (Hook.) Coult. & Rose var. foeniculacea (Nutt.) Mathias. Pteryxia terebinthina var. foenicula- cea in Idaho and Wyoming, and P. terebinthina var. californica (Coult. and Rose) Mathias in California (Fig. 2a-c). A visit to the Wyoming locality adjacent to Yellowstone National Park on 24-VI-2000 found it de- ee by road construction. Hereatter this species will be referred to as Depressaria sp. “A.” Females are closest in appearance to D. multifidae Clarke. We have 40 reared only female specimens, and it is for this reason that we do not formally describe this species here. This putative species can be differentiated from D. lepto- taeniae Clarke, D. multifidae, and D. pteryxiphaga Clarke by the absence of a sclerotized, folded struc- ture on the anterior margin of the female eighth ab- dominal sternum, and by the absence of heavy sclero- tization of the ductus bursae. Maculation also differs somewhat from other members of the douglasella- group. Scales on the vertex of Depressaria sp. A are buff to rust-colored, and lighter-colored on the head, thorax, costal portion of the base of the forewings, and tegulae, than on the abdomen. The remainder of the forewing is covered with grey-brown, brown, and buff- tipped-brown scales. A white spot formed by 1-3 white scales surrounded by grey-brown and brown scales is located at 1/2 the length of the wing in the fold. This spot is absent in some specimens. A second white spot composed of 3-5 white scales surrounded by grey-brown and brown scales is located immedi- ately distal to first near the end of the cell. Larvae of D. togata Walsingham were observed to feed and lightly web in the developing umbels of L. brandegei (Coult. & Rose) Macbr. We have never ob- served D. togata feeding on the leaves of this host plant. Larval D. multifidae were most frequently observed feeding in and webbing the developing umbels of L. grayi Coult. & Rose. Rarely, other host plants were used, such as P. terebinthina (Hook.) Coult. & Rose and L. columbianum Math. & Const. Larvae collected from L. columbianum stopped feeding and died be- fore or shortly after pupation. After feeding in the um- bels of L. grayi and P. terebinthina, D. multifidae lar- vae usually moved to the leaves where they tied together the ultimate divisions of the leaflets to form a small tube from which they fed on adjacent leaflet ma- terial. Most individuals pupated in litter at the bottom of the rearing containers, but some pupated in hollow peduncles of L. grayi placed in the containers. We observed D. pteryxiphaga larvae feeding in the leaflets and umbels of P. terebinthina. The larvae ap- peared to move to the leaves from umbels as later in- stars. Like D. multifidae, the larvae form tubes of web- bing from which they feed, and into which they retreat when disturbed. These tubes sometimes form exten- sive networks of light webbing involving several leaves and occasionally an umbel. We observed abundant early instar D. leptotaeniae feeding gregariously in the developing umbels of L. dissectum (Nutt.) Math. & Const. As larvae mature, they disperse to feed on the leaves where they form small tubes of webbing that incorporate the ultimate segments of the leaflets. They do not form extensive JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY tubes of webbing as D. multifidae and D. pteryxiphaga do. Occasionally, larvae continue to feed in the um- bels, tying together several rays and feeding on the de- veloping flowers or young, green fruits. Typically, nearly mature larvae move to the leaf axils where they feed lightly, deposit some frass, and pupate. The pupa is always oriented with the ventral surface downward and is covered with a small amount of silk that serves to anchor it in place. Reared larvae rarely pupated in broken L. dissectum peduncles or in litter at the base of the plant in the rearing container. We made many observations of pupal exuviae in leaf axils in the field. Near Twin Falls, Idaho, D. leptotaeniae is exception- ally abundant on L. dissectum in basaltic lava flows. Here, we observed thousands of D. leptotaeniae pupae and pupal exuviae in leaf axils in late May 1998. Larvae of D. angelicivora Clarke begin life feeding in the developing umbels and leaves, of Angelica arguta Nutt. Their feeding often results in distortion of the emerging umbels and leaves making them easy to spot. Most larvae complete feeding by the time the umbels are fully expanded. Occasionally, larvae can be found feeding on the developing (still green) seeds or leaves, but apparently only when flowers are not avail- able. In such situations we have observed the larvae to tie several developing fruits together with webbing to form a small tube from which they reach to feed in a manner similar to D. pteryxiphaga. Nearly mature lar- vae were observed to wander from the developing meristem of the host plant to debris in the bottom of the rearing container where they pupated. Betina-group Larval D. betina Clarke have been observed feeding in the umbels of L. triternatum (Pursh.) Coult. & Rose. At the one site where they were observed, late instars were inconspicuous, usually involving only three or fewer rays of the host umbel in their webbing. Tubes of webbing were not formed. The only adult we reared pupated in litter at the bottom of the rearing container. Pastinacella-group A single adult specimen of a strikingly-colored, ap- parently undescribed Depressaria was reared from a larva collected on L. bicolor var. leptocarpum Lo- matium bicolor (S. Watson) J. Coulter & Rose var. lep- tocarpum (Torrey & A. Gray) M. Schlessman south of the town of Davis Creek, Modoc County, California. The larva was feeding in a webbed mass of leaf and umbel material and was the only larva of its kind ob- served. Because we have only a single specimen we do not formally describe it here. This female specimen is VOLUME 57, NUMBER 1 dramatically distinct from other members of the pasti- nacella-group. For convenience, we will refer to this specimen hereafter as Depressaria sp. “B”. Depres- saria sp. B most resembles D. juliella Busck, D. dau- cella Denis and Schiffermiiller, and D. eleanorae Clarke. It can be differentiated from them by the col- oration of forewing and tarsomere maculation, the ab- sence of a folded structure on the anterior margin of the eighth abdominal sternum, and the wide ostium bursae. Dorsum of head with rust to salmon scales. A tuft of red-rust scales protrude from posterior edge of each eye. Tegulae are salmon-colored, and nearly the same color as the thorax. The forewing is covered with off-white, rust, and salmon scales. Longitudinal streaks of dark rust scales are scattered across the surface of the forewing, mostly parallel to the wing margin. All scales appear pearlescent in reflected light. The forewing fringe is a uniform rusty salmon. The legs have off-white scales at the distal end of each tar- somere, and appear banded. The eighth abdominal sternum is weakly pigmented throughout. The ostium bursae is wider than in other Depressaria species. Larvae of Depressaria juliella Busck were found feeding on the flowers of Cicuta maculata L. tying the rays of the inflorescence together, but not noticeably distorting the umbel. The two specimens reared pu- pated in debris at the bottom of the rearing container. We collected larvae of Depressaria daucella while they were feeding on an umbel on Oenanthe sarmen- tosa J. S. Presl. We have not observed leaf-feeding. Larvae do not noticeably distort the umbels and pro- duce relatively little webbing. Depressaria pastinacella was abundant throughout the western United States wherever either of its com- mon host plants, Heraclewm lanatum and Pastinaca sativa L., (local and introduced, respectively) were found. Larvae were not found on any other plant species, (but no extensive effort was made to look for them on other known host genera such as Angelica). Depressaria pastinacella was by far the most fre- quently encountered Depressaria species. Other Observations Larvae whose behavior and ecology resembled those of known Depressaria and Agonopterix were ob- served, but not collected because of small population sizes or occurrence in national parks. Surprisingly, no parasitoids were reared from any Depressaria species; however, during the course of this study several Ichneumonidae were reared from Sparganothis Hiibner species (Tortricidae) that co-oc- curred with D. multifidae on P. terebinthina. In fact, Sparganothis co-occurred with Depressaria spp. on L. 4] dissectum and Pteryxia terebinthina at several study sites. Papilio indra (Papilionidae), and Agonopterix spp. (Elachistidae: Depressariinae) were less fre- quently encountered than Depressaria or Sparga- nothis, but larvae occurred on nearly an identical suite of host plants as the douglasella-group of Depressaria. Epermenia Hiibner spp. (xarmomdae) were occa- sionally encountered as larvae feeding in the seeds of L. dissectum and L. triternatum. Adults were occa- sionally seen on the flowers of Achillea millefolium L. (Asteraceae). Relatively few other lepidopteran larvae were observed. Crab spiders (Thomisidae) were often abundant on flowering Lomatium but were not ob- served to take Depressaria larvae or adults as prey. On several occasions predacious Hemiptera were ob- served feeding on Depressaria larvae that were still in their webbed tubes. Substantial changes in land use have occurred in some parts of the western United States since the works of Clarke (1933, 1941, 1947, 1952). Several sites, including the type locality for D. whitmani and most former collection locals near the Pacific Coast and in former dry grasslands, have been destroyed or seriously degraded by agriculture, grazing, and other forms of development. DISCUSSION AND CONCLUSIONS The host plant usage patterns documented for De- pressaria in the literature accurately reflect contempo- rary patterns of host usage by North American De- pressaria to the extent that they were encountered in this study. The relatively narrow search image we formed for potential Depressaria host plants Grhich included only select Asteraceae and Apiaceae) con- tributed to this view of patterns of host plant utiliza- tion. Our survey data are also complementary to the ecological work of Thompson and Moody (1985) and Thompson (1983a, b). Future surveys should include other potential host plant genera and families. Our ob- servations and our surv ey results suggest that most De- pressaria species are more widespread than existing published records indicate (Hodges 1974). Potential host plant species are widely distributed, and relatively abundant throughout much of the western United States, and considerable additional rearing is necessary from throughout the region to document the species richness of Depressaria and the systematic relation- ship of their hosts. ACKNOWLEDGMENTS We would like to thank Katherine McKenna for her assistance in the field. Arthur Zangerl, Terry Harrison, and Stephen Downie pro- vided helpful comments and guidance. Conversations with Walter Fertig, Ron Hartman, and John Thompson proved immensely help- 42 ful in locating extant potential host-plant populations to be surveyed, and provided other useful suggestions. David Adamski provided helpful editorial assistance with early drafts of this paper. National Science Foundation grant DEB 9903867, the University of Illinois Program in Ecology and Evolutionary Biology, and the Herbert H. Ross Memorial Fund partially funded this work. LITERATURE CITED BERENBAUM, M. R. 1981. 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Collec. 117:1—20. CRONOQUIST, A., N. H. HOLMGREN & P. K. HOLMGREN. 1997. Inter- mountain flora. Vol. 3, Part A. Rosidae other than Fabales. NYBG Press, New York. 446 pp. HANNEMANN, J. H. 1953. Naturliche Gruppierung der europais- chen Arten der Gattung Depressaria s. |. (Lepidoptera: Oe- cophoridae). Mitt. Zool. Mus. Berl., 29:270-372. HENDRIX, S. D. 1984. Reactions of Heracleum lanatum to floral herbivory by Depressaria pastinacella. Ecology 65:191—197. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Hopces, R. W. 1974. Gelechioidea Oecophoridae. In Dominick, R. B., D. C. Ferguson, J. G. Franclemont, R. W. Hodges & E. G..Munroe [eds.], The moths of America north of Mexico. Fas- cicle 6:2. E. W. Classey, London. fase. 6.2:i-x + 142 pp. Hopces, R. W. [gpD.]. 1983. Check list of the Lepidoptera of America north of Mexico; xxiv + 1-284. E. W. Classey Limited and Wedge Entomological Foundation, University Press, Cam- bridge, Great Britain. PLUNKETT, R. & S. R. Downie. 1999. Major lineages within Api- aceae subfamily Apioideae: a comparison of chloroplast restric- tion site and DNA sequence data. Am. J. Bot. 86(7):1014—1026. PowELL, J. A. & P. A. OpLER. 1996. Moths of western North America. 4. Distribution of “Oecophoridae” (sense of Hodges 1983) of western North America. Contributions of the C.P. Gillette Museum of Arthropod Diversity. Colorado State Uni- versity, Fort Collins, Colorado. 63 pp. THompson, J. N. 1983a. Selection of plant parts by Depressaria multifidae (Lepidoptera: Oecophoridae) on its seasonally- restricted hostplant, Lomatiwm grayi (Umbelliferae). Ecol. En- tomol. 8:203-211. . 1983b. The use of ephemeral plant parts on small host plants: How Depressaria leptotaeniae (Lepidoptera: Oe- cophoridae) feeds on Lomatium dissectum (Umbelliferae). J. Anim. Ecol. 52:281-291. . 1998. Coping with multiple enemies: 10 years of attack on Lomatium dissectum plants. Ecology 79:7 255-2554. TuHompson, J. N. & M. E. Moopy. 1985. Assessing probability of interaction in size-structured populations: Depressaria attack on Lomatium. Ecology 66:1597—1607. THompson, J. N. & P. W. Price. 1977. Plant plasticity, phenology, and herbivore dispersion: wild parsnip and the parsnip web- worm. Ecology 58:1112-1119. ZANGERL, A. R. & M. R. BEREMBAUM. 1993. Plant chemistry and insect adaptations to plant chemistry as determinants of host plant utilization patterns. Ecology 74:478-504. Received for publication 19 November 2001; revised and ac- cepted 3 September 2002. Journal of the Lepidopterists’ Society 57(1), 2003, 43-46 NOTES ON THE LIFE HISTORY OF EANTIS THRASO (HESPERIIDAE: PYRGINAE) IN ECUADOR HAROLD F. GREENEY Yanayacu Biological Station and Center for Creative Studies, Cosanga, Ecuador c/o Carrion No. 21-01 y Juan Leon Mera, Quito, Ecuador AND ANDREW D. WARREN! Department of Entomology, Oregon State University, Corvallis, Oregon 97331-2907, USA ABSTRACT. The early stages and larval behavior of Eantis thraso (Hiibner) in eastern Ecuador are described. All larval instars were found to build leaf shelters on the host plant, which are described and illustrated. A summary of larval host plants for species in the genus Eantis Bois- duval is provided. Cultivated Citrus L. species are reported as the local larval host plants for E. thraso in eastern Ecuador. Additional key words: Warren (1996) resurrected the generic name Eantis Boisduval from synonymy with Achlyodes Hiibner for taxa including species and subspecies of Evans’ (1953, 1955) Achlyodes mithridates Fab. group, after a phylo- genetic analysis of morphological characters observed in Achlyodes. The genus Eantis is entirely Neotropical in its distribution and includes 9 superficially similar species. Material studied by Warren (1996) indicated a geographical range for E. thraso from extreme south- em Chiapas, Mexico, east to southern Belize, and south throughout tropical Central and South America; recent color illustrations of adult males appear in Lewis (1987:80), Brown (1992:176) and Warren (1996). Eantis thraso is replaced in most of Mexico and Texas by the similar E. tamenund W. H. Edwards. Food plant records for Eantis are mostly from the family Rutaceae, including cultivated Citrus L. plants (Panton 1898, Smyth 1919). Native hosts in most areas also appear to be Rutaceae, especially Zanthoxylum L. (Bruner et al. 1945, Poey 1832, Janzen & Hallwachs 2003). Wolcott (1923, 1951) described the larva and pupa of Eantis minor (Comstock) from Puerto Rico on cultivated Citrus and native Zanthoxylum species. Eantis tamenund has been reared in Texas on native Zanthoxylum species (Kendall 1965). Hayward (1941, 1948) reported E. thraso from various Zanthoxylum species and Moss (1949) found it on one Zanthoxylum species. Biezanko et al. (1974) found E. thraso on four species of Citrus and two species of Zanthoxylum. Most recently, Janzen and Hallwachs (2003) found E. thraso on six species of Zanthoxylum in northwest- ern Costa Rica. Despite the fact that larvae of E. thraso have been observed and reared several times, the most detailed description of its larva avail- able is that provided by Moss (1949), who described the " Research Associate, Museo de Zoologia, Facultad de Ciencias, Universidad Nacional Auténoma de México, Apdo. Postal 70-399, México, D.F. 04510 Mexico. larvae, larval food plants, larval shelters, life cycles, pupae. mature larvae of E. thraso as being “plain green with a rotund brown head.” Here, we provide a more detailed description of the early stages of E. thraso in Ecuador, as well as notes on its larval shelter-building behavior. MATERIALS AND METHODS The majority of observations were made between September and November of 1997 in north-eastern Ecuador at the La Selva Lodge Research Station, 75 kilometers E.S.E. of Coca, Garzacocha, Sucumbios Province, at 250 meters elevation. For a detailed de- scription of this site, see DeVries et al. (1999). Subse- quently, several larvae were found and reared under similar conditions at the Sacha Lodge Research Sta- tion located 10 kilometers up river from La Selva. All larvae and pupae were collected from cultivated Cit- rus trees located along the edge of seasonally inun- dated forest. Over 30 individual larvae of various in- stars were encountered on the host plants and transferred to the lab for rearing. Head capsule width measurements were taken from shed head capsules using an ocular micrometer. Voucher material includ- ing preserved larvae of all instars together with their associated shelters is deposited in the collection of the senior author. RESULTS Early stages. No eggs were encountered. First in- star (n = 7). Head capsule black to dark brown, mod- erately heart-shaped; body parallel sided, slightly flat- tened, roughly dome shaped in cross section, entirely pale olive-green to clear green with some variation due to gut contents, dorsum bare but with a ventro-lateral fringe of minute, pale setae and four long, stiff, pale setae along the margin of the anal plate. Second instar (n = 6) as described for first instar. Third instar (n = 13) head capsule caramel-brown, fading gradually to 44 Dee Fic. 1. Shelters and head capsule of Eantis thraso from eastern Ecuador: A, Larval shelter type built by first through third (or rarely through fifth) instar larvae; B, Larval shelter type built by many fourth and fifth instar larvae; C, Head capsule of fifth instar larva. dark brown basally around ommatidia and mouthparts, heart shape more pronounced, average width 1.41 mm (n = 2); body as described for first instar but some indi- viduals with a faint supraspiracular line of yellow hatch marks from T2 to A8. Fourth instar (n = 14) average head capsule width 2.62 mm (n = 3), body as described for third instar, but yellow hatch marks more pro- nounced, and anal plate still with four stiff, pale setae, which are approximately the same size as in the third instar, so now appear proportionately smaller. Fifth in- star (n = 9) head capsule (Fig. 1C) now with distinct black area basally giving a bearded appearance, aver- age width 4.37 mm (n = 4); body as described for fourth instar (also see photos by Janzen & Hallwachs 2003). Immediately upon molting, the fresh head cap- sules of last instar larvae are pale lime green and slowly change to an ivory color and finally to caramel over the course of an hour. Pupa (n = §) stout with a short blunt horn arising between eyes and projecting foreword and slightly upward, ground color entirely lime-green; four tiny black dots behind the head form a small crescent on the dorsum of the prothorax, four additional small brown spots located on dorsal abdomen just anterior to cremaster. Cremaster dark brown, entire pupa covered with a light dusting of white waxy flocculence except for two small bare patches in the shape of crescents along the costal edge of wing pads. Approximately 24 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY hours before eclosion, the pupa turns generally more yellowish, with the wing pads gaining an orange-brown cast and the eyes becoming dark brown. Larval shelters. Larvae of all instars formed a shel- ter made by modifying the leaves of their host plant. Terminology for discussion of shelters follows Greeney and Jones (in press). First instar shelters (n = 7) were roughly triangular or trapezoidal shaped sections of leaves cut from the leaf margin and flipped onto the dorsal surface of the leaf (Fig. 1A). Two major cuts in the leaf were made that ended towards the central por- tion of the leaf and angled to a narrow shelter bridge. This is termed a two-cut, stemmed fold, Group III, type 10 shelter (Greeney & Jones in press). Second in- star larvae (n = 6) were found in shelters as described for the first instar. No abandoned first-instar shelters were found at the study sites, suggesting that larvae re- mained in the shelter built during the first instar. Third instar shelters (n = 13) were similar to those described for the first instar, but were larger. Three of 14 fourth instar larvae had built a third shelter formed by silking together two leaves so that the dorsal surface of one leaf contacted the ventral surface of another leaf and formed a pocket (Fig. 1B). This is termed a two-leaf pocket, Group I, type 2 shelter (Greeney & Jones in press). The remaining eleven fourth instar larvae were encountered in the second larval shelter as described for the third instar. Fifth instar (n = 9) larvae remained in shelters built during the fourth (or third) instar. Two pupae were found in the field, and both were inside the last feeding shelter built. The pupae were attached, face down, to the ventral surface of the shelter by heavy silking at the cremaster, and were supported by a band of silk across the mid-thorax. DISCUSSION Despite the relative abundance of larvae at the study site, adult E. thraso were not common. Those encoun- tered were exclusively associated with disturbed areas. Other species of Rutaceae, such as Zanthoxylum amer- icanum Mill and Ptelea trifoliata L., have been sug- gested as larval host plants for Eantis from other re- gions (Kendall 1965, Kendall & McGuire 1975). It is unknown if these genera are utilized in our area, but the occurrence of adults associated with disturbed ar- eas, where Citus is often abundant, suggests that Citws is an important larval host in eastern Ecuador. On the other hand, Janzen and Hallwachs (2003) reported on 201 rearings of Costa Rican E. thraso, all from Zan- thoxylum species. The construction of larval shelters by hesperiid lar- vae has been known for many years (e.g., Scudder 1889) and has since been reported for a wide variety of species (e.g., Moss 1949, Miller 1990, Atkins et al. VOLUME 57, NUMBER 1 1991). Within the Lepidoptera, species of many taxa- nomic groups build shelters of some type (e.g., Miller 1983, DeVries 1987, Jones 1999), and there are at least 10 distinct types built by the Hesperiidae alone (Greeney & Jones in press). All of those recorded for the Hesperiidae involve modifying the host plant with cuts and/or silk. Ontogenetic changes in the form of these shelters are also well documented (Graham 1988, Miller 1990). Eantis thraso larvae form shelters of two basic types. The first, involving cutting of the leaf (Fig. 1A), is a common type seen in many other species (Young 1991, 1993, HFG pers. obs.). The sec- ond, involving the silking together of two separate leaves (Fig. 1B), is likely utilized in this case due to size constraints imposed by the growing larvae and the relatively small leaves of the Citrus host. While no oviposition events were observed and no eggs were found in the field, it is likely that E. thraso oviposit on the meristem leaves of their host. All early instars were found on pale, fresh leaves, and later instars were typically on or near new growth. This preference for meristem tissue is known for other rutaceous feeders (Vaidya 1969, Young 1993, HFG pers. obs.), as well as many species of butterflies feeding on a variety of host plants (DeVries 1987). The value of detailed natural history studies in cre- ating and testing phylogenetic hypotheses has been noted by other authors (Hennig 1966, DeVries 1987). In general, the morphological and ecological attributes of hesperiid larvae are very poorly known, and the tax- onomy of most tropical skipper groups remains con- fused; much additional work is needed in these areas. In the face of ever increasing rates of habitat destruction, we hope this study may encourage others to continue publishing observations on the life history of this and other poorly known (but frequently encountered) taxa. ACKNOWLEDGMENTS This research was funded in part by a grant from the Azar Foun- dation. HFG thanks the owner and staff of Sacha Lodge for their generosity and support. Phil DeVries, George Austin, and Meg Jones provided useful comments on the manuscript and have gener- ously provided information on the biology of various skippers. Fi- nally, Ryan Hill and Nicole Gerardo provided a great deal of assis- tance in the field. Thanks also to Roy Kendall, Richard Peigler, and Andrew Atkins for providing some literature cited herein. Addition- ally, we acknowledge the PBNHS for their ongoing support of our natural history studies. LITERATURE CITED Atkins, A., R. Mayo & M. Moore. 1991. The life history of Signeta tymbophora (Meyrick and Lower) (Lepidoptera: Hesperiidae: Trapezitinae). Aust. Ent. Mag. 18:87—90. BIEZANKO, C. M., A. RUFFINELLI & D. Linp. 1974. Plantas y otras sustancias alimenticias de las orugas de los lepid6pteros Uruguayos. Revista do Centro de Ciéncias Rurais 4:107-148. Brown, K.S. 1992. Borboletas da Serra do Japi: diversidade, habi- tats, recursos alimentares e variacdo temporal, pp. 142-187. In Morellato, L. P. C. (org.), Histéria Natural da Serra do Japi, Ecologia e Preservagao de uma Area Florestal no Sudeste do Brasil. Universidade Estadual de Campinas, Brazil. 321 pp. BRUNER, S. C., L. C. SCARAMUZZA & A. R. OTERO. 1945. Catalogo de los insectos que atacan a las plantas econdmicas de Cuba. Ministerio de Agricultura, La Habana, Cuba. DeVries, P. J. 1987. The butterflies of Costa Rica and their natural history. Vol. 1. Papilionidae, Pieridae, and Nymphalidae. Princeton University Press, Princeton, New Jersey. 327 pp. DeVries, P. J., T. R. WALLA & H. F. GREENEY. 1999. Species diver- sity in spatial and temporal dimensions of fruit-feeding butterflies from two Ecuadorian rainforests. Biol. J. Linn. Soe, 68:333-353. Evans, W. H. 1953. A catalogue of the American Hesperiidae indi- cating the classification and nomenclature adopted in the British Museum (Natural History). Part II] (groups E, F, G), Pyrginae, section 2. British Museum, London. 246 pp. 1955. A Catalogue of the American Hesperiidae indicating fhe classification and nomenclature adopted in the British Mu- seum (Natural History). Part IV (groups H to P), Hesperiinae and Megathyminae. British Museum, London. 499 pp: GranaM, A. J. 1988. The life history of a semi-arid population of Croitana croites (Hewitson) (Lepidoptera: Hesperiidae: Trapezitinae). Aust. Ent. Mag. 15:123-126. GREENEY, H. F. & M. T. JonEs. (in press). Shelter building in the Hesperiidae: a classification scheme for larval shelters. J. Res. Lep. Haywarp, K. J. 1941. Plantas alimenticias de hespéridos argenti- nos. Revta Soc. Ent. Argent. 11:31—36. . 1948. Insecta, Lepidoptera (Rhopalocera), familia Hes- periidarum, subfamilia Pyrrhopyginarum et Pyrginarum. In De- scole, H. R. (ed.), Genera et Species Animalium Argentinorum. Guillermo Kraft, Buenos Aires. Vol. 1. 389 pp. HENNIG, W. 1966. Phylogenetic systematics. University of Illinois Press, Urbana. JANZEN, D. H. & W. Hatiwacus. 2003. Philosophy, navigation and use of a dynamic database (“AGC caterpillars SRNP”) for an in- ventory of the macrocaterpillar fauna and its foodplants and parasitoids, of the Area de Conservaci6n Guanacaste (AGC), northwestern Costa Rica. http://janzen.sas.upenn.edu Jones, M. T. 1999. Functions of shelter building in the Hesperi- idae. Unpublished Masters Thesis, Georgetown University, Washington, DC. KENDALL, R. O. 1965. Larval food plants and distribution notes for twenty-four Texas Hesperiidae. J. Lep. Soc. 19:1-33. KENDALL, R. O. & W. W. Mccuire. 1975. Larval foodplants for twenty-one species of skippers (Lepidoptera: Hesperiidae) from Mexico. Bull. Allyn Mus. 27:1-7. Lewis, H. L. 1987. Butterflies of the World. Harrison House, New York. 312 pp. MILLER, C. G. 1990. The life history of Chaetocneme denitza (He- witson) (Lepidoptera: Hesperiidae: Pyrginae). Aust. Ent. Mag. 17:97-100. MILLER, W. E. 1983. Eucosmomorpha albersana (Hiibner), a Palaearctic species, collected in North America (Tortricidae, Grapholitini). J. Lep. Soc. 37:88-89. Moss, A. M. 1949. Biological notes on some Hesperiidae of Para and the Amazon (Lep. Rhop.). Acta Zool. Lilloana 7:27-29. PANTON, E. S. 1898. The life history of some Jamaica Hesperiidae. J. Inst. Jamaica 2:435-441. Poey, F. 1832. Centurie de lépidoptéres de lle de Cuba, con- tenant la description et les figures coloriées de cent espéces de papillons nouveaux ou peu connus, répresentés d’aprés nature, souvent avec la chenille, la chrysalide, et plusieurs détalis mi- croscopiques. J. Albert Mercklein, Paris. SCUDDER, S. H. 1889. The Butterflies of the Eastern United States and Canada, with special reference to New England. Vol. 2, Ly- caenidae, Papilionidae, Hesperiidae. S. H. Scudder, Cam- bridge, Massachusetts. xii + 767-1774 pp. SmyTH, H. E. G. 1919. Dominio de insectos en los citricos en Puerto Rico. Revta. Agric. Puerto Rico 3:39—50, 55-62. Vaipya, V. G. 1969. Inv vestigations on the role of visual stimuli in the egg-laying and resting behavior of Papilio demoleus L. (Pa- pilionidae, Lepidoptera). Anim. Behav. 17:350—355. 46 WarrEN, A. D. 1996. Phylogenetic revision of the skippers of the mithridates species group, and their replacement in Eantis Bois- duval, 1836 (Lepidoptera: Hesperiidae: Pyrginae). Entomology Department Honors Thesis, Comell University, Ithaca, New York. Wo tcotTt, G. N. 1923. “Insecate Portoricensis.” A preliminary an- notated check-list of the insects of Puerto Rico, with descrip- tions of some new species. J. Dept Agric. Puerto Rico 7:1-313. . 1951. The insects of Puerto Rico. J. Dept Agric. Puerto Rico 32:417—748. JOURNAL OF THE LLEPIDOPTERISTS’ SOCIETY YouNG, A. M. 1991. Notes on the natural history of Quadrus (Pythonides) contuberalis (Hesperiidae) in Costa Rica. J. Lep. Soc. 45:366-371. . 1993. Notes on the natural history of Achlyodes selva (Hesperiidae) in Costa Rica. J. Lep. Soc. 47:323-327. Received for publication 16 January 2002; revised and accepted 22 August 2002. Journal of the Lepidopterists’ Society 57(1), 2003, 47-53 PROMYLEA LUNIGERELLA GLENDELLA DYAR (PYRALIDAE) FEEDS ON BOTH CONIFERS AND PARASITIC DWARF MISTLETOE (ARCEUTHOBIUM SPP.): ONE EXAMPLE OF FOOD PLANT SHIFTING BETWEEN PARASITIC PLANTS AND THEIR HOSTS KAILEN A. MOONEY Department of EPO Biology, University of Colorado [Boulder], Boulder, Colorado 80309-0334, USA, Email: kailen.mooney@colorado.edu ABSTRACT. Larvae of Promylea lunigerella glendella Dyar (Pyralidae, Phycitinae) feed on Arceuthobium vaginatum susp. cryptopodum (Hawks.) (Viscaceae), the Southwestem dwarf mistletoe, a parasite of Pinus ponderosa (Laws.) scopulorum (Pinaceae) at the Manitou Experi- mental Forest, U.S.D.A. Rocky Mountain Research Station, Woodland Park, Colorado. A previous food plant record for P. lunigerella describes the larvae as feeding on a variety of conifers. A careful evaluation of this record suggests it is reliable, and I conclude that P. lunigerella is actively shifting between dwarf mistletoe and conifer feeding, or has done so recently. My review of the literature on food plant use by lepidopteran her- bivores of dwarf mistletoe and their relatives suggests that food plant shifts between parasitic plants and their hosts, and vice versa, have oc- curred multiple times and may be common among taxa that feed on parasitic and parasitized plants. These findings support a model of food plant shifting in which the close proximity necessarily maintained by parasitic plants and their hosts provides an ecological opportunity that fa- cilitates food plant shifts between these taxonomically and chemically very dissimilar plants. Finally, I describe the life history of P. lunigerella larvae and compare them to those of Dasypyga alternosquamella Ragonot (Pyralidae), a closely related phycitine that also feeds on dwarf mistle- toe at this same location. Additional key words: Mitowra (Lycaenidae), Filatima natalis (Gelechiidae), Chionodes (Gelechiidae), Euthalia (Nymphalidae). Insect herbivores, including lepidopterans, often specialize on individual species or groups of closely- related food plants (Ehrlich & Raven 1964, Holloway & Hebert 1979, Vane-Wright & Ackery 1988). The evolutionary and ecological mechanisms by which such specialist herbivores might switch to novel food plants have received considerable attention (Holloway & Hebert 1979, Denno & McClure 1983, Futuyma & Slatkin 1983, Strong et al. 1984, Vane-Wright & Ack- ery 1988). From these studies comes the prediction that food plant switches are most likely to occur be- tween pants that are similar in phenotypic characters of importance to herbivores such as tissue chemistry. Closely related plants are likely to share such charac- ters due to common ancestry, but taxonomically distant plants may share such characters due to convergence (Judd 1999). There has been relatively little said about how specialist herbivores might shift between taxo- nomically and phenotypically distinct plants, except to predict that such events are not likely to be common. Promylea lunigerella glendella Dyar (Pyralidae, Phycitinae) was first described by Ragonot (1887) and the subspecies by Dyar (1906). The species range stretches from coastal British Columbia to California and east to Colorado. Larvae in British Columbia have been reported to feed as solitary defoliators on conifers including grand fir (Abies grandis (Doug. ex. D. Don) Lindl. (Pinaceae)), Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco (Pinaceae)) and western hemlock (Tsuga heterophylla (Raf.) Sarg. (Pinaceae )) (Prentice 1965). I report here on a population of P. lu- nigerella in Colorado that feeds on Arceuthobium vaginatum (Willd) Presl susp. cryptopodum (Engelm.) Hawksw. & Wiens (Viscaceae), the Southwestern dwarf mistletoe parasitizing ponderosa pine (Pinus ponderosa Laws. scopulorum (Pinaceae)). This novel food plant record suggests a recent or ongoing food plant shift in this species, despite the fact that dwarf mistletoes (Arceuthobium spp.) and conifers differ substantially in chemistry (Buckingham 1994) and are taxonomically unrelated. Dwarf mistletoes (Arceuthobium spp.) are common parasites of conifers in North America, and they are fed upon by a number of specialist herbivores, includ- ing several species of Lepidoptera. Dwarf mistletoes are obligate parasites, and for this reason these plants occur in closer physical association than plants without host-parasite relationships. Consistent and close phys- ical association of taxonomically and chemically dis- tinct plants may lead to rates of herbivore food plant shifting higher than that found between plant taxa lacking this physical proximity (Holloway & Hebert 1979, Chew & Robbins 1988) due to what Strong et al. (1984) call increased “ecological opportunity.” An op- portunity-based model of food plant shifting predicts that herbivores shifting between parasitic plants and the hosts of those plants should be common. To test this hypothesis I reviewed the food plant lit- erature for dwarf mistletoe herbivores and their close relatives to identify what evidence there is to support the hypothesis that food plant shifts between parasitic plants and their hosts, and vice versa, are common. As part of this review, I also carefully inspected the previ- ous report of P. lunigerella feeding on conifers (Pren- tice 1965) to assess its reliability. Finally, I provide nat- ural and life history data on the larval stages of P. lunigerella and compare these larvae to Dasypyga al- ternosquamella Ragonot (Pyralidae), another phycitine herbivore of Southwestern dwarf mistletoe that occurs sympatrically with P. lunigerella. MATERIALS AND METHODS Life history of P. lunigerella. 1 conducted the field and laboratory work for this project at the Mani- tou Experimental Forest, an administrative unit of the U.S. Department of Agriculture Forest Service Rocky Mountain Experiment Station located in Woodland Park, Colorado (39°06’00”N, 105°05’00”W). Manitou includes several stands of ponderosa pines (Pinus pon- derosa var. scopulorum Laws.(Pinaceae)) parasitized by Southwestern dwarf mistletoe (A. vaginatum subsp. cryptopodum Hawks. (Viscaceae)). This field site and the natural history of dwarf mistletoe are described more fully in Mooney (2001). In a previous report (Mooney 2001), I described the natural- and life-history of D. alternosquamella Ragonot (Pyralidae, Phycitinae), a common herbivore of dwarf mistletoes throughout western North America (Heinrich 1920, Reich 1992). It was while conducting this work that I became aware that P. lu- nigerella was also feeding on dwarf mistletoe. Because D. alternosquamella and P. lunigerella are both phyci- tine pyralids, it was only after rearing larvae through pupation that I became aware that some of the animals with which I was working were not, in fact, D. alter- nosquamella. Consequently, the life history data re- ported here are not as complete as they would be had I expressly set out to study P. lunigerella. I collected Southwestern Dwarf Mistletoe from the field between 30 June and 1 August 1999 in individual plastic bags and brought them into the lab on eight separate occasions. Individual plants ranged from 3-10 cm in height and in most cases only one or two plants were taken from any single host-pine. I observed larval feeding in the field, and in most cases the presence of larvae within these plants was indicated by their frass within and surrounding dwarf mistletoe shoots. Larvae were isolated from these plants using a dissecting mi- croscope. In no instance was pine foliage or branch tis- sue collected, and all larvae were on dwarf mistletoe plants at the time of collection. I reared P. lunigerella individually in clear plastic petri dishes lined with filter paper in a laboratory facil- ity. The larvae were fed small (2-5 cm) shoots of dwarf mistletoe collected from the same general location as the larvae themselves, and they were replenished with fresh plant material approximately every third day. In all cases the larvae readily fed upon the dwarf mistletoe. I wetted the filter paper linings of each petri dish on a daily basis. The lab building was neither heated nor cooled, and I stored the petri dishes in the open and near a window where they received indirect sunlight. I JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY measured larval head capsule widths daily, and resting body lengths at the time of molting using a stereomi- croscope with an ocular micrometer. Comparison between species. Because these two pyralids are relatively close taxonomically, the larvae can be difficult to distinguish in the field. Anticipafing that characters allowing such discrimination may be useful, I formally tested for differences in head cap- sule width and resting body lengths between the two species using the data presented in this paper on P. lu- nigerella and data on D. alternosquamella from Mooney (2001). Reliability of previously published food plant record. It is possible that the previous claim of P. lu- nigerella feeding on conifers (Prentice 1965) is erro- neous and that in fact the larvae were feeding on dwarf mistletoe in those trees. To evaluate this possibility, I carefully inspected the methods and dataset presented by Prentice (1965). I then consulted Hawksworth et al. (1996) and summarized the ranges for species of dwarf mistletoe known to parasitize the conifers from which P. lunigerella were reportedly collected. Dwarf mistle- toes are of great commercial importance as parasites of conifers in North America and have been called “the single most destructive pathogen of commercially valuable coniferous timber trees in . . . western Canada and western United States” (Hawksworth et al. 1996). For these reasons, they have been thoroughly studied, and the compendium by Hawksworth et al. (1996) is widely accepted as the authoritative source of infor- mation about the geographic distributions of these parasites and the coniferous hosts they use. By cross referencing data from Hawksworth et al. (1996) and Prentice (1965) I assessed the likelihood that dwarf mistletoes occurred on the conifers from which P. lu- nigerella larvae were collected. Literature review. Lepidopteran larvae known to specialize on dwarf mistletoe (Arceuthobium spp.) are the following: Mitoura spinetorum Hewitson (Ly- caenidae), Mitoura johnsoni Skinner, Filatima natalis Heinrich (Gelechiidae), D. alternosquamella Ragonot (Pyralidae) (Stevens & Hawksworth 1970, Hawksworth et al. 1996, Mooney 2001), and now P. lunigerella. In order to identify possible examples of food plant shifts between dwarf mistletoes and conifers I conducted a literature review to identify whether the relatives of any or all of these taxa include conifer feeders. Although examples of sister taxa feeding on dwarf mistletoe and conifers provides evidence for a recent food plant shift, more data are needed to infer the direction of the food plant shift. Such sister taxa examples by them- selves not indicate whether the shift was from conifers to dwarf mistletoe, or vice versa. VOLUME 57, NUMBER 1 49 TaBLE 1. Mean values for head capsule width, pre- and post-molt body lengths of resting larvae, and instar duration for Promylea lunigerella. Sample sizes and standard errors follow each measurement. x head capsule X post-molt body X pre-molt body X instar duration Instar width mm (N, SE) length mm (N, SE) length mm (N, SE) days (N, SE) 1 —_ ies ie 2 2 = = 2.09 (1, —) = 3 0.31 (3, 0.0008) Dil (il, ) 2.71 (3, 0.14) 9.0 (2, 3.0) 4 0.44 (9, 0.0016) 2.72 (3, 0.14) 3.71 (6, 0.25) 8.0 (4, 1.2) 5 0.58 (11, 0.0020) 3.72 (6, 0.25) 5.84 (8, 0.49) 8.3 (7, 0.68) 6 0.76 (16, 0.00003) 5.85 (8, 0.49) 9.12 (3, 1.16) 12.2 (10, 1.00) RESULTS Life history of P. lunigerella. I reared 16 P. lu- nigerella larvae through pupation, although none of these were collected as eggs. One larva passed through five instars before pupating, but I believe this species normally has six instars for several reasons. Dasypyga alternosquamella has six larval instars (Mooney 2001) and Dasypyga and Promylea are likely sister genera (Heinrich 1956). The head capsule width and length of this earliest P. lunigerella larvae were nearly identical to those of a second instar D. alternosquamella. The last three larval instars of P. lunigerella are significantly smaller than the last three larval instars of D. alter- nosquamella (see below). For P. lunigerella to have only five instars would require that this species hatch at a size 30% larger than the relatively closely related D. alternosquamella, but pupate at a size only half that of D. alternosquamella. Following this assumption of six larval instars, the 16 larvae I reared through pupation were collected from the field in the following life-stage distribution: One second instar, three third instar, five fourth instar, three fifth instar, and four sixth instar. Head capsule widths and larval resting lengths for P. lunigerella are presented in Table 1 according to this assumption, and the same data for D. alternosquamella from Mooney 2001 are presented in Table 2. Promylea lunigerella and D. alternosquamella were collected at the same time and from the same dwarf mistletoe plants. A comparison of the instar distribu- tions from these collections (Fig. 1) suggests that the time of emergence and oviposition of P. lunigerella is substantially earlier than that of D. alternosquamella, which occurs in mid-June (Mooney 2001). The median and modal life-stage for P. lwnigerella was fourth instar larvae and for D. alternosquamella was egg-first instar larvae, i.e., the former precedes the later by three to four instars. Based on instar duration data I estimate P. lunigerella emergence precedes D. alternosquamella by approximately three weeks, i.e., P lwnigerella emerges in late May. Comparison between species. There were suffi- cient sample sizes to compare larval lengths, head cap- sule widths, and instar duration between fourth, fifth, and sixth instar P. lunigerella and D. alternosquamella. I tested for differences between species in these three characters using separate one-way ANOVAs for each instar. I accounted for the increased likelihood of type I error with multiple tests using a Bonferroni adjust- ment (Zar 1999). Promylea lunigerella was significantly smaller in length than D. alternosquamella in fourth (F 141 = 32.98, p < 0.0001), fifth (F, ,, = 18.60, p = 0.0005) and sixth (F,, = 13.13, p = 0.011) instars and had significantly smaller head capsule widths in fifth (F, ,, = 11.13, p = 0.0037) and sixth (F, ,, = 389.36, p < 0.0001) instars at the Bonferroni adjusted alpha of 0.016. There were not significant differences in fourth instar head capsule widths (F, ,, = 0.69, p = .487), nor in duration of fourth (F, ,, = 2.42, p = 0.1483), fifth (F, , = 1.77, p = 0.2035), and sixth (F, ,. = 0.71, p = 0.4104) instar larvae. I reared 25 larvae through pupation for the life his- tory work described here and in Mooney 2001. Of TABLE 2. Mean values for head capsule width, pre- and post-molt body lengths of resting larvae, and instar duration for Dasypyga alter- nosquamella. Sample sizes (N) are given in column two. Standard errors follow each measurement. Post-molt body length for instar one is size at time of hatching. Reproduced from Mooney (2001). X head capsule X post-molt body X pre-molt body X instar duration Instar N width mm (SE) length mm (SE) length mm (SE) days (SE) 1 5 0.15 (0.005) 1.19 (0.048) 1.61 (0.093) 7.33 (0.558) 2 8 0.20 (0.004) 1.62 (0.093) 2.30 (0.088) 6.5 (0.563) 3 9 0.29 (0.010) 2.31 (0.088) 3.25 (0.124) 6.38 (0.263) 4 ) 0.43 (0.012) 3.26 (0.124) 5.36 (0.288) 6.33 (0.471) 5 9 0.64 (0.011) 5.37 (0.288) 8.25 (0.310) 7.11 (0.351) 6 9 0.96 (0.111) 8.26 (0.310) 16.56 (1.034) 14.78 (0.760 50 i D. alternosquamella [ | P. lunigerella proportion egg, 1st 2nd 3rd 4th 5th 6th larval instar Fic. 1. Distributions of life-stages of P. lunigerella (N = 16 lar- vae) and D. alternosquamella (N = 9) collected between 30 June and 1 August 1999. these, 16 were P. lunigerella and nine were D. alter- nosquamella. These data suggest a relative abundance of approximately 2:1. The precision of this estimate is reduced by the following facts: (1) the two species were at different stages in their phenology and likely had ex- perienced different rates of mortality prior to my col- lections, (2) differences in larval sizes due to phenolog- ical differences likely resulted in unequal rates of detection during larval collection, and finally, (3) not all larvae collected survived through pupation and the two species may have suffered different rates of mortality in the laboratory. Reliability of previously published food plant record. The P. lunigerella host plant data from Pren- tice (1965) are summarized in Table 3. A total of 347 larvae were found feeding on Abies amabilis (Dougl.) Forbes (Pinaceae), A. grandis (Pinaceae), Picea sitchensis (Bong.) Carr. (Pinaceae), Pseudotsuga men- ziesii and Tsuga heterophylla in the southern coastal area of British Columbia near Vancouver (“coastal B.C.”) and in interior B.C. near Lillooet (“interior B.C.”). These larvae were from 118 separate collec- tions, where each collection is from a separate locality, but distance between localities is unclear. The number of collections and the number of larvae from the coastal and interior regions were not specified. Using dwarf mistletoe species range data from Hawksworth et al. (1996) I determined which species of dwarf mistletoes parasitize the conifers listed by Prentice (1965), and whether the parasite range ex- tends to either coastal or interior B.C. Of the six dwarf mistletoe species parasitizing these five conifers, only A. tsugense (Rosendahl) G. N. Jones occurs in British Columbia, and its range is limited to the coastal region. Furthermore, while A. tsugense commonly parasitizes JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Abies amabalis, it very rarely parasitizes A. gandis and T. heterophylla (Hawksworth et al. 1996). Cross refer- encing these data on dwarf mistletoe ranges and larval host plant records (Table 3) demonstrates that a mini- mum of 46 larval collections (number of larvae is not determinable) were from trees on which there could not have been dwarf mistletoe. If I discount the possi- bility that larvae were collected from A. tsugense on its rare hosts then 110 of the 118 collections were made from trees without dwarf mistletoe. I therefore con- clude that most, and probably all, of Prentice’s records of P. lunigerella feeding on conifers are reliable food plant records. Literature review. Mitoura johnsoni Skinner (Ly- caenidae) and M. spinetorum Hewitison both feed on dwarf mistletoes while M. gryneus Hiibner and several species in the sister genus Callophrys (C. eryphon Boisduval, C. niphon Hiibner, C. lanoraicensis Shep- pard, C. hesseli Rawson & Ziegler) are conifer feeders (Scott 1986). Given that there are no dwarf mistletoe feeders reported in Callophrys, it would be reasonable to assume that the ancestral character for Mitoura is conifer feeding and that either one or two shifts from conifers to dwarf mistletoe have occurred. Filatima natalis Heinrich (Gelechiidae) is a dwarf mistletoe feeder (Heinrich 1920, Stevens & Hawks- worth 1970, Hawksworth et al. 1996) while several species of Chionodes Hiibner (Gelechiidae) feed on conifers (Heinrich 1920, Hedlin et al. 1981). While these species are not congeners, there is evidence to suggest that Filatima and Chionodes are sister taxa (R. Hodges pers. com.). There is not a great deal of infor- mation on food plants for other species of Filatima, but at least some feed on Salix (Karshold & Razowsky 1996). Feeding within Chionodes is diverse (Hodges 1999). Without an accurate phylogeny of this clade, and more complete food plant records, it is difficult to ascertain whether the taxonomic proximity of conifer and dwarf mistletoe feeding is the result of a past food plant shift or simply a coincidence. The dwarf mistletoe herbivores discussed here, P. lunigerella and D. alternosquamella, are both phycitine pyralids. There were a sufficient number of shared characters for Heinrich (1956) to at least preliminarily group these genera together: Heinrich’s key separates the genera within a single couplet and they are treated on adjacent pages in his text (Heinrich 1956). While no phylogenetic work has been done on these groups, more recent inspection of genital characters support Heinrich’s groupings (H. Neunzig pers. com.). The only food plant records within these two genera are those already discussed, i.e., P. lwnigerella, which feeds on both conifers and dwarf mistletoe, and D. alter- VOLUME 57, NUMBER 1 Dill TABLE 3. Conifer species from which Prentice (1965) reports P. lunigerella were isolated in coastal and interior British Columbia, the dwarf mistletoes (Arceuthobiwm spp.) known to parasitize those conifers (Hawksworth 1996), and whether the dwarf mistletoes ranges include the re- gions where larvae were found (Hawksworth 1996). “Coastal” refers to southern coastal BC including Vancouver, “interior” refers to the Lillooet area. The 118 larval collections (2 = 347 larvae) were made from separate localities from 1950-1957. Neither the number of collections from coastal vs. interior B.C., nor the larvae per collection were determinable. Conifer species of larval collections Abies Abies Picea Dwarf mistletoe amabilis grandis A. abietinum x x A. abietis-religiosae x x A. douglasii A. microcarpum x A. pusilum x A. tsugense x xi P. lunigerella Collections 8 53 1* sitchensis Dwarf mistletoe range Pseudotsuga Tsuga coastal interior menziesii heterophylla BC? BC? no no no no x no no no no no no x yes no 45* 11 D3, = UNS} *These collections were made from trees outside of the range of any possible dwarf mistletoe parasitism. nosquamella, a dwarf mistletoe feeder (Heinrich 1956). These records suggest that the congeners of these species may also be conifer and/or dwarf mistle- toe feeders. Based on the fact that D. alternosquamella is a dwarf mistletoe feeder, it appears the ongoing shift observed in P. lunigerella is from an ancestral condi- tion of dwarf mistletoe feeding to a derived condition of conifer feeding. DISCUSSION Both P. lunigerella and D. alternosquamella were abundant and occurred sympatrically at the Manitou Experimental Forest. This is somewhat surprising as it would seem that competitive exclusion should prevent two species of such close taxonomic relation and ecol- ogy from occurring sympatrically in the same habitat (Hardin 1960). The two species do differ significantly in size, and possibly this difference facilitates their co- existence. The previous record of P. lunigerella feeding on conifers is reliable, as are my observations of the species feeding on dwarf mistletoe. It is notable that these two accounts are separated by several thousand kilometers and multiple decades. These data suggest that either a food plant shift is actively occurring within this species, or perhaps that P. lunigerella is ac- tually two geographically separated, cryptic species that are more easily diagnosed by dietary preference than morphology. My review of the dietary literature suggests that shifts in feeding between parasitic plants and the hosts of those plants, and vice versa, have occurred multiple times and may be common among lepidopteran taxa that feed on parasitic and parasitized plants. Every one of the five species of Lepidoptera known to feed on dwarf mistletoe has a relative in the same or sister genus that feeds on conifers. In three of those cases (the two Mitoura and P. lunigerella) a food plant shift almost certainly occurred. The evidence for a shift in the gelechiids is suggestive but far from clear. The evidence to-date suggests that the shift in Mi- toura was from conifer to dwarf mistletoe, while the shift in the phycitine pyralids was from dwarf mistletoe to conifer. Holloway and Hebert'’s (1979) review of the Canadian Forest Insect Survey Data (e.g., Prentice 1965) suggested that forest lepidopterans feeding on conifers are less specific in their food plant choice than an- giosperm-feeding species. This suggests that switches from conifers to dwarf mistletoes may be more common. While I made no attempt to review the literature beyond those species feeding on dwarf mistletoes, in doing this work I became aware of another example in a different parasitic plant-host system: The nymphalid Euthalia lubentina Cramer feeds on several species of the mistletoe Loranthus (Loranthaceae) (Wynter- Blyth 1957) parasitizing Anacardiaceae, including mango Mangifera indica L. and Anacardium occiden- tale. These two Anacardiaceae species are fed upon by Euthalia aconthea garuda Moore (Corbet et al. 1978). A more exhaustive literature search would likely reveal more such examples. Despite the high degree of chemical dissimilarity and taxonomic distance between conifers and dwarf mistletoe, food plant shifting appears to have hap- pened repeatedly. These data provide support for a model of opportunity-based food plant shifting in which consistent physical association between plants may facilitate such shifts (Fig. 2). While the parasite- host relationship between dwarf mistletoes and conifers guarantees an unusual degree of close and consistent physical association, other associations might be predicted to produce the same phylogenetic pat- terns of food plant use. In their discussion of ecologi- cal opportunity and host shifting, Strong et al. (1984) 52 parasitic plants and their hosts HIGH c= al a“ ~~ a -3 oo) & i 6 2 rs [J s g a Fic. 2, Schematic model demonstrating how (1) phenotypic dif- ferences between food plants and (2) physical association between food plants affect the (3) likelihood of specialist herbivores shifting between those food plants. Food plant shifts are most likely between plants that are phenotypically similar and between plants that are consistently in close physical association. Dwarf mistletoes and conifers are phenotypically very distinct, but are in close physical as- sociation. cite Winter's (1974) findings of food plant shifts by in- sects from Myricaceae and Ericaceae moorland plants to the conifers with which they are frequently associ- ated. Strong et al. (1984) also cite multiple examples of laboratory studies in which, following initially high rates of mortality, insects shifted and adapted to novel and often dissimilar food plants. For example, Gould (1979) was able to induce phytophagous mites to shift from Curcurbitaceae to Fagaceae. Chew and Robbins (1988) review literature suggesting that lycaenid and ri- odinid mutualisms with ants have resulted in shifts to feeding on the lichens frequently associated with these ants, and to carnivorous feeding on the ants themselves and on ant-tended homopterans. Shifting among food plants has been an active topic of evolutionary and ecological research, but to-date there has been little work suggesting the mechanisms by which food plant shifts occur among dissimilar plants. While parasitic plants are but a small proportion of the flora available to lepidopteran larvae, the unusu- ally consistent physical association these plants must maintain with their hosts make these systems ideal for investigating the role of physical proximity among plants in food plant shifts by specialist herbivores. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY ACKNOWLEDGMENTS This research was supported in part by funds provided by the Rocky Mountain Research Station, Forest Service, U.S. Department of Agriculture. Brian Geils (U.S.D.A Rocky Mountain Research Sta- tion) provided valuable advice on dwarf mistletoe-related matters. M. Deane Bowers (University of Colorado, Boulder) provided facili- ties for, and guidance on rearing larvae. Herbert Neunzig identified P. 1. glendella. Ronald Hodges helped with information on gelechiid taxonomy and host plants use. Kathy Thomas provided field and lab- oratory assistance. Wayne Sheppherd and Steve Tapia (U.S.D.A. Rocky Mountain Research Station, Manitou Experimental Forest) provided logistical assistance for this work. LITERATURE CITED BUCKINGHAM, J. 1994. Dictionary of natural products. 1st ed. Chap- man & Hall, New York. 8 vols. CHEw, F. S. & R. K. Rospins. 1988. The Biology of Butterflies, pp. 65-78. In Vane-Wright, R. I. & P. R. Ackery (eds.), The biology of butterflies. Princeton University Press, Princeton, New Jer- sey. 429 pp. Corset, A. S., H. M. PENDLEBURY & J. N. ELior. 1978. The but- terflies of the Malay peninsula. 3rd ed. Malayan Nature Society, Kuala Lumpur. 578 pp. DENNO, R. F. & M. S. McCLurE. 1983. Variable plants and herbi- vores in natural and managed systems. Academic Press, New York. 717 pp. Dyar, H. G. 1906. Descriptions of new American moths. J. New York Ent. Soc. 14:30-31. EHRLICH, P. R. & P. H. RAVEN. 1964. Butterflies and plants: a study in coevolution. Evolution 55:247—270. Futuya, D. J. & M. SLATKIN. 1983. Coevolution. Sinauer Associ- ates, Sunderland, Massachusetts. 555 pp. GouLD, F. 1979. Rapid host range evolution in a population of the phytophagous mite Tetranychus urticae Koch. Evolution 33:791-802. Harbin, G. 1960. The competitive exclusion principle. Science 131:1292-1297. HAWKsWoRTH, F. G., D. WIENS, B. W. GEILS & R. G. NISLEY. 1996. Dwarf mistletoes: biology, pathology, and systematics. U.S. Dept. of Agriculture Forest Service, Washington, District of Co- lumbia. 410 pp. HEDLIN, A. F., H. O. I. Yates, D. C. Tovar, B. H. EBEL, T. W. Ko- ERBER & E. P. MERKEL. 1981. Cone and seed insects of North American conifers. Canadian Forestry Service, U.S. Forest Service, Secretaria de Agricultura y Recursos Hidraulicos Mex- ico, Ottawa. 112 pp. HEINRICH, C. 1920. On some forest Lepidoptera with descriptions of new species, larvae, and pupae. Proceedings of the U.S. Na- tional Museum 57:53-92. . 1956. American moths of the subfamily Phycitinae. Smith- sonian Institution, Washington, District of Columbia. 581 pp. Hopces, R. W. 1999. Fascicle 7.6, Gelechioidea: Gelechiidae (Part). Wedge Entomological Research Foundation, Washing- ton, District of Columbia. 339 pp. Hottoway, J. D. & P. D. N. HEBERT. 1979. Ecological and taxo- nomic trends in macrolepidopteran host plant selection. Biol. J. Linnean Soc. 11:229-251. Jupp, W. S. 1999. Plant systematics: a phylogenetic approach. Sin- auer Associates, Sunderland, Massachusetts. 464 pp. KARSHOLD, O. & J. RAZOWSKY (EDS.). 1996. The Lepidoptera of Eu- rope: a distributional checklist. Apollo Books. Mooney, K. A. 2001. The life history of Dasypyga alternosquamella Ragonot (Pyralidae) feeding on the Southwestern dwarf mistle- toe (Arceuthobium vaginatum) in Colorado. J. Lep. Soc. 55:144-149. PRENTICE, R. M. 1965. Microlepidoptera. Canadian Dept. of Agri- culture, Forest Biology Division, Ottawa. 840 pp. Raconot, N. 1887. Diagnoses of North American Phycitidae and Galleriidae. Published by the author, Paris. 20 pp. VOLUME 57, NUMBER 1 REICH, R. 1992. Voracious moth larvae feed heavily on lodgepole pine dwarf mistletoe shoots in Prince George Forest Region. British Columbia Ministry of Forests, Victoria, British Colum- bia. 22 pp. Scott, J. A. 1986. The butterflies of North America: a natural his- tory and field guide. Stanford University Press, Stanford, Cali- fornia. 583 pp. STEVENS, R. E. & F. G. HAwKsworTH. 1970. Insects and mites as- sociated with dwarf mistletoe. USDA Forest Service Research Paper, Washington, District of Columbia. 12 pp. STRONG, D. R., J. H. Lawron & R. SOUTHWOOD. 1984. Insects on plants: community patterns and mechanisms. Harvard Univer- sity Press, Cambridge, Massachusetts. 313 pp. 53 VANE-WRICHT, R. I. & P. R. AcKERY. 1988. The Biology of butter- flies. Princeton University Press, Princeton, New Jersey. 429 pp. WINTER, T. G. 1974. New host plant records of Lepidoptera asso- ciated with conifer afforestation in Britain. Entomologists’ Gaz. 25:247-258. WynTeER-BLyTH, M. A. 1957. Butterflies of the Indian region. Ist ed. Bombay Natural History Society, Bombay. 523 pp. Zar, J. H. 1999. Biostatistical analysis. 4th ed. Prentice Hall, Upper Saddle River, New Jersey. 663 pp. Received for publication 15 April 2002; revised and accepted 5 September 2002. Journal of the Lepidopterists’ Society 57(1), 2003, 54-61 A CONTRIBUTION TO THE SYSTEMATICS OF THE COPAXA SEMIOCULATA SPECIES-GROUP (SATURNIIDAE), WITH NOTES ON THE EARLY STAGES, AND A DESCRIPTION OF COPAXA LUNULA, NEW SPECIES Kirpy L. WOLFE! 3090 Cordrey Drive, Escondido, California 92029, USA, email: kirwolfe@pacbell.net CLAUDE LEMAIRE? La Croix des Baux, F-84220 Gordes, France, email: Lemaire.C@wanadoo.fr ANGELA AMARILLO S. Department of Entomology, University of Kentucky, $225 Agricultural Sciences Central North, Lexington, Kentucky 40546-0091, USA, email: aamarill@hotmail.com AND CHRISTOPHER A. CONLAN! 11513 Hadar Drive, San Diego, California 92126, USA; email: conlan@adne.com ABSTRACT. Copaxa semioculata is re-described. Recent collecting has demonstrated that a population from western Ecuador was misidentified as C. semioculata semioculata by Lemaire (1975, 1978), and that this population plus C. semioculata orientalis, of eastern and west- er Ecuador and central and western Colombia, are actually conspecific and specifically distinct from C. semioculata, which occurs only in the eastern Andes of South America from Venezuela to Peru. The male of C. semioculata differs noticeably from C. orientalis, new status by its smaller size, narrower forewings, variable color, dark antennae, differences in the genitalia and especially hours of nuptial flight. Males are di- urnal and have rarely been collected at lights. Male and female genitalia are figured and immature stages are described and illustrated in color. Larvae fed in the laboratory on Persea americana (Lauraceae). Copaxa orientalis is hereby elevated to full specific rank. Additionally, a new species closely related to C. semioculata is described from Bolivia and Peru. RESUMEN. Se describe Copaxa semioculata de nuevo. Mediante muestreo realizado recientemente se ha demostrado que una poblacién del oeste del Ecuador fue malidentificada como C. semioculata semioculata por Lemaire (1975, 1978), y que esta poblacién mas C. semioculata orientalis, del este y el oeste del Ecuador y el centro y el oeste de Colombia, en realidad son de la misma especie y son distintas a C. semiocu- lata, la cual se encuentra en los Andes orientales de Venezuela al Pert. E] macho de C. semioculata difiere notablemente de C. orientalis, es- tatus nuevo por su tamafio menor, alas delanteras mas delgadas, color variable, antenas més oscuras, diferencias en los genitales y en especial las horas de su vuelo nupcial. Los machos son diumos y rara vez se han colectado con luces. Se ilustran los genitales del macho y de la hembra y se describen y presentan fotografias en colores de los estadios inmaduros. En el laboratorio las larvas se alimentaron con Persea americana (Lauraceae). Se eleva Copaxa orientalis a pleno rango especifico. Adicionalmente se describe una especie nueva de Bolivia y el Pert, de prox- ima afinidad con C. semioculata. Additional key words: Bolivia, Colombia, Ecuador, immature stages, lunula, Neotropical, orientalis, Persea, Peru, Venezuela. The genus Copaxa (Walker 1855) comprises more was retained by Packard (1914) and Bouvier (1936). than 36 species of often large and colorful moths, dis- Curiously, Draudt (1929) separated semioculata from tributed from Mexico to Argentina. These were di- sapatoza and placed semioculata in Saturniodes vided by early authors into three genera, with Copaxa where it remained for some years. Michener (1952) containing the majority of species. Among the high al- unified Saturniodes and Sagana under Copaxa but titude Andean species, some were placed in Sagana retained the three names as subgenera. Lemaire Walker (1855) and others, along with several Mexican (1978) demonstrated problems with Michener’s species, in Saturniodes Jordan (1911). The Andean model and discarded the subgenera, synonymizing all species are a poorly-studied group generally restricted in Copaxa. to often cold, wet and steep habitats between 2000- Most of the high altitude Andean Copaxa species 4000 m, from Venezuela to Bolivia. are characterized by lunate or modified lunate hyaline The genus Sagana was proposed for Sagana sapa- discal spots on all four wings. Until now, only three toza (Westwood 1853) from Colombia, and was sub- easily distinguishable species of the Copaxa semiocu- sequently used to harbor the closely related but lata complex were recognized: C. sapatoza, C. semioc- slightly larger Sagana semioculata R. Felder & Ro- ulata and C. herbuloti. C. sapatoza has wide, squared genhofer (1874) from Venezuela. This arrangement lunate discal spots, C. semioculata, a variable moth en- compassing several hidden species, has lunate spots. ' Research Associate, Natural History Museum of Los Angeles a : : ae County, Califormia, USA. Copaxa herbuloti Lemaire (1971), described from a > Correspondant du Musetim national d’Histoire naturelle, Paris. single male from northwestern Peru and obviously dis- VOLUME 57, NUMBER 1 tinct in the genitalia, has widely distorted hyaline dis- cal spots on the forewing. C. orientalis Lemaire (1975), a large, dark species, was described from the eastern Andes of Ecuador as a subspecies of C. semi- oculata. Recent collecting has yielded additional phe- notypes of the C. semioculata species group, calling into question the previous identification of the nominotypical taxon of C. semioculata. A taxonomic problem began with Felders’ and Ro- genhofer's description of Copaxa semioculata from an unspecified number of female specimens from “Venezuela.” Lemaire (1978:197) designated as lecto- type a specimen preserved in The Natural History Mu- seum (BMNH, London) (by way of the Felder and W. Rothschild collections). Examined by KLW and CL, it provides no precise locality data on the label. Sonthon- nax’s (1901) citation of a male and female with wingspan of 12 cm, from “Bogota, Venezuela” is erroneous on two counts, as Bogota is in Colombia and the size is much too large for C. semioculata. When Lemaire de- scribed C. semioculata orientalis no male specimens of C. semioculata were known from Venezuela or Colom- bia. A large series of male and female specimens from western Ecuador, collected at lights and preserved in the BMNH, appeared to CL to match the original de- scription and illustrations and he erroneously assigned the specimens to the nominotypical subspecies in the description of the new subspecies orientalis and in the revision of the genus (Lemaire 1978). The preponderance of small females of C. semiocu- lata attracted to lights in eastern Ecuador aroused our suspicion that the true male of this species might be diurnal, and that Lemaire’s “semioculata” of western Ecuador were misidentified. Evidence of this began to emerge with the net capture by Amarillo of a small or- ange male flying slowly and low to the ground in full sunlight at 1630 h in January, 1992 at 2850 m in Iguaque National Wildlife Sanctuary in Boyaca De- partment, northeast of Bogota, Colombia. A female was later collected at lights. Returning to Iguaque in April 1998, and April 2000, KLW and AAS captured seven additional females. While examining public and private collections in Venezuela and Colombia, KLW found a female speci- men of C. semioculata from Tachira, western Venezuela (2425 m) and two male specimens of the same species from Colombia. Both males were cap- tured flying at noon above 2000 m, one on the Venezuelan-Colombian border and the other near Bo- gota, by J. F. Le Crom (pers. com.), who regularly sees it flying high above the ridges east of the city. In spite of the vagueness of the type locality “Venezuela,” it can be assumed, based on the biogeo- graphical data, especially elevation, that the lectotype of C. semioculata originates from the Mérida Cordillera or from the Province of Tachira in western Venezuela near Colombia. Thus, this lectotype and the two above specimens from Bogota and from the Venezuelan/Colombian border can be considered as conspecific, which resolves the long perplexing prob- lem of the identity of C. semioculata and of the identi- fication of the corresponding male. With the additional specimens, we are confident that the male is diurnal. The name Copaxa semiocu- lata semioculata should no longer be applied to the western Ecuadorian population, in which the male is nocturnal. Both previous subspecies should, pending further research, be referred to Copaxa orientalis (Figs. 7, 8), which is hereby elevated to full species sta- tus. The usually much larger C. orientalis has not been found in Venezuela, and in Colombia has been col- lected only in the southern, central and western Andes. Copaxa semioculata (R. Felder & Rogenhofer) Redescription. Male (Figs. 1, 2). Head brown, orange beige or black, eyes large. Antennae with brownish yellow shaft and dark gray rami, quadripectinate. Thorax variable, with indistinct yellow collar. Tibia pink with long beige hairs, tarsi bright pink. Abdomen vari- able, lighter ventrally. Forewing length 35-50 mm, falcate; apex rounded. Background color usually orange brown of varying inten- sity from light orange beige to dark brown or gray, more or less suf- fused with dark gray or black scales in the median and especially the distal outer border; tornus lighter, yellowish; ante- and postmedial black or brown lines often blurry and indistinct. Apical spot small, gray with white on apical edge; trace of white second spot caudad to first. Lunate hyaline discal spot broad, usually bordered first nar- rowly black then broader dark yellow and again narrowly black. Forewing ventrally dark or light, similar to dorsal color but lighter, with dark band along indistinct postmedial line and on border. Hindwing same color as forewing but most of costal area from base to border pale, often tinged pink on forward basal area; brown ante- medial and undulating postmedial lines enclose broad darker area encompassing discal lunate spot; spot narrower but bordered as in forewing; submarginal band an indistinct series of U-shaped gray or black dashes bordered faintly white on outer edge. Ventrally similar to forewing. Male genitalia (Fig. 17) similar to C. orientalis (Fig. 21) (these il- lustrated as C. semioculata semioculata by Lemaire, 1978:197, Figs. 156, 157), with long hooks on each arm of the transtilla, but differ- ent in having a triangular, instead of round, juxta and narrower, more pointed apices of the valves. The vesica evaginates dorsally. Female (Fig. 3). Head brown, palpi brown. Antennae dull yel- low, bipectinate. Thorax anteriorly dark gray with yellow tuft collar, otherwise beige, tinted rose, yellow, brown or gray. Tibia and tarsi pinkish beige. Abdomen beige, lighter laterally and darker and in some specimens pinker ventrally. Forewing length 41-55 mm, wings broadly rounded; ground color light beige, in some specimens darker, with pink, yellow, brown or gray tint; central band and bor- der medium brown; antemedial and wavy postmedial lines dark gray; costal border dark gray at base with long white hairs; wing base, median area and vague submarginal band suffused with dark gray and reddish brown scales, the submarginal band outwardly suf- fused with white; single apical spot dark gray bordered outwardly white. Hindwing colored similarly to forewing but with lighter mar- gins, submarginal band of dark gray, outwardly white U-shaped dashes; irregular area between antemedial and postmedial lines as in 56 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Fics. 1-16. Adults of Copaxa semioculata, Copaxa lunula, new species, Copaxa orientalis, new status, immature stages of C. semiocu- lata and larva of C. lunula. 1. Copaxa semioculata, male, brown phenotype, COLOMBIA, Boyaca, Santuario Nacional de Flora y Fauna de Iguaque, 2990 m, 22-24 April 2000, leg K. & S. Wolfe. 2. C. semioculata, male pale phenotype, COLOMBIA, Boyaca, Iguaque National Wildlife Refuge, VOLUME 57, NUMBER 1 male; forward area of wing pale, pink in some specimens. Discal spots as in male, surrounded by yellow and gray or black. Ventrally similar, but suffused with white scales except on margin; antemedial and postmedial lines medium brown. Female genitalia (Fig. 18) similar to C. orientalis. Diagnosis. Males of C. semioculata can be distin- guished from C. orientalis (Fig. 7) by smaller size, much narrower forewings and dark antennae. Males of C. semioculata apparently search for females for about one hour at midday and remain in copula until late afternoon or evening. KLW noted three males fly- ing from 1120 h to 1205 h (Iguaque, April 2000, un- published obs.). In all known cases, males were col- lected either during daytime with nets or at lights just at dusk. In two cases, males were attracted to lights with a virgin female of C. sapatoza, with which one copulated, resulting in infertile eggs (Diego Bonilla P. pers. com.). Females of C. semioculata are smaller than C. orientalis (Fig. 8) and are strictly nocturnal, arriving at lights throughout the night. In C. orientalis, in which the antennae are yellow in both sexes, flight is nocturnal, with males attracted to lights between 1930 h and 2115 h (KLW, CL, AAS, CAC, pers. observ.). Distribution. All known specimens of C. semiocu- lata originated in the eastern Andes between 2150-3430 m, in forest on both slopes, from western Venezuela, Colombia, Ecuador and northern Peru. Lo- calities where it has been collected include: VENEZUELA, Tachira, PAramo Tama, Betania, 2425 m, 16-10 Mar 1983, Exp. Instituto Zoologia Agricola, Fac. Agronomia; COLOMBIA, (near Venezuelan bor- der), 22 Dec. 1993, leg. LeCrom (netted at noon); COLOMBIA, Boyaca, Santuario Nacional de Flora y Fauna de Iguaque, El. 2990 m, 24-26 Apr 1998, at MV & UV lights, leg K. Wolfe, A. Amarillo, C. Sarmiento; COLOMBIA, Cundinamarca, Represa E] Sisga, 4 Jan 1968, J. Cayon; COLOMBIA, Cundinamarca, 3300 m, 12 Nov 1995, leg LeCrom; COLOMBIA, Cundina- marca, Villa Punzén, 2900 m, May 2001, leg D. Bonilla; COLOMBIA, Cundinamarca, Bogota, Rd. Bogotéa— Tunja, Villa Pinzon, 3100 m, Sep 1999, leg D. Bonilla; COLOMBIA, Cundinamarca, Rd. Bogotéa—Tunja, Chaconta, 2600 m, Jul 2001, leg T. Decaéns & D. Bonilla; ECUADOR, Cotopaxi, ca. 25 km NE of Lata- cunga, MV light 1930 h, el. 3151 m, 10 Mar 1995, K. << | Ol Wolfe & S. Smoot; ECUADOR, Napo, Papallacta, 2800 m, 2 Mar 1992, Wm. Kelly; ECUADOR, Napo, rd. Baeza to Tena, S. of Cosanga, 2150 m, N. Vene- dictoff, 23 Mar 1976; ECUADOR, Napo, Cosanga to Tena km 7, 2350 m, 19 Jul 1990, leg. D. Herbin & J. Haxaire; ECUADOR, Morona Santiago, 44 km on Rd. Gualaceo-Limon, El. 2300 m, 4 May 2000, GPS = 03°01.00S x 078°34.83W, K. & S. Wolfe, C. & M. Con- lan; PERU, Amazonas, Achuras 3100 m, 10 May 1999. Copaxa orientalis and C. semioculata are probably sympatric in parts of their range, but semioculata prefers dense humid forest below the treeline whereas orientalis ranges at higher altitudes, near the tree line and in alpine shrubbery. Copaxa orientalis has been collected in COLOMBIA in the following locations: eastern slope of Central Cordillera in departments of Caldas (Parque Nevado) and Tolima (Anaime Re- serve); Western Cordillera in Valle (Anchicaya Alto) and Narifio ( Cumbal, Chachaguf, La Laguna, Ipiales, 86 km S of Pasto). In ECUADOR, specimens of C. orientalis exist from Carchi, near Tulcdn; the eastern Cordillera in Napo near Cotopaxi border, Rd. Salcedo to Napo, km 49, 3500 m (type locality); western Cordillera in Pichincha, old Rd. Quito to Santo Domingo de los Colorados, km 26, 3200 m; and Tun- gurahua (west of Ambato). Both C. semioculata and C. orientalis fly during much of the year. However, males of C. semioculata do not seem to be active during pe- riods of heavy, all-day cloudiness, and females are not usually found during the following nights. IMMATURE STAGES Females captured at lights in Colombia oviposited in paper bags. Larvae were subsequently reared in the laboratory on Persea americana Mill. (Lauraceae). Some larvae initially accepted Quercus sp. (Fagaceae), the only other hostplant accepted among several of- fered, but were later moved to Persea. Larvae were consistent in color, pattern and spination with other Copaxa species, most resembling larvae of the canela- lavendera group (Wolfe 1993). More than 20 cocoons with pupae were obtained but only one female emerged. Remaining pupae died within a year. Egg (Fig. 9): 1.9 mm long x 1.6 mm wide x 1.3 mm 2900 m, 5 January 1992, leg A. Amarillo. 3. C. semioculata, female, COLOMBIA, Cundinamarca, Bogota, Rd. Bogota - Tunja, Villa Pinzon, 3100 m, September 1999, leg T. Decaéns & D. Bonilla. 4. Copaxa lunula, paratype male, orange phenotype, BOLIVIA, Cochabamba Prov., dwarf cloud forest 1 km E La Siberia, 3050 m, 17°47.63S x 064°44.70W, 12 November 1999: ex-2 at lights; leg. K. Wolfe & C. Conlan, reared on Persea by C. Conlan. 5. C. lunula, paratype male, dark phenotype, ibid. 6. C. lunula, paratype female, ibid. 7. Copaxa orientalis, male, COLOMBIA, Tolima, Municipio Cajamarca, Anaime Reserve, el. 3310 m, 29 March 1995, K. Wolfe, S. Smoot, A. Amarillo, C. Sarmiento. 8. C. orientalis, fe- male, ibid. 9. C. semioculata eggs. 10. C. semioculata 1st instar larva. 11. C. semioculata 2nd instar larva. 12. C. semioculata 3rd instar larva. 13. C. semioculata 4th instar larva. 14. C. semioculata 5th instar larva. 15. Copaxa lunula 5th instar larva. 16. C. semioculata cocoon with pupa. Illustrations by KLW. 58 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 17-21. Genitalia of Copaxa semioculata, C. lunula and C. orientalis. 17. C. semioculata, male genitalia with aedeagus separated. 18. C. semioculata, female genitalia, with damaged corpus bursae. 19. C. orientalis, male genitalia with aedeagus separated. 20. C. lunula, male genitalia with aedeagus separated. 21. C. Junula female genitalia. Genitalia illustrations by KLW. thick; broad transparent dark brown on both faces with large, dark micropile at one end; deposited flat, singly or in short strings of 2-5. Eggs maintained at 21°C + 3° required ca. 18 days to hatch. Larva: Most larvae hatched between dawn and noon, the majority midmorning. Larvae fed for ca. 55 days before spinning cocoon. First instar (Fig. 10): Head: 1 mm wide; mahogany brown with long translucent setae, area of stemmata and mandibles black. Body: 10 mm max. length; lemon yellow, broad notal plate mahogany brown, narrow dorsal stripe and three interrupted and irregular sub- dorsal and lateral stripes black; scoli with broad base salmon, dorsal setae dark brown, lateral long hairs lighter. Thoracic legs black; abdominal prolegs, paranal lobes and ventral body greenish white. VOLUME 57, NUMBER 1 Second instar (Fig. 11): Head: 1.6 mm wide, ma- hogany brown with white clypeus. Body: 15 mm max. length; yellowish green, dorsum white to greenish white; narrow black middorsal stripe almost disappear- ing on central area of some segments; single black sub- dorsal zigzag stripe interrupted by subdorsal scoli; scoli mostly golden orange, except reddish brown dor- sal prothoracic scoli and notum. Thoracic legs brown, anal legs greenish white, anal area light reddish brown. Dorsal prothoracic and ninth abdominal segment scoli, subdorsal second thoracic and ninth abdominal seg- ment scoli, and all lateral scoli with central seta con- sisting of long hair with wide lanceolate tip. Third instar (Fig. 12): Head: 2 mm wide, dull green. Body: 20 mm max. length; green mottled olive, darker ventrally, no dorsal stripes. Bases of dorsal scoli slender, elongated, scarlet with narrow yellow band at base. Cen- tral spine of dorsal and lateral (but not most subdorsal) scoli with long, lanceolate tipped shiny black hairs as in second instar; segments 6-11 with wide white forward arching spines originating just anteriorly of each dorsal and subdorsal scolus; numerous tiny white fan-shaped setae scattered over integument. On segments 5-10 a black diagonal slash bordered yellow on upper posterior side hides yellow spiracles. Thoracic legs, abdominal and anal feet brown, rest of paranal lobes yellowish green. Fourth instar (Fig. 13): Head: 3.1 mm, green. Body: 36 mm max. length; color, pattern and spination with central long, lanceolate tipped hairs generally similar to third instar except scoli now submerged, scolic spines tiny, thin; forward arching wide dorsal and narrower subdorsal spines now deep pink; a white prothoracic collar band; spiracles yellow. Feet and paranal area as in third instar. Fifth instar (Fig. 14): Head: 5.2 mm, green. Body: 65 mm long x 12 mm thick after feeding is completed. Color and spination as in fourth instar; curved setae on feet white. Cocoon (Fig. 16): Medium brown, double walled, open mesh, shiny. Pupa: 23-26 mm long x 9 mm—12 mm thick, light brown, smooth, with cremaster of single, short spine. Copaxa lunula Wolfe & Conlan, new species Description. Male (Figs. 4, 5): Head variable shades of brown, eyes large. Antennae with brownish yellow shaft and dark gray or black rami, quadripectinate. Thorax variable, with indistinct yellow collar. Tibia pink with long beige hairs, tarsi bright pink. Abdomen variable, lighter ventrally. Forewing length 38-40 mm, falcate; apex rounded. Background color orange to dark brown or gray, darker be- tween ante- and postmedial lines and toward apex. Tornus and apex lighter, apical spot black bordered white on outer edge; lunate hya- line discal spot bordered black then faintly yellow. Underside as above but lighter. Hindwing same color as forewing but lighter. Black undulating antemedial and postmedial lines enclose broad darker area and lunate spot notably ringed by yellow bordered with 59 black. Submarginal band a bold series of black U-shaped dashes. Ventrally similar to forewing. Male genitalia (Fig. 19) similar to those of C. semioculata (Fig. 17) but with bilateral long curved sclerotized spines on the ventral rim of the juxta. Aedeagus is more robust than in C. semioculata and vesica evaginates laterally. Female (Fig. 6): Head greenish gray, palpi dark brown. Anten- nae dull yellow, bipectinate. Thorax anteriorly olive to yellow, re- mainder olive. Tibia and tarsi pink. Abdomen olive, darker in some individuals. Forewing length 43-54 mm, wings broadly rounded; ground color gray, mostly greenish with yellow on the borders, some specimens almost black. Antemedial and postmedial lines black. Two distinct apical black spots, discal spot as in male. Hindwing color similar to forewing but more rosy or mauve. Discal spot and lines as in male. Ventrally lighter. Female genitalia (Fig. 20) do not differ obviously from C. semioculata. Types. Holotype ¢: BOLIVIA, Cochabamba Dept., dwarf cloud forest 1 km E La Siberia, 3050 m, 17°47.63S x 064°44.70W, 12 Nov 1999; leg. K. Wolfe & C. Conlan; ab ovo., ex-° at lights, reared in CA on Persea americana by C. Conlan; em.24 Jul 2000. Allo- type °: Same locality data as holotype, wild-caught. Paratypes (2 d and 3 2): 2 d, same locality and data as holotype, em. 2 2 24 Jul 2000; 1 2°, same data as allo- type; 1 2°, BOLIVIA, Cochabamba Dept., lower cloud forest E of Pojo, 2700 m, 17°46.12S x 065°42.04W, 1 Nov 1999; at MV & UV lights, leg. K. Wolfe & C. Con- lan; 1 2, BOLIVIA, La Paz Dept., Rd. La Paz— Coroico, 2615 m, 07 Dec 1991, leg. G. Lecourt & T. Decaéns. The holotype and allotype are placed in the collec- tion of the Muséum national d’Histoire naturelle, Paris, France. Paratypes will remain in the following collections: K. Wolfe, 1 ¢, same data as holotype; 1 2, same data as holotype; 1 2, BOLIVIA, Cochabamba Dept., lower cloud forest E of Pojo, 2700 m, 17°46.12S x 065°42.04W: C. Conlan 1 6 same data as holotype; T. Decaéns, 1 2°, BOLIVIA, La Paz Dept., Rd. La Paz—Coroico, 2615 m. Etymology. This species is named for the translu- cent lunate discal spots on all four wings. Diagnosis. This new species is closely allied with C. semioculata, with which it shares size, shape, mark- ings, similar variable colors, habitat and midday nup- tial flight in the male. In the eastern Andes of central Peru it can only be distinguished with certainty from C. semioculata by dissection of the genitalia, which present two obvious long spines on the ventral rim of the juxta, completely lacking in C. semioculata. Distribution. As in C. semioculata, known specimens originate in eastern Andes in forest from 2000-3050 m, from north-central Peru to central Bolivia. In Cochabamba Department, females we captured at lights in dwarf cloud forest at 3050 m oviposited in pa- per bags. Larvae were reared and three adult males of two color morphs were obtained. Although close to C. semioculata, the genitalia are easily distinguished. 60 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY ‘os ee ef é ) (oo) 2 fo) OZ 4 Ohne! Oem 2 Days after infestation JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY WO 2 4 6 6 @ 12 4 Days after infestation Fic. 3. Proportion (+SE) of initial monarch larvae remaining strictly on potted milkweed plants (circles and solid lines) or within pots (tri- angles and dashed lines), i.e., collectively on plants, pots, and soil within pots, from 0-14 days post infestation at 0, 1, 2, and 8 m from a Bt corm field (Bt11). Predicted lines are based on the Weibull model. milkweed east of the corn (1.12 + 0.28 grains cm~), with intermediate densities on milkweed south and west of the corn (3.34 + 1.47 and 1.63 + 0.28 grains cm~, respectively), statistically not different from den- sities on the north and east. Pollen densities differed significantly (df for error = 32; F = 8.14; p = 0.0004) on milkweed 0 and 1 m from corn (4.15 + 1.28 and 3.86 + 1.32 grains cm”, respectively) compared to plants 2 and 8 m from corn (1.58 + 0.39 and 0.49 + 0.17 grains cm”, respectively). Eight days after larval infestation, the maximum mean pollen density was 2.9 + 0.74 grains cm. Pollen densities differed significantly (df for error = 32; F = 4.50; p = 0.0096) on milkweed north of the corn (1.10 + 0.38 grains em) compared to milk- weed east or west of the corn (0.52 + 0.10 and 0.34 + 0.17 grains cm™, respectively), with milkweed on the south (0.71 + 0.11 grains cm~) being indistinguishable from the other directions. Pollen densities differed sig- nificantly (df for error = 32; F = 9.88; p = 0.0001) on milkweed at 1 m (1.33 + 0.34 grains cm~) compared to TABLE 3. Weibull model parameters (b = mortality rate and c = shape) for proportion of larvae remaining strictly on potted milkweed plants or collectively from plants, soil, and pots from 0-14 days post infestation, 2000. Location b (+SEM) c (+SEM) F 1 0 meters Plants 7.02 (+£0.78) a 0.92 (£0.17) a 127.01" 0.78 Plants, soil and pots 10.73 (41.76) a 0.68 (+0.15) a 142.59* 0.80 1 meter Plants 4.45 (40.41) a 1.19 (40.17) a 136.95* 0.80 Plants, soil and pots 8.42 (41.33) a 0.71 (40.17) a 94.81* 0.73 2 meters Plants 5.78 (+0.44) a 1.55 (+0.24) a 170.47* 0.83 Plants, soil and pots 8.27 (+0.76) b 1.12 (40.21) a 147.91* 0.81 8 meters Plants 2.67 (40.40) a 0.75 (40.13) a 62.38* 0.64 Plants, soil and pots 3.38 (40.78) a 0.49 (+0.12) a 49.86* 0.59 For each regression, there were 2 df for the regression and 70 df for residual. Means within a column for each distance followed by the same letter are not significantly different; 95% confidence intervals (constructed using Welch’s unpaired ¢) for the difference in parameter estimates between treatments included zero. *p < 0.001. VOLUME 57, NUMBER 2 1.0 0.8 0.6 0.4 0.2 0.0 3d,0m 7d,0m 1.0 0.8 0.6 0.4 0.2 0.0 3d,1m 7d,1m 1.0 0.8 0.6 0.4 0.2 0.0 1.0 0.8 0.6 0.4 0.2 0.0 3d,2m 7d,2m Proportion of total larvae Pill ah 3d,8m 7d,8m ima ae Instar Instar Fic. 4. Instar distribution (+SD) of monarch larvae remaining, near Bt corn (Bt11), strictly on potted milkweed plants (black) and larvae that have left milkweed plants, but remained on the pots or on the soil within the pots (gray) at 0, 1, 2 and 8 m on 3 and 7 days post infestation in 2000. milkweed 0, 2 and 8 m from corn (0.70 + 0.17, 0.40 + 0.09 and 0.23 + 0.08 grains cm™, respectively). DISCUSSION In 1999 and 2000, we were unable to detect an ad- verse effect of Bt corn on monarch larvae. Survival was similar for larvae near Bt and non-Bt cornfields, and the rate of mortality was lower (i.e., larger Weibull b value) for larvae on milkweed near the edge of a Bt cornfield compared to larvae on milkweed farther from a field. Stanley-Horn et al. (2001) and Zanger! et al. (2001) were also unable to detect any adverse im- pact of Bt corn on monarch larvae under field condi- tions. The likely mechanisms by which Bt corn might adversely affect monarchs depend upon larval con- sumption of corn pollen or other corn tissue, such as anthers (Hellmich et al. 2001). The present study re- ports a maximum Bt corn pollen density of 10.7 grains cm~ on milkweed leaves. Our observed pollen densi- ties, along with mean pollen densities reported in Jesse and Obrycki (2000), Wraight et al. (2000), Stanley- Hom et al. (2001), Zangerl et al. (2001) and Pleasants 97 et al. (2001), are far below the least observable effect concentration of 1000 grains cm for the commonly grown Bt events Mon810 and Bt11 (Sears et al. 2001). The lack of an observed adverse effect of Bt corn on monarch larvae in the present studies is similar to re- sults obtained by Wraight et al. (2000) working with black swallowtails, Papilio polyxenes Fabricius, near Bt corn. However, not all Lepidoptera are equally sus- ceptible to Bt toxins. For example, Bt sweet corn pro- vides greater control of European corn borer than corn earworm, Helicoverpa zea (Boddie) (Burkness et al. 2001). In addition, Wagner et al. (1996) observed differential susceptibilities among non-target forest Lepidoptera to foliar applications of Bt insecticide. Over time, environmental factors (e.g g., rain and wind) may reduce densities of corn pollen (Pleasants et al. 2001) on the relatively smooth upper surface of milkweed leaves (Bhowmik 1994). Multiple rainfall events that occurred during the course of our experi- ments likely contributed to the low pollen densities we observed. A single rainfall event can remove up to 86% of the corn pollen from a milkweed leaf (Pleasants et al. 2001). The fact that rainfall events occurred and likely reduced the pollen loads on the milkweed leaves does not weaken the significance of our experiments. On the contrary, our results suggest that any potential adverse effect of Bt corn on monarch larvae can be substantially mitigated under field conditions. Our results from the 2000 study indicate that the rate of larval mortality is lower (i.e., larger Weibull b parameter) on milkweed at the edge of a Bt cornfield compared to milkweed farther from the field. The sur- vival of larvae at different locations relative to the cornfield edge (e.g., on the field edge, within the field or outside the field) has been variable among studies. Lower survival was found at the edges of corn fields in the Iowa II and New York studies of Stanley-Horn et al. (2001) compared to milkweed at various locations within and outside cornfields. Oberhauser et al. (2001) reported higher larval survival (i.e., larger Weibull b values) on milkweed in cornfields compared to milk- weed on cornfield edges. In contrast, in the Iowa I study of Stanley-Horn et al. (2001), higher larval sur- vival was found on milkweed at cornfield edges com- pared to milkweed within cornfields. The observed increase of larval mortality rate (i.e., decrease in Weibull b parameter) and apparent shift of mortality to a less-strong type HI survivorship curve (i.e., increase in Weibull c parameter), due to not counting larvae that have moved off of experimental plants, may lead to biased estimates of mortality. Monarch larvae will spend up to 17.5% of daylight hours off of milkweed plants (Rawlins & Lederhouse 98 1981). Individual monarch larvae have been difficult to follow for more than one week (Borkin 1982). Larvae, as early as second instars (Borkin 1982), leave what seem to be suitable host plants for no apparent reason (Borkin 1982, Urquhart 1960). We observed that most larval movement from potted milkweed plants began by 7 days post infestation, which was about the third instar and onward. Larvae that were observed off of the potted milkweed plants in this study were pre- dominantly in the third or later instars. Researchers conducting survivorship studies in the field must be aware of the potential for monarch larvae to leave host plants. Biased underestimates of larval survival will re- sult from counting larvae that have disappeared from the host plants as mortality. Conversely, monarch larval movement may also confound results when survivor- ship studies are conducted in small field cages. Small field cages could preclude normal larval movement from plants and thereby result in overestimates of sur- vivorship compared to open field studies. ACKNOWLEDGEMENTS We thank K. Oberhauser for providing monarch larvae, R. Moon and G. Oehlert for helpful suggestions, and M. Zalucki, T. Shelton, F. Gould, and R. Hellmich for reviewing an earlier draft of this man- uscript. We also thank E. Burkness, P. K. O'Rourke, S. Wold, A. Genetzky, E. Rye, and K. Bennett for assistance in the field. This re- search was supported by the Minnesota Department of Agriculture, Biological Control Program and the University of Minnesota Agri- cultural Experiment Station, University of Minnesota. LITERATURE CITED Baker, D. B. 1983. Climate of Minnesota: part X1V—wind clima- tology and wind power. Technical Bulletin. Agricultural Exper- iment Station, University of Minnesota. 48 pp. BHowmik, P. C. 1994. Biology and control of common milkweed. Rev. Weed Sci. 6:227-250. BorKIN, S.S. 1982. Notes on the shifting distribution patterns and survival of immature Danaus plexippus (Lepidoptera: Danaidae) on the food plant Asclepias syriaca. Great Lakes En- tomol. 15:199-206. BURKNESS, E.. C., W. D. HUTCHISON, P. C. BOLIN, D. W. BARTELS, D. F. WarNocK & D. W. Davis. 2001. Field efficacy of sweet corm hybrids expressing a Bacillus thuringiensis toxin for management of Ostrinia nubilalis (Lepidoptera: Crambidae) and Helicoverpa zea (Lepidoptera: Noctuidae). J. Econ. Entomol. 94:197-203. ENVIRONMENTAL PROTECTION AGENCY. 1998. Guidelines for eco- logical risk assessment. Federal Register 63:26846-26924. FEDERICI, B. A. 1993. Insecticidal bacterial proteins identify the midgut epithelium as a source of novel target sites for insect control. Arch. Insect Biochem. Physiol. 22:357—371. HELLMICH, R. L., B. D. SIEGFRIED, M. K. SEARS, D. E. STANELY- Horn, H. R. Martina, T. SPENCER, K. D. BIDNE, M. J. DANIELS & L. C. Lewis. 2001. 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USA 98:11913-11918. OEFHLERT, G. W. 2000. A first course in design and analysis of ex- periments. W. H. Freeman and Company, New York. 659 pp- OsTLIk, K. R., W. D. HuTcHison & R. L. HELLMICH. 1997. Bt com and the European corn borer. University of Minnesota, St. Paul, Minnesota. 16 pp. PILCHER, C. D., M. E. Rick, J. J. OBRyYcKI & L. C. Lewis. 1997. Field and laboratory evaluation of transgenic Bacillus thuringiensis corm on secondary lepidopteran pests (Lepi- doptera: Noctuidae). Econ. Entomol. 90:669-678. PINDER, J. E., J. G. WEINER & M. H. SmitH. 1978. The Weibull distribution: a new method of summarizing survivorship data. Ecology 59:175-179. PLEASANTS, J. M., R. L. HELLMICH, G. P. DivELy, M. K. Sears, D. E. STANLEY-Horn, H. R. MATTIa, J. E. FosTer, P. CLARK & G. D. JoNEs. 2001. Corn pollen deposition on milkweeds in and near cornfields. Proc. Nat. Acad. Sci. USA 98:11919-11924. RAWLINS, J. E. & R. C. LEDERHOUSE. 1981. 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USA 98:11931-11936. Urqunart, F. A. 1960. The monarch butterfly. University of Toronto Press, Toronto. 361 pp. WAGNER, D. L., J. W. PEACOCK, J. L. CARTER & S. E. TALLEY. 1996. Field assessment of Bacillus thuringeinsis on nontarget Lepi- doptera. Environ. Entomol. 25:1444-1454. WOLFENBARGER, L. L. & P. R. PHIFER. 2000. The ecological risks and benefits of genetically engineered plants. Science 290:2088-2093. Waraicut, C. L., A. R. ZANGERL, M. J. CARROLL & M. R. BEREN- BAUM. 2000. Absence of toxicity of Bacillus thuringiensis pollen to black swallowtails under field conditions. Proc. Nat. Acad. Sci. USA 97:7700-7703. ZALUCKI, M. P. 1981. Temporal and spatial variation of parasitism in Danaus plexippus L. (Lepidoptera: Nymphalidae: Danainae). Aust. Entomol. Mag. 8:3-8. ZALuCKI, M. P. & L. P. Brower. 1992. Survival of first instar lar- vae of Danaus plexippus L. in relation to cardiac glycoside and latex content of Asclepias humistrata. Chemoecology 3:81-93. ZALUCKI, M. P., L. P. BROWER & A. ALONSO-M. 2001. Detrimen- tal effects of latex and cardiac glycosides on survival and growth of first instar Danaus plexippus feeding on the sandhill milkweed Asclepias humistrata. Ecol. Entomol. 26:212-224. ZALucKI, M. P., A. R. CLARKE & S. B. MALCoM. 2002. Ecology and VOLUME 57, NUMBER 2 behaviour of first instar larval Lepidoptera. Annu. Rev. Ento- mol. 47:361-393. ZALUCKI, M. P. & R. L. Kircuinc. 1982. Temporal and spatial vari- ation of mortality in field populations of Danaus plexippus L. and D. chrysippus L. larvae (Lepidoptera: Nymphalidae). Oe- cologia 53:201—207. ZANGERL, A. R., D. MCKENNA, C. L. WRAIGHT, M. CARROLL, P. FI- 99 CARELLO, R. WARNER & M. R. BERENBAUM. 2001. Effects of exposure to event 176 Bacillus thuringiensis corn pollen on monarch and black swallowtail caterpillars under field condi- tions. Proc. Nat. Acad. Sci. USA 98:11908-11912. Received for publication 21 April 2002; revised and accepted 26 Sep- tember 2002. Journal of the Lepidopterists’ Society 57(2), 2003, 100-106 DESCRIPTION OF A NEW GENUS FOR “EUPTYCHIA” PECULIARIS (NYMPHALIDAE: SATYRINAE): IMMATURE STAGES AND SYSTEMATIC POSITION ANDRE VICTOR LUCCI FREITAS Museu de Hist6ria Natural and Departamento de Zoologia, Instituto de Biologia, Universidade Estadual de Campinas, CP 6109, Campinas, Sao Paulo 13083-970, Brazil. Email: baku@unicamp.br ABSTRACT. Based on distinct character states in life history and adult morphology, the monotypic genus, Taydebis, new genus with “Eu- ptychia” peculiaris Butler as the type species is described. Analysis of the morphological characters and comparisons with four nearby genera suggest that the genus is closely aligned to or should be placed near Taygetis Hiibner and Pseudodebis Forster. Additional key words: _ life history, Poaceae, Pseudodebis, Taydebis, Taygetis. Within the Neotropical Nymphalidae, the subfamily Satyrinae is one of the most poorly understood groups, with many systematic problems and undescribed species, a fact often noted in the literature (Forster 1964, Miller 1968, DeVries 1987:257, Freitas 2002). Adult characters have been useful for understanding relationships in some cases (Forster 1964, Miller 1968), but have been insufficient to resolve some sys- tematic problems in the subfamily. Since Miiller (1886) early stages have been shown as a useful source of characters in butterfly systematics (Kitching 1985, Brown & Freitas 1994, Freitas et al. 1997, Penz 1999) including for the Satyrinae (Singer et al. 1983, DeVries et al. 1985, Freitas 2002, Freitas et al. 2002). “Euptychia” peculiaris Butler 1874 is a problem species from southeastern Brazil. This species occurs at moderate elevations (SO0-1700 m) and is known from only a few localities along the Mantiqueira moun- tains and the Serra do Mar in the states of SAo Paulo and Santa Catarina (Campo Alegre and Lages) (see list below). The record of Hayward (1973:256) from Mi- siones, Argentina, requires further confirmation. The present paper illustrates and describes the criti- cal morphological characters that distinguish this taxon, such as the wing venation and male genitalia. For the first time, the early stages are illustrated and described in detail. A comparative discussion of sys- tematic relationships of “E.” peculiaris within the Satyrinae is presented and a new genus, Taydebis, is described. MATERIALS AND METHODS Adults and immatures of “E.” peculiaris were stud- ied at six different localities in SAo Paulo State, SE Brazil: banks of the Rio Tieté (Mogi das Cruzes, 700-800 m), Morro Grande Forest Reserve (Cotia, 850-950 m), Nucleo Santa Virginia (Sao Luis do Paraitinga, 900-1100 m), Campos do Jordao State Park (Campos do Jordao, 1500-1700 m), Intervales Park (Capao Bonito, 900-1100 m) and Grota Funda Municipal Park (Atibaia, 900-1000 m). Fertile eggs were obtained from wild-captured fe- males that were confined in plastic bags. Larvae were reared in plastic containers cleaned daily, with fresh plant material provided every two or three days (fol- lowing Freitas 1991). Observations and data were recorded on behavior and development times for all stages. Dry head capsules and pupal castings were re- tained in small glass vials. When there was sufficient material, immatures were fixed in Kahle solution (AVLF collection). All measurements were made using a microscope fitted with a calibrated micrometric ocu- lar. Egg size is presented as length and diameter, and head capsule size is the distance between the most ex- ternal ocelli (as in Freitas 1991). Taxonomic nomen- clature follows Miller (1968) as modified by Harvey (1991), who treated the group as a subfamily, down- ranking Miller’s subfamilies and tribes to tribes and subtribes, respectively. Nomenclature of wing veins follows Miller (1969), and of body setae follows Hin- ton (1946). Taydebis Freitas, new genus (Figs. 1, 2, Table 1) Type species: Euptychia peculiaris Butler, 1874. Diagnosis. Eyes hairy, reddish brown. Labial pal- pus one and a half times as long as head, brown with light brown hairs. Antenna (8.5-9.5 mm) up to 0.4 times the length of the costa; shaft dark brown dor- sally, orange brown ventrally, sparse scaled dorsally; club not conspicuously developed, including eleven segments, with apical portion (last five segments) dark brown. Wing venation very similar to Pseudodebis and Taygetis (Fig. 2). Both wings extremely rounded api- cally (Figs. 1, 2). Description of adults. Male. Forewing length 20-23 mm, hindwing length 16-19 mm (n = 15). Body dark brown, abdomen ventrally light brown. Upperside ground color of wings medium brown, without marks, except for a dark brown zigzag sub marginal line on both wings, and a light marginal line on the hindwing. Un- derside ground color lighter brown, three-tone: forewing discal area darker, hindwing distal half lighter. Two prominent scalloped brown lines crossing both wings 35% and 60% out from base; sub marginal region of forewing with a diffuse darker brown area with four VOLUME 57, NUMBER 2 101 Fic 1. Adult male (top) and female (bottom) of Taydebis peculiaris from Parque Estadual de Campos do Jordao, SP. minute light blue centered black ocelli bordered with orange in spaces R5-M1, M1-M2, M2—M3 and M3-Cul; sub marginal area of hindwing with two prominent light blue centered black ocelli with orange margins in spaces Rs-M1 and M1-M2, minute similar ocelli in spaces M2—M3 and M3-Cul, somewhat larger in Cul—Cu2 and Cu2-1A. A dark brown zigzag sub marginal line and a light marginal line are present on both wings. Male genitalia (Fig. 2) with an elon- gated saccus, well developed tegumen and long pointed uncus. The gnathos appears as two long pointed processes. Valvae trapezoidal ending with a single well developed point. Aedeagus with one large cornutus. Additional morphological characters (legs and labial pal- pus) are shown in Fig. 2. Female. Forewing length 22-24 mm, hindwing length 17-22 mm (n = 6). Body dark brown, ventral abdomen light brown. Gen- eral color and patter very similar to but in general lighter than that of males. Wings more rounded than in males. Variation. Variation in the dorsal wing surfaces is very low, with most variation being recorded on the underside. The size of the ocelli is variable in both sexes, and in some individuals only the two prominent ocelli of the hindwing can be seen without magnification. The wing patter is also variable, being weakly marked in some few individuals from Campos do Jord&o. Some females have the under- side ground color much more yellowish, especially in the sub mar- ginal and anal areas. R3 R4 R2 R5 R1 sc M1 A " M3 Cul Cu2 2A Sc+R1 Rs h M1 M2 M3 2A Cut Cu2 1A i> JOURNAL OF THE LEPIDOPTERISTS SOCIETY 4 ) Fic 2. Morphological characters of Taydebis peculiaris. A, Male wing venation—hindwing above and forewing below; B, Male midleg; C, Female foreleg: D, Male foreleg; E, Male labial palpus; F, Lateral view of male genitalia; G, Aedeagus in lateral view. Description of early stages. The following descriptions are based on immatures reared from Santa Virginia and Morro Grande. The typical features of the immatures are very similar at these two sites. However, the number of instars was variable with four instars in Morro Grande (Table 2) and five in Santa Virginia (see discussion). Females laid individual eggs when confined in plastic bags, suggest- ing the oviposition of isolated eggs as the usual situation in nature. Egg. Spherical, light green, without visible ridges or marks under the optic microscope. Height and diameter 0.92 mm (n = 30). Du- ration: 5-7 days. First instar (Figs. 3a, b, 4). Head capsule black, with enlarged chalazae, bearing a pair of short scoli on vertex, each with two long narrow setae ending into a fine point. Third stemma larger than the other stemmata. Head capsule width 0.66-0.72 mm (mean = 0.69 mm, SD = 0.017, n = 30): scoli 0.08—0.12 mm (mean = 0.11 mm, SD = 0.013, n = 30). Body beige, becoming light green after feeding, smooth, with many weak sranntie longitudinal stripes and a pair of short caudal filaments. Setae XD, D, SD and L thickened with clubbed tips; body chaetotaxy illustrated in Fig. 4. Maximum length 5 mm. Duration: 8-9 days. Second instar 8 3c). Head black with two div erging scoli on vertex. Head capsule width 0.90-1.02 mm (mean = 0.97 mm, SD = 0.039, n = 26): scoli 0.28—0.40 mm (mean = 0.34 mm, SD = 0.031, n = 26). Body slender, light green with many longitudinal white stripes; caudal filaments short. Maximum length 10 mm. Duration: 7-8 days. Third instar. Head black with green front and two short black diverging scoli on the vertex. Head capsule width 1.20-1.42 mm (mean = 1.32 mm, SD = 0.053, n = 18); scoli 0.40-0.56 mm (mean = 0.46 mm, SD = 0.050, n = 18). Body dark green with many longitu- dinal yellow stripes; caudal filaments short. Maximum length 16 mm. Duration: 6-8 days. Fourth instar. Head green with a pair of short scoli with red tips. Head capsule width 1.70-1.90 mm (mean = 1.78 mm, SD = 0.055, n = 17); scoli 0.58—0.74 mm (mean = 0.63 mm, SD = 0.044, n = 17). Body emerald green, with many longitudinal thin yellow and light green stripes; caudal filaments short. Maximum length 22 mm. Duration: 9-10 days. Fifth (last) Instar (Fig. 3d—f). Head the same as in previous in- star. Head capsule width 2.45-2.75 mm (mean = 2.54 mm, SD = 0.090, n = 10); scoli 0.75-—0.88 mm (mean = 0.82 mm, SD = 0.039, n = 10). Body color same as fourth instar. Maximum length 33 mm. Duration: 9-10 days. Pupa (Fig. 3g—h). Entirely green, elongated, smooth, with short ocular caps and slightly projecting alar caps bordered with a thin yel- low line. Total length 12-14 mm. Duration 10 days (n = 7). Etymology. The name is a reduced combination of Taygetis and Pseudodebis, possibly the two most closely related genera. 103 2 ea a ea 2 =) Z, t 19 is2} 2 =) = e) > viyV—8 ‘dS ‘oepro[ op sod st xt 19 Sat Sa Il €1 poounouo.d you peounouoad jou poounouoad jou poounouo.d jou poounouo.d jou poounouoad jou poounouord yoounouoad AYysys I BYST poounouord y9ounoUOAd ANYsIs I pYys!| poounouord poounouord poounouord ) Lh ‘qs ‘seurduey, papuNoI 9104s PEI ROU poyutod ‘payesuoje poyutod ‘payesuoye 9} uIOd ‘poyesuoyo SOC I poepunog ‘j.10ys poyutod ‘payesuoye poyutod ‘payesuoye poqutod ‘payesuoje pozutod ‘peye3uoja prorq oys JOpUNOL ‘j.10Ys Perl 10 poyutod ‘payesuoye poqutod ‘poysuoje prorq jysrexys Jopues ‘poamo Iopusys jysre.ys proiq yYsre.ys Jopusys ‘WYyste.4s Japuays jyste.ys Japueys Fyste.ys Jopusys WY sTe.s prorg ‘WYsreys Jopus]s “peamo Iopusys ‘poeamoa peorq ‘peamo Lopusys ‘WYyste.s Ppayiyysue] UUM], OVI SNOOVS ‘ sOyeUsS Jo odeyg snoun jo odeyg ‘RIOUSS Poel YI sigaphyy, yo suosuredu0D GI 8 VI 9 Ol Ol Gl GG €L il SI €6 al WPMypsus] ‘snsvopoy puo yeastp oy} FW peamo yystwe.ys Wysre.ys yYysmw.ys Wysre.ys PYysre.1ys Wysre.ys ysrerys Ysre.1ys Wysre.ys yysrens poamo A]3u0.1)s Wysre.ys AARA\ AAR AA AACA AAR A\ KAVA AAG AAG Aaeas A YSIS Aneas Ap ysis Aaea ApYSiys AARM JOU snsropor jo odeys UIS.ICUI SUIMpUIT] ‘T ATAVL, pepunor pepunor popuno. aynor a}eoun.y aor aor a}eoun.9 anor poepunol pepunor pep unLOL pepu nol xode SUIMOLO ‘dS ‘Oqriq Op OLL0JN—6 “LIN “PIS9L0] oT 9 ‘dS ‘wee[—sg ‘oy ‘osmyeumeyy, Pysorwy—F ‘oa ‘soreyury—e ‘Oy ‘na1e[—zZ ‘s0ded smpj—y «(pizerg ur sonreooy [[e) feLteyeur Jo vo.M0G Amey Auey sey asieds Moy Auey Arey Arey uon.tod peta] ut suey osaeds ytoys AOA Arey Aney Auey Arey rey asieds moj Aurey SoA wash "y, gpDajauad syashvysog sppunjg visalioyy Y Sh Ona syashny, ppiaghs syashyy, 7 Sh. ,DYao syanhny Diausau syaohy puanydh syashny, sayon) syashny, 50 in Lytrosis; there are oblique (black) lines on AT (below spiracle) and A8 (above and below spiracle) that are more apparent in Lytrosis than any of the Eu- chlaena that we have examined (six eastern species); the D setae are approximately one-half of the spiracu- lar height (in Ewchlaena the D setae are subequal to the spiracular height); and lastly, both the paraprocts and anal proleg are proportionately larger in Lytrosis. The close pairing of the two SD setae and the proxi- mate grouping of the L setae on TI also may be diag- nostic for Lytrosis. This condition does not occur in Euchlaena marginaria (Minot) (McGuffin 1981) or Euchlaena serrata (Drury) (DLW specimens). Rindge (1971) noted that adults of Lytrosis permag- naria possessed the most primitive features of any of the four members in the genus: i.e., the male has a metatibial hairpencil and the vesica has separate spines that are exerted on the right, anterior to the apex of the aedeagus. The unremarkable larval mor- phology of L. permagnaria supports Rindge’s posi- tion—its body is unwarted and more closely resembles that of a Euchlaena than either L. unitaria or L. sinu- osa (Figs. 21-26). Given Lytrosis permagnarias overall scarcity in the East, we are puzzled by its abundance at Goshen, Vir- ginia. Nothing impresses us as exceptional about the locality and indeed we probably would have passed on blacklighting at the site, had we not known that L. per- magnaria had been collected along the road in previ- ous years. The Goshen colony strikes us as undistin- guished botanically; woody plants growing in the vicinity of our sheets and traps include Acer rubrum, Amelanchier sp., Carya sp., Cornus sp., Nyssa sylvat- ica, Platanus occidentalis, Quercus rubra, Quercus alba, Sassafras albidum, and Tsuga canadensis. Both Lytrosis unitaria and L. sinousa fly with L. permag- naria at Goshen during early June. J. R. Heitzman (pers. com.) informs us that all four Lytrosis species may fly sympatrically in the Ozarks. Lytrosis permagnaria has been reported to be lo- cally common in northeastern Georgia by James Adams and in Cheaha State Park, Alabama by Tim McCabe. Both of these localities and Goshen are low elevation or foothill Appalachian forests—to the best of our understanding, no unusual plant is common to JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY the three sites. In captivity L. permagnaria larvae ac- cepted Quercus alba, Q. ilicifolia, Q. rubra, and a Carya species. Survivorship was higher on Querucs, perhaps because picked oak foliage holds up longer. Lytrosis unitaria, the best known member of the genus, has been recorded from Acer, Amelanchier, Crataegus, Pinus strobus, Prunus, Quercus, Rosa, and Vaccinium (McGuffin 1981, Wagner et al. 2002, DLW unpublished data). Wild hosts are unknown for Lytro- sis sinuosa, but captive individuals have been reared from both Acer negundo and Quercus (Wagner et al. 2002). Host data for the related, and more well studied genus, Euchlaena, indicate that its members are widely polyphagous on woody plants (McGuffin 1981, Handfield 1999, Wagner et al. 2002). It seems unlikely that the moth’s scarcity will be explained by an unusual host association—we leave it to others to discover why such a widespread, unspecialized feeder, remains one of the East’s rarest moths. ACKNOWLEDGMENTS The late Douglas Ferguson provided many helpful comments on a previous draft, encouragement, and companionship at Goshen. John Richard Heitzman shared several useful observa- tions on Lytrosis species in Missouri. Shawn Kennedy prepared the larval habitus, pupal drawings, and assisted with the prepara- tion of the figures; Virge Kask the photographic plate. Dale Schweitzer kindly sent us the livestock of Lytrosis sinuosa. The fig- ured Lytrosis unitaria caterpillar was collected by Keith Hartan and photographed by Valerie Giles. Support for this paper came from the U.S. Department of Agriculture, Forest Service, Forest Health Technology Enterprise Team, Cooperative Agreements 01- CA-11244225-215. LITERATURE CITED FEeRGuSON, D. C. 1983. Geometridae, pp. 88-107. In Hodges, R. W. et al. (ed.), Check list of the Lepidoptera of America north of Mexico. E. W. Classey Ltd. and The Wedge Entomolgical Research Foundation. Cambridge Univ. Press, Cambridge, United Kingdom. Fores, W. T. M. 1948. The Lepidoptera of New York and neigh- boring states. II. Geometridae, Sphingidae, Notodontidae, Ly- mantriidae. Memoir 274. Comell University Agricultural Ex- periment Station, Ithaca, New York. 263 pp. HANDFIELD, L. 1999. Le guide des papillons du Québec. Broquet Inc., Boucherville, Quebec, Canada, 982 pp. McGuFrrFin, W. GC. 1981. Guide to the Geometridae of Canada (Lepidoptera). III. Subfamily Ennominae, 3. Memoirs of the Entomological Society of Canada No. 117. 153 pp. RINDGE, F. 1971. A revision of the moth genus Lytrosis ( Lepidoptera, Geometridae). American Museum Novitates 2474. 21 pp. WAGNER, D. L., D. C. FERGUSON, T. L. MCCABE & R. C. REARDON. 2002. Geometroid caterpillars of northeastern and Appalachian forests. USFS Technology Transfer Bulletin, FHTET-2001-10. 239 pp. Received for publication 3 August 2002; revised and accepted 19 De- cember 2002. Journal of the Lepidopterists’ Society 57(2), 2003, 113-120 FIRST RECORD OF LARVAL ENDOPHAGY IN EULIINI (TORTRICIDAE);: A NEW SPECIES OF SETICOSTA FROM COSTA RICA JoHN W. BROWN Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, ‘/o National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0168, USA. E-mail: jbrown@sel.barc.usda.gov AND KENjI NISHIDA Escuela de Biologia, Universidad de Costa Rica, 2060 San José, Costa Rica. E-mail: knishida@cariari.ucr.ac.cr ABSTRACT. Seticosta rubicola, new species, is described and illustrated from Costa Rica. The species is assigned provisionally to Seti- costa on the basis of superficial similarities to other species in the genus (e.g., forewing length and pattern, long antennal cilia in the male, and extremely elongate labial palpi in both sexes), as well as features of the genitalia (e.g., “trifurcate” uncus). The absence of long, strong setae from the costa of the valva in the male genitalia is the only character contradicting this placement, and the setae are assumed to be lost secondarily. Seticosta is assigned to Euliini on the basis of the shared possession of a uniquely derived foreleg hairpencil in the male. Larvae of the new species are endophagous feeders in the stems of Rubus spp. (Rosaceae), which represents the first reported case of gall-inducing or stem-boring in Kuliini. Larvae also have been reported as borers in the fruit of Rubus. The early stages of S. rubicola, the first reported for the genus, are de- scribed and illustrated. They are unusual in the possession of several features more characteristic of Olethreutinae than of Tortricinae. The species has been recognized as a pest of quarantine significance by the Ministerio de Agricultura, Costa Rica. RESUMEN. Una nueva especie de mariposa nocturna, Seticosta rubicola, ha sido descrita e ilustrada desde Costa Rica. La especie ha sido asignada provisionalmente al género Seticosta, basandose en similitudes superficiales con otras especies del género, asi como en caracteres de la genitalia. Algunas de estas similitudes son: longitud y el patrén de colores de las alas anteriores, cilio antenal largo en machos, palpos labiales extremadamente largos en ambos sexos, y presencia de un penacho de pelos en las patas anteriores de los machos. La ausencia de setas gruesas en la costa de la valva en la genitalia del macho es la tinica caracteristica que contradice esta ubicacién taxondémica, y se asume que las setas se perdieron secundariamente. El género Seticosta esta asignado dentro de Euliini basandose en la presencia de un penacho de pelos distintivo en las patas delanteras del macho. Las larvas de esta nueva especie son formadoras de agallas en los tallos de especies de Mora (Rubus; Rosaceae). Los estadios tempranos de la Seticosta rubicola, primer registro para el género, son descritos e ilustrados. Estos son inusuales debido a la pos- esion de varios caracteres que son distintivos de Olethreutinae mas que de Tortricinae. Additional key words: Neotropical, systematics, Rubus spp., life history, pest species, Seticosta rubicola, taxonomy, parasitoid, Bassus new species, biological control. Although endophagous feeding and gall-induction is stages, and comment briefly on its unusual life his- not unusual in the subfamily Olethreutine (Tortrici- tory. dae), it is relatively rare in Tortricinae, where it is re- stricted primarily to the tribe Cochylini. Hence, it is AUATIBRUALS ARID BME TEIODS fairly surprising that during investigations on gall- Adults were borrowed from or examined at the fol- inducing Lepidoptera in Costa Rica, the second author lowing institutions: Instituto Nacional de Biodiversi- discovered several species of Tortricinae causing galls dad, Santo Domingo de Heredia, Costa Rica (INBio); in Rubus species (Rosaceae). One in particular, an un- Essig Museum of Entomology, University of Califor- described species provisionally assigned to Seticosta, is nia, Berkeley, California, U.S.A. (UCB); Museo de In- especially unusual in its larval chaetotaxy and other sectos, Escuela de Biologia, Universidad de Costa features. Larvae identical to these, and assumed to be Rica, San José (UCR); National Museum of Natural conspecific, also have been intercepted by the U.S. History, Smithsonian Institution, Washington, D.C., Department of Agriculture’s Plant and Animal Health U.S.A. (USNM); and Vitor Becker personal collection, Inspection Service at U.S. ports-of-entry within the fruit Planaltina, Brazil (VBC). Dissection methodology fol- of Rubus spp. from Guatemala. In addition, this species lows that summarized in Brown and Powell (1991). II- recently was identified as a pest of quarantine signifi- lustrations of genitalia are photographs of slide mounts cance by the Ministerio de Agricultura, Costa Rica. taken with a SONY DKC5000® digital camera and en- Food plants of the tribe Euliini, to which Seticosta hanced using Adobe Photoshop® and Adobe Illustra- belongs, were reviewed by Brown and Passoa (1998), tor® software. Forewing measurements were made who identified no previously recorded endophagous- with the aid of an ocular micrometer mounted in a feeding species in the tribe. We take this opportunity Wild M3Z dissecting microscope under low power to describe and illustrate this new species from Costa (x10-16). Terminology for wing venation and genitalic Rica, present details on the morphology of the early structures follows Horak (1984); terminology for larval 114 features follows Brown (1987). Abbreviations and sym- bols are as follows: FW = forewing; HW = hindwing; DC = discal cell; n = number of specimens examined; X = mean; ca. = circa (approximately); Est. = Estacion; r.f. = reared from. Larvae were obtained primarily during field work conducted between February 2000 and June 2001, along dirt trails near La Georgina, Villa Mills (3000-3100 m) and Estacién Biolégica Cerro de la Muerte (3050-3100 m) at Cerro de la Muerte, Cartago and San José provinces, Costa Rica. The veg- etation of the region is referred to as Tropical Montane Cloud Forest. During the dry season, which lasts from December/January through April, rain is infrequent, although humidity remains high, and dense fog is com- mon in the afternoons. During the wet season, which lasts from April through November/December, heavy rains are common; average annual rainfall is 2812 mm. Daily average temperature is 10.9°C, but tempera- tures can be as low as —3°C during the dry season (Kappelle 1996). During field work, individuals of various species of Rubus were examined for galls (e.g., larval frass and swollen parts of stems). When discovered in the field, some larvae, along with their galls and additional freshly-cut stems of the food plant, were placed in plastic bags and taken to the laboratory where they were either stored in an air-conditioned room (approx- imately 16-18°C) at Museo de Los Insectos, Universi- dad de Costa Rica, in San Pedro (1150 m), or placed in a refrigerator (6.2°C) and removed and kept at ambi- ent temperature (approximately 23°C) for 8 hours each day. Other active galls were reared under field conditions in plastic bags at the station, where temper- atures ranged from 11—18°C during the day and 1-3°C at night. Examples of galls, larvae, and pupae were preserved in 75% EtOH and are deposited in the USNM and UCR. Adult specimens were pinned, and pupal shells were saved in gelatin capsules pinned along with the adult moths. Parasitoids were submitted to Michael Sharkey for identification. SYSTEMATICS Seticosta rubicola Brown & Nishida, new species (Figs. 1-8) Diagnosis. In forewing pattern, S. rubicola is ex- tremely similar to many other species of Seticosta, in- cluding S. aeolozona (Meyrick) (Clarke 1958), S. arachnogramma (Meyrick) (Clarke 1958), S. tridens Razowski, and S. tambomachaya Razowski. These JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY species typically have a somewhat uniform tan, brown- ish, or reddish ground color divided by a light subter- minal fascia paralleling the termen, and a similarly col- ored diagonal subbasal fascia, extending outward from the costa. Additional external features that S. rubicola shares with other Seticosta are the extremely elongate labial palpi in both sexes, the long antennal cilia of the male, and the male foreleg hairpencil (presumably lost secondarily in related genera such as Anopinella Pow- ell, Strophotina Brown, and Punctapinella Brown). The male of S. rubicola lacks the dense patch of strong setae from the costa of the valva characteristic of all other species of Seticosta, and it is assumed that this character is lost secondarily. The male of the new species possesses a pair of lateral processes near the distal end of the uncus giving it a trifurcate appear- ance, which appears to define a species group within Seticosta that includes S. arachnogramma (Clarke 1958), S. tridens (Razowski 1988), S. cerussograpta Razowski, and one or more undescribed species; this character state is less developed or absent in other species such as S. homosacta (Meyrick) and S. sagmat- ica (Meyrick). Description. Adult. Male (Fig. 3). Head: Frons smooth-scaled, pale cream; vertex slightly roughened, pale cream; labial palpus ex- tremely elongate, all segments combined ca. 3 times horizontal di- ameter of compound eye, pale cream on inner surface, pale cream scales tipped with brown on outer surface; antennal cilia 4—5 times width of flagellomere. Proboscis present, presumably functional. Thorax: Forewing length 8.0-11.6 mm (x = 9.9; n = 11); ground color brick red, with diffuse area of darker scaling near middle of wing; costa with short, irregular, transverse, white and brown striae; a white fascia parallel to termen, overscaled with yellow-green; a second white fascia with yellow-green overscaling extending out- ward from costa ca. 0.2 distance from base to apex; a small blotch of white with yellow-green overscaling at lower half of base of FW; aforementioned fasciae and basal blotch connected by narrow line along lower edge of FW; fringe brick red. Underside grayish. Hind- wing white, with faint, pale gray mottling. Abdomen: Somewhat shiny cream white; an indistinct brownish dot near mid-venter of A3-7. Genitalia as in Fig. 1 (photograph of JWBrown slide 1260; 5 preparations examined). Uncus bearing a pair of subdistal pointed processes, giving a trifurcate appearance; socii moderately short, digitate, sparsely setose; gnathos weak, broadly u-shaped, without conspicuous terminal process at junction of arms; transtilla moder- ately large, slightly narrowed and sclerotized near middle, where it bears microtrichiae; valva thick, somewhat swollen, weakly lanceo- late, with rounded apex, sacculus weakly developed, confined to basal one-third of valva, cuculluslike region of dense, large setae in ventral half beyond sacculus and in apical region, costa with basal ex- cavation bearing tiny setae. Aedeagus moderately small, curved, at- tenuate distally, with rounded phallobase and protruding lobe at ductus ejaculatoris; cormnuti absent. Female. Head, Thorax: Essentially as described for male, except antennal cilia unmodified (inconspicuous). Forewing length 9.2-11.1 mm (x = 10.5; n = 8). Abdomen: Essentially as described for male. Genitalia as in Fig. 2 (photograph of UCB slide 2516; 12 preparations examined). Papillae anales slender; apophyses anteri- ores and posteriores elongate, posteriores ca. 1.2 longer than anteri- ores; sterigma slightly variable, either totally unsclerotized or with posterior edge bearing a pair of weak subventral sclerotizations; os- tium extremely simple, not surrounded by sclerotization; ductus VOLUME 57, NUMBER 2, 115 Fics. 1-3. Setisota rubicola. 1, Adult male; 2, Male genitalia, valvae spread, aedeagus removed; 3, Female genitalia. bursae moderately long, frail, slender at ostium, gradually widening anteriorly, with ductus of accessory bursae originating ca. 0.1 dis- tance from ostium to junction with corpus bursae; corpus bursae somewhat rounded-triangular, with ductus seminalis originating in posterior third of corpus; corpus bursae with dense, extremely minute spinules throughout. Larva (Figs. 4-8). Based on two fourth instars and one third in- star collected 9 May 2001, two second instars collected 11 April 2000, one fifth instar collected 31 May 2001, and two fifth instars collected 20 July 2001, at Estacién Biologica Cerro de la Muerte, 3050 m, Provincia San Jose, Costa Rica, on Rubus vulcanicola (Donn. Sm.) Rybd. General: Length 12-13 mm (fifth instars only); head black (in early instars) to orange with conspicuous black genal and stemmatal patches (later instars); body maroon (paler in ma- ture larva of each stage), with moderately large, conspicuous, darker, brownish pinacula; prothoracic and abdominal shields brownish yellow to reddish brown, with pattern of pale brown specks; integument strongly granular; spiracles moderately large, rounded, those on T1 and A8 larger than others. Thorax: Protho- racic shield with broad, translucent region immediately anterad of line formed by XD1, XD2, and SD1; L-group trisetose on T1, with pinaculum irregularly oblong-round, situated mostly ventrad of spiracle, L1 roughly equidistant from L2 and L3; SV-group on T1-3 is 2:1:1; meso- and metathorax weakly annulate dorsally, both segments with first annulation bearing an extra SD1 seta (=MD1), an extra pair of L setae (=MSD1, MSD2), an extra SV seta 116 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY ioe Sd Fe oY ie hare OO eon b Fics. 4-8. Larval chaetotaxy of Seticosta rubicola. 4, Head and thorax, lateral view; 5, First and second abdominal segments, lateral view; 6, Fourth and fifth abdominal segments, lateral view; 7, Seventh, eighth, ninth, and tenth abdominal segments, lateral view; 8, Seventh, eighth, and ninth abdominal segments, ventral view, anterior end at top. VOLUME 57, NUMBER 2 (=MV1), and an extra V seta on smaller, less conspicuous pinacula. Abdomen: D1 pinacula usually with a deep notch at ventro- anterior margin, at least on A2—5 (sometimes on more segments); extra, tiny D seta (=MD1) and V seta (=MV3) situated near ante- rior edge of segments Al—9; SD1 located dorsad of spiracle on Al-7, with tiny SD2 remote, ventro-anterad, usually without pinaculum; L1 and L2 on same enlarged pinaculum on Al-8; SD1 anterad of spiracle on A8; D2 setae usually on common dorsal pinaculum on A8; D2 setae always on common dorsal pinaculum on AY; D1 and SD1 on common pinaculum on AQ; L-group trise- tose on AQ, usually with all setae on same pinaculum; SV-group on A1,2,7,8,9 is 2(3):3:2(3):2:1; V setae ca. 2 times farther apart on A9 than on A8; anal comb present, with 3-6 teeth; crochets in biordi- nal circle, 22—30 (in third and fourth instars) to 28-38 (in fifth in- stars) on prolegs on A3-6, 14-21 on A10 (extremely variable from instar to instar). Pupa (Fig. 9). Based on two preserved in alcohol and three exu- viae (one male, two females). Typically tortricine, fusiform, 7.5-8.5 mm in length, 2.1-2.3 mm in width. Head and thorax typical for the family, as described elsewhere (e.g., Horak 1998). Abdomen with Al lacking dorsal spines; A2 with double row of weak dorsal spines; A3-A8 with double row of sparse, strong dorsal spines; segment A9 with four large dorsal thorns. Posterior end of abdomen bluntly rounded; cremaster absent; A10 with a pair of posterolateral thorns; four long hooked setae on A10, two at posterior end, two posterolat- erad. Holotype. 3, Costa Rica, Cartago Province, Parque Nacional Tapanti, E] Guarco, San Isidro, Est. Esperanza, 2600 m, May 2001 (R. Delgado, INBio). Paratypes (29 d, 16 2). COSTA RICA: Cartago Province: 1 km NE Cerro Asuncion, Cerro de la Muerte, 3100 m, 2 Mar 1985, D. Janzen & W. Hallwachs (1 d, INBio). 7.5 km S Ojo de Agua, 9°15’N, 84°48’W, 2682 m, 16 Jun 1973, Erwin & Hevel (1 6, USNM). Villa Mills, 3100 m, 9 Jul 1993, E. Phillips (2 3, INBio), 3 Jul 1999, E. Phillips & J. Powell (1 2°, UCB). Rio Macho, Est. Ojo de Agua, 3000 m, 25-26 May 1997, B. Gamboa (2 °, INBio & USNM). Pension La Georgina, Cerro de la Muerte, S border Cartago Province, 3000 m, 23-25 May 1985, J. Powell & P. Opler (1d, 2 2°, UCB). Cerro de la Muerte, 3100 m, 17 Sep 1999 (3 d, 3 2), 1-2 Sep 2000 (2 4, 4 9, VBC), V. Becker. 7 km SE El Canon, 2500 m, 28 May 1985, black- light, J. Powell (1 2, UCB), 28 May 1985, ex-loose bark of live tree, J. Doyen (1 2, UCB). Parque Nacional Tapanti, El Guarco, San Isidro, Est. Esperanza, 2600 m, May 2001, R. Delgado (4 d, 1°, IN- Bio & USNM). El Guarco, Macizo de la Muerte, Sector de Esper- anza, 2600 m, Jun 2001 (1 2, INBio), Oct 2001, R. Delgado (1 4, IN- Bio). El] Guarco, Villa Mills-CATIE, 2840 m, 26-28 Oct 2000, R. Delgado (2 d, INBio). R.F. Rio Macho, E] Guarco, Macizo de la Muerte, Sector de Esperanza, 2600 m, Aug 2001, R. Delgado (1 ¢, INBio). Heredia Province: Est. Barva, Braulio Carrillo N. P., 2500 m, Nov 1989, G. Rivera (1 d, INBio), Nov 1989 (1 3, INBio), May 1990 (1 3, INBio), A. Fernandez. Mount Poas [2350 m], no date (1 9°, USNM), Wm. Schaus. Lim6n Province: Bratsi, Valle del Silencio, 2472 m, 11-12 Oct 2000, R. Delgado (1 d, 1 2, INBio). San José Province: San Gerardo de Dota, Cerro de la Muerte, 2430 m, 23 Aug 1981 (1 d, INBio), 23 Dec 1981, D. Janzen & W. Hallwachs (2 3, 2 2, INBio), 20 Feb 1996, D. & J. Powell (1 3, UCB), 5 Jul 1999, E. Phillips & J. Powell (2 6, UCB). Est. Cuerci, por Quebrada los Leones, 4.5-4.6 km E Villa Mills, 2500-2700 m, 21-26 Sep 1995 (1 3, INBio), 21-24 Oct 1995 (1 4, INBio), 25 Nov 1995 (1 3, INBio), 10-12 Oct 1996 (1 ¢, INBio), 7-10 Dec 1996, A. Picado (1 ¢, 2 2, INBio & USNM), 12-15 Jul 1996, B. Gamboa (2 9, INBio). Cerro de la Muerte, Villa Mills, 3000 m, 25 Mar 2000, r.f. Rubus braecipus, K. Nishida (1 2, UCR). Cerro de la Muerte, Est. Biologia Cerro de la Muerte, 3100 m, em: 4 Jul 2001, rf. Rubus vulcanicola, K. Nishida (1 2°, UCR). GUATEMALA: Volcan Santa Maria, [no date] (2 2, USNM), Schaus & Bames. Heu., Bulej, 2000 m, 15°27’N, 91°35’W, 25 Jul 2000 (1 3, VBC), V. Becker. Additional Specimen Examined. COSTA RICA: San José Province: Pérez Zeledén, 2260 m, em: May 2000, r.f. Rubus sp. (cul- tivo de mora), K. Nishida (1 2, UCR). Fic. 9. Pupa of Seticosta rubicola. 9, Venter (a), dorsum (b). Distribution. Seticosta rubicola is known primarily from the high elevations (2000-3100 m) in the central cordillera of Costa Rica, including the provinces of Cartago, Heredia, Limon, and San José. The majority of the specimens are from the vicinity of Cerro de la Muerte, a high elevation cloud forest. Based on a few specimens records cited above and larvae intercepted at U.S. ports-of-entry on Rubus sp., the species also occurs in Guatemala. Etymology. The specific epithet is derived from the association of the larvae with Rubus. Remarks. The larva and pupa of Seticosta rubicola are the first reported for the genus. At least three char- acters of the larva are more typical of Olethreutinae than Tortricinae: (1) the occurrence of SD1 and D1 on a shared pinaculum on AQ; (2) a bisetose SV-group on AT (although it was trisetose on one of eight larvae ex- amined); and (3) SD2 on a pinaculum separate from that of SD1 on Al-8. The widely separated V setae on AQ are unusual for Tortricinae as well, although this condition is present in almost all Sparganothini (MacKay 1962) and Cochylini. Other unusual features of the larva include the extra SD, L, SV, and V setae on the meso- and metathorax; the extra D and V setae on A1-8; the notched D2 pinacula of A2—5, characteristic 118 of the Cryptophlebia-Ecdytolopha group of genera (Olethreutinae: Grapholitini) (Adamski & Brown 2001); and the position of the L pinaculum on the pro- thorax, i.e., mostly ventrad of the spiracle. Based on previous studies on the early stages of Euliini, it ap- pears that both the “olethruetine” and “tortricine” con- ditions of SD1+D1 on A9 occur in this tribe, i.e., either on a shared (olethreutine condition) or on sep- arate pinacula (tortricinae condition). Both states are reported to occur in Proeulia Obraztsov and Anopina Obraztsov (Brown & Powell 2000). The pupa of Seti- costa rubicola, lacking a distinct cremaster and with fewer spines in the ventral rows, is also somewhat olethreutinelike and dissimilar to that of all other Eu- liini reported thus far (ie., Accuminulia Brown, Anopina Obraztsoyv, Chileulia Powell, Cuproxena Brown & Powell, and Dorithia Powell). BIOLOGY The eggs of S. rubicola are unknown. Larvae were discovered boring in stems of Rubus eriocarpus Liebm. and, more frequently, Rubus vulcanicola, in- ducing a fusiform gall (Fig. 10). The size of larva- containing late stage galls on the latter species is ca. 5-6 mm wide and ca. 12-15 mm long; the stem width at the base of the gall is ca. 3 mm. Galls often were sit- uated near or between nodes of young parts of the stems, with one to four galls per stem. A single larva was found per gall chamber. At the base of the gall there is an opening (>3 mm in diameter) from which the larva ejects frass, head capsules, and other debris (Fig. 10). Apparently this opening represents the point at which the larva enters the stem (Figs. 10-12); it is usually located at the base of a leaf petiole or a shoot axis, facing upward. The opening is characterized by a patch of larval frass and debris, including head cap- sules and bits of the plant tissue, all of which are at- tached by silk. The scraps of plant tissue are made by the larva excavating the stem and by larval regurgita- tion. Occasionally, some larval frass is retained within the gall chamber (Fig. 13). Within the galls of early in- stars there usuaily is a silk-lined shelter, woven with frass and bits of the plant tissue. Dissection of galls on Rubus vulcanicola revealed that the tissue surrounding the larval chamber is ap- parently parenchyma tissue. This tissue, upon which the larva feeds, is light green and consists of dense cells, resembling tissue in apical parts of the plants. In contrast, other parts of the stem were filled with white spongy tissue (Figs. 11, 13). The gall chamber was sur- rounded with irregular tissue (calluslike growth) or ir- regularly consumed tissue. The surface of the gall chamber has a brownish tint (Fig. 13) and is loosely JOURNAL OF THE LEPIDOPTERISTS SOCIETY covered with silk. When galls of later instar larvae were cut open, the larvae immediately began to seal the opening with silk, incorporating frass and debris. These galls were approximately 20 mm in length with a maximum radius of about 4 mm. In contrast, larval chambers of Seticosta rubicola on Rubus eriocarpus reached a length of approximately 40 mm, although the swelling of the stem was less conspicuous than that of galls on R. vulcanicola. The initiation of stem-boring can be detected by the presence of a small amount of frass near the stem apex (Fig. 12). The swelling or ini- tiation of gall-formation can be detected less than two weeks after the initiation of boring. When reared in plastic bags, immature larvae left the original galls and moved to the extra stems that were included in the bag. Larvae usually bored into the stem from the cut surface (Fig. 15). Three larvae completed development feeding on the stem tissue by boring (parenchyma and apparently some vascular tis- sues). Densely spun silk (denser than the silk spun in the gall chamber) was present on the chamber floor. The larvae bored the stem continuously, resulting in chambers slightly greater than 4 cm in length (n = 15). In response to probing with forceps, larvae regurgi- tated brown liquid. Larvae also often responded to “irri- tation” by moving the head and the caudal part of the ab- domen up and down a few times for about two seconds. We assume that under natural conditions pupation takes place outside of the gall chamber since pupae were found in none of the older gall chambers we dis- sected (n = 50). Under laboratory conditions, most lar- vae left their galls and pupated in the plastic bag with- out spinning cocoons (n = 7). However, one larva pupated inside the gall chamber, spinning a thin co- coon; and two pupated in the split part of the stem, spinning cocoons with bits of the plant tissue (Fig. 15). In the latter two cases, the larvae initially left the gall chamber, presumably searched for an appropriate pu- pation site (i.e., wandered around in the plastic bag), and finally returned to the gall or split part of the stem. This behavior suggests that the larvae were searching for a narrow or concealed space within which to pu- pate. Larval development from the beginning of the third instar to pupation took about 20 days in the re- frigerator (n = 4); the pupal stage required about 35 days (n = 1). Two specimens of a parasitoid wasp, Bassus nr. cin- gulipes Sharkey (Braconidae: Agathidinae), were reared from a larva of S. rubicola. An additional female of this parasitoid was captured while it investigated a larva on R. vulcanicola. The second author reared a single female of S. rubi- cola from cultivated blackberry (mora), Rubus prae- VOLUME 57, NUMBER 2 119 15 Fics. 10-15. Galls of Seticosta rubicola on Rubus vulcanicola. 10, Gall swelling, opening “decorated” with larval frass and debris; 11, Lat- eral section illustrating parenchyma tissue and spongy stem tissue; 12, Initiation of boring near stem apex; 13, Fifth instar boring in cut stem; 14, Fifth instar; 15, Cocoon spun in split part of stem. cipuus L. H. Bailey, and larvae of S. rubicola have the fruit of Rubus sp. imported from Guatemala. been reported as a serious pest of this crop in Pérez Hence, larvae occasionally may be responsible for Zeled6n, Costa Rica (Ruth Leén pers. com.). Parts of damaging fruit as well as stems. the stems where galls were present showed splitting In general, gall-inducing species usually require a tissues. Larvae identical to those from Costa Rica have specialized food source (i.e., a specific part of gall tis- been intercepted by APHIS at U.S. ports-of-entry on sue commonly called nutritive tissue) in order to com- 120 plete development (Dreger-Jauffret & Shorthouse 1992). Based on gall structure and larval behavior, S. rubicola may be a stem-borer behaving like a gall- inducer, or a gall-inducer behaving like a stem-borer. The swellings found on the stems of Rubus spp. prob- ably are induced by the mechanical damage caused by larval feeding and/or silk deposition in the chamber. The densely spun silk in the stem chamber may indi- cate that larvae responded to non-regrowing stem tis- sue, tried to induce regrowth of the tissue, or the stem tissue did not dissolve the silk. No species of Euliini previously have been reported to have endophagous-feeding larvae. While some species are known to attack fruit (e.g., Proeulia Clarke, Chileulia Powell, Accuminulia Brown), larvae of these taxa are assumed (or are reported) to feed externally on the surface of the fruit. During the preparation of a systematic treatment of Anopinella Powell, a close relative of Seticosta, we discovered a species in that genus that has been reared from the fruit of Styrax (Styracaceae) and a second species from a fungus gall on Inga longispina (Fabaceae). In addition, the closely related genus Apolychrosis Amsel is reported to feed on the seeds of pine cones (Pogue 1986, Brown & Pas- soa 1998). These limited data suggest the possibility that this clade within Euliini may be adapted to inter- nal or endophagous feeding, a unique adaptation within the Tortricinae, excluding Cochylini. ACKNOWLEDGMENTS We thank Jenny Phillips (INBio) and Jerry Powell (UCB) for al- lowing us to examine specimens in their care. The following provided information regarding the pest status of the new species in Costa Rica: Soffa Angulo Mora, Vanessa Solis Calvo, Lisseth Navarro Mora, and Ronaldo Vega, Colegio Técnico Professional de San Pablo de Léon Cortés. Illustrations of male genitalia, female genitalia, and pupa were provided by Linda Lawrence, USDA Systematic Ento- mology Laboratory, Washington, D.C. Illustrations of the larva were drawn by David Adamski, USDA Systematic Entomology Labora- tory. The photograph of the adult was taken by Karie Darrow, De- partment of Systematic Biology, National Museum of Natural His- tory, Washington, D.C. We thank Michael J. Sharkey, University of Kentucky, Lexington, for identifying the parasitoid wasps, and Ruth Le6n G., San José, Costa Rica, for sharing information and speci- mens. The following provided helpful reviews of the manuscript that enhanced its clarity and quality: Paul E. Hanson (UCR); William E. Miller, University of Minnesota, St. Paul, Minnesota, U.S.A.; Nor- man Woodley, USDA Systematic Entomology Laboratory, National JOURNAL OF THE LEPIDOPTERISTS SOCIETY Museum of Natural History, Washington, D.C.; and Jerry Powell, University of California, Berkeley, U.S.A. We thank Federico Valverde for allowing K. Nishida to use Estacién Bioldgica Cerro de la Muerte. Instituto Nacional de Biodiversidad, Santo Domingo, Costa Rica, and the NSF grant ALAS IV (Arthropods of La Selva) to Jack Longino, Evergreen State College, Olympia, Washington, pro- vided funds for J. Brown to visit Costa Rica on several occasions. LITERATURE CITED ADAMSKI, D. & J. W. Brown. 2001. Revision of the Ecdytolopha Group of genera (Lepidoptera: Tortricidae: Grapholitini) in the New World. Entomol. Scand. Suppl. 58:1-86. 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 Chrysoxena group of genera (Tortricidae: Tortricinae: Euliini). Univ. Calif. Publ. Entomol. 111. 87 pp. + 17 pp. figs. . 2000. Systematics of Anopina Obraztsov (Lepidoptera: Tortricidae: Euliini). Univ. Calif. Publ. Entomol. 120. 128 pp. + 32 pp. figs. Brown, R. L. 1987. Tortricidae (Tortricoidea), pp. 419-433. In Stehr, F. (ed.), Immature insects. Vol. 1. Kendall/Hunt Publ. Co., Dubuque, Iowa. CLARKE, J. F. G. 1958. Catalogue of the type specimens of mi- crolepidoptera in the British Museum (Natural History) de- scribed by Edward Meyrick, Vol. 3. Published by the Trustees of the British Museum, London. DREGER-JAUFFRET, F. & J. D. SHORTHOUSE. 1992. Diversity of gall- inducing insects and their galls, pp. 8-33. In Shorthouse, J. D. & O. Rohfritsch (eds.), Biology of insect-induced galls. Oxford University Press, New York. Horak, M. 1984. Assessment of taxonomically significant struc- tures in Tortricinae (Lep., Tortricidae). Mitt. Schweiz. Entomol. Ges. 57:3-64. . 1998. Tortricoidea, pp. 199-215 In Kristensen, N. (ed.), Lepidoptera, moths and butterflies. Volume 1: Evolution, Sys- tematics and Biogeography. Handbook of Zoology 4(35). Arthropoda: Insecta. Walter deGruyter, Berlin and New York. KAPPELLE, M. 1996. Los Bosques de Roble (Quercus) de la Cordillera de Talamanca, Costa Rica: Biodiversidad, Ecologia, Conservaci6n y Desarrollo. Universidad de Amsterdam/ Insti- tuto Nacional de Biodiversidad. xvi + 319 pp. MacKay, M. R. 1962. Larvae of the North American Tortricinae (Lepidoptera: Tortricinae). Canad. Entomol. Suppl. 28. 182 pp. PocuE, M. 1986. A revision of the neotropical genus Apolychrosis Amsel with descriptions of new species (Lepidoptera: Tortri- cidae), pp. 19-28. In Cibrian-Tovar, D., B. H. Ebel, H. O. Yates & J. T. Mendez-Montiel (eds.), Cone and seed insects of the Mexican conifers. Southeastern Forest Experiment Sta- tion. RAzowskI, J. 1988. New genera and species of the Neotropical Archipini (Lepidoptera, Tortricidae). Acta Zool. Cracov. 31:387-422. ; Received for publication 19 April 2002; revised and accepted 22 Oc- tober 2002. Journal of the Lepidopterists’ Society 57(2), 2003, 121-136 REVISION AND PHYLOGENETIC ANALYSIS OF ACCINCTAPUBES SOLIS (PYRALIDAE: EPIPASCHIINAE) WITH A LARVAL DESCRIPTION OF AN AVOCADO-FEEDING SPECIES M. ALMA SOLIS Systematic Entomology Laboratory, PSI, ARS, USDA, Smithsonian Institution, P.O. Box 37012, National Museum Natural History, E-517, MRC 168, Washington, D.C. 20013-7012, USA. Email: asolis@sel.barc.usda.gov AND LINDA STYER Department of Entomology, University of California at Davis, Davis, California 95616, USA. Email: Imstyer@ucdavis.edu ABSTRACT. Accinctapubes Solis from the tropical Westem Hemisphere is revised. Four species of Accinctapubes with overlapping dis- tributions are recognized: the type species, A. albifasciata (Druce), A. apicalis (Schaus), A. chionophoralis (Hampson), and from Costa Rica A. amplissima Solis & Styer, new species. Stericta leucoplagialis var. purusalis is anew synonym of A. albifasciata. Accinctapubes anthimusalis (Schaus) is transferred to Quadraforma Solis, new combination. The larva of A. albifasciata that feeds on avocado is described and illustrated for the first time. A dichotomous key for the species is presented. A phylogenetic analysis of 17 morphological characters resulted in one tree with a consistency index of 0.89. Additional key words: Accinctapubes adults are hairy, noctuid-like pyra- loids with heavy bodies and forewings in length from 1 to almost 2 cm (Fig. 1). These moths can be collected at lights in the Caribbean and from Veracruz, Mexico to Bolivia, Brazil, and Paraguay between elevations of 400 to 2500 meters. Very little is known about the bi- ology of this genus, although larvae of A. albifasciata are known to feed on Lauraceae, specifically avocado trees. During a study of the Pococera complex in the Epi- paschiinae, Solis (1993) discovered five New World species that were apparently a natural group in the Old World genus Stericta Lederer. Dissection and study of Stericta divitalis Guenée, the type species, showed that these species did not belong in Stericta, and re- quired a new genus, for which the name Accincta- pubes was chosen. Accinctapubes includes four species: A. albifasciata (Druce), A. apicalis (Schaus), A. chionopheralis (Hampson), and A. amplissima Solis & Styer, a new species from Costa Rica. The status of A. anthimusalis is revised. A series of specimens col- lected in the Dominican Republic that are probably another new species of this genus were discovered too late to be included in this study and will be dealt with in another paper. MATERIALS AND METHODS Pinned specimens were examined with an incandes- cent light source. Male and female genitalic dissec- tions were prepared following Clarke (1941), using chlorazol black as a staining agent. Wings were stained with Eosin-Y. Both genitalia and wings were mounted permanently in Canada balsam. All slide preparations Persea americana, Ocotea veraguensis, Lauraceae, Caribbean, South America, Costa Rica. were examined with dissecting and compound micro- scopes. Wing length measurements are from the cen- ter of the axillary area to the apex of the forewing. Wing width measurements are from anal angle to the apex of the forewing. Long series of specimens for study of variation within Accinctapubes were collected by D. Janzen and W. Hallwachs and by parataxono- mists of the Instituto Nacional de Biodiversidad (IN- BIO), Costa Rica. Neotropical specimens were also ex- amined at the following museums: American Museum of Natural History, New York City, USA (AMNH); Bo- hart Museum, University of California at Davis, Cali- fornia, USA (UCDC); California Academy of Natural Sciences, San Francisco, California (CAS); Canadian National Collection, Ottawa, Canada (CNC); The Nat- ural History Museum (BMNH), London, England; Carnegie Museum, Pittsburgh, Pennsylvania, USA (CMNH); Los Angeles County Museum of Natural History, Los Angeles, California, USA (LACM); Trans- vaal Museum, Pretoria, South Africa (TMSA); National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA (USNM); Naturhistorisches Museum Wien, Austria (NMW): Museum fiir Natur- kunde der Humboldt-Universitét, Berlin, Germany (ZMHB); Zoologische Staatssammlung, Munich, Ger- many (ZSMC). The deposition of types is indicated by acronyms following the locality data. Specific locality, collector, and date of collection are reported as written on the label. Morphological characters for the analysis included five characters from the head, eight from the wings, and four from the genitalia (Table 1). The character matrix (Table 2) consisted of five taxa and 17 un- JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fic. 1. A. albifasciata perching. Photo by Kjell Sandved taken in Venezuela (specific locality unknown). ordered binary characters. The matrix was analyzed using PAUP 4 (Swofford 1998). Each character trans- formation series is polarized, that is, the direction of the supposed evolution of a character from a plesiomor- phic (=state 0) to apomorphic (=state 1) condition is de- termined. This was accomplished with the outgroup method using Carthara Walker. All species of Carthara were combined and used as a single outgroup. The hypothesized sister group to Accinctapubes is the Cecidipta-Roeseliodes clade (Solis 1993). However, this clade was not chosen as the outgroup because it is highly derived with many reduced morphological char- acters and the sister group relationship between Accinc- tapubes and the Cecidipta-Roeseliodes clade is not well supported. These two groups share only one homoplas- tic character, the presence of vein CuP in the forewing, a character also found in Deuterollyta Lederer. Instead, Carthara, a genus outside this group with few autapo- morphies and located in a less derived position of the tree (Solis 1993) was chosen as the outgroup for this study. It shares several homoplastic characters with Ac- cinctapubes. In the forewing the origin of vein R, is proximal to the discocellular cell and 3A is coincident with 1A+2A and in the hindwing Sc+R, are separate. In the male genitalia the saccus and a sclerotized structure at the base of the uncus are present, the length of the medial lobe at the base of the valva is rectangular, twice as long as wide, and the juxtal arms extend beyond the costa of the valva. The ductus ejaculatorius is subtermi- nal at the anterior end of the aedeagus. RESULTS The phylogenetic analysis generated one parsimo- nious tree with a length of 19 steps, a consistency in- dex of 0.89, and retention index of 0.60 (Fig. 2). The resulting cladogram indicates that A. albifasciata and A. chionophoralis are sister species based on the pres- ence of white scales in the antemedial area of males (character 7). Accinctapubes apicalis is the sister species to these two taxa; they share an M,/M, vein junction of the hindwing that is contiguous (not stalked) with the outer margin of the discal cell and a signum shaft that is greater than 0.3 mm (character 14). Based on this analysis the members of this genus share a male scape extension that is greater than 3.5 mm (character 1); male maxillary palpus with the 3rd segment less than two-thirds as long as the 2nd seg- ment (character 2); apex of the 2nd segment of the male maxillary palpus round (character 3); length of the 2nd segment of the male maxillary palpus less than 0.15 mm with the 3rd segment (character 4); apex of the 3rd segment of the male labial palpus pointed VOLUME 57, NUMBER 2 TABLE 1. Characters and states used in construction of cladogram for Accinctapubes. 1. Male scape extension length 2. 3rd segment of male maxillary palpus 3. Apex of 2nd segment of male maxillary palpus 4. Length of 2nd segment of male maxillary palpus 5. Apex of 3rd segment of male labial palpus 6. Patch of a thick, sclerotized hooked setae on apex of male forewing 7. Color of antemedial area in male forewing 8. Outer margin of discal cell of male forewing 9. Male and female entire forewing apical area white 10. Width of male forewing 11. Sc+R, vein of male and female hindwing point where M, splits from R 12. Location of M,/M,, vein junction of male and female hindwing 13. Apex of male frenulum 14. Length, apex to base, of signum 15. Shape of signum shaft 16. Setae of papillae anales 17. Medial lobe of juxta 0 less than or equal to 3.5 mm 1 greater than 3.5 mm 0 as long as 2nd segment 1 less than 2/3 as long as 2nd segment 0 pointed ; 1 rounded 0 greater than or equal to 0.15 1 less than 0.15 mm 0 rounded 1 pointed 0 absent 1 present 0 same as basal color of wing 1 white 0 bent inwards medially 1 linear and angled upward 0 absent 1 present 0 less than or equal to 0.9 em 1 greater than 0.9 cm 0 curves toward costa in the area prior to the 1 not curved in this area 0 distal from outer margin of discal cell 1 contiguous with outer margin of discal cell 0 tapered 1 bulbous 0 less than or equal to 0.3 mm 1 greater than 0.3 mm 0 curved 1 straight 0 simple 1 spatulate, then terminally branched 0 absent 1 present, elongated (character 5); outer margin of the discal cell of the male forewing linear and angled upward (character 8); Sc+R, of hindwing not curved in the area prior to the point where M, splits from R (character 11); apex of the male frenulum bulbous (character 13); setae of the papillae anales spatulate, then terminally branched (character 16); and medial lobe of the juxta present and elongate (character 17). KEY TO SPECIES OF ACCINCTAPUBES SOLIS 1. Junction of M,/M, in hindwing not contiguous (stalked) with outer margin of discal cell (Fig. 27); forewing width greater dna 09) orn (URES, ©, IO) o.2oc2s0s000ccee000008 amplissima — Junction of M,/M, in hindwing contiguous (not stalked) with outer margin of discal cell (Fig. 24); forewing width less imam 0.9 oan (Pigs, BO; 7) ooccccscecvecssnboogn00scanee 2; 2. Male scape length less than or equal to 3.5 mm (Figs. 11, 15); outer margin of the male forewing discal cell bent inwards at center forming two points extending distally (Fig. 24) BN ie a8 Eberly an hore a care RRC a clUS ec i ane albifasciata — Male scape length greater than 3.5 mm (Figs. 12, 13); outer margin of male forewing discal cell linear and angled upward Geo ee rer ee ae 8 3. Patch of thickened, dark setae present on apex of forewing; male and female forewing apical area to postmedial line not HOE (UU YO) ees ead aia ome an Re orem chionophoralis — Patch of thickened, dark setae on apex of male forewing ab- sent; male and female forewing with apical area to post- marshall lime wimmtee (TENG, @) 2 2coccccusosasaunogncane apicalis SYSTEMATICS Accinctapubes Solis, 1993 Accinctapubes Solis 1993:48. Type species: Cecidiptera |sic] albifasciata Druce, 1902:325, by original designation (TMSA). Type local- ity: Sarayacu, Ecuador. Diagnosis. Accinctapubes is defined by two autapo- morphies, a male frenulum that is bulbous at tip (Fig. 24) and ovipositor lobes with some spatulate setae that are bifurcate or trifurcate distally (Fig. 37). Identification synopsis. Accinctapubes can be identified by the forewing pattern (Figs. 3-10) with the postmedial line curving toward the outer margin at M.. Species exhibit sexual dimorphism in wing pattern and antennae. In the forewing of both sexes, the me- dian line and reniform spot are not prominent, and, in the hindwing, the postmedial line is prominent from the costal margin to A,. Distribution. Caribbean, southern Mexico to Ar- gentina and Brazil. 124 Biology. Only the biology of A. albifasciata is known and the caterpillars feed on leaves of Lau- raceae. Accinctapubes albifasciata (Druce) (Figs. 1, 3-5, 11, 15, 20, 24, 25, 29-30, 37, 41, 42) Cecidiptera |sic] albifasciata Druce, 1902:325. Stericta leucoplagialis Hampson, 1906:143; Holland & Schaus, 1925:115; Solis, 1993:71; 1995:89. Stericta leucoplagialis var. purusalis Holland & Schaus, 1925:115, new synonym. Stericta albifasciata; Dyar, 1912:66; Holland & Schaus, 1925:114; Kaye & Lamont, 1927:125. Jocara ban Dyar, 1916:37; Solis, 1993:71; 1995:89. Accinctapubes albifasciata; Solis, 1993:71; 1995:89. Diagnosis. Antemedial area white near costa with a tuft of white scales on posterior margin of discal cell (Figs. 3, 4) and male scape shortest in the genus, barely reaching thorax (Figs. 11, 15). Redescription. Male: Head (Figs. 11, 15): frons brownish red, green scales behind ocellus and chaetosema. Antenna with each seg- ment brown distally, brownish red basally; male scape length 2.25 mm (n = 7), anteriorly reddish, posteriorly reddish scales with brown tips, with longer, straight red scales mediolaterally throughout. Labial palpus green. Thorax (Figs. 3, 4): collar reddish. Tegula basally red- dish with brown-tipped scales, dorsally with reddish scales, posteri- orly scales are tipped dark brown and appear as two dark spots. Legs (Fig. 20): forecoxa basally reddish, distally light green, forefemur basally light reddish, all other segments and legs basally dark brown, peppered with green scales and white distally. Wings (Figs. 3, 4, 24, 25): forewing length 1.1—1.2 cm, width 0.55-0.8 cm (n = 10). Basal area near costa greenish, reddish near anal margin. Antemedial line indistinct. Antemedial area white near costa with a tuft of white scales on posterior margin of discal cell. Medial line faintly white. Medial area reddish with more green near costa and 1A+2A. Post- medial line faintly dark brown basally, white distally. Postmedial area brownish-red. Terminal line dark brown. Underside reddish white along costa, dark brown posteriorly until CuA,, white to posterior margin. Postmedial band light brown. Hindwing: beige, marginal shade darker brown separated from dark brown postmedial line by light brown scales, reddish scales on some veins. Anal area with long, straight, faintly pink scales. Underside with costa to M, reddish white, remainder white. Abdomen: yellowish, peppered with red and brown scales. Male genitalia (Figs. 29, 30): uncus length = 1.04 mm (n = 7), width at tip = 0.22 mm (n = 7). Female: Head (Fig. 5): same as male, except scape simple. Tho- rax (Fig. 5): female with collar and tegula entirely reddish. Legs: same as male. Wings (Figs. 3-5, 24-25): forewing length 1.2-1.4 cm, width 0.55-0.8 cm (n = 10). Female similar to male, but basal color brownish red, antemedial area near costa mostly reddish with a few TABLE 2. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fic. 2. Hypothesis of phylogenetic relationships of Accinca- pubes species. Numbers on the left refer to characters, numbers on the right refer to character states. white scales, a long tuft of brown scales on posterior margin of discal cell, antemedial line distinct, brown basally, white distally. Hindwing: Female sometimes light brown throughout and anal area always with long, straight scales reddish brown. Abdomen: Female genitalia (Fig. 37): signum curved, length from apex to base 0.52 mm (n = 1). Larval description (Figs. 41, 42). Length 23 mm (last instar) (n = 2). Head beige and yellow ventrally, reddish anteriorly. Epicranial suture present. Ventral margin of frons and clypeus yellow. Labrum white with brown ventral margin. Capsule area on either side of clypeus highly sclerotized. T1-3 and Al-10 integument smooth, pinacula dark brown. Pinacula ring at base of SD1 on A8. Protho- racic shield beige with 6 complete longitudinal lines and an incom- plete line extending to SD1 from anterior margin and less brown Matrix of characters and taxa used in the cladistic analysis of Accinctapubes (numbers for characters correspond with those used in the text). All species of Carthara were combined and used as the outgroup. (? = missing data.) Characters O01 02 03 04 05 06 07 Carthara 0 0 0 0 0 0 0 albifasciata 0 1 1 1 1 0 1 apicalis 1 iL 1 1 1 0 0) chionophoralis 1 1 1 1 1 1 1 amplissima 1 1 I 1 1 0 0) Taxa 08 09 10 ILI 12 13 14 15 16 17 = jj SS SHE o © io orole> a) oOrrFe oS el il le) OrRrHO — SS & 8) io) eee RO VOLUME 57, NUMBER 2 125 Fics. 3-10. Adults of Accinctapubes. 3, A. albifasciata 3, forewing length = 1.3 cm; 4, A. albifasciata 3, forewing length = 1.4 cm, note dif- ference in hindwing pattem between 3 and 4; 5, A. albifasciata 2, forewing length = 1.2 cm; 6, A. apicalis 3, forewing length = 1.5 cm; 7, A. chionophoralis 3, forewing length = 1.4 cm; 8, A. chionophoralis °, forewing length = 1.4 cm; 9, A. amplissima d, forewing length = 1.7 cm; 10, A. amplissima °, forewing length = 1.8 cm. 126 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 11-14. Lateral view of adult male heads (eye diameter = 1.5 mm). 11, A. albifasciata; 12, A. apicalis; 13, A. chionophoralis; 14, A. amplissima. VOLUME 57, NUMBER 2 12 iD) Vy 18 “I Fics. 15-19. Frontal view of dissected adult heads. 15, A. albifasciata , scape length = 2.25 mm; 16, A. apicalis 3, scape length = 4.5 mm; 17, A. chionophoralis 4, scape length = 4.22 mm; 18, A. amplissima d, scape length = 4.6 mm; 19, A. amplissima 8. 128 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 20-23. Lateral view of male forelegs excluding coxae. 20, A. albifasciata, tibial length = 2.0 mm; 21, A. apicalis, tibial length = 2.2 mm; 22, A. chionophoralis, tibial length = 2.0 mm; 23, A. amplissima, tibial length = 2.5 mm. margin of prothoracic shield on each side. T1 with white and dark brown mottling anterior to thoracic legs; dark brown line to 2 L setae below and anterior to spiracle. T1—-3 legs sclerotized dark brown; 2 dark brown longitudinal lines between thoracic legs. V1 on small, dark brown pinacula. T2-3 with D1-2 and SD1-2 on same pinaculum lo- cated on 2nd and 3rd longitudinal lines. SV1 with one seta. Al—8 with L1 and L2 ventral to spiracle; SD1 pinaculum on 4th longitudinal line; both D1 and D2 setae on a continuous pinaculum on 2nd longitudinal line. Al—A8 with 5th longitudinal line anterior to L1 and L2. Al and A8 with longitudinal lines 2-5 coalesced into a dark brown square. Al1-6 with 3 SV setae and A7-9 with 2 SV setae. Al-9 with V1 on small, dark brown pinacula, medially with light brown maculation and lighter colored, almost white, medial line. Al—-2 with SV setae on dark brown pinacula, and dark brown maculation continuing to posterior part of segment. A3-6 with proleg dark brown. V1 on A7 twice as far apart as on A9. Al-8 and SD2 absent. Spiracle on A8 at least twice as large and slightly more dorsal than other abdominal spiracles. A9 with L1 and L2 setae on same pinaculum, L3 separate, but in an an- teroventral line; D1, D2, and SD1 on separate pinacula. A10 com- pletely dark brown ventrally; dorsally both D1 and D2 setae together on separate dark brown maculations. SD1 and SD2 on both sides on separate dark brown maculations. Prolegs with crochets biordinal in a circle, longer crochets 4-5 times as long as short crochets. Biology. Accinctapubes albifasciata has been reared on avocado (Lauraceae) (Dyar 1912). Kaye and Lamont (1927) reported A. albifasciata as a pest VOLUME 57, NUMBER 2 129 Fics. 24-28. Forewing and hindwing venation. 24, A. albifasciata 4; 25, A. albifasciata 2; 26, A. chionophoralis 3; 27, A. amplissima d, 28, A. amplissima °. species on Persea americana Miller (Lauraceae), the common avocado, in Trinidad. Larvae live gregariously in nests made by webbing leaves and branches to- gether with a tough silk. The following account by D. Janzen and W. Hallwachs (pers. com.) describes the life history of A. albifasciata from northwestern Costa Rica. This species feeds exclusively on mature green leaves of the only native lauraceous tree in its habitat, Ocotea veraguensis (Meissn.) Mez (Lauraceae); feed- ing larvae may be found in any month of the year. The five to fifteen last instar larvae occasionally cluster to- gether in loose structures of living leaves and webbing. When ready to pupate, the larvae drop from the web to the leaf litter and spin a silken cocoon with dirt and leaf parts glued to the outside. Distribution. Southern and western Mexico to Brazil, and from the Caribbean known only from the Dominican Republic and Trinidad. 130 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 29-32. Male genitalia. 29, A. albifasciata; 30, A. albifasciata aedeagus; 31, A. apicalis; 32, A. apicalis aedeagus. Type material examined. Cecidiptera [sic] albifasciata Druce, holotype 3, Sarayacu, Ecuador, C. Buckley (TMSA). Stericta leu- coplagialis Hampson, holotype 4, Guyana, Mapiri, Stewart (BMNH). Jocara ban Dyar, holotype °, Teapa, Tabasco, Mexico, Dec. 13, R. Miiller Collector, Type no. 19285 (USNM). Other material examined. ARGENTINA: Misiones, 13/3/1909, 1 6. BELIZE: Cayo: Mtn. Pine Ridge, 1000’, Falls, Lin- wood C. Dow, 28-VI-1990, 1 2. Stann Creek: Middlesex, E. C. Welling, August 5, 1964, 1 d; August 7, 1964, 1 2. BOLIVIA: Rio Songo, 750 m, Coll. Fassl, 1 d; Boliviae Andes, 1 6; Boliviae Cordil- lieres, 1 d. La Paz: Yungas de la Paz, 1000 m, 1 3, Santa Cruz: Buena Vista, 1 d; Alt. 400 m, J. Steinbach, Aug 1914, 4 d; Sept 1914, 4 3; Nov 1914, 1 6; Dec 1914, 1 d; May 1915, 2 6: Mar 1915, 1 4; Prov. del Sara, 450 m, J. Steinbach, July 1914, 1 6; R. Yapacani, Alt. 600 m, J. Steinbach, Sept 1914, 1 3: Feb 1915, 1 6. BRAZIL: Ama- zonas: Amathura, 1 4; Sao Paulo de Olivenga, November-Decem- ber, 1 d; Reserva Ducke, km. 26, Manaus-Itacoaiara Highway, E. G., I., & E. A. Munroe, 14-22 May 1972, 1 3. Bahia: Camaca, V. O. Becker, 21-30. ix. 1991, 1 °. Espirito Santo: Linhares, 40 m, V. O. Becker, 16-18. IX. 1991, 1 2. Para: Unt. Amaz. Taperinha b. Santarem, Zerny, 1-10. VII ’27, 5 6; 21-31. VI. 27, 6 3,12. Hyu- tanahan, Rio Purus, S. M. Klages, Jan 1922, 2 6; Feb 1922, 4 d; March 1922, 5 d; Apr 1922, 1 d. Rio de Janeiro: Mangaratiba, 150 m, V. O: Becker, 20. i. 1993, 1 d. Rondénia: 62 km S. Ariquemes, Fazenda Rancho Grande, 165 m el., Ron Leuschner, 14—25 Nov 1993, 1 ¢. Santa Catarina: Jaragud, Fr. Hoffman, 1 6. Sao Paulo: Bertioga, 5 m, V. O. Becker, 7-9. x. 1996, 2 4; Sao Paulo, Alto de Serra, R. Spitz, 20. V. 24, 1 6. COLOMBIA: Valle: Anchicaya, 250 VOLUME 57, NUMBER 2 131 Fics. 33-36. Male genitalia. 33, A. chionophoralis; 34, A. chionophoralis aedeagus; 35, A. amplissima; 36, A. amplissima aedeagus. m, J. Bolling Sullivan, Feb. 3, 1989, 1 6. COSTA RICA: 1 ¢. Ala- juela: Fca. La Campana, El] Ensayo, 7 km NW Dos Rios, DH Janzen & W. Hallwachs, 15-17 Mar 1986, 2 d, 1 2; Finca San Gabriel, 16 km ENE Quebrada Grande, 650 m, I Gauld & J. Thompson, 11-15 Jun 1986, 2 d; DH Janzen & W. Hallwachs, 8 Feb 1983, 1 d: 11 Nov 1983, 1 6d; 9 Mar 1984, 1 d: Estacion Pitilla, 9 km S. Santa Cecilia, 700 m, M. Espinosa, M. Espinosa, Jun 1988, 2 é: Espinosa & Chaves, Jul 1988, 2 6; DH Janzen & W. Hallwachs, 20 Nov 1987, 2 6; 4 km W. Sta. Cecilia, 300 m, DH Janzen & W. Hallwachs, 17 Apr 1983, 1 d. Cartago: Turrialba, R. Saunders, III- 21-63, 1 d; V. O. Becker, 600 m, 1 d, 1 2; Pejibaye, at blacklight in cut over forest near river, W. E. Steiner, 22-24 March 1987, 1 2; Juan Vinas, June, 1 ¢. Guanacaste: Santa Rosa National Park, DH Janzen & W. Hallwachs, 2 larvae, voucher number 82-SRNP-675; Rincon Nat. Pk., 4 km E. Casetilla, 750 m, DH Janzen & W. Hallwachs, 27 Dec 1981, 6 6; 23 Feb 1982, 1 d; 14 Feb 1983, 1 d; 11 Apr 1983, 1 d; 6 Jun 1981, 1 6; 7 km Southeast South Heda. Inno- centes, 550 m, DH Janzen & W. Hallwachs, 18 Apr 1985, 7 ¢, 4 2: Estacion Mengo, SW side V. Cacao, 1100 m, DH Janzen & W. Hallwachs, 24 Jan 1987, 2 d; 3 Jan 1987, 2 d; 24 Jan 1987, 4 d; 10 February 1988, 1 9; Mar 1988, 5 6; 13-26 Jun 1987, 3 é: Jun 1988, 1 d; La Luz, W. side V. Cacao, 1000 m, DH Janzen & W. Hallwachs, 3-8 Aug 1986, 4 d; La Mariksa, Hda. Orosi, 550 m, 2-5 Jun 1986, W. Hallwachs & DH Janzen, 3 9; 17 Jan 1986, 7 d, 1 9; W. of Car- mona, Nicoya, 600-700 m, DH Janzen & W. Hallwachs, 19 Aug 1982, 2 6; SSW side Cerro E] Hacha, 300-400 m, DH Janzen & W. Hallwachs, 26-30 Jul 1986, 1 3; Fea. Biesnan, Colonia Refug. Los Angeles, 11 km E. Quebrada Grande, 500 m, DH Janzen & W. Hallwachs, 13 Jun 1985, 2 4, 12; Estac. Cacao, SW side Volcan Ca- cao, 1000-1400 m, R. Blanco & C. Chaves, Sep 1989, 1 d; Santa Rosa National Park, 300 m, DH Janzen & W. Hallwachs, 1-15 Jan 1982, 46: DH Janzen, 12 Dec 1978-10 Jan 1979, 1 2; 1 Jan 1979, 1 2; DH Janzen & W. Hallwachs, 10-20 Mar 1982, 1 6; 9-17 Mar 1981, 2 6; DH Janzen & W. Hallwachs, 28-31 Jul 1979, 3 d; 28-31 Jul 1979, 1 2; DH Janzen & W. Hallwachs, 4-6 Jul 1980, 1 6; 10-12 Jul 1980, 1 6; 13-15 Jul 1980, 1 6; DH Janzen, 1-8 Aug 1979, 1 °; DH Janzen, 7-9 Nov 1979, 1 2; 10-12 Nov 1979, 1 6; 13-15 Nov 1979, 1 6; 16-18 Nov 1979, 2 d, 1 2: 20-22 Nov 1979, 2 2: 23-25 Nov 1979, 2 6, 3 2: 26-28 Nov 1979, 1 2: 4-6 Dec 1979, 3 6; 7-9 Dec 1979, 1 2; 12-14 Dec 1979, 5 d: 16 Dec 1978, 1 2; 18-20 Dec 1978, 1 9: 21-24 Dec 1979, 2 6: La Florida, 500 ft., Wm. Schaus, 1 ¢. Heredia: La Selva Biol. Station, Puerto Viejo de Sarapiqui, 40 m, M. M. Chavarria, May 1986, 1 d; 24 Feb-4 Mar 1987, 1 d; Mar 1987, 5d, 1 9; Apr 1987, 4 ¢; May 1987, 1 2: Jun 1987, 1d: Sep 1986, 1 2: Oct 1987, 1 6; Nov 1987, 1 6; Dec 1987, 2 6; M. M. Chavarria & A. Chacon, Feb 1986, 1 6; M. M. Chavarria, A. Chacon, W. Hallwachs & D. Janzen, 11 Jan 1986, 1 J; DH Janzen & W. Hallwachs, 14-15 Nov 1982, 2 d; La Selva Field Sta. near Puerto Viejo, W. E. Steiner, J. M. Sweringen & J. M. Mitchell, 21-28 March 1988, 1 d; Chila- mate, C. V. Covell, Jr., 100 m, 12-VHI 1986, 1 3; September 8 1988, 1 6; 13-VIII 1986, 1 3. Limon: 9.4 km W. Bribri, Suretka, DH Janzen & W. Hallwachs, 9-11 Jun 1983, 8 9; Cerro Tortuguero, P. N. Tortuguero, 100 m, Dec 1989, 1 d; Brade, 1 d; Sixola River, March, Schaus & Barnes, 4 6; Hacienda La Suerte/Tapezco, 29 air miles W.Tortuguero, elev. 40 m, lat. 10°27’N, long 83°47’, JP & KE Don- ahue, CC Hair, NK Moore, MA Hopkins, 13-31 Aug. 1979, 4 d; Tor- tuga Lodge N. of Tortuguero, el. ca. 20’, Julian P. Donahue, 23/30 Sept. 1977, 1 d; Hac. Tapezco, 29 air km W of Tortuguero, el. 40 m. J. Donahue, D. Panny, D. Moeller, & C. Lewis, 6-23. iii 1978, 5 d,1 2. Puntarenas: Corcovado N. P., 100 m, G. Fonseca, Est. Sirena, Feb 1990, 2 3, 3 2: Apr 1989, 1 3; Nov 1989, 1 2: Dec 1989, 1 2: DH Janzen & W. Hallwachs, 5-11 Jan 1981, 2 6; 23 Mar 1984, 1 9; 15-25 Mar 1981, 1 6; 19-27 Mar 1981, 1 ¢; 1 May 1984, 2 3; 10-12 Aug 1980, 8 6; C. Chavez & R. Aguilar, Feb 1990, 1 4; Isla del Cano, I. Chacon, 12 Mar 1986, 1 6; Monteverde, DH Janzen & W. Hallwachs, 15-16 May 1980, 1 d; 20-21 Jul 1982, 1 6; DH Janzen, 25-26 Jun 1979, 1 3; 10-11 Dec 1979, 2 6: 8-10 Dec 1978, 3 d: Fila Esquinas, 35 km S. Palmar Norte, 150 m elev., 7-8 Jan 1983, 1 d, 1 2; Estacién Quebrada Bonita, R. B. Carara, 50 m, R. Zuniga, Oct 1989, 1 6; Boca de Barranca, Hogue & Dockweiler, 12-14 june 1972, 1 4; Monteverde, E. Giesbert, June 30, 1978, 1 6. DOMINI- CAN REPUBLIC: Dajabon: 13 km S. Loma de Cabrera, Don & Mignon Davis, ca. 400 m, 20-22 May 1973, 1 d. El Seibo: 15 km S. Miches, Don & Mignon Davis, ca. 500 m, 31 May 1973, 4 ¢. La Vega: Constanza, 1164 m, Hotel Nueva Suiza, Don & Mignon Davis, 29 May 1973, 3 d; Convento, 12 km S of Constanza, Flint & Gomez, 13 June 1969, 1 6. ECUADOR: Carchi: Maldonado, 1500 m, V. O. Becker, 9-11. i. 1993, 1 4; Chical, 1200 m, J. Rawlins, R. Davidson, 11 July 1983, 2 d; 15 July 1983, 1 4; 2 July 1983, 1 6; 1250 m, 0-56N, 78-11W, J Rawlins, R. Davidson, 14 July 1983, 3 4, 19. Paramba, Rosenberg, 1050 m, 4 4. Cotopaxi: Las Pampas, Casa Ce- sar Tapia, S 00°25.5°W 78°57.5’, 1200 m, 20-IV-2000, at light UV/MV, 1 d. Esmeraldas: 5 km E. Alto Tambo, 900 m, Jan Hill- man, 8 Dec 1995, 1 d; Rio Durago, 27 km W Alto Tambo, 200 m, Jan Hillman, 5 Dec 1995, 1 5. Loja: Zamora, 1895, 2 4; July 1896, 1 d; Environs de Loja, 1889, 5 d; E] Monje near Loja, 1893. Morona- Santiago: Macas, | 6. Pichincha: Tinalandia, el. 700 m., C. V. Cov- ell, Jr., 5.24.1983, 1 d; 5-18-1985, 1 d; 16 km E Santo Domingo de Los Colorados, el. 600 m, 5-11 May 1990, R. H. Leuschner, 1 ; 17 km SE Sto. Domingo de los Colorados, 3000’, blt It, Oct 21, 1988, 1 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 37-40. Female genitalia. 37, A. albifasciata, note inset a magnified trifurcate seta from ovipositor lobes; 38, A. apicalis; 39, A. chionophoralis; 40, A. amplissima. 2: E of Santo Domingo de los Colorados, Jeffrey A. Smith, 6-11 May 1990, 2 6. FRENCH GUIANA: Saint Laurent du Maroni: St. Jean du Maroni, 1 d, 1 9; St. Laurent, 2 km S. Rte. 1 at pk 244 W. Cayenne, 50 m, Apr 27, 1994, 1 3; Mana River, May 1917, 5 }. Cayenne: Pied Saut, Oyapok River, S. M. Klages, Dec 1917, 1 3: Febr 1918, 4.4; March 1918, 6 6. GUATEMALA: Cayuga, Schaus & Bames, April, 1 3; May, 1d; June, 1 4, 1 2; September, 1 6; October, 1d; Quirigua, Schaus & Barnes, Jan, 2 4; April, 2 9; May, 3 9; August, 3 2; September, 2 4; 2 9; October, 1 6; December, 3 2; no date, 19; Purulha, Schaus & Barnes, April, 1 9; May, 1 9; Rio Dulce, UV, I-16- 1986, 1 2; Below San Lorenzo nr. Pasabien River, 300 m, P. T. Dang, 16-20-X1.1986, 3 6, 1 2. GUYANA: 1916, Franz Knudsen, 3 ¢. a | | VOLUME 57, NUMBER 2 nf prothorax mesothorax metathorax abdominal metathorax segment 1 abdominal segment 2 133 abdominal abdominal 4 1 segment 6 segments 8-10 42 abdominal segment 3 Fics. 41-42. A. albifasciata last instar larva. 41, Lateral view. 42, Ventro-lateral view of thoracic segment 3 and abdominal segments 1-3. HONDURAS: San Pedro Sula, Mountain, blacklight, Robert D. Lehman, 8-IV-1972, 1 d; 15-VII-1972, 1 6. MEXICO: Chiapas: Esmeralda, 19-XI-30, 1 6. Oaxaca: Metates, 2600’ at UV light, John Kemner, 4 March 1992, 1 6. San Luis Potosi: 2 mi. N Tamazun- chale, 400’, Duckworth & Davis, August 2, 1963, 1 9; Valles, V-18- 1952, 1 d. Sinaloa: Coopala, 605 m, C. L. Hogue, 28 Dec.=1 Jan 78-79, 3 3. Oaxaca: Mo Cuo (Cerro Pelon), Mpio. Yolox, 2150 m, E. C. Welling, Sept. 17, 1962, 7 2. Tabasco: Teapa, R. Miiller, Feb- ruary 1914, 1 d. Veracruz: Misantla, R. Miiller, May 1910, 1 6; May 1912, 2d; August 1912, 1 }; August 1915, 16; June 1911, 1d; June 1912, 1 6: November, 1910, 1 3. Yucatan: Chichen Itza, [X-12-1952, 1d. NICARAGUA: Matagalpa: Fuente Pura, 12km N Matagal pa, 1500m, E. van den Berghe, 10 Jan 1997, 1 ¢. PANAMA: 1 ¢, 1 2; Canal Zone: Rincon, Reared from avocado leat? webbing caterpilla, J. Zetek & J. Molino, Aug 15, 1921, 2 4, 3 2; Coco Solo, 1946-1947, 6 d; Barro Colorado Isl., VII-24-63, 1 d; C. W. & M. E. Rettenmeyer, 16. XI.1956, 2 6. TRINIDAD: Wm. Schaus, | 2; St. Joseph, nests of larvae on avocado, December & January, 1 d, 1 2; Arima Valley, 6-II 1950, 1 d; 2-V 1953, 1 d; 29-IV 1951, 1 d; 20-V 1951, 1 d; 4-IIT 1953, 1 6. VENEZUELA: Amazonas: Cerro de la Neblina, Basecamp, 0°50'N, 6°9’44"W, 155 m, canopy, D. Davis & T. McCabe, 1-10 Mar. 1984, 1 d; Rio Mavaca Cp. 65°06’W 2°2’N, 150 m, III-16/27- 1989, 1 5. Suere: Caripito, 3-VI 1942, 1 ¢. Tachira: Btto. Junin, ex follage de aguacate, E. Rubio, 30-XI-1972, 1 d. Trujillo: Valera, 1d, 1 9. Yaracuy: Hacienda Tropicale, ca. 10 km S San Felipe, 10°17'30”S 68°40°W, elev. 100-1400 meters, Kareoleles & Witham, 26 Jan—23 Feb. 1993, 2 9. Accinctapubes apicalis (Schaus) (Figs. 6, 12, 16, 21, 31, 32, 38) Jocara apicalis Schaus, 1906:141. Stericta apicalis Schaus, 1912:669; Holland & Schaus, 1925:116. Cecidipta elphegealis Schaus, 1934:109; Solis, 1993:71; 1995:89. Accinctapubes apicalis Solis, 1993:71. Diagnosis. Entire apical area to postmedial line white on forewing (Fig. 6). Redescription. Male: Head (Figs. 12, 16): frons brownish red, more green scales behind ocellus and chaetosema. Antenna with each segment brown distally, white basally; male scape length is 4.5 mm (n = 1), anteriorly reddish, posteriorly greenish aan increas- ingly longer mediolateral white scales distally. Labial palpus mostly green. Thorax (Fig. 6): collar green. Tegula basally dark brown, dis- tally light green; dorsally wath light aaltiah scales. Legs (Fig. 21): forecoxa basally dark brown, distally white, forefemur basally white, distally dark brown, all other segments and legs basally dark brown, peppered with green scales awl white distally. Wings (Fig. 6): forewing length 1.3-1.5 cm, width 0.7-0.85 cm (n = 10). Basal area greenish white. Antemedial area greenish white, a long (half the length of basal area) tuft of green scales on posterior margin of dis- cal cell, reddish green posterior to tuft. Antemedial line white. Me- dial area greenish white, reddish between CuA, and CuA,, white at 1A+2A. Postmedial line dark brown basally, sahttie distally. Terminal line dark brown. Apical area entirely white from terminal line to postmedial line. Underside reddish white along costa, dark brown posteriorly until CuA,, white to posterior margin. Postmedial band light brown. Hindwing: light brown, marginal : shade darker brown separated from dark awa postmedial tHe by light brown scales, reddish scales on some veins. Anal area with long, straight reddish scales. Underside with costa to M, reddish white, ennetatioe white. Abdomen (Fig. 6): white, peppered with black dorsally, white with yellow ventrally. Male genitalia (Figs. 31, 32): uncus length 1.0 mm (n = 1), width = 0.2 mm (n = 1). Female: Head: similar to male except female scape simple. Thorax: female similar to male but tegula entirely light green. Legs: similar to male. Wings: forewing length 1.4-1.7 cm, width 0.7-0.85 cm (n = 10). Female forewing similar to male, but basal color greenish brown, a long (half the length of basal area) tuft of yellow scales on posterior margin of discal cell, antemedial line al- most invisible. Hindwing: female dark brown. Female anal area with long, straight scales reddish brown. Abdomen: similar to male. Fe- male genitalia (Fig. 38): signum length from apex to base 0.45 mm (= 1): 134 Biology. Unknown. Specimens have been collected at elevations of 700 m to 3800 m. Distribution. Southern Mexico south to Brazil. Type material examined. Jocara apicalis Schaus, holotype 4 Orizaba, Mexico, Coll. Wm. Schaus, Type no. 9623 (USNM) [mis- takenly identified as a female in the description]. Stericta apicalis Schaus, holotype 4, Jan[uary], Juan Vinas, C[osta] R [ica], Type no. 17673 (USNM). Cecidipta elphegealis Schaus, holotype 2, St. Catha- rina, [Brazil], F. H. Hoffman, Type no. 34506 (USNM). Other material examined. BOLIVIA: Rio Songo, 750 m, Coll. Fassl, 1 °. COSTA RICA: Sitio, May, 1 2. Alajuela: Estacién Pitilla, 9 km S. Santa Cecilia, 700 m, Janzen & Hallwachs, 18 May 1988, 1 4; P. Rios, C. Moraga & R. Blanco, Mar 1990, 1 d; Finca San Gabriel, 16 km ENE Que. Grande, D. Janzen & W. Hallwachs, 9 Mar 1984, 1 d. Cartago: Juan Vinas, June, Feb, 2 d, 1 °. Guanacaste: 4 km E. Casetilla, Rincon Nat. Pk. Gate, 750 m, Ds H. Janzen & W. Hallwachs, 14 Feb 1983, 1 6; 27 Dec. 1981, 2 4; 22 May 1982, 4d; 11 April 1983, 1 2; Rincon Nat. Pk., 19 Nov 1979, D. H. Janzen. Here- dia: E] Angel Waterfall, 8.2 km downhill Vara Blanca, 1350 m, D. Janzen & W. Hallwachs, 5 Aug 1981, 1 d; 3 Jan 1981, 1 d; Braulio Carrillo, 1100 m, vii 1981, V. O. Becker, 1 ¢. Puntarenas: Mon- teverde, D. H. Janzen, 8-10 Dec 1978, 4 4; 25-26 June 1978, 1 2; D. H. Janzen & W. Hallwachs, 15-16 May 1980, 2 2; I-20-1961, 1 2; 35 Km NE of San Vito at Las Alturas Field Station, 4800 ft., June 20, 1992, 1 2; June 26, 1992, 1 9; July 2, 1992, 1 2; Tuis, Aug. 29. 08, 1 é: Puntarenas, Finca Las Cruces, 6 km S San Vito, Eric Fisher, 21-25 August 1976, 1 d. San Jose: Estacién Zurqui (El Tunel), Par. Nac. Braulio Carrillo, 1500 m, W. I. y A. Chacon, Aug 1985, 1 d; Sept. 1985, 1 d; La Montura, 1100 m, E. H. Janzen & W. Hallwachs, 17 Dec. 1981, 1 °. ECUADOR: Tungurahua: Bajios, Julian Donahue, 30 June 1980, 2 6. Pichincha: Chiriboga, Reserva Botanico Palmeras, 1900 m, J. Hillman, 2 Dec. 1995, 1 d. Cafiar: Cuenca Trail above Huigra, Alt, 4-5000 ft., W. J. Coxey, III. 26. 1933, 1 2; Dos Puentes, Alt. 1700 ft., W. J. Coxey, Jan 1929, 1 9. GUATEMALA: Chejel, June, 1 3; Purulha, Schaus & Barnes, July, 1 4, Quirigua, Dec, 1 2; Volcan Sta. Maria, June, July, Nov, 4 °. MEXICO: Chiapas: Santa Rosa, V-1932, 1 d. Puebla: Orizaba, Dognin Collection, 2 9 Oaxaca: Vista Hermosa, Mpio. Comaltepec, 1450 m, E. C. Welling, Sept. 22, 1962, 1 9; Sept. 24, 1962, 7 °; Mo Cuc (Cerro Pelon), Yolox, 2150 m, E. C. Welling, Sept. 17, 1962, 1 2; 24 mi S Juchatengo, E. Fisher, P. Sullivan, 9 Aug. 1970, 1 ; Sierra Juarez, Gulf slope, 4600’, at UV light, John Kemner, 8 April 1992, 1 6. Veracruz: Misantla, R. Miiller, Sept. 10, 1 d; S. Tiago, Tuxtla, 800 m., V. O. Becker, 30. x - 2. XI. 1973, 1 6, 1 9. Guerrero: 26 km NW E!] Paraiso, 1800 m, R. Davidson, J. Rawlins, 8 Aug 1986, 1 6. NICARAGUA: Matagalpa: Fuente Pura, 1600 m, van den Berghe, 3 XII 1994, 3 d; 26 x 1995, 1 3; 27 XII 1994, 1 2; 12 km N Matagalpa, 1500 m, van den Berghe, 10 Apr 1996, 1 d; 10 Jan 1997, 1 d. PANAMA: Chiriqui: Lagunas de Chiriqui, 750 m, UV, 6-20-94, 1 6. VENEZUELA: Las Quigas, Este- ban Valley, 1 2. Cojedes: Aroa, 1 2. Lara: Yacambu Natl. Pk., 1560 m. 13 km SE Sanare, cloud forest, 1560 m, J. Heppner, 28-31 Jul 1981, 1 9. Aragua: Rancho Grande, 1100 m, R. W. Poole, Aug. 22-31 1967, 4 6: 1100 m, E. & I. Munroe, 19 Feb. 1971, 1 d; 19-24 Feb. 1971, 7 3; 1100 meters, J.C. & K.G. Shaffer, 16 June 1973, LG Accinctapubes chionophoralis (Hampson) (Figs. 7, 8, 13, 17, 22, 26, 33, 34, 39) Stericta chionophoralis Hampson, 1906:143; Holland & Schaus, 1925:115. Accinctapubes chionopheralis [sic]; Solis, 1993:71; 1995:89. Diagnosis. This species can be distinguished by an apical area with a cluster of broad red scales extending towards the outer margin over a cluster of hooked se- tae extending basally in the male (Figs. 7, 26). JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Redescription. Male: Head (Figs. 13, 17): frons reddish, green scales behind ocellus and chaetosema. Antenna with each segment reddish brown; male scape length 4.22 mm (n = 5), anteriorly green- ish, peppered with dark brown scales, posteriorly reddish with longer red scales mediolaterally throughout. Labial palpus mostly green, 1st segment with dark brown scales distally. Thorax (Figs. 7, 8): collar light green, and white distally red scales; tegula with white and distally red scales throughout; posteriorly scales are tipped dark brown and appear as two dark spots. Legs (Fig. 22): forecoxa basally dark brown, distally white, forefemur basally white, distally dark brown, all other segments and legs basally dark brown, peppered with green scales and white distally. Wings (Figs. 7, 8, 26): forewing length 1.2-1.4 em, width 0.7-0.8 cm (n = 10). Basal area reddish, dark brown on costa. Antemedial area white, a long (half the length of basal area) tuft of green scales tipped with dark brown on posterior margin of discal cell, reddish brown posterior to tuft, a short row of dark brown scales an- terior to tuft. Antemedial line absent. Medial area reddish green be- tween CuA, and CuA,,. Postmedial line dark brown basally, white dis- tally. Tema line ahak brown. Apical area with a cluster of broad red scales extending towards outer margin over a cluster of straight scales extending towards posterior margin. Underside reddish white along costa, anak brown posteriorly until CuA,, white to posterior margin. Postmedial band light brown. Hindwing: beige, marginal shade darker brown separated from dark brown postmedial line by light brown scales, reddish scales on some veins. Anal area with long, straight red- dish scales. Underside with costa to M, reddish white, remainder white. Abdomen (Figs. 7, 8): white, peppered with black dorsally, white with yellow ventrally. Male genitalia (Figs. 33, 34): uncus length = 1.1 mm (n =5), width = 0.21 mm (n = 5). Female: Head: similar to male except female scape simple. Thorax (Fig. 8): similar to male. Legs: similar to male. Wings (Fig. 8): forewing length 1.3-1.5 cm, width 0.7-0.8 cm (n = 10). Female similar to male, but basal color light green, short row of dark brown scales anterior to tuft not as prominent, antemedial line slightly more visible than in male, apical area without a cluster of broad and straight red scales, and postmedial line more prominent throughout its length. Hindwing: female dark brown. Wing underside of female similar to male but anal area with long, straight scales reddish brown. Abdomen (Fig. 8): similar to male. Female genitalia (Fig. 39): signum length from apex to base 0.40 mm (n = 1). Biology. Unknown. Distribution. Costa Rica south to Brazil and Peru. Type material examined. One 2, 32 mm, Brazil, Organ Mts., Wagner; 3 d, Sapucay, Paraguay, Foster (BMNH). The original de- scription is from a type series of 4 specimens, therefore one male specimen labeled Sapucay, Paraguay, Foster is here designated lecto- type in order to fix the concept of the name and to ensure universal and consistent interpretation of the same. Other material examined. BOLIVIA: Rio Songo, 750 m, Coll. Fassl, 3 6; Boliviae Cordilleres, 1 ¢; Boliviae Andes, 1 3. La Paz: Yun- gas de la Paz, 1000 m, 2 d. Santa Cruz: Sta. Cruz de la Sierra, 450 m, I. Steinbach, Aug 1913, 2 d; Jan 1915, 1 d; R. Yapacani, Steinbach, 9 3, Buena Vista, 2 6; Dec 1914, 1 d; Sept. 1914, 4 d; Aug.1914, 3 d; P. del Sara, Steinbach, Jan 1913, 1 2; Nov 1917, 1d; Dec 1917, 1 4; Nov 1913, 1 3; no date, 4 d; Proy. del Sara, 450 m, J. Steinbach, June 1909, 1 ¢. BRAZIL: Amazonas: Rio Manués, 1 2; San Antonio, Rio Madeiras, 1 6; Ponte Nova, Rio Xingu, d; Reserva Ducke, km. 26, Manaus-Itacoaiara Highway, E. G., I., & E. A. Munroe, May 16-21, 1972, 11 3. Bahia: Morro do Chapeut, 1400 m, V. O. Becker, 23-24. iv. 1991, 1 &. Espirito Santo: Linhares, 40 m., V. O. Becker, 05-09. iv. 1992, 1 2. Maranhao: Acailandia, 150 m, V. O. Becker & G. S. Dubois, 19-27. xi. 1990, 1 2; RO, Vilhena, 600 m, V. O. Becker, 10-13. iv. 1996, 1 6. Matto Grosso: Chapada, 15-268, 55—45W, 450-750 m, Herbert H. Smith, 13 ¢. Para: Nova Olinda, Rio Purus, S. M. Klages, May 1922, 1 6; Hyutanahan, Rio Purus, S. M. Klages, 1 3; Feb 1922, 1 4, 19. Parana, Marumbi, 500 mts., V. O. Becker, 16. XII 1969, 1 J; Campo do Tenente, 800 mts., V. O. Becker 21-1-1974, 1 2: Guaratuba, 600 m, V. O. Becker, 5.VII.1975, 1 6; Curitiba, 920 VOLUME 57, NUMBER 2 m., V. O. Becker, 28-IV.1975, 1 2; Nova Teutonia, Fritz Plaumann, 1 3; 5. X. 1939, 1 d; Castro, 4 d, 1 2. Rio de Janiero: 1 d, 1 2; Nova Friburgo, 600m, 10. ii. 1993, 1 2; Rio de Janeiro, Holland Collection, Nov, 2 2; Campo Bello, Zikan, 1 d. Rio Grande do Sul: Guarani, 29.iv.31, 1 6. Rondénia: Cacaulandia, 140 m, V. O. Becker, xi. 1991, 46d: xi. 1994, 4 d, 3 2; 15-18. x. 1993, 1 d; 15-20. iv. 1996; 62 km S. Ariquemes. Faz. Rancho Grande, 165 m., Ron Leuschner, 18-29 Sept. 1996, 1 d, 1 2; 60 km S. Ariquemes, C. V. Covell, Jr., March 17-22, 1991, 1 6. Santa Catarina: Rio Vermelho, S. Bento do Sul, 850 m, V. O. Becker, 24.1.1974, 2 2; Hansa Humboldt, 1 6; Blume- nau, Pohl, 1 6; Blumenau, Nosswitz, 1 d; Jaragud, Fr. [Ske anen, 3 2: Corupa, A. Maller, IX 1956, 1 3; V 1956, 1 6; XII 1957, 1 6; V 1957, 1 d; VII 1957, 1 6. Sao Paulo: R. Spitz, 1 d; Alto de Serra, R. Spitz, 17. 11.24, 1 d:; Sado Paulo, V. O. Becker, 29. i. 1993, 1 d; Est. Biol. Bora- ciela nr. Salesopolis, 850 m, E. G., I., & E. A. Munroe, 24-26 IX 1971, 1 3. COLOMBIA: Vista Nueva near Santa Maria Mts., M. A. Carriner, Nov. 4 26, 1 6. COSTA RICA: Cartago: aluee Vinas, Schaus & Bames, Feb., May, 2 3: Turrialba, E. L. Tord 3 —5 XI 1967, 1 d: 600 m, V. O. Becker, 25. XII. 1972, 1 d, 12. Alajuela: 9 km S. Sta. Cecilia, M. Espinosa, June 1988, 4d, 1 2; Espinosa & Chaves, Jul 1988, 5 d, 1 9; A. Chacon & M. Espinosa, Feb. 1988, 1 d, 1 2; Janzen & Hallwachs, 18 May 1988, 6 d, 2 9; D. H. Janzen & W. Hallwachs, 20 Nov 1987, 5 d, 2 2; GNP Biodiversity Survey, Mar 1989, 2 ¢; Jul 1988, 1 6; May 1988, 1 d; Nov 1988, 1 d, 2 2; Res. For. de San Ramon, 5 km N. Col. Palmarena, 900 m, I. & A. Chacon, July 1986, 1 d; Finca San Gabriel, 16 km East Northeast Queb. Grande, D. H. Janzen & W. Hallwachs, 9 Mar 1984, 1 d; 11-15 June 1986, 2 4, 1 2°; 11 Nov 1988, 3 d; Finca La Campana. Guancaste: 4 km E. Casetilla, Rincon Nat. Pk., 750 m, D. H. Janzen & W. Hallwachs, 27 Dec 1981; 22 May 1982, 2 d; 6 June 1981, 1 d; 25 Jan 1982, 1 d; Feb 14 1983, 1 9; Estaci6n Mengo, SW side Volcan Cacao, 1100 m, D. H. Janzen & W. Hallwachs, Mar 1988, 1 d, 2 2; E] Ensayo, 7 km NW Dos Rios, D. H. Janzen & W. Hallwachs, 15-17 Mar 1986, 1 d. Puntarenas: Las Cruces Biol. Sta. San Vito, 1200 m, I. Chacon, 16-26 Mar 1988, 1 d; July 11-16, 1988, 1 ¢. Limon: 9.4 km W. Bribri, Suretka, 200 m, D. H. Janzen & W. Hallwachs, 9-11 June 1983, 1 6. ECUADOR: Napo: Parque Nacional Yasuni, 80 km S. PUCE station, Ginta Road, Jan Hillman, 15 May 1996, 1 ¢. FRENCH GUIANA: Saint Lau- rent du Maroni: Piste Paul Isnard, 5.15-53.50, Morton S. Adams, 17-18 January 1985, 1 d; Mana River, May 1917, 5 d. Cayenne: Pied Saut, Ovapok River, S. M. Klages, Mar 1918, 5 d, 1 9; Feb 1918, 1d. GUYANA: Omai, 1 ¢. PARAGUAY: Amambay: Parq. Nac. Cerro Cora, M. Pogue & M. Solis, 7-10 April 1986, 3 6. PERU: Cuzco: Pilcopata, 600 m, premontane moist forest, J. B. Heppner, 11-14 XII 1979, 1 P. Hudnuco: Tingo Maria, 24. XI. 46, 1 d; 28. X. 46, 1d. Junin: Satipo, April, 1 d; Upper Rio Tapiche, 9 XI 26, 1 d; 10 XI 26, 14. Loreto: Upper Rio Marajfion, 1. I. 25, 1 d; Iquitos, 22 XI ’27, 1 d; Middle Rio Ucayali, 19-21 XII’26, 1 5. Madre de Dios: Tambopata Reserve, Laguna Chica, 12°51’S 69°18’W, 200 m, at light, 7 Dec 1996, 1 d. VENEZUELA: Las Quigas, Esteban Valley, 1 d, 19. Aragua: Rancho Grande, 1100 m, SS & WD Duckworth, 16-19. I. 66, 1 5; Duckworth & Dietz, 10-21. II. 69, 2 6; R. W. Poole, Aug. 1-7, 1967, 2 4; blacklight, cloud forest, J. B. Heppner, 20-31 II 1978, 4d, 1 9; 25-26 I 1978, 1 d: 22-93 I 1978, 1 d; 15-16 III 1978, 2 3; 1-3 IV 1978, 12: 1100 m, J.C. & KG. Shaffer, 16 June 1973, 1 4; 10 June 1973, 1 d, 1 9; 19 June 1973, 19; E. &. I. Munroe, 19-26 Feb 1971, 1364. Accinctapubes amplissima Solis & Styer, new species (Figs. 9, 10, 14, 18, 19, 23, 27, 28, 35, 36, 40) Diagnosis. Large wing size of both sexes (Figs. 9, 10). Female genitalia with a straight signum shaft (Fig. 40). Description. Male. Head (Figs. 14, 18): frons gray with longer, reddish scales dorsally; beige scales behind ocellus and chaetosema. Antenna brown with male scape extension mostly beige with a few 135 reddish scales, interspersed with gray. Length of male scape 4.6 mm (n = 1). Labial palpus medially beige, laterally peppered with dark brown and red scales. Thorax (Fig. 9): collar dorsally green, reddish laterally, dark brown ventrally. Tegula with bands of dark brown, green, and beige; remainder of thorax mostly beige with some red- dish tipped scales. Legs (Fig. 23): basal half of forecoxa dark brown, distal half with beige scales; forefemur with beige scales laterally, dark brown scales medially; foretibia and tarsus with siematine bands of beige and dark ee scales. Mid- and hindleg coxae and femora with beige scales; tibiae and tarsi similar to foreleg. Entire hindleg banded with alternating beige and dark brown. Wings (Figs. 9, 27, 28): forewing length 1.7 cm, width 0.95 cm (n = 4). Basal area dark brown anteriorly, speckled brown and beige posteriorly. Ante- medial line white, bordered by brown, becoming more distinct to- wards posterior edge. Antemedial line variable, from very distinct to indistinguishable. Medial area with green, beige, red, and brown scales. Some specimens with dark brown patch from anterior edge of postmedial line to midpoint between antemedial and postmedial lines extending posteriorly for two-thirds of wing width. Postmedial line white bordered by brown, curving toward outer margin at M,. Patches of dark brown raised scales posterior to discal cell and along outer margin of discal cell yellow with reddish tips basally and dark brown distally. Terminal line dark brown. Apical area partially white, not extending to postmedial line. Underside with anterior half brown from base to postmedial band, and posterior half beige. Post- medial band beige, with brown border distally; red scales from post- medial line to outer margin. Hindwing: beige, costal margin brown, postmedial line beige bordered by brown. Abdomen (Fig. 9): light brown and beige. Male genitalia (Figs. 35, 36): uncus extends be- yond valva; length 1.1 mm, width 0.3 mm (n = 1). Female: Head (Fig. 19): similar to male except female scape simple. Thorax (Fig. 10): female similar to male but tegula without distinct bands; color variable from dark brown to beige. Legs: fe- male foreleg with only % basally dark brown, remainder white with a few dark brown scales; other legs similar to male but with reddish scales intermingled throughout. Wings (Figs. 10, 28): forewing length 1.7-1.9 cm, width 0.95-1.0 cm (n = 6). Similar to male. Hind- wing: similar to male. Abdomen (Fig. 10): similar to male. Female genitalia (Fig. 40): signum cone shaped with straight shaft; length from apex to base 0.25 mm (n = 1). Biology. Unknown. Specimens have been collected at elevations above 2400 m. Distribution. Costa Rica. Type material. Holotype 4, Costa Rica, Heredia Province, Braulio Carrillo National Park, Estaci6n Barva, 2500 meters, No- vember 1989, L-N 233400, 523200, G. Rivera, CR1000-089184 [INBIO]. Paratypes: 1 d, Heredia Province, Estacion Barva, Braulio Carrillo N.P., 2500 m, Oct. 1989, G. Rivera; 1 ¢ and 5 2. San José Province, San Gerardo de Dota, Cerro de la Muerte, 2430 m, 1981, D.H. Janzen and W. Hallwachs. USNM genitalia slide numbers 104, 238; 104,239; 106,253. Paratypes deposited in INBIO, USNM, BMNH. Etymology. The species name amplissima is de- rived from the Latin meaning “largest.” The name refers to the largest wing size in Accinctapubes. Accinctapubes anthimusalis (Schaus) Stericta anthimusalis Schaus, 1925:34. Accinctapubes anthimusalis; Solis, 1993:71; 1995:89. Dissection and study of the type of Stericta an- thimusalis Schaus deposited at the Camegie Museum, placed in Accinctapubes by Solis (1993), showed that it belongs to Quadraforma Solis, new combination. 136 Quadraforma is defined by a rectangular medial lobe at the base of the valva and the tegumen sclerite with the tip as broad as the base in the male genitalia, and a tubular second segment in the male labial palpus (So- lis 1993). ACKNOWLEDGMENTS We thank the individuals in charge of the collections mentioned above, as well as V. O Becker, C. V. Covell, Jr., R. Leuschner for the use of their private collections. Dan Janzen and Winnie Hallwachs generously provided the immature larvae and biological informa- tion. Susan Escher illustrated most of the line drawings and Linda Lawrence illlustrated some of the line drawings and provided some of the photographs. L. Styer conducted part of this study as an in- term at the National Museum of Natural History Research Training Program funded by a National Science Foundation grant to the Smithsonian Institution. We thank the following for helpful com- ments on the manuscript: A. Konstantinov and D. Smith, Systematic Entomology Laboratory, PSI, USDA, J. Shaffer, Department of Bi- ology, George Mason University, B. Landry, Muséum dhistoire na- turelle, Geneva, Switzerland, and C. Penz, Dept. of Invertebrate Zo- ology, Milwaukee Public Museum, Milwaukee, Wisconsin. LITERATURE CITED CLARKE, J. F.G. 1941. The preparation of slides of the genitalia of Lepidoptera. Bull. Brooklyn Entomol. Soc. 36:149-161. Druce, H. 1902. Descriptions of some new species of Lepi- doptera. Ann. Mag. Nat. Hist. (7) 9:321-334. Dyar, H. G. 1912. Note ona Stericta from Tropical America. Proc. Entomol. Soc. Wash. 14:66. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY . 1916. Descriptions of new Lepidoptera from Mexico. Proc. Entomol. Soc. Wash. 51:1-37. Hampson, C. F. 1906. New Thyrididae and Pyralidae. Ann. Mag. Nat. Hist. (5) 17:112-147. HOLLAND, W. J. & W. ScHaus. 1925. I. The Epipaschiinae of the Western Hemisphere: a synonymic catalog of the species hith- erto described, with figures of many, which have not heretofore been depicted. Ann. Cam. Mus. 16:49-130. Kaye, W. J. & N. Lamont. 1927. A catalogue of the Trinidad Lepi- doptera: Heterocera. Mem. Dept. Agric. Trinidad & Tobago 3:124-125, ScHaus, W. 1906. Descriptions of new South American Moths. Proc. U.S. Natl. Mus. 30:85-141. . 1912. New species of Heterocera from Costa Rica, XVI. Ann. Mag. Nat. Hist. Ser. (8) 9:656-671. . 1925. New species of Epipaschiinae in the Carnegie and U.S. National Museums. Ann. Carn. Mus. 16:9-48. . 1934. New species of Heterocera from Tropical America. Ann. Mag. Nat. Hist. (10) 14:79-115. Souis, M. A. 1993. A phylogenetic analysis and reclassification of the genera of the Pococera complex (Lepidoptera: Pyralidae: Epipaschiinae). J. New York Entomol. Soc. 101(1):1-83. . 1995. Epipaschiinae, pp. 89-93 In Heppner, J. B. (ed.), Check list. Part 2: atlas of Neotropical Lepidoptera. Brill/Flora and Fauna Books, Gainesville, Florida. 243 pp. SwoFFoRD, D. L. 1998. PAUP*, Phylogenetic analysis using parsi- mony (*and other methods). Version 4. Sinauer Associates, Sunderland, Massachusetts. Received for publication 30 January 2002; revised and accepted 18 December 2002. Journal of the Lepidopterists’ Society 57(2), 2003, 137-143 THE EFFECTS OF A FALL PRESCRIBED BURN ON HEMILEUCA EGLANTERINA BOISDUVAL (SATURNIIDAE) PAUL M. SEVERNS! U.S. Army Corps of Engineers, Willamette Valley Projects, Box 429, Lowell, Oregon 97452, USA ABSTRACT. Autumn prescribed burning is often used to manage a rare wet prairie plant community endemic to the Willamette Valley in westem Oregon, USA. A local race of day flying Saturniid moth, Hemileuca eglanterina, was used to investigate the effects of a prescribed burn on adult, larval, and egg mass abundance conmmnicl with an adjacent oma Symmae area. Adult male moths were not more frequently encountered in the bumed habitat but female H. eglanterina laid more than twice as many egg masses in the burned compared to the unburned habitat in the burn year. Furthermore, females laid significantly more egg masses on the burn edge in the bum year (p < 0.001), suggesting that H. eglante- rina chose to oviposit on burned host plants over unburned host plants. Egg masses laid before the prescribed burn did not survive the fall fire, demonstrating that the management practice is catastrophic for the immature population. Although fire can substantially reduce immature Lep- idoptera populations, some species living in ecosystems that had a frequent historic fire return interval may benefit from the ecological release caused by a prescribed bum. Fires consuming entire habitat parcels of fragmented ecosystems may lead to population Inoféllenedlts and an in- creased frequency of inbreeding. Conservative prescribed burning g practices with unburned refugia may be the most effective way to manage for the conservation of rare grassland plant communities and ‘inet insect fauna. Additional key words: _ fire, grasslands, maternal investment, fire-adaptation, buckmoth. Many grassland ecosystems historically experienced wild or anthropogenic fires that maintained floral structure and community composition (Vogl 1974). Grasslands the world over have suffered substantial re- ductions in area from urbanization, agricultural devel- opment, habitat fragmentation, and successional change following the suppression of wildfires. Pre- scribed burning is frequently employed to manage grasslands for rare plants and maintain a primarily herbaceous plant community by restoring a past eco- logical process (Leach & Givnish 1996, Pendergrass et al. 1998a, b). Fires, prescribed or wild, are generally catastrophic for immature insects that live above or near the ground level (Fay & Samenus 1993, Schultz & Crone 1998) and may also kill adult insects that are weak fliers (Morris 1975, Panzer 1988). The effect of prescribed burning on insect abundance often differs between insect families and even among individual species of the same genus (Crawford & Harwood 1964, Cancelado & Yonke 1970, Bertwell & Blocker 1975, Evans 1984, Benzie 1986, Siemann et al. 1997, Blanche et al. 2001, Panzer & Schwartz 2001), suggesting that some species benefit from fire while others do not. Lepidoptera communities also appear to have fluctuat- ing or unpredictable adult abundance between burned and unbumed treatments (Swengel 1996, 1998, Fleish- man 2000, Panzer & Schwartz 2001), intimating that Lepidoptera response to fire may be species specific. Swengel (1998), Panzer and Schwartz (2001), and Siemann et al. (1997) all mention that prairie inhabit- ing insects, especially prairie endemics, are likely to be adapted to cope with fires. I investigated the effects of a prescribed burn on Hemileuca eglanterina Boisduval ‘Current address: Department of Botany and Plant Pathology, Oregon State University, Corvallis, Oregon 97331, USA. Email: severnsp @science.oregonstate.edu (Saturniidae), a dayflying moth of western North America, which occupies a unique wet prairie ecosys- tem in the Willamette Valley of western Oregon, USA. Historically, the Willamette Valley was burned on nearly an annual basis by Native Americans to increase native food crops and aid in hunting (Boyd 1986). Wet prairie fires are typically low intensity and burn quickly over the grassland consuming the low levels of avail- able fuel (pers. obs.), which is generally true for most grasslands (Agee 1993). Translocation of heat from a wet prairie burn rarely reached soil depths >6.0 cm (Pendergrass 1995). Because of the historical role that anthropogenic fires had in maintaining the Willamette Valley prairie flora, autumn prescribed burns are em- ployed to manage the remnant prairie plant communi- ties (Pendergrass et al. 1998b). The effect of pre- scribed burning on the floral community has been studied intensively by Pendergrass (1995) and Taylor (1999), but the consequence of fall fires on wet prairie insects has not yet been investigated. I chose H. eglanterina as a study species because: (1) it lays eggs in masses that are conspicuous (Fig. la); (2) it is monophagous at the study site; and (3) the lo- cal race of H. eglanterina appears to be ecologically and temporally restricted to the wet prairie. Temporal difference in flight times of two to three weeks and el- evation separates the wet prairie from the montane moth populations. Moreover, the wet prairie popula- tions appear to be ecologically restricted to the wet prairie because I have not located H. eglanterina in the nearby oak woodlands, upland prairie, or riparian ar- eas surrounding occupied wet prairie sites. H. eglante- rina are considered polyphagous throughout their range, accepting host species from the Salicaceae, Rosaceae, Rhamnaceae, and Aceraceae (Ferguson 1971), but use only Rosa nutkana in the wet prairie, 138 Fic. 1. Immature lifestages of wet prairie H. eglanterina. a, Egg mass laid on the apical end of Rosa nutkana; b, A group of late early 3rd instar larvae. despite the presence of other reported larval host species (pers. obs.). Owing to a frequent historical fire return interval in the Willamette Valley, it is possible that native wet prairie Lepidoptera have developed adaptations re- lated to fire survival and behaviors that exploit vacant ecological niches created by the fire. This paper re- ports the behavioral and life history response of H. eglanterina to a fall prescribed burn that bisected a wet prairie moth population. I monitored the adult, larval, and egg populations to describe the demo- graphic differences of each life stage in the burned and unburned habitat. Specifically, I tested the hy- pothesis that H. eglanterina adults were differentially attracted to burned prairie and that egg masses were adapted to fire survival. MATERIALS AND METHODS Study site. Willamette Valley wet prairie is a sea- sonally inundated grassland ecosystem currently exist- JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Pei 2nd and ing in fragments that total <1% of its historical ex- panse. Due to the sizable loss of habitat, Willamette Valley wet prairie represents one of the most endan- gered ecosystems in the U.S. (Noss et al. 1995). The wet prairie ecosystem contains four endemic plant species listed as either threatened or endangered (Oregon Natural Heritage Project 2001), and it is dominated by tufted hairgrass (Deschampsia cespitosa L., Poaceae), camas lily (Camassia quamash Pursh, Agavaceae), dwarf wooly sunflower (Eriophyllum lana- tum Pursh, Asteraceae), Hall’s aster (Aster hallii Cronq., Asteraceae), and nootka rose (Rosa nutkana Presl., Rosaceae). I selected Amazon wet prairie Research Natural Area (RNA), in the southern Willamette Valley ap- proximately 10 km west of Eugene, Oregon, USA (Fig. 2) as the study site because of its relatively large size and the integrity of the native plant community. In Oc- tober 1998, the U.S. Army Corps of Engineers burned 16.2 ha of a 33 ha wet prairie parcel to control exotic VOLUME 57, NUMBER 2 139 | Fic. 2. Map of Oregon and the relative location of the prescribed burn study site. The insert shows the burn edge eight months following the prescribed burn. plants and encourage the native wet prairie plant com- munity. The other half of the parcel was left unburned. Study species. Hemileuca eglanterina is a large, di- urnal moth species ranging west of the Rocky Moun- tains from southern California, USA to southern British Columbia, Canada. Adults fly in early July through the middle of August in the Cascade and Coast Range mountain populations >1500 m eleva- tion, but fly from mid August through late September in Willamette Valley wet prairie (~100 m elevation). Wet prairie H. eglanterina oviposit eggs in early Sep- tember and remain in diapause until the beginning of April. Instars 1-3 are gregarious on Rosa nutkana (Fig. 1b) but disperse in the 4th instar. The larvae are armed with urticating spines that can range from mildly irri- tating to as painful as a honeybee (Apis mellifera L.., Apidae) sting when pressure is applied to the spines (pers. obs.), which is common in the genus (Ferguson 1971). Adult abundance. To sample adult abundance I placed two macroplots, each 0.41 ha and marked with 2 m tall metal rebar sections, in the center of the pre- scribed burn and control treatments. For one hour during the peak flight period, 1100-1400 h, on three separate occasions in two preburn years and the burn year from mid to late August, adult moths flying through the burn and control macroplots were cap- tured and marked on the ventral hindwing with a per- manent marker, then released. All adults flying through the macroplots were counted whether they were marked or captured. I used the number of adult fly-throughs as a relative abundance index to identify any adult bias for burned or unburned habitat. In ad- dition to direct adult observations, the location of egg masses with respect to the burn treatment was used as an indicator for the presence of adult female moths. The burn year is defined as the first calendar year from the time of the burn, October 1998—October 1999, the preburn year as October 1997—September 1998, and the postburn year being from November 140 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TaBLE 1. Adult, larval, and egg population data collected from the bummed and unburned treatments for H. eglanterina in the Amazon RNA study site. * = significantly different when p < 0.05. Demographic measure # of adults observed/macroplot Preburn year | Preburn year 2 Burn year # of larvae/macroplot Preburn year Burn year Postburn year # of egg masses laid on the burn edge Burn year Postburn year # of egg masses for the entire bumed and unburned area Burn year Postburn year 1999-October 2000. H. eglanterina fly from mid Au- gust through late September at the study site and laid eggs before the prescribed burn. Larval abundance. To detect larval population dif- ferences in the burned and unburned habitat, I di- rectly counted all of the H. eglanterina larvae in the two macroplots that were used for the adult sampling. I counted larvae in the first week of May of the pre- burn, burn, and postburn years. Egg mass abundance and fire adaptation. | searched for egg masses during late April and early May of the bum and postburn years in the entire study site. Line transects approximately 15 m apart were walked throughout the entire burn (~16 ha) and con- trol (~16 ha) areas, inspecting each Rosa nutkana plant for early instar larvae. The number of egg masses en- countered and the number groups of Ist instar larvae for which no egg masses could be found were com- bined into a total egg mass census for the burned and unburned areas. To determine if egg masses were adapted for fire survival, I located five egg masses and followed their fate immediately following the bum. I noted any qual- itative differences in the egg masses before and after the burn. Female preference for oviposition was measured by the number of egg masses laid on the burned or con- trol side of the burn edge. The burn edge (Fig. 2), ap- proximately 500 m long, was sampled +30 m on each side of the edge for the entire length of the burn boundary. The amount of host plant appeared to be more or less equivalent on both sides of the burn edge. Adult, larval and egg mass analysis. The number of egg masses found on the burned side of the edge was compared to the number of masses found on the unburned edge among years and the burn treatment Burn treatment Unbumed (control) Statistical analysis Chi-square test 34 Q7 Prel/Pre2 p> 0.05 39 36 Pre2/Bum p > 0.05 48 Al Prel/Burn p> 0.05 Chi-square test 64 136 Pre/Burn __p < 0.0001* 0) 370 Burn/Post p < 0.0001* 258 185 Pre/Post p < 0.001* Fisher Exact test 10 0 p < 0.001* I 5 Chi-square test 39 17 p > 0.05 34 24 using a Fisher's exact test. Differences in adult abun- dance, the number of larvae in each macroplot, and the number of egg masses laid in the burned and un- burned areas were assessed among years and between burn treatments by Chi-square tests. Host plant response and analysis. Host plant re- sponse to the prescribed burn was measured in the first and second week of June in the burn and post- burn year. I estimated R. nutkana cover in sixteen 30 m long x 1 m wide belt transects randomly located in each burn and control macroplot. Cover measure- ments were made in 1 m? subplots for ease and accu- racy, then weighted and added together to yield cover for the belt transect. Host plant stature was divided into height classes, 0-25 cm and >25-50 cm, and the cover of each height class was visually estimated to the nearest 1%, for 1-10% cover, and in 5% increments thereafter. R. nutkana cover estimations were per- formed by the author for consistency between and within years. Host plant height class cover differences between treatments (burn vs. control) within years were analyzed using a Mann-Whitney U-test. All sta- tistical analyses were performed with the NCSS (2000) statistical package. RESULTS Adult, larva, and egg mass populations. Adult abundance did not differ between burned and unburned prairie between the two preburn years and the burn year (Table 1). The number of egg masses censused in the burn year from the burned (16.2 ha) and unbumed (16.2 ha) habitats was nearly twice as high in the bumed area compared to the unburned area in the bum year (Table 1), suggesting that female moths preferred to lay eggs on burned rose. A similar distribution of egg mass number occurred in the postburn year, however there were no VOLUME 57, NUMBER 2 statistical differences in egg mass number within burn treatments and between years (Table 1). Egg mass num- ber from the burn edge habitat was significantly differ- ent (p < 0.001) between the bum and postburn year among the burn and unbumed edge treatment, indicat- ing that females preferred to oviposit on the burned plants in the burn year (Table 1). The fire destroyed the five egg masses found imme- diately following the burn. Many of the eggs appeared to have boiled and then ruptured from the heat of the burn, giving the appearance that they had hatched. Larva number in the burned macroplot of the burn year was lower in the burn year compared to all other years (Table 1), indicating that none of the egg masses laid before the burn produced Ist instar larvae. Host plant response. In the burn year, the amount of R. nutkana in the <25 cm height class did not statis- tically differ between the control and burn plots (mean cover = 0.22 m?/transect +0.06 Mann-Whitney U-test p = 0.85), but there was significantly more rose from the 25-50 cm height class in the control plot compared to the burn plot (burn = 0.21 m*/transect + 0.1 m*; un- burned = 0.62 m?/transect + 0.1 m* Mann-Whitney U-test p = 0.018). The postburn year experienced no significant rose quantity differences between treat- ments in the <25 cm height class (mean cover = 0.174 m?/transect + 0.05 m* Mann-Whitney U-test p = 0.895) and the 25-50 cm height classes (mean cover = 0.645 m7/transect + 0.20 m? Mann-Whitney U-test p = 0.11). These comparisons suggest there was more H. eglanterina host plant available in unburned areas dur- ing the burn year when females laid eggs. DISCUSSION Adult abundance, measured by the number of indi- viduals observed flying through the macroplots in the burn and unburned areas, was not significantly differ- ent between years or the burn treatment (Table 1), ar- guing against the hypothesis that adult moths prefer recently burned prairie to adjacent unburned prairie. However, females laid more than twice as many egg masses in the burned compared with the unburned prairie during the burn year (Table 1), implying there was a burn bias that was not detected through direct adult observations. Examination of the egg mass place- ment on the burn edge, where moths were assumed to have made a choice to oviposit between burned or un- burned plants, suggested that reproductive effort was directed towards the burned plants in the burn year (Table 1). Maternal preference for the burned area and the burn edge in the burn year supports the hy- pothesis that female H. eglanterina were attracted to the burned area. 141 The discrepancy between adult and egg mass abun- dance may be explained by the gender of adult moths surveyed by each method. In the combined 18 hours of adult sampling among the three years, no females were detected. In fact, over the last five years of visit- ing the study site I observed only three females amongst hundreds of male observations. During the two preburn years and the burn year adult sampling effort, male H. eglanterina patrolled the study site in roughly circular flight patterns (presumably searching for females) over large areas of the entire prairie, en- compassing both the burned and unburned areas. Since egg mass counts and adult abundance appeared to effectively measure the relative occurrence of the two genders, it is not surprising that the results are in- consistent with each other, especially if there are be- havioral differences between genders. Differences in the effect of prescribed burning on adult abundance and the number of egg masses laid demonstrates the need to sample multiple lifestages within a species to estimate the effects of prescribed burning. For example, if only adult abundance was used to determine the effects of fire, I would have con- cluded there were no effects on the population. Con- versely, if burn affinity was based solely on egg mass number, I could have inferred that adult distribution was biased towards the burn area. Basing the effects of the prescribed burn on larval abundance would have yielded a conclusion that the fire was catastrophic. Many studies often rely heavily on adult abundance to describe the effects of fire on Lepidoptera (Swengel 1996, 1998, Fleishman 2000, Huebschman & Bragg 2000, Panzer & Schwartz 2001). However, in this study direct adult H. eglanterina observations yielded a non-significant treatment response (Table 1), sug- gesting that adult observations of vagile lepidopterans may not be adequate to assess the effects of fire on study populations. Sampling all lifestages and monitor- ing abundance may narrow the variability of results in any community study, but measuring abundance of multiple lifestages is time consuming. Perhaps focus- ing on a subset of specialist, generalist, widely distrib- uted, and locally restricted species may yield general- izable trends concerning the effects of fire on Lepidoptera natural history, conservation, and com- munity response. Although H. eglanterina egg masses showed no evi- dence of being resistant to fire, there may be an ad- vantage to insect species that colonize recently burned areas in an ecosystem that experiences frequent fires. Larvae feeding on plants in a burned area may experi- ence higher quality food (McCullough & Kulman 1991, Stein et al. 1992) which could result in a rapid population size increases if survivorship and fecundity is increased by food quality. Recently burned habitat should also have a number of exposed niches, from fire induced mortality on immature insects, that can tem- porarily be exploited by opportunistic species. Fur- thermore, females choosing to oviposit in a burned area may impart increased survival to their progeny. A recently burned area would tend to have a low fuel load than an area that has not been burned, resulting in a fire that is not as hot as the original one, and per- haps increased survival of egg masses. Two egg masses that produced numerous Ist instar larvae were found on the burn edge where the fire was extinguished, sug- gesting egg masses may be resilient to slightly elevated temperatures above the ambient. Lepidoptera conservation and_ prescribed burning. The number of larvae observed in preburn, burn, and postburn years within the macroplots demonstrated that fire was lethal for egg masses laid immediately before the burn and larvae did not move into the burned area (Table 1). Complete mortality of eggs and larvae from prescribed burning has been al- luded to in other studies (Siemann et al. 1997, Swen- gel 1998, Panzer & Schwartz 2001) and demonstrated directly in gall forming wasps (Fay & Samenus 1993) and a Lyceanid butterfly (Schultz & Crone 1998). The mortality of immature lifestages inspires criticism for the effects of prescribed fire on rare Lepidoptera while managing for plant communities (Pyle 1997, Schlict & Orwig 1999). In cases where entire prairie fragments are burned, the high mortality of immature life stages may indeed be a cause for concern, as high immature mortality rates may result in population bot- tlenecks. Increasingly smaller population sizes in but- terfly populations have been linked to an increase in the risk of population extinction (Nieminen et al. 2001). However, when land parcel subdivisions are not frequently burned, lepidopteran populations may be less likely to experience a catastrophic loss of individu- als affecting the overall population fitness. This study suggests that prescribed burning has the potential to limit or encourage the H. eglanterina population depending on the size of the burn, the presence of a colonizing population, and bum fre- quency. Unfortunately, I was unable to measure sur- vivorship and vigor of larvae in the field to determine the effects of the burn on the intrinsic rate of popula- tion growth, and this information is essential to deter- mining if the maternal bias for burned plants has an adaptive value to the population. Without survivorship estimates, it can not be known if a burn area acts as a population source or sink, and a strong argument for or against prescribed burning cannot be given. Schultz JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY and Crone (1998) recommended burning for a Willamette Valley upland prairie endemic butterfly, Icaricia icarioides fenderi Macy. They proposed that a rotation of small scale prescribed burns within the but- terfly’s habitat could maximize the growth rate of the butterfly population while still managing for invasive plant species and the native plant community. Less de- structive methods of managing grasslands, such as mowing, may also be a viable management practice combined with rotations of smaller burns, as H. eglanterina egg masses were observed to survive a fall mowing event (pers. obs.). ACKNOWLEDGMENTS I would like to thank Kat and Jim Beal for their support of this project and access to the study site, Cody Litster and Brady Metzger for helping with adult surveys, Alice Moran-Tacey for aiding with egg mass counts, Suzy Holmes for her help counting larvae, and Wes Messinger, George T. Austin, Paul Z. Goldstein, and Philip J. De- Vries for providing valuable comments on this manuscript. I would also like to thank P. J. DeVries and C.M. Penz for their advice and encouragement. LITERATURE CITED AGEE, J. K. 1993. Fire ecology of pacific northwest forests. Island Press, Washington D.C., USA. 493 pp. BENZIE, J. A. H. 1986. The distribution, abundance, and the effects of fire on mound building termites (Trinervitermes and Cu- bitermes spp., Isoptera: Termitidae) in northern Guinea sa- vanna, West Africa. Oecologia 70:559-567. BERTWELL, R. L. & H. D. BLockErR. 1975. Curculionidae from dif- ferently managed tallgrass prairie near Manhattan, Kansas. J. Kansas Entomol. Soc. 48:319-326. BLANCHE, K. R., A. N. ANDERSEN & J. A. Lupwic. 2001. Rainfall contingent detection of fire impacts: responses of beetles to ex- perimental fire regimes. Ecol. Appl. 11:86-96. Boyp, R. 1986. Strategies of Indian burning in the Willamette Val- ley. Can. J. Anthropol. 5:65-85. CRAWFORD, C. S. & R. F. Harwoop. 1964. Bionomics and control of insects affecting Washington grass seed fields. Technical Bul- letin of the Agricultural Experimental Station, Washington State 44:]—25. EVANS, E. W. 1984. Fire as a natural disturbance to grasshopper as- semblages of tallgrass prairie. Oikos 43:9-16. Fay, P. H. & R. J. SAMENUS. JR. 1993. Gall wasp (Hymenoptera: Cynipidae) mortality in a spring tallgrass prairie fire. Environ. Entomol. 22:1333-1337. FERGUSON, D. C. 1971. The moths of America north of Mexico: fascicle 20.2A Bombycoidea, Saturniidae (part). E. W. Classey and R. B. D. Publications Inc. 153 pp. FLEISHMAN, E. 2000. Monitoring the response of butterfly com- munities to prescribed fire. Environ. Management 26:685—695. HUEBSCHMAN, J. J. & T. B. BRAGG. 2000. Response of regal fritil- lary (Speyeria idalia Drury) to spring burning in an eastern Ne- braska tallgrass prairie, USA. Nat. Areas J. 20:386-388. Leacu, M. K. & T. J. GivnisH. 1996. Ecological determinants of species loss in remnant prairies. Science 273:1555-1558. McCuLLoucu, D. G. & H. M. Kutman. 1991. Differences in fo- liage quality of young jack pine (Pinus banksiana Lamb) on burned and clearcut sites: effects on jack pine budworm (Cho- ristoneura pinus pinus Freeman). Oecologia 87:135-145. Morris, M. G. 1975. Preliminary observations on the effects of burning on the Hemiptera (Heteroptera and Auchenorhyncha) of limestone grassland. Biol. Cons. 7:311—319. NCSS. 2000. Number cruncher statistical software. Kaysville, Utah. VOLUME 57, NUMBER 2 NIEMENEN, M., M. C. SINGER, W. FORTELIUS, K. SCHOPS & I. HAN- SKI. 2001. Experimental confirmation that inbreeding depres- sion increases extinction risk in butterfly populations. Amer. Nat. 157:237-244. Noss, R. F., E. T. LAROE, & J. M. Scorr. 1995. Endangered ecosystems of the United States: a preliminary assessment of loss and degradation. National Biological Service, Biological Report 28, Washington, D.C. OREGON NATURAL HERITAGE PROGRAM. 2001. Rare, threatened, and endangered plants and animals of Oregon. Oregon Natural Heritage Program, Portland, Oregon. 94 pp. PANZER, R. 1988. Managing prairie remnants for insect conserva- tion. Nat. Areas J. 8:83—-90. PANZER, R. & M. ScHwartz. 2000. Effects of management burning on prairie insect species richness within a system of small, highly fragmented reserves. Biol. Cons. 96:363—369. PENDERGRASS, K. L. 1995. Vegetation composition and response to fire of native Willamette valley wetland prairies. M.Sc. Thesis, Oregon State University, Corvallis, Oregon. 241 pp. PENDERGRASS, K. L., P. M. MILLER & J. B. KAUFFMAN. 1998a Pre- scribed fire and the response of woody species in Willamette Valley wetland prairies. Restoration Ecol. 6:303-311. PENDERGRASS, K. L., P. M. MILLER, J. B. KAUFFMAN & T. N. Kaye. 1998b. The role of prescribed burning in maintenance of an endangered plant species, Lomatium roles Ecol. Appl. 9:1420-1429. 143 PyLE, R. M. 1997. Buming bridges. Wings 20:22-23. SCHLICT, D. W. & T. T. Ornwic. 1992. The last of the lowa skippers. Amer. Butterflies 7:4—13. SCHULTZ, C. B. & E. E. CRONE. 1998. Burning prairie to restore butterfly habitat: a modeling approach to management tradeoffs for the Fender's blue. Restoration Ecol. 6: 944-959, SIEMANN, E., J. HAARSTAD & D. TILMAN. 1997. Short-term and long-term effects of burning on oak savanna arthropods. Amer. Midland Nat. 137:349-361. STEIN, S. J., P. W. Prick, W. G. ABRAHAMSON & C. F. SAccut. 1992. The effect of fire on stimulating willow regrowth and subse- quent attack by grasshoppers al elk. Oikos Bx 190-196. SWENGEL, A. B. 1996. Effects of fire and hay management on abundance of prairie butterflies. Biol. Cons. 76:73-85. . 1998. Effects of management on butterfly abundance in tallgrass prairie and pine barrens. Biol. Cons. 83:77-89. TayLor, T. H. 1999. Long-term vegetation response to fire of Willamette Valley wet prairie species. M.Sc. Thesis, University of Oregon. VocL, R. J. 1974. Effect of fire on grasslands, pp. 139-194. In Ko- zlowski, T. T. and C. E. Ahlgren (ed.), Fire and ecosystems. Ac- ademic Press, New York. — Received for publication 15 April 2002; revised and accepted 3 Jan- uary 2003. GENERAL NOTES Journal of the Lepidopterists’ Society 57(2), 2003, 144-147 NOTE ON THE DISCOVERY OF THE LARVA OF CUCULLIA SIMILARIS (NOCTUIDAE, CUCULLIINAE) Additional key words: — Cucullia montanae, Cucullia asteroides, Chrysothamnus, Grindelia, Aster. During the period 20 July-20 August 1992, we had the opportunity to examine some aspects of the flora and entomological fauna of the western part of the USA. As a result, we discovered, in the eastern sub- urbs of Provo (Utah), two bright colored (yellow- green) larvae, in the last instar, basking in sunlight and feeding on gray rabbitbrush, Chrysothamnus nauseo- sus (Pall. ex. Pursh.) Britt. (Asteraceae). Initially, it ap- peared evident that these larvae belonged to the Cu- culliinae subfamily, because of some resemblance to the European species Cucullia asteris Denis & Schif- fermiiller. Furthermore, they also look like the Ameri- can Cucullia species of the asteris group whose larvae have been identified, such as Cucullia asteroides Guenée (Dethier 1944, Crumb 1956, Stehr 1987, Poole 1995), Cucullia montanae Grote. (Crumb 1956, Poole 1995) or Cucullia postera Guenée subspecies omissa (Crumb 1956) but, according to Poole (1995), misidentified: the described larvae corresponding rather to a mixture of Cucullia florea Guenée, Cucul- lia postera Guenée and Cucullia obscurior Smith. In the following days the same species was collected again, first, on Chrysothamnus nauseosus near of the Timpanogos caves (Utah) and near Silver Lake (Oregon), and second, on Douglas rabbitbrush Chryso- thamnus viscidiflorus (Hook.) Nutt. (Asteraceae) (Munz & Keck 1973) near Cedar City (Utah). The description of C. montanae Grote given by Crumb (1956) from larvae collected on Grindelia species (Asteraceae) in western Washington is not con- sistent with the larvae collected by us in Utah and Ore- gon on Chrysothamnus sp. However, Cook (1935) has indicated in his Montana list that he collected some C. montanae larvae on Chrysothamnus sp. at Three Forks and Hamilton (Montana); his short description “a green and white striped worm” does not exactly fit that of Crumb (1956), nor that of the species found by us on Chrysothamnus sp. Identification of the larvae collected on Chrysothamnus sp. Assuming that at least two differ- ent Cucullia species can live on Chrysothamnus and Grindelia, we decided to search Grindelia squarrosa (Pursh) Dunal for the presence of a second larva. Un- fortunately, we have not been able to discover any lar- vae on this plant in Utah during the period of 20 July to 20 August 1992. Consequently, we decided to wait for the eclosion of the adults in order to identify the species collected on Chrysothamnus. Several adults eclosed in July 1993 suggesting that this species is one- brooded. At that time the only available drawings of the adults were those presented by Hampson (1906) and Seitz (1919-1944). They did not allow us to clearly distinguish between C. similaris and C. montanae. In 1993, Dr. J. D. Lafontaine (Agriculture Canada, Ottawa) kindly supplied us very precise information on the habitus and the genitalia of the closely related C. similaris and C. montanae. It became clear after ex- amination of the imago and of the male genitalia that the species living on Chrysothamnus in Utah and Ore- gon was C. similaris (C. similaris always showing the presence of a single large cornutus in the vesica in- stead of generally two differently sized cornuti in C. montanae). Later, this was unambiguously confirmed with the appearance of Poole’s book (1995) which gives, not only good color and black and white pho- tographs of adults and both male and female genitalia, but also very interesting maps of distribution for these two species. For instance, the latter show that C. simi- laris is more frequently recorded in Utah than C. mon- tanae (for the latter species a single data point on the distribution map indicates that this species is probably not very common in this state). Our records of C. sim- ilaris in Utah and Oregon are consistent with these distribution maps. Description of Cucullia similaris larva. The full grown larvae are about 40 mm long; head whitish green with two darker shades running across; conspicuous frontal triangle blue green; at moderate magnification numerous small light freckles are visible (Fig. 1). The ground color is green. The main features of this larva are a set of conspicuous longitudinal yellow, green and white stripes (Figs. 2, 3). The middorsal, nearly continuous bright yellow stripe is superim- posed on a larger white stripe overflowing on each side. Between this white stripe and the spiracles there is a set of five stripes: the first and fifth have a dark green color and are partially bordered by traces of thin black lines (only visible at moderate magnification; Fig. 4). The third stripe appears lighter green than the other two. The second and fourth stripes are bright yellow (bordered by white in the upper part) and white, respectively. The spiracles, which are cream encircled with black, are connected to the traces of black lines located at the ventral border of the fifth stripe (dark green). Under the spiracles there is a broad festooned yellow stripe, which is followed by a white stripe. Description of Cucullia montanae larva. From the descrip- tion by Crumb (1956), C. montanae larva appears, at first glance, of a green ground color with conspicuous longitudinal black lines. The greenish white head is strongly marked with large black freckles. The spiracles are white. “A bright yellow continuous middorsal VOLUME 57, NUMBER 2 145 Fics. 1-9. Larvae of Cucullia; ordered sequentially from left to right, top to bottom. 1, Enlarged head of Cucullia similaris larva (last instar), (VII-1992), vicinity of Timpanogos caves (Utah). 2, Cucullia similaris (lateral view) penultimate instar on Chrysothamnus nauseosus (VIII-1992), vicinity of Timpanogos caves (Utah). 3, Cucullia similaris (dorsal view) penultimate instar on Chrysothamnus nauseosus (VIII-1992) vicinity of Tim- panogos caves (Utah). 4, Cucullia similaris; enlarged view of Fig. 2. 5, Cucullia montanae (lateral view), last instar on Grindelia integrifolia, green form (IX-2002), shore of Hood canal (western Washington). 6, Cucullia montanae (dorsal view), last instar on Grindelia integrifolia, green form (IX-2002), shore of Hood canal (western Washington). 7, Cucullia montanae; enlarged view of Fig, 5. 8, Cucullia montanae larva (lateral view), last instar pho- tographed on Aster sp., pink form (IX-2002), shore of Hood canal (western Washington). 9, Cucullia montanae; enlarged view of Fig. 5. 146 stripe and a broad subventral stripe yellow dorsally and white on ventral third” are present. Between these two yellow stripes there is a set of “3 longitudinal darker stripes, the median lighter than the others, all bordered by black lines which tend, in the darker stripes, to be broadened about midway of each segment.” However, Crumb does not indicate the exact color of these three stripes (presumably green), and that of the associated two spaces, preventing us from having a clear idea of the appearance of this species. In the second fortnight of September 2002, following the infor- mation given by Crumb (1956) we collected some C. montanae lar- vae in western Washington, upon a salt-tolerant Grindelia deter- mined later with the help of the book of Hitchkock et al. (1955), as Grindelia integrifolia D.C. (Asteraceae). Figures 5-9 show the two chromatic forms of the larva: green and pink. The three stripes are green as expected (or pink), whereas, the two spaces (Figs. 5-9) be- tween these stripes are bright yellow and dirty white, respectively; ie., nearly of the same colors encountered for stripes 2 and 4 in C. similaris larvae. In some cases, the dirty white stripe is partially or totally invaded by the ground color especially by the pink ground color, which sometimes also partially invades the yellow stripes and the spiracles (Fig. 8). An unique specimen of C. montanae larva has been collected upon a salt-tolerant Aster sp. (Asteraceae) and bred on this plant which appears as an occasional food plant. R. W. Poole (1995) indicates, in a comment of his map of distribution, that C. montanae is more commonly found in dry places and at moderate el- evations (7000-8000 feet, ~2130-2440 m). It seems this is more likely connected to xerothermic preferences of the food plants as for instance the resin weed (Grindelia squarrosa), rather than that of the moths themselves (since the larvae collected by Crumb, like those collected by us, have been found at sea level). In captivity, C. montanae larvae have accepted seeds of Chrysothamnus viscidiflorus, showing this species is clearly oligophagous. The latter result makes the observation by Cook (1935) more credible. However, until now, no C. montanae larvae have been found either on Chrysothamnus sp. or on Grindelia sp. in Central Washington in the second fortnight of September. This may be because of inadequate period of collection in this drier and warmer region than western Washington. Further discussion is un- warranted before obtaining new data. Comparison of the larvae of Cucullia similaris with the nearest species: Cucullia montanae and Cucullia asteroides. From the descriptions of C. as- teroides larva given by Crumb (1956), Dethier (1944), Stehr (1987), Poole (1995), the drawings of Dethier (1944) and the black and white photograph of Stehr (1987) and our own photographs of the two species C. similaris and C. montanae larvae, the following re- marks can be made. The larvae of the three species have in common: (1) the presence of an almost continu- ous middorsal yellow stripe, and between this stripe and the spiracles there is a set of five stripes which are green, yellow or white; (2) the presence of two broad yellow and white subspiracular stripes; and (3) the spir- acles are of a light color encircled with black. Cucullia similaris larvae may be distinguished from the other two species by the nearly complete absence of black lines at the borders of the five stripes lying be- tween the middorsal yellow stripe and the spiracles (traces of black lines are only visible under moderate magnification on the borders of the darker green stripes 1 and 5; Fig. 4). On the other hand, six black lines bordering the five lateral stripes are visible to the JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY naked eye in C. montanae (conspicuous continuous lines) and in C. asteroides larvae (more or less inter- rupted lines). The black markings (resulting of the broadening of two black lines), are completely absent in C. similaris larvae. These markings are present only in the subdorsal region in C. montanae larvae, while in C. asteroides they are larger in size in the subdorsal region and are also pre- sent in the spiracular stripe, around the spiracles. In addition, in C. similaris larvae, the yellow stripes are, at least for the middorsal and subdorsal stripes, su- perimposed on a white stripe overflowing on one or both sides. Until now, only the green ground color has been ob- served in C. similaris larvae, whereas the other two species have both green or pink/brown ground colors. The head of the three species is similar in the ground color but whereas C. similaris larvae have inconspicu- ous numerous light freckles (only visible at moderate magnification), C. asteroides larvae have small brown freckles and C. montanae larvae display strong large black freckles. Although the discrimination between C. montanae and C. asteroides larvae is mainly obtained by compar- ison of the extension of the black lines and markings, other differences in the latter two species may be evi- denced. They are, according to Crumb (1956), the pres- ence in C. montanae larvae, in the bluish green (or pink) venter, of a faint “black line along the base of prolegs” (becoming double between the prolegs; Fig. 9), and in the midventer of traces of two black lines, whereas in C. asteroides larvae the venter is also green (or brown) “with traces of a white stripe in the line of spiracles”. The clues given by Crumbs (1956) and Poole (1995) based on the data of Cook (1931) considering Chrysothamnus nauseosus as a host plant for C. mon- tanae need further verifications. Cucullia similaris larva and, at least, two of its food plants must be re- garded as well known, now more than a century after the description of this species by Smith. We thank Dr. J. D. Lafontaine, Centre for Land and Biological Resources Research, Agriculture Canada, (Ottawa) for the commu- nication of very pertinent information allowing us to distinguish the adults of C. similaris from C. montanae. LITERATURE CITED Cook, W. C. 1930. An ecologically annoted list of the Phalenidae of Montana. Can. Entomol. 62:265-277. Crump, S. E. 1956. The larvae of the Phalaenidae. U.S. Dept. Agric. Tech. Bull. 1135:59-63. DETHIER, V. G. 1944. Observations on the life history of Cucullia asteroides Gn. Can. Entomol. 76:161—162 (black and white drawings of Cucullia asteroides). Hampson, G. F. 1906. Catalogue of the Lepidoptera Phalaenae in the British Museum, 6:1-689, pl. XCVII and XCVII. VOLUME 57, NUMBER 2 HITCHCOCK, C. L., A. CRONQUIST, M. OWNBEY & J. W. THOMPSON. 1955. Vascular plants of the Pacific Northwest. Part 5. Com- positae. University of Washington Press, Seattle. Munz, P. A. & D. D. Keck. 1973. A California flora. University of California Press, Berkley. POOLE, R. W. 1995. Noctuoidea, Noctuidae (part) in Dominick, R. B. et al. (ed.), The moths of America north of Mexico, fase. 26. 1. SEITZ, A. 1919-1944. Die Gro}-Smetterlinge der Erde. Abteilung II. Amerikanischen Faunengebietes, Band 7 Eulenartige Nachtfalter. Stuttgart. Alfred Kernen. 508 pp., 96 plates. STEHR, W. F. 1987. Immature insects. Kendall Hunt Publishing Company, Dubuque, Iowa, USA. p. 565 (Fig. 26.397, black and white photograph of Cucullia asteroides). J. C. Pemir AND M. C. Petit, 2 Rue du Maréchal Juin, F-45100 Orléans, France Email: jc.petit@ mageos. com Received for publication 21 February 2002; revised and accepted 25 November 2002. Journal of the Lepidopterists’ Society 57(2), 2003, 147-149 147 NOTE ADDED IN PRESS: While this manuscript was in press we have collected the following additional information. First, a photo showing the green form of Cucullia montanae’s larva upon Grindelia integrifolia D.C. taken by Jeremy B. Tatum, B.C., Canada, is avail- able on the web site entitled “Butterflies and moths of Southem Vancouver Island” at the address: http://alpha.furman.edu/~snyder/ snyder/lep/intern.htm. This is, to our knowledge, the first photo- graph of Cucullia montanae’s larva ever published. It also confirms the identity of the main food-plant. Second, according to M. Hre- blay and L. Ronkay: “The palearctic Cucullia ledereri Staudinger 1892, known from Kamchatka by its holotype female only”, has for “closest relative Cucullia similaris 1892, they may represent two dif- ferent populations of the same species!” This quote is from Moths of Nepal. Part 5. Tinea. Vol. 15 (supplement), pp. 174-175. In Tashiro Haruta (ed.). The Japan Heterocerist’s Society, Tokyo, 1998. A simi- lar view concerning the relationship between the two species is given in the Illustrated catalogue of Noctuidae in Korea by V. S. Kononenko, S. B. Ahn, L. Ronkay, Insects of Korea, Series 3, Park Kyu-Tek, Korea 1998. It will be interesting to find the male and the larva of Cucullia ledereri in order to know if they show any signifi- cant differences with Cucullia similaris. NOTES ON THE COMMON PALM BUTTERELY, ELYMNIAS HYPERMNESTRA UNDULARIS (DRURY) (SATYRINAE) IN INDIA Additional key words: Genitalia, toothed brachia, angular appendices, signa, genital plate. Hemming (1967) clarified that E. jynx Hiibner (=Papilio undularis Drury) is the type-species of the genus Elymnias Hiibner, which remained without a valid type-species for some time (Hemming 1943). Unlike other satyrines, palm butterflies often are brightly colored and generally resemble danaines, which they mimic in one or both the sexes. According to Bingham (1905), Evans (1932), Talbot (1947), Pin- ratana (1988), and Corbet and Pendlebury (1992), the species referable to the genus Elymnias differ from other satyrine genera in having a hind wing predis- coidal cell. Of the eleven species from India, three, ie., E. hypermnestra (Linnaeus), E. malelas (Hewitson), and E. patna (Westwood), have been reported from Northwest India. However, in recent surveys, only E. hypermnestra could be located and reexamined. This reexamination revealed that the male and female gen- italia possess certain unique taxonomic characteristics. The genitalia are described here, along with remarks on the distribution of the species. Elymnias hypermnestra undularis (Drury) Male genitalia (Figs. 1-5). Uncus long, slightly curved, longer than tegumen, distal end sharply pointed; brachia very thin, long, slender, upwardly tumed, distal end with minute teeth, strongly sclerotized; tegumen broader dorsally, narrower ventrally; appen- dices angulares long, broad proximally, narrow, hooked distally; vin- culum much longer than tegumen, slightly curved inwardly, broader in the middle; saccus short, tubular, distal end rounded; valva some- what boat-shaped, costa and sacculus not demarcated, harpe strongly sclerotized, narrow, knife-like, with inner margin dentate, pilose; juxta squarish plate-like, weakly sclerotized; aedeagus tubu- lar, slightly squeezed in the middle, subzone smaller than suprazone, ductus ejaculatorius entering dorsad. Female genitalia (Figs. 7, 8). Corpus bursae cylindrical, mem- branous; signa represented by two scobinate patches which run along whole length of corpus bursae; ductus bursae moderately long, broader anteriorly, narrower posteriorly; ductus seminalis originate from ductus bursae near base of corpus bursae; central process of lamella antevaginalis very small, roughly triangular, lateral flaps long, membranous except on their inner margin; lamella postvaginalis re- duced, with small oval plates; apophyses anterioris wanting; apophy- ses posterioris moderately long, slender, membranous; papilla analis guttiform, pilose. Length of forewing. Male: 34.0-36.0 mm (n = 10); Female: 40.0-42.0 mm (n = 5). Material examined. Himachal Pradesh: 4 d, 3 2, 1.xi.91, Paonta Sahib, 850 m, Sirmaur. Assam: 2 6, 2 9, 8.v.95, Vasistha, 213 m, Guwahati. Sikkim: 2 4, 30.ix.95, Rangpo, 600 m; 2 9, 4.x.95, Jorethang, 630 m. Remarks. Among fifty-four satyrine species for which the male genitalia have been examined, certain structures, such as toothed brachia and angular appen- dices, are unique to E. hypermnestra. Similarly, the fe- male genitalia have a unique signa and genital plate, both conspicuous structures not encountered in any other satyrine examined so far. The account of the male and the female genitalia are described for the first time. 148 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 1-7. Elymnias hypermnestra undularis (Drury). 1, Male genitalia (lateral view). 2, Valva (inner view). 3, Juxta. 4, Aedeagus (lateral view). 5, Aedeagus (dorsal view). 6, Female genitalia (ventral view). 7, Corpus bursae (dorsal view). Abbreviations: AED: Aedeagus, APX.ANG: Appendix angulares, BR: Brachium, CO: Costa, CRP.BU: Corpus bursae, DU.BU: Ductus burse, DU.E]J: Ductus ejaculatorius, DU.SEM: Duc- tus seminalis, LA.AV: Lamella antevaginalis, LA.PV: Lamella postvaginalis, O.B: Ostium bursae, P. A: Papilla analis, PO.APO: Apophysis poste- rioris, SA: Saccus, SBZ: Subzonal portion of aedeagus, SIG: Signum, SL: Sacculus, SPZ,: Suprazonal portion of aedeagus, TEG: Tegumen, UN: Uncus, VIN: Vinculum, VLV: Valva. In addition to the genitalic characteristics, it is ob- served that the hind wing prediscoidal cell has an addi- tional prominent vein. An obscure black androconial patch near the base of the forewing space 1A+2A, above, reported for E. hypermnestra (Corbet & Pendlebury 1992), is lacking in E. hypermnestra undu- laris. As well, there is a nacreous area on the forewing underside and another on the dorsal surface of the hind wing costal margin, which also has a pair of hair tufts, all of which agree with the observations made by Pin- ratana (1988) and Corbet and Pendlebury (1992). Mackinnon and de Niceville (1897), while reporting on this species from North West India (Mussoorie and Dehradun, below 909 m ASL) remarked that it is not a common species in this area. Though no specimens could be collected from the localities mentioned above, four males and three females were collected from Paonta Sahib (399 m ASL), forty-two kms from Dehradun. Marshall and de Niceville (1883) reported that E. undularis is the common Elymnias in North West India, where it is found in the warm valleys of the outer Himalayas as far east as Mussoorie. Contrary to Wynter-Blyth (1957), our surveys indicate that the species is not common in North India. Females mimic Danaus plexippus (Linnaeus) and D. chrysippus (Lin- naeus) at the above mentioned localities. VOLUME 57, NUMBER 2 The author is grateful to the Indian Council of Agricultural Re- search, New Delhi for funding the project on Butterflies. LITERATURE CITED BincuaM, C. T. 1905. Butterflies. Fauna Br. India, l:iv 511, pls. 1-10. CorseET, A. S. & H. M. PENDLEBURY. 1992. The butterflies of the Malaya Peninsula. Eliot, J. N. (ed.). 4th ed. Malaya Nat. Soc.:viii + 595 pp., 69 pls. Evans, W. H. 1932. The identification of Indian butterflies. 2nd ed. revised. Madras, Bombay nat. Hist. Soc., 1932:x + 454 pp., 32 pls., 9 figs. HEMMING, A. F. 1943. The types of genera established by Double- day (E.) in the genera of diurnal Lepidoptera and by Westwood (J.O.) in the continuation thereof: a correction regarding the genus Corybas Westwood, (1850). J. Soc. Bibl. Nat. Hist. 1(12):470. . 1967. The generic names of the butterflies and their type species (Lepidoptera: Rhopalocera). Bull. Br. Mus. Nat. Hist. (Entomol.) Suppl. 9:509. MACKINNON, P. W. & L. DE NICEVILLE. 1897. List of butterflies from Mussoorie and Dun Valley. J. Bombay Nat. Hist. Soc. 11:205-221, 368-389, 585-603. Journdl of the Lepidopterists’ Society 57(2), 2003, 149-150 149 MARSHALL, G. F. L. & L. DE NICEVILLE. 1883. The butterflies of In- dia, Burmah and Ceylon. A descriptive handbook of all the known species of Rhopalocerous Lepidoptera inhabiting that re- gion, with notices of allied species occurring in the neighbouring countries along the border; with numerous illustrations. Vol. 1. Part II. Calcutta Central Press, Calcutta. vii + 327 pp. PinraTANA, B. A. 1998. Butterflies in Thailand, Satyridae, Libythei- dae and Riodinidae. The Viratham Press, Bangkok. 6:vii + 61 pp., 44 pls. TaLBoT, G. 1947. The fauna of British India. Including Ceylon and Burma. Butterflies. Vol. 2. Taylor and Francis, London, 506 pp. WynTER-BLYTH, M. A. 1957. Butterflies of the Indian region. Bom- bay Nat. Hist. Soc., 1957:xx + 523 pp., 72 pls. NARENDER SHARMA, Department of Entomology and Apiculture, Dr. Y.S. Parmar University of Horti- culture and Forestry, Nauni, Solan 173 230 Himachal Pradesh, India Received for publication 14 October 2002; revised and accepted 3 December 2002. OXYPOLIS RIGIDIOR, A NEW LARVAL FOOD PLANT RECORD FOR PAPILIO POLYXENES (PAPILIONIDAE) Additional key words: black swallowtail, Wisconsin. Oxypolis rigidior (L.) Raf. is yet another larval food plant in the family Apiaceae for Papilio polyxenes Fabr. (Papilionidae). This Nearctic swallowtail has long been known to develop on various native and exotic species in Apiaceae now found in its range (Scudder 1889, Scott 1986). The genus Oxypolis has been reported in this context, with O. filiformis (Walt.) Britt. (Tietz 1952) and O. canbyi (Coult. & Rose) Fern. (Scott 1986) included in lists of suitable food plants. These two species grow in the southeastern United States (Mathias & Constance 1944-45). Oxypolis rigidior is a native species that grows in swamps, marshes, ditches and wet prairies from coastal New York to Minnesota, south to Florida and Texas (Gleason & Cronquist 1991). Fifteen caterpillars were collected from O. rigidior inflorescences bearing young fruits. These included second, third and fourth instars, taken at 3 sites in Grant, Juneau and Marquette Counties, in southern Wisconsin, in early September, 1999 and 2001. These sites support native, wet prairie vegetation as defined by Curtis (1959). Caterpillars were reared to pupation on developing fruits of O. rigidior in the lab; though foliage was also provided, it was scarcely eaten. Pupae were caged in a garage over winter and then returned to the lab. One caterpillar died, 2 pupae died, 10 pu- pae each yielded single adults of Trogus pennator (Fabr.) (Ichneumonidae) and 2 pupae yielded adults of P. polyxenes asterius Stoll. The exotics Daucus carota L. and Pastinaca sativa L.., both ubiquitous along roadsides throughout southern Wisconsin, are also suitable to these larvae (Scudder 1889). I have reared Wisconsin larvae, taken off these plants, on their foliage. In response to roadside mowing, these exotics may provide forage well into autumn. But in the historically natural regime of these wet prairies, O. rigidior provides forage later in the year than do other suitable native plants on these 3 sites—Cicuta maculata L., Heracleum lanatum Michx., Sium suave Walter and Zizia aurea (L.) Koch. (Scott 1986). Voucher specimens are in the Insect Research Collection of the University of Wisconsin—Madison. I thank Dan Young and Mike An- derson for donating space in which rearing could be done, John Luhman for determining the wasps and J. Mark Scriber and an anonymous second reviewer. 150 LITERATURE CITED Curtis, J. T. 1959. Vegetation of Wisconsin. University of Wiscon- sin Press, Madison, Wisconsin. 657 pp: GLEASON, H. A. & A. CRONQuIST. 1991. Manual of vascular plants of northeastern United States and adjacent Canada. 2nd ed. New York Botanical Garden, Bronx, New York. 910 pp. Matuias, M. E. & L. ConsTaNcE. 1944-45. Family 2. Umbellif- erae, pp. 43-295. In Rickett, H. W. (ed.), North American flora. Vol. 28B. Umbellales, Commales. New York Botanical Garden, Bronx, New York. Scott, J. A. 1986. The butterflies of North America. Stanford Uni- versity Press, Stanford, California. 583 pp. SCUDDER, S. H. 1889. The butterflies of the eastern United States and Canada with special reference to New England. Vol. 2. Publ. by the author, Cambridge, Massachusetts. 1774 pp. Journal of the Lepidopterists’ Society 57(2), 2003, 150-152 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TiETZ, H. M. 1952. The Lepidoptera of Pennsylvania, a manual. Penn. State College School of Agriculture, Agric. Expt. Station, State College, Pennsylvania. 194 pp. ANDREW H. WILLIAMS, Dept. of Entomology, Uni- versity of Wisconsin, Madison, Wisconsin 53706, USA. Email: awilliam@facstaff.wisc.edu. Received for publication 29 March 2002; revised and accepted 4 De- cember 2002. THE CORRECT TYPE LOCALITY OF CYANIRIS LADON VAR. QUESNELLITI COCKLE, 1910 (LYCAENIDAE), WITH DESIGNATION OF A LECTOTYPE Additional key words: Celastrina, nigrescens, lucia, British Columbia. The original description of “Cyaniris ladon, Cramer, var. Quesnellii” was based on two specimens taken “at Bala Lake, Quesnelle, northern B.C.” (Cockle 1910). Cockle also stated that he thought it would “prove a lo- cal race which will be found abundant in the Ques- nelle Valley”. We recently had the opportunity to ex- amine the type specimens of these butterflies in the Canadian National Collection of Insects and Arthro- pods (Agriculture Canada, Ottawa, Ontario, Canada). The two specimens and their data labels are shown in Fig. 1, with lectotype and paralectotype designations provided below. The two specimen data labels are in different hand- writings. “J.M. Anderson” on the paralectotype label is written as if it is a signature and the date is written in full. The label on the lectotype is printed, the date uses Roman numbers for the month, and part of the data on the other label is omitted. This suggests that J. M. Anderson wrote the paralectotype label, and someone else wrote the other when the specimens were pinned. Dr. Fletcher is more likely than Cockle for the second label, because of Cockle’s error in reading “Aubau” Lake as “At Bala” Lake (below). The spelling of the lake name on the label of the paralectotype can readily be seen to be “Au Baw” Lake, with the alternative name of “Graveyard Lake”. “Ah” is “Mr.” in Chinese, hence the lake name referred to the Chinese Mr. Baw or Bau (alternative spellings). For many years he prospected and worked gold claims on and around what are now known as Ahbau Creek and Ahbau Lake in the summer, and trapped in the area during the winter. Apparently Cockle misread “Au Baw” as “At Bala”. Ahbau Creek was labeled on maps as Graveyard Creek until 1921 (Janet Mason pers. com.), hence the alternative name Graveyard Lake. Ahbau Lake is about 40 km (25 miles) northeast of the modern town of Quesnel, apparently contradict- ing the “35 miles N.W.” indicated on the specimen la- bel. However, Ahbau Lake is 35 miles northwest of Quesnelle Forks, a settlement (now historic site) at the junction of the Cariboo River with the Quesnel River. Ahbau Lake is at elevation 2950 feet, not 2480 feet, but such errors in elevation were common at that time. Ahbau Lake is not in the Quesnel River valley, as implied by Cockle, and is in what is now considered to be central, rather than northern, British Columbia (“northern” is of course a relative term). Ahbau Creek is part of the Cottonwood/Swift River watershed, the watershed immediately north of the Quesnel River watershed. The correct type locality is therefore “[Ah- bau] Lake, [elevation 2950 feet], [latitude 53°14’, lon- gitude 122°07’,] 35 miles northwest of Quesnelle [Forks], B.C.[, Canada]”, with interpolated and cor- rected data shown in brackets and the coordinates be- ing for the outlet at the south end of the lake. There is a second locality label attached to one spec- imen, specifying Kaslo as the collection site. The date on this label is in a different handwriting than the date on the other two data labels, indicating that a third person wrote it. Celastrina ladon lucia (the true iden- VOLUME 57, NUMBER 2 151 Fic. 1. The type specimens of Cyaniris ladon, Cramer, var. Quesnellii Cockle, 1910. tity of the types, see Fig. 1) does not occur near Kaslo, so the label must be in error. Perhaps it was intended to indicate that the specimens were part of Cockle’s collection (Cockle lived in Kaslo). This extra label may have contributed to the erroneous association of the name quesnellii with nigrescens Fletcher, 1903, which is discussed below. The two specimens from which Cockle described quesnellii had labels indicating J.M. Anderson col- lected them on 8 June 1907. One of the labels reads “Cyaniris Quesnellii Type F & C.” The designation of “Type” on this label has no bearing on the question of the type status of the specimen, even though Cockle may have written the label. The International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature 1999) requires that the designation of a type must occur in the original de- scription, and use of the word “type” on a specimen la- bel does not make that specimen the holotype (Article 72.4.7). The two specimens are syntypes, rather than having the status of holotype and paratype as indicated on the existing specimen labels, because Cockle did not specify a single “type” or “holotype” in the original description. Accordingly, under Article 74 of the Inter- national Code of Zoological Nomenclature we hereby designate one specimen (the one with the existing “type” label) as the lectotype and the other as the paralectotype, as shown in Fig. 1. The taxonomic pur- pose of this lectotype designation (ICZN Article 74.7.3) is to clarify that the name quesnellii is correctly associated with lucia, rather than with nigrescens, and to provide future opportunity to determine whether quesnellii is correctly placed as a synonym of lucia Kirby, 1837. Also of interest is the phrase “F & C”. This indicates that Cockle (or the person who wrote the label) con- sidered quesnellii to have been described by two people, with the initials presumably being an abbrevi- ation of “F|letcher] & C[ockle]”. Cockle had submit- ted the specimens to “the late Dr. Fletcher”, who had provided comments on them, but the original descrip- tion is clearly that of Cockle alone and hence Cockle is the sole author. Perhaps Cockle wrote the labels while Dr. Fletcher was still alive, with the intention that they would co-author the description, but then assumed sole authorship after Dr. Fletcher's death. Blackmore (1920) lists Lycaenopsis pseudargiolus race nigrescens form quesnelli [sic]. McDunnough (1938) follows Blackmore in listing “form quesnelii [sic] Cockle” under “Lycaenopsis pseudargiolus nigrescens. , with “maculata-suffusa Cockle” as an infrasubspecific synonym. Comstock and Huntington (1963) list ques- nellii with the correct spelling, and cite McDunnough’s taxonomic placement. Dos Passos (1964) apparently copied McDunnough (1938) in placing “form quesnelii [sic] (Cockle), 1910” as a synonym of Celastrina argio- lus nigrescens (Fletcher), 1903. The listings by Black- more, McDunnough and dos Passos had several errors. First, they use two incorrect spellings of the taxon name. Second, quesnellii was clearly described not as a form but as a geographically defined variety (=sub- species). This is indicated by Cockle’s statement “there is every reason to think that if this variety is found to be (as I think) a distinct local race, it should be entitled to 152 a specific name”. Hence quesnellii is an available species-group name under the International Code of Zoological Nomenclature (1999). Third, the type spec- imens, and all the numerous specimens of Celastrina that Guppy has collected in the vicinity of Quesnel, are clearly referable to lucia (Kirby), 1837 and not to ni- grescens (Guppy collected the nearest nigrescens 120 km south of Quesnel at Williams Lake in 2002). Miller and Brown (1981) repeated the error of placing ques- nellii as a synonym of nigrescens rather than lucia, but corrected the spelling and correctly treated the name as an available species-group name. Guppy and Shep- ard (2001) placed quesnellii as a synonym of C. ladon lucia, and abbreviated the type locality to “Quesnel, B.C.” because at the time Guppy had not seen the specimen labels and hence could not determine the lo- cation of “Bala Lake”. An additional name is mentioned by Cockle (1910), in the sentence “I submitted them [the specimens of quesnellii| to the late Dr. Fletcher, who wrote me that, had they been taken in Ontario, he would have named them ‘maculata-suffusa .” Clearly this name is not be- ing formally applied to the specimens in question, not even by Dr. Fletcher. It is clear that Cockle used the name quesnellii instead of the name maculata-suffusa, not in addition to that name. McDunnough (1938), Dos Passos (1964) and Miller and Brown (1981) were in error to list “maculatasuffusa (Cockle)” as a syn- onym of quesnellii. The name maculatasuffusa has no standing even as an infrasubspecific name, and should Journal of the Lepidopterists’ Society 57(2), 2003, 152-153 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY be omitted from checklists and other publications. We thank Janet Mason, Provincial Toponymist, Base Mapping & Geomatic Services Branch, BC Ministry of Sustainable Resource Management for information on the historical names of Ahbau Creek and Ahbau Lake, and the suggestion that “Quesnelle” may re- fer to Quesnelle Forks. LITERATURE CITED BLACKMORE, E. H. 1920. The Lycaenidae of British Columbia. Proceedings of the Entomological Society of British Columbia (1919)14:5-11. COCKLE, J. W. 1910. Notes on a few butterflies found at Kaslo and in northern British Columbia. Can. Entomol. 42(6):203-204. Comstock, W. P. & E. I. HUNTINGTON. 1963. An annotated list of the Lycaenidae (Lepidoptera: Rhopalocera) of the Western Hemisphere (continued). J. New York Entomol. Soc. 21:45-57. Dos Passos, C. F. 1964. A synonymic list of the Nearctic Rhopalo- cera. Lep. Soc. Mem. 1:i-v, 1-145. Guppy, C. S. & J. H. SHEPARD. 2001. Butterflies of British Colum- bia. UBC Press Vancouver, B.C. 414 pp: INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE. 1999. International code of zoological nomenclature. 4th ed. The International Trust for Zoological Nomenclature, London. xxix + 306 pp. McDunnoucu, J. H. 1938. Check list of the Lepidoptera of Canada and the United States of America. Part 1. Macrolepi- doptera. Mem. Calif. Acad. Sci. 1:3-272 (1-3 (corrigenda)). MILLER, L. D. & F. M. Brown. 1981. A catalogue/checklist of the butterflies of America north of Mexico. Lepid. Soc. Mem. 2:i-vii, 1-280. CrisPIN S. Guppy, 4627 Quesnel-Hydraulic Road, Quesnel, BC V2] 6P8 Canada AND NORBERT G. KONDLA, P.O. Box 244, Genelle, BC VOG 1G0, Canada Received for publication 5 March 2002; revised and accepted 20 September 2002. FIRST REPORT OF THE PALAEARCTIC DICHRORAMPHA ACUMINATANA (LIENIG & ZELLER) INNORTH AMERICA (TORTRICIDAE) Additional key words: immigrant, holarctic, Olethreutinae, Dichrorampha petiverella, D. vancouverana. In the course of an ongoing inventory of the moths of Steuben, Washington Co., Maine, a single specimen of the Old World olethreutine Dichrorampha acumi- natana (Lienig & Zeller) was captured in 2001, evi- dently a first record for North America. The specimen, a fresh male (Figs. 1, 3), was taken on a door screen at approximately 1600 h EDST on 15 June at 44°30'22”N, 67°59/28”W. Nothing is known of its ori- gins, but as a reported root feeder on Chrysanthemum leucanthemum L. and C. segetum L. (Asteraceae) (Bentinck & Diakonoff 1968, Kuznetsov 1987), it can be presumed to have developed on naturalized food- plants present within 1-2 km of the collection site. Initial identification of the specimen was based on figures of wings and genitalia in Bentinck and Diakonoff (1968) and Kuznetsov (1987), and con- firmed by comparison with authentic Palaearctic spec- imens listed below. The species is distinguished from similar Nearctic forms by the acuminate shape of its forewing (signalized in its name), the continuous pale band in its terminal fringe, its diffuse dorsal patch, its broad cucullus with blunt ventral cusp, and its bifid aedeagus terminating in a distinctive open trough (Figs. 1-4). It belongs in the nominate subgenus in lacking anellar lobes but possessing a male forewing costal fold. VOLUME 57, NUMBER 2 Fics. 1-4. Dichrorampha acuminatana. 1, Wings of male from Steuben, ME. 2, Wings of male from Apetlon, Austria. 3, Genitalia of male from Steuben, ME. 4, Genitalia of male from Apetlon, Austria. Further details are in the Specimens examined section of the text. The species is widely distributed in western and central Europe (Razowski 1996). Two Palaearctic con- geners, D. vancouverana McDunnough (=D. gue- neeana Obraztsov) and D. petiverella (L.), were previ- ously reported in Maine (Roberts 1991), and subsequent collecting there has revealed well estab- lished populations of these species along the immedi- ate coastline wherever undisturbed stands of their na- tive or naturalized foodplant Achillea millefolium L. (Asteraceae) occur. With captures of D. vancouverana in the Pacific Northwest (Miller 1999), coastal distri- bution patterns of the two holarctic congeners con- tinue to suggest they are immigrants, although the possibility cannot be ruled out that they represent spotty relicts of circumpolar distributions. Specimens examined. 4, Steuben, ME (Fig. 1), M. A. Roberts, 15/06/2001, genit. slide prep. MAR2027M (Fig. 3), forewing length 7.0 mm, in M. A. Roberts collection, Steuben, ME; 3, Wangeroog, Ostfries. Inseln [Germany], 07/09/1949, E. Jiackh, genit. prep. on pin, forewing length 6.0 mm; ¢, Kel- heim, Obfrk. [Germany], 03/08/1952, Jackh, genit. prep. on pin, forewing length 6.0 mm; d, Hannover, Misb Moor [Germany], 29/05/1931, genit. slide prep. WEM 612011, forewing length 6.5 mm; ¢, Apetlon, Burgenland [Austria] (Fig. 2), 11/09/1971, E. Jackh, Journal of the Lepidopterists’ Society 57(2), 2003, 153-156 genit. slide prep. WEM612012 (Fig. 4), forewing length 5.5 mm. The four Palaearctic specimens are in the U.S. National Museum of Natural History (USNM), Washington, D.C.; we thank J. W. Brown for loaning them. LITERATURE CITED BENTINCK, G. A. & A. DtaKonorr. 1968. De Nederlandse bladrollers (Tortricidae). Monogr. Nederlands. Entomol. Ver. No. 3. 201 pp. Kuznetsov, V. I. 1987. Family Tortricidae (tortricid moths), pp. 279-956. In Medvedey, G. S. (ed.), Keys to the insects of the European part of the USSR. U.S. Dept. of Agr. & Natl. Science Found. 991 pp. [translation] MILLER, W. E. 1999. A new synonymy in Dichrorampha that re- veals an overlooked immigrant record for North America (Tor- tricidae). J. Lepid. Soc. 53:74-75. RAZowsKI, J. 1996. Tortricidae, pp. 130-157. In Karsholt, O. & J. Razowski (eds.), The Lepidoptera of Europe: a distributional checklist. Apollo Books, Stenstrup. 380 pp. Roserts, M. A. 1991. Two Palearctic species of Dichrorampha dis- covered in Maine (Tortricidae). J. Lepid. Soc. 45:169-171. MICHAEL A. ROBERTS, 367 Village Road, Steuben, Maine 04680, USA. Email: maroberts@acadia.net AND WILLIAM E. MILLER, Department of Entomology, Uni- versity of Minnesota, St. Paul, Minnesota 55108, USA. Email: milleO14@umn.edu Received for publication 8 April 2002; revised and accepted 11 No- vember 2002. HOST PLANT ASSOCIATIONS OF WESTERN SPECIES OF PAPAIPEMA (NOCTUIDAE) WITH PARTICULAR REFERENCE TO THE APIACEOUS PLANTS Additional key words: host plants, Apiaceae, Asteraceae, polyphagy. The genus Papaipema Smith (Noctuidae) is the largest noctuid genus endemic to North America has long been a favorite among students of lepidopteran life history (e.g., Kwiat 1916, Hessel 1954). With 46 described species and at least 5 undescribed species of which we are, Papaipema is the fifth most speciose noctuid genus on this continent (Hodges 1983), super- seded only by the Holarctic genera Acronicta Ochs. (n = 81 Nearctic species), Catocala Schrank (n = 110), Lacinipolia McDunnough (n = 57), and Schinia Hiib- ner (n = 123 species in North America) (Hodges 1983). Papaipema currently includes 46 valid de- scribed species, at least five undescribed species (Quinter, in MS), and two valid subspecific entities, 154 Papaipema baptisiae baptisiae (Bird) and Papaipema b. limata Bird (E.L. Quinter, in Hodges 1983). Papaipema and its relatives form a putatively mono- phyletic clade of endophagous plant borers in the Apameini (sensu Hodges 1983). Whereas most of the species in this tribe are associated with monocotyledo- nous plants, species of Papaipema feed and specialize on members of between 22 and 25 plant families (Goldstein 1999). Though well studied, a number of questions remain concerning host plant associations in this group, especially among the relatively few western species (Papaipema attains its highest regional diver- sity in the eastern United States). In this paper, we present life history data based on recent collecting and rearing efforts for species belonging to the Papaipema birdi (Dyar 1908) and Papaipema harrisi Grote species complexes, and summarize the known host as- sociations for the remaining western Papaipema species and those associated with Apiaceae regardless of geography. Our observations bear on the evolution of umbellifer-feeding in Lepidoptera, and Papaipema in particular and possibly the role of coumarin com- pounds in mediating the evolution of host association (e.g., Berenbaum 1981, 1983). We also discuss collect- ing and rearing efforts on eastern umbel-feeding Pa- paipema species. All larvae encountered were reared on artificial diet, and adult specimens deposited at AMNH and FMNH. Apiaceous host plant records for Papaipema species. A few species of Papaipema are known to feed on apiaceous plants, and although at least three of these (P. birdi, P. harrisii and P. eryngii Bird) appear to be specialists on Apiaceae, the others exhibit a broader range of apiaceous and non-apiaceous host use. The P. birdi complex includes an eastern species (P. birdi) and two western species (P. pertincta Dyar and P. in- sulidens |Bird]), all of which are associated with the Apiaceae (=Umbelliferae). Papaipema birdi has been considered oligophagous specialist on apiaceous plants, its primary host being the water hemlock Cicuta maculata L. Prior to the present study, other host records included Sium suave Walt. (Apiaceae), and “other umbellates” (Hessel 1954:60; treating P birdi as a synonym of P. marginidens, of which there are no known host records), as well as several astera- ceous plants (Kwiat 1916). The two other species in the P. birdi complex, P. pertincta and P. insulidens, each of which have been recorded from both apia- ceous and non-apiaceous plants, are western species apparently separated by the Cascade Mountains, with P. pertincta to the west and P. insulidens to the east. A host of P. insulidens was described by Bird (1921, 1931) as a species of Senecio (Asteraceae). In the field JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY notes of his son (archived at the American Museum of Natural History), the late Junius Bird, the host plant was described as a “large, Dill-like weed,” suggesting an apiaceous plant. The published association of P. per- tincta with Lupinus polyphyllus Lindley (Fabaceae) (see Bird 1926) is curious because only two other Pa- paipema species are associated with fabaceous plants: the western Papaipema angelica with Psoralea macrostachya DC., and the eastern Papaipema bap- tisiae with Baptisia tinctoria (L.). Outside the Papaipema birdi complex, umbellifer- feeding occurs in Papaipema eryngii, a threatened species restricted to prairie wetlands where it special- izes on Eryngium yuccifolium, and in the P. harrisi group, comprising P. harrisi and P. verona Smith). Host records for P. harrisi, whose distribution suggest an association with Heracleum lanatum Michx. (Api- aceae) along the Atlantic Coast and an association with Angelica atropurpurea L. (Apiaceae) westward follow- ing the Great Lakes (Kwiat 1916, Hessel 1954, Jones & Kimball 1943, Quinter unpublished data). Both of these host species are apiaceous plants. In the North- east, it is thought that P. birdi and P. harrisi segregate themselves according to host plant, with P. birdi con- fining itself to Cicuta maculata and P. harrisi to Angel- ica atropurpurea (see Kwiat 1916, Hessel 1954). Pa- paipema verona, for which we do not report novel host records, is a western species recorded primarily from species of the umbel genus Heracleum. Recent field collections. During 1995, we exam- ined several eastern USA sites for larvae of both P. birdi and P. harrisi. Visits to wetlands in western Con- necticut and Massachusetts with dense populations of Angelica atropurpurea yielded only two Papaipema larvae (Papaipema harrisi has become decidedly rare in New England and is considered extirpated from Massachusetts). However, visits to a calcareous sedge- meadow complex in Otsego County, New York yielded more than two dozen Papaipema larvae from both An- gelica atropurpurea and Cicuta maculata. All larvae collected from C. maculata and A. atropurpurea at the upstate New York site proved to be P. birdi. Although reports of “other umbellates” than Cicuta maculata and Sium suave occur in the literature (e.g., Kwiat 1916), our collections appear to be the first documen- tation of Angelica atropurpurea as a host for P. birdi. Although Kwiat (1916) reported non-apiaceous hosts for P. birdi, it is conceivable that the taxonomic confu- sion that typically surrounds Papaipema has resulted in erroneous reporting of hosts subsequent to that publi- cation. Our findings in the northwestern United States ex- tended the known host ranges of P. pertincta and P. in- VOLUME 57, NUMBER 2 155 TABLE 1. Collecting information and host associations of western Papaipema species discovered during this study. Species Locality Oregon: Tillamook Co.: Rt. 101, 1 mi S. of Wheeler Oregon: Clatsop Co.: Rt. 101, 8 mi. S. of Astoria (at jet. Rts. 101 & 30) Life stage Host plant Dates P. pertincta 4 larvae Heracleum maximum 8 July 1995 9 larvae, 1 pupa Cicuta douglasii 8 July 1995 Oregon Tillamook Co.: Rt. 101, 8 larvae Senecio vulgaris 9-10 July 1995 1-2 mi. N. of Manzanita 8 larvae Heracleum maximum 4 larvae Ligusticum apifolium 1 larva Daucus sp. 8 larvae Cirsium sp. 3 larvae Erechtites minima Oregon: Lincoln Co:, E. Devil’s 1 larva Cicuta douglasii 11 July 1995 Lake Rd., 0.7 mi E. of Jct. Rt. 101 Oregon: Tillamook Co.: Rt. 101, 12S. of Jet. Rt. 22 6 larvae, 1 pupa Heracleum maximum 11 July 1995 Heracleum maximum Oregon: Lincoln Co: Three Rocks 1 larva Rd., 1.5 mi W. Rt. 101 P. sauzalitae Rd., 1.5 mi W. Rt. 101 Washington: Whitman Co: Steptoe Butte, el. 2500’—3000’ P. insulidens sulidens. These represent the least well-known Pa- paipema species for which published host records ex- ist; their close resemblance to P. birdi as well as the in- formal description by Junius Bird of a “large, dill-like” host for P. insulidens suggested that apiaceous plants might fall within the host spectra of one or both of these two western species in the birdi complex. An ad- ditional southwestern species, Papaipema angelica Smith, 1899 has remained uncollected in recent decades despite our knowledge of its life history and host affiliation (Bird 1931). Although several dozen specimens of P. pertincta exist in the Oregon State University insect collection (which now includes the private collection of the late Elmer Griepentrog), we have been unable to verify the association of P. pert- incta with any species of Lupinus. We thoroughly ex- amined the botanical holdings at OSU to identify sites likely to support strong populations of various western lupines, but we found no Papaipema at any of these, and apparently no western collectors have observed or reared P. pertincta from Lupinus since Bird’s (1926) second-hand account of the association. However, sev- eral dozen P. pertincta were reared from a variety of plants, mostly apiaceous, at six sites in Tillamook, Clat- sop, and Lincoln counties (Table 1). Like its eastern relative P. birdi, P. pertincta appears to feed primarily in apiaceous plants; but unlike its eastern associate, P. pertincta also feeds in non-apiaceous plants. In three weeks of field work in eastern Washington and Idaho, we failed to collect P. insulidens from its recorded host, Senecio hydrophilus Nutt. We exam- ined sites suggested by the literature, museum labels, and the hand-written field notes and sketches of Ju- Oregon: Lincoln Co: Three Rocks 1 larva 13 larvae 13 July 1995 Cirsium sp. 13 July 1995 Heracleum maximum 15 July 1995 nius Bird indicating large apiaceous host plants in Whitman County, Washington. Near localities visited by Junius Bird, several stands of Conium maculatum L. (Apiaceae), which matched his description and sketch, were checked without success. Thirteen larvae of P. insulidens were discovered and reared from Her- acleum maximum Bartr. (Apiaceae) at Steptoe Butte at an elevation of 2500-3000’. Although we could not ver- ify many published host associations of the two western members of the Papaipema birdi complex, we did take them on other hosts, apiaceous and otherwise. One possibility is that the dill-like host plant of P. insulidens referred to by Junius Bird was not apiaceous at all, but the introduced tansy ragwort Senecio vulgaris, one of the host plants from which we reared P. pertincta. The reported host associations of the western P. sauzalitae (Grote) are atypically diverse for Pa- paipema, and include members of the asteraceous plant genera Arctium, Cirsium, and Cynara as well as Castilleja (Scrophulariaceae), and Rumex (Polygo- naceae) (Crumb 1956). Peter McEvoy (pers. com.) of Oregon State University reports an association of P. sauzalitae with Senecio (Asteraceae) as well. Our collecting efforts yielded but a single specimen, from the exotic Cirsium vulgaris. However, California ma- terial at the Essig Museum includes specimens from In- verness Park (Marin Co.) where larvae were observed in Heracleum maximum, Artemisia douglasiana Bess. in Hook. (Asteraceae), and Ribes sp. (Grossulariaceae), suggesting that P. sauzalitae may be polyphagous (J. Powell pers. com.). If this is the case, the member species of each of the primary umbellifer-feeding Pa- paipema species groups (the harrisi-verona-sauzalitae 156 complex and the birdi-pertincta-insulidens complex), have broadened their host usage to include both com- posites and umbels on the west coast. The association of P. pertincta with apiaceous and non-apiaceous plants is noteworthy for two reasons. First, this species parallels P. insulidens in having a wider range of non-apiaceous recorded hosts than ex- pected, given the apparently tighter associations of their eastern relatives (P. birdi and P. sauzalitae, re- spectively) with umbels. Second, based on available DNA sequence data, P. pertincta is nearly indistin- guishable from P. birdi (Goldstein 1999). We can not, therefore, rule out the possibility of P. pertincta’s feed- ing facultatively on Lupinus, though we were unable to recover larvae from any fabaceous plants and it is clear that the species is not thus restricted. Although both facultative and obligate association with asteraceous plants is common among Papaipema species, umbel-feeding is less common. The parallel variation in host breadth among umbel-feeding Pa- paipema species is thus noteworthy, and suggests a profitable line of inquiry for further work on host use specialization in this group. We thank P. McEvoy, E. Griepentrog, and J. Lattin for access to specimen material and assistance in locating collecting sites. J. Pow- ell loaned material from the Essig Museum and shared his observa- tions based on collecting and rearing efforts. We thank Kenneth and Elaine Bernard for lodging and permission to collect in their farm in Otsego Co., NY. This work was supported by an NSF doctoral dis- sertation improvement grant, an EPA graduate fellowship, and an AMNH Theodore Roosevelt Memorial Fund travel grant to PZG. LITERATURE CITED BERENBAUM, M. 1981. Patterns of furanocoumarin distribution and insect herbivory in the Umbelliferae: plant chemistry and com- munity structure. Ecology 62:1254-1266. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY . 1983. Coumarins and caterpillars: a case for coevolution. Evolution 37:163-179. BirD, H. 1921. New species and life histories in Papaipema Sm. (Lepidoptera). No. 20 Can. Entomol. 53:79-81. . 1926. New life histories and notes in Papaipema (Lepi- doptera). No. 24. Can. Entomol. 58:249-284. . 1931. The Papaipema species of the Pacific Coast (Lepi- doptera). Can. Entomol. 63:183-187. Crump, S. E. 1956. The larvae of the Phalaenidae. U.S. Dept. Agric. Tech. Bull., no. 1135. U.S. Government Printing Office, Washington, D.C. 356 pp. GOLDSTEIN, P. Z. 1999. Molecular systematics and the macroevolu- tion of host plant use in the endophagous moth genus Papaipema Smith 1899. Ph.D. Dissertation, University of Connecticut. HESSEL, S. A. 1954. A guide to collecting the plant-boring larvae of the genus Papaipema (Noctuidae). Lepid. News 8:57-63. Hopces, R. W. et al. (eds.). 1983. Check list of the Lepidoptera of America north of Mexico. London, E. W. Classey Limited and The Wedge Entomological Research Foundation. 284 pp. Jones, F. M. & C. P. KIMBALL. 1943. The Lepidoptera of Nan- tucket and Martha's Vineyard Islands, Massachusetts. Publica- tions of the Nantucket Maria Mitchell Association IV. Nan- tucket. 217 pp. Kwiat, A. [=A. K. Wyart]. 1916. Collecting Papaipemae (Lep.). Entomol. News 27:228-234. QuinTER, E. L. 1983. Papaipema, pp. 138-139 (in part). In Hodges, R. W. et al. (eds.), Check list of the Lepidoptera of America north of Mexico. London, E. W. Classey Limited and The Wedge Entomological Research Foundation. PAUL Z. GOLDSTEIN, Division of Insects, Field Mu- seum of Natural History, 1400 S. Lake Shore Drive, Chicago, Illinois 60605, USA and Committee on Evo- lutionary Biology, University of Chicago, Culver 401, 1025 East 57th Street, Chicago, Illinois 60637, USA AND ERIC L. QUINTER, Division of Invertebrate Zool- ogy, American Museum of Natural History, Central Park West @ 79th, New York, New York 10024, USA Received for publication 25 August 2002; revised and accepted 20 December 2002. BOOK REVIEWS Journal of the Lepidopterists’ Society 57(2), 2003, 157 THE BUTTERFLIES OF CASCADIA, by R. M. Pyle, Seattle Audubon Society; Cloth ISBN: 0-914516-13-2; Price: $29.95 Bob Pyle has produced another book detailing his love affair with butterflies. The Butterflies of Cascadia, is a newly eclosed field guide derived from long ram- bles in those emerald mountains, boreal meadows, and rocky fields that form the author's back yard, and it ra- diates the spirit of a butterfly enthusiast and naturalist. There are three parts. The first contains a short ex- planation of “Cascadia” and its mosaic of habitats (eco- geographic provinces), a short history of the butterfly pioneers in the Pacific Northwest, followed by a brief ‘how to use this book’. The second and most ample part is, of course, the species accounts. Like most field guides, each species is treated in telescopic manner to facilitate field identification and provide a snippet of natural history information. Next to the individual ac- count are color portraits that were photographed in the field. But here the similarity to other field guides ends. Nearly all species accounts are unique by having Pyle’s eclectic anecdotes to accompany them. Overall this renders a bucolic flavor such that the reader can almost smell the mold, pine needles or sagebrush of the Pacific Northwest, and take part in its butterfly his- tory. Such lagniappe! Moreover Pyle manages to navi- gate, with considerable élan, the turbid debates over collecting versus watching, and the chloroform of nomenclature squabbles. The excellent color photos from nature and the lucid writing make the book both pleasing to the eye, and readable into the bargain. Well done! The final part consists of a checklist (complete with little boxes to tick off) followed by lists of refer- ences, organizations, a glossary, data for each color photo, and an index of butterfly names. In summary, The Butterflies of Cascadia will help ensure that butterflies of the Pacific Northwest stay in the public eye, and it will be an important tool for pro- fessional entomologists and conservation biologists. This sturdily bound book deserves to be on the shelves of anyone who is interested in butterflies, the Pacific Northwest, or just fun reading. P. J. DEVrIES, Center for Biodiversity Studies, Mil- waukee Public Museum, 800 West Wells St., Milwau- kee, Wisconsin 53233, USA Journal of the Lepidopterists’ Society 57(2), 2003, 157-158 THE SATURNIIDAE OF AMERICA .. . LES SATURNIIDAE AMERICAINS. VOLUME 4: HEMILEUCINAE, by Claude Lemaire. 2002. Three parts, hardbound separately, 1388 pages, 140 color plates, 21 cm by 30 cm, ISBN 3-931374-08-4. Published by Goecke & Evers, Sportplatzweg 5, Keltern 75210, Germany; website: www.insecta.de; Price 340 euros (about US$365). For the specialist, this book will be seen as the de- finitive treatment of the subfamily Hemileucinae; nothing else has ever come close or probably ever will. The Hemileucinae are famous for caterpillars with stinging spines and moths with bright and contrasting colors, often with eyespots, typically represented by species of Automeris, and long favored by collectors. Rare and unique moths from southern Chile, the is- land of Hispaniola, the cerrados of central Brazil, and the high Andes of Peru and Ecuador, are shown in color for the first time. Specimens of some Automeris are bigger than our polyphemus moth (Antheraea polyphemus) and Europe’s peacock moth (Saturnia pyri). Serious taxonomic errors, even by recent au- thors, have been exposed and corrected. Because this work took many years to prepare, many amateur and professional lepidopterists have eagerly anticipated it. The publication consists of three hardbound books. The text is in English, with a French summary for each genus and species. The smooth covers are a light greenish yellow, with a color image of a different hemileucine on the front cover of each. Since this work is to be cataloged as the “volume 4” continuation of Lemaire’s previous three volumes on subfamilies of American Saturniidae (Saturniinae in 1978, Arsenuri- nae in 1980, and Ceratocampinae in 1988), the present three books are labeled parts A—C, instead of volumes 1-3. These parts cannot be purchased separately, which is entirely appropriate. Part A consists of the preface, foreword, introductory sections, and text treatments of 31 genera including Lonomia, Col- oradia, Hemileuca, Automeris, Hylesia, and several others, running from pages 1-688. The lengthy pref- ace by Daniel Janzen offers biographical notes about Lemaire and some colorful commentary on the multi- faceted value of his published works. Part B completes the treatments of the remaining 18 genera, and has an exhaustive bibliography, 185 pages of distribution maps, and 214 pages of drawings showing genitalia, wing venation, antennae, and legs, running from pages 689-1388. Part C contains 126 color plates of pinned 158 adults (all shown life size), and 14 color plates of im- mature stages, mostly mature caterpillars, including 23 taxa that occur in the United States. Many of the lar- vae have colors and patterns as striking as the wings of the moths. The text is printed on thin paper, and the plates are shown on much heavier paper. Overall, the quality of the paper, binding, and color reproduction is of high quality, and attractively presented. The Ger- man publishers and the French photographers and computer technologists should be commended for the final results. One powerful lesson this book provides to lepi- dopterists in North America is that the Hemileucinae we see in Canada and the United States, or find in our books about Nearctic saturniids, represent only a small part of the diverse hemileucine fauna that exists from Baja California Norte southward. In this work, 46 new species are described and named. As now defined by Lemaire, the subfamily Hemileucinae consists of 49 genera and 670 species. (No Hemileucinae occur in the Old World.) The dazzling genus Automeris com- prises over 135 species, making it the largest of the subfamily. As I view plate after plate of the not-so- dazzling Hylesia, I am amazed that Lemaire was able to sort out such a taxonomic nightmare. Many species of Hylesia look alike, and for some species, it was a challenge to associate the males and the females, some of which were assigned different specific epithets by various authors. Some species are wide ranging, others are known from a single collecting site. Many names have been assigned to synonymies, but other Hylesia Lemaire found to be long represented in museum col- lections, yet new to science, including the one that ranges far north in Mexico close to the Arizona border. To me it looked like so many other Hylesia, but now when seen on Lemaire’s color plates, I just might be able to recognize it in collections or even in the field. Perhaps only a taxonomist can fully appreciate the Herculean task required to assemble this monograph. In terms of sheer compilation, it represents far more of a final product than most doctoral dissertations. Lemaire has been blessed with a unique combination of accessibility to specimens, both by purchase from dealers and exchange with field collectors, and by fre- quent visits to the museums that hold the type speci- mens for the majority of the names in the group. Added to this is Lemaire’s ability to access and inter- pret the historical and current literature in several lan- guages, and a work ethic and dedication to purpose that few taxonomists can match. He also has made sev- eral collecting expeditions to South America, espe- JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY cially in the Andes, and has reared numerous Hemileucinae from eggs, so his study goes far beyond analysis of the literature and pinned moths. Lemaire has handled the complex of populations that we call Automeris io with great skill, by demonstrating how we are seeing evolutionary processes frozen in time, which sometimes makes it difficult for our artificial classifications (i.e., names) to represent real species and populations within them. The classic treatise by Charles Michener (1952, Bull. Amer. Mus. Nat. Hist. 98:335-501) made subgenera a central theme, affect- ing almost every genus of Hemileucinae. I was glad to see that Lemaire has recognized the unnecessary com- plexity of subgenera by not using them, except in the genus Meroleuca, which may be provisional until more is known. The fact that some of the Meroleuca have wingless females accounts for their rarity in collec- tions. Lemaire’s compilation is remarkably free of er- rors, and I would categorize him as a perfectionist. I should point out here that I reviewed the manuscript of the work and have considered Lemaire a friend for 30 years. But those who may question the objectivity of this book review will agree with the praise recorded here if they objectively examine Lemaire’s treatise on Hemileucinae. For those who collect and rear Saturniidae, this set of books will be indispensable. All known host plants are cited for each species, and the descriptions and fig- ures will enable one to identify virtually any specimen that they acquire. The meticulously assembled sec- tions of “Material Examined” are especially valuable, as they give complete label data for hundreds of spec- imens, enabling future collectors to know exactly where and when to find each species. Sitting and view- ing the dozens of color plates will be an immense plea- sure for any saturniid lover, each moth with its story about crypsis, camouflage, mimicry, and/or warning coloration. There are hundreds of species of these moths, dramatically demonstrating the concept that biodiversity is greatest in the tropics. To summarize, I will end by saying that this is a fine and impressive piece of work, and that Lemaire’s taxo- nomic decisions are all firmly supported. The beautiful plates document the ever-increasing improvements in color technology and reproduction. This work will be the crown jewel of many personal libraries, but it should also be widely available in libraries. RICHARD S. PEIGLER, Department of Biology, Uni- versity of the Incarnate Word, 4301 Broadway, San Antonio, Texas 78209-6397, USA VOLUME 57, NUMBER 2 Journal of the Lepidopterists’ Society 57(2), 2003, 159-160 A REVISION OF THE SILKMOTH GENUS SAMIA, by Richard S. Peigler and Stephan Naumann. University of the Incarnate Word, 2003. 227 pages, 10 maps, 146 color and 82 B&W illustrations, in English but with summaries also in German, Bahasa Indonesian, Chinese, Japanese, and French. Softcover, $ 36.00 + $4.00 s&h. Available from: University of the Incarnate Word, atten. Margaret Preston, 4301 Broadway, San Antonio, Texas 78209-6397 USA. ISBN 0-9728266-0-2 Saturniids suffer from their popularity. As young collectors many of us were obsessed with the saturni- ids, and perhaps also swallowtails, for their size and beauty and ease of rearing. With time most broadened their interests or concentrated on some group of spe- cial interest, and came to view the silkmoths as too well-known and perhaps too gaudy to be taken seri- ously other than as natural history teaching tools. In recent years this popularity has been turned around to accent the importance of the saturniids in studies of ecology, evolution and systematics, and we should be grateful with the publication of this book that some have not lost their early obsession. (I should know!) The two authors have spent considerable time and ef- fort to travel, collect and rear many of the taxa, and to pore over private and museum collections for addi- tional information. The result is this fine book which offers to Lepidopterists, and natural historians in gen- eral, a diversity of topics illustrated by these beautiful creatures. Among the Saturniidae, the genus Samia is notable not just for the striking display of pinks, browns, and olive tones in the adult, but also for the domestication and commercial exploitation of a few species for their silk, and the less successful attempts to introduce these forms in various locales around the world. It is in this context, as an introduced exotic confined to an ur- ban ecosystem, that we know our own S. cynthia. But this familiarity is also a distortion, and as the authors point out, the natural history of most of the 19 (yes, there are 18 other species) is relatively unknown. Like Hyalophora cecropia, S. cynthia probably exists in low-density, dispersed populations in the wild state (not in the U.S.), but can be locally abundant in an ur- ban or suburban setting. The book is divided into chapters on taxonomy, phy- logeny, aberrations, Samia in art and culture, bio- chemistry and physiology, cytogenetics, biology (in- cluding parasitoids and diseases), ecology and rearing, sericulture (a special interest of Peigler’s and well- presented), and biogeography, in addition to the 159 species treatment. No taxa are newly described, but wangi, kohli, abrerai, and naessigi were described by the here in 2001, and peigleri, yayukae, naumanni, and treadawayi were named in honor of these workers in the 1990s by various authors. While some of the chapters are understandably brief summaries and could perhaps have been combined, the citation list is quite extensive and the list of topics should attract the interest of a wide audience, both amateur and profes- sional. The treatment of known life histories is exhaus- tive, and I found the implications of host plant use for phylogenetic relationships particularly fascinating. The only criticism of the production I might have would be the small type and wide line width, making reading sections such as “Material Examined” a bit difficult. The color plates are of excellent color and density (al- though somewhat small but keeping cost down), and the photos of larvae are especially striking. See Peigler and Wang (1996) for additional illustrations in larger format of some of the species. I would have liked to have seen a more expanded discussion of biogeography (only two pages), includ- ing an updated discussion with accompanying maps of general saturniid biogeography in the realm of the Wallace Line, from the Malay Peninsula to Irian Jaya and New Guinea. These authors are in a position to build on the cited earlier works of Barlow, Holloway and Peigler himself to produce such a work, probably more extensive than appropriate for this specialized book. Those interested in phylogeny and saturniid evolution should read the discussion of the possible ancestral relationship within the Attacini of Samia to the African genus Epiphora. As cited by the authors, I also highly recommend the popular work by van Oost- erzee (1997) on the biogeography of the Malay Arch- ipelago. Judging from the list of synonymies given for each species (113 for cynthia over 4 pages), Samia taxon- omy was a mess until this publication. Curators will ap- preciate this material, and the general reader will also find sections on types, geographical distribution, diag- nosis, descriptions, discussion (largely on life history), and material examined. The Samia are characteristi- cally conservative in adult wing characters, and some of the species look very similar until genitalia or imma- tures are examined. Some of the species are wide- spread and variable, such as canningi, kohli, and wangi, leading to considerable past Ron and me dentit: cation, which the authors correct. Other species are in- sular and endemic or otherwise quite distinct. In this regard, the authors recognize no subspecies, a taxonomic category they adamantly oppose on philo- sophical grounds, and prefer instead to either raise to 160 full species status distinct allopatric populations, or to lump geographic variants under a single species with accompanying descriptive discussion. Although this treatment employs the dichotomous splitting of the Phylogenetic Species Concept (PSC), no formal cladistic analysis is given, probably because DNA or protein samples were unavailable (although Peigler (1989) used morphological and life history characters in his work on the genus Attacus). The Biological Species Concept (BSC) is explicitly rejected as out- dated and its use by Tuskes et al. (1996) criticized in their treatment of North American saturniids. While controversy over species concepts is widespread and useful, the authors’ position here illustrates three in- teresting ironies regarding saturniid taxonomy. First, the PSC is a species concept, not a speciation concept, that stresses pattern of descent over genetic processes in populations. Yet, the allopatric mode of speciation underlying the PSC is pure Mayr in form, but unlike current application of the BSC tends to downplay the blending and intergradation among geo- graphic forms that so often characterizes Lepidoptera. While the lack of totally effective reproductive isola- tion often seen among closely related taxa makes ap- plication of the traditional BSC difficult and arbitrary , the tendency of the PSC to oversplit into full species (or amplify species count—depending on your philos- ophy [Avise 1997, 2000]) is also a valid criticism. So, for all its faults, the subspecific category is still used to represent geographic variation. Second, Tuskes et al. (1996) briefly described cladis- tic methods under the PSC and proposed their appli- cation to saturniid taxonomy (see Regier et al. 2002 as a recent example), although this discussion wasn’t JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY mentioned by Goldstein (1997) (cited by Peigler and Naumann) in his review of the Tuskes book. Finally, and most important, a justification for the application of the BSC to the saturniids is that, unlike many other Lepidoptera or animals in general, they can be easily collected, mated, and reared in the lab, mak- ing experimental hybridization possible in investigating concepts of genetic cohesion and species boundaries. “Hybrids of Samia” is the concluding chapter in this re- vision, and the statement that hybridization “can yield valuable data on genetics and phylogeny” suggests that the cooperative use of both cladistic analysis and exper- imental hybridization can lead to a better understand- ing of the two key aspects of a biological species—a phylogenetic history and genetic processes among populations within a geographic range. LITERATURE CITED AVISE, J. 1997. Phylogenetics and the origin of species. Proc. Natl. Acad. Sci. 94:7748-7755. AVISE, J. 2000. Cladistics in wonderland. Evol. 54:1828-1832. GOLDSTEIN, P. Z. 1997. Review of: Wild Silk Moths of North America., by P. M. Tuskes et al. J. N.Y. Entomol. Soc. 105:121—-125. VAN OOSTERZEE, P. 1997. Where worlds collide: the Wallace line. Cornell Univ. Press. PEIGLER, R. S. 1989. A revision of the Indo-Australian genus Atta- cus. Lepidoptera Research Foundation. PEIGLER, R. S. & H. Y. WANG. 1996. Saturniid moths of southeast- ern Asia. Taiwan Museum, Taipei. REGIER, J. C., C. MittEr, R. S. PEIGLER & T. P. FRIEDLANDER. 2002. Monophyly, composition, and relationships within Sat- urniinae (Lepidoptera: Saturniidae): evidence from two nuclear genes. Insect Syst. & Evol. 33:9-21. TuskEs, P. M., J. P. TurrLe & M. M. Couns. 1996. The wild silk moths of North America. Cornell Univ. Press. MICHAEL M. 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POSITION ~ André Victor Lucci F reitas - THE LARVA AND PUPA OF Lamiosis PERMAGNARIA Pack: (Geomermpae) GENERAL Notes NOTE ON THE DISCOVERY OF THE LARVA OF CUCULLIA SIMILARIS (Noctua ( ‘Petit and M. C. Petit eres raat _ THE CORRECT TYPE LOCALITY OF CYANIRIS LADON VAR. QUESNELLI pede, 1910 WITH DESIGNATION OF A LECTOTYPE Cee ay Cone and Norbert ¢ G. Age Michael A. Roberts and William ay “Miller - Host PLANT ASSOCIATIONS OF WESTERN SPECIES OF PAPAIPEMA (Nocrumae) wir PB REFERENCE TO THE APIACEOUS PLANTS Paul Z. Goldstein and Eric L. Quint / AMERICA Te Book Reviews | Tue Burrerriies or Cascapia PJ. DeVries ee n---- < THE SATURNIIDAE OF AMERICA . ... Les SaruRNIIDAE AMERICAINS. VOLUME a “Henney Richard S. Peigler ee - A Revision oF THE SILKMOTH Genus Samia... Michael M. 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The additional cost for members outside the U.S. is to cover mailing costs. Journal of The Lepidopterists’ Society (ISSN 0024-0966) is published quarterly by The Lepidopterists’ Society, % Los Angeles County Museum of Natural History, 900 Exposition Blvd., Los Angeles, CA 90007-4057. Periodicals postage paid at Los Angeles, CA and at additional mailing offices. POSTMASTER: Send address changes to The Lepidopterists’ Society, % Natural History Museum, 900 Exposition Blvd., Los Angeles, CA 90007-4057. Cover illustration: Caterpillars of Morpho theseus (Nymphalidae). Photo by P. J. DeVries. JOURNAL OF THE LEED O BT RIS TS SOCIETY Volume 57 2003 Number 3 Journdl of the Lepidopterists’ Society 57(3), 2003, 161-167 MID-WINTER FORAGING OF COLONIES OF THE PINE PROCESSIONARY CATERPILLAR THAUMETOPOEA PITYOCAMPA SCHIFF. (THAUMETOPOEIDAE) T. D. FITZGERALD Department of Biological Sciences, State University of New York at Cortland, Cortland, New York 13045, USA AND XAVIER PANADES I BLAS University of Bristol, Department of Earth Sciences, Wills Memorial Building, Queens Road, Bristol, England ABSTRACT. The pine processionary caterpillar Thawmetopoea pityocampa Schiff. (Thaumetopoeidae) overwinters as an active larva. Field recordings made at our study site in Catalonia (Spain) during mid-winter show that the caterpillar is remarkable in its ability to locomote and feed at temperatures well below those at which the activity of most other insects is curtailed. Colonies initiated foraging bouts in the evening, 93.1 + 35.2 minutes after the end of civil twilight and retumed to the nest the following morning, 42.9 + 24.9 minutes before the onset of civil twilight. Despite an overnight mean minimum temperature of 3.8 + 0.25°C during the study period, caterpillars were active each night and did not become cold-immobilized until the temperature fell below —2°C. During the daytime, the caterpillars sequester themselves seat their nests and on sunny days are able to elevate their body temperatures by conducting heat from the structures. The mean difference between the daily high and low nest temperature was 30.9 + 0. 9°C. The maximum nest temperature recorded was 38°C. Salient features of the biology and ecology « of T. pityocampa are compared to those of other central place foragers in an attempt to elucidate the factors that may underlie the evo- lution of foraging schedules in social caterpillars. RESUMEN. La procesionaria del pino, Thaumetopoea pityocampa Schiff. (Thaumetopoeidae), permanece activa durante el invierno. Los registros de campo obtenidos en el presente estudio en Catalufia (Espafia), durante la parte media del invierno, muestran que esta larva es asom- brosa por su habilidad para desplazarse y alimentarse a temperaturas muy por debajo de aquellas a las que la actividad de la mayoria de los in- sectos es impedida. Las colonias inician su periodo de forrajeo en la noche, 93.1 + 35.2 minutos después de la penumbra civil y regresan a sus nidos a la mafiana siguiente, 42.9 + 24.9 minutos antes del inicio de la penumbra civil. A pesar de que la temperatura minima durante la noche en el periodo de estudio fue de 3.8 + 0.25°C, las larvas estuvieron activas cada noche y no se inmovilizaban por el frfo hasta que la temperatura descendia por debajo de los -2°C. Durante el dia, las larvas se mantenian dentro de sus nidos, dependiendo de la habilidad de estas estructuras para absorber la energia solar para elevar su temperatura corporal. En el interior de los nidos, el diferencial promedio entre la temperatura max- ima y minima fue de 30.9 + 0.9°C. La temperatura maxima registrada dentro de los nidos fue de 38°C. Las caracteristicas sobresalientes de la biologia y ecologia de esta larva procesionaria son compar: adas ¢ con las de otras especies de forrajeo central, en un intento por dilucidar las fac- tores que subyacen en la evolucién de los patrones de forrajeo de larvas sociales. Additional key words: _ processionary behavior, trail following, activity patterns, thermal regulation. Thaumetopoea pityocampa Schiff., the pine proces- ing the establishment of their permanent nests (Fabre sionary caterpillar, is distributed throughout much of 1916), there have been no studies of the foraging be- southern Europe where the larvae feed gregariously havior of larvae in midwinter, nor are there any long on the needles of pine (Pinus spp.). Colonies develop term records of foraging behavior for any time of the from egg masses of 70 to 300 eggs (Dajoz 2000). The year. Of particular interest is the question of whether siblings at first build and abandon a series of loosely the caterpillars forage on evenings when overnight spun nests but in the third instar establish a perma- temperatures approach freezing. Of parallel interest is nent nest and become central place foragers (Halperin the role that the nest might play in enabling the cater- 1990). In Catalonia (Spain), the larval stage typically pillars to process food in their guts at low ambient extends from August until April of the following year temperatures. Several investigators have made spot and the caterpillars overwinter as active larvae. Al- measurements of nest temperatures and have reported though it is known that the larvae feed at night follow- that when irradiated by the sun the structures achieve temperatures as much as 17°C in excess of the ambi- ent temperature (Breuer et al. 1989, Demolin 1969, Breuer & Devkota 1990, Halperin 1990), but there have been no continuous measurements of nest tem- peratures in midwinter. The recent availability of small, portable data loggers has made possible the un- interrupted recording of physical and behavioral data heretofore not feasible in remote locations, and a de- tailed database of ecologically relevant aspects of the foraging behavior of social caterpillars has begun to ac- cumulate (Fitzgerald et al. 1989, Fitzgerald & Under- wood 1998a, b, Ruf & Fiedler in press). We undertook the present study of the pine processionary caterpillar to add to this database and, more specifically, to inves- tigate the mid-winter foraging behavior of the insect. We monitored both the daily temperature cycles of the nests and the foraging and resting cycles of the cater- pillars. MATERIALS AND METHODS Study site. Studies of nest temperature and colony activity patterns were undertaken during February 2001 in a mountainous region near La Moixeta, (Baix Penedés County), Catalonia (Spain) (41°21’N, 001°31’E), eleva- tion approximately 400 m. The canopy of the study area consisted almost entirely of pure stands of Pinus halepensis and P. pinea. Climate records. Seasonal climate records for 1999-2001 were obtained from the Catalonia Meteo- rology Service. Data are from Font-Rubi (Alt Penedés County, elevation 409 m) the nearest govern- ment maintained weather station, approximately 8.5 km from the La Moixeta study site. The hot summers typical of the Mediterranean climatic zone are moder- ated by elevation at the La Moixeta site, and the months of November through February are corre- spondingly cooler with midwinter temperatures ap- proaching, but only occasionally falling below, freezing. Orientation of nests at the field study site. The positions of 157 T. pityocampa nests relative to the car- dinal compass points were plotted at the study site to determine if the nests are positioned to take advantage of solar radiation. The nests occurred naturally on ei- ther P. halepensis or P. pinea. Trees were divided into quadrants each centered about a cardinal compass di- rection and the position of each nest on a tree assigned to one of the quadrants. Heat gain in nests under controlled conditions. Laboratory studies were conducted to determine how nests gain and maintain heat when exposed to a radi- ant heat source. Four empty nests of different sizes were maintained in a temperature controlled chamber with an ambient temperature of 6.0 + 2.0°C and irra- JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY diated with a 250-W infrared lamp situated 0.5 m from the nests (Breuer & Devkota 1990, Fitzgerald & Un- derwood 2000). Temperature probes were inserted at the centers of the irradiated and shaded sides of the nests, approximately 2 cm below the surface. The tem- perature of each nest was measured at 1 minute inter- vals for approximately 135 minutes after which the heat source was extinguished and additional measure- ments made until the nests cooled to ambient temper- ature. Temperature measurements were automatically written to data loggers (Onset Computer Co., accuracy + 0.2°C ) and the data downloaded with BoxCarPro Software (Onset Computer Co.). Temperature measurement of nests at the field study site. The internal temperatures of five nests of the pine processionary were monitored in the field from 17-26 February. Temperature probes were in- serted approximately 8 cm below the upper surfaces in areas of nests occupied by caterpillars. Ambient tem- peratures were measured in shaded areas near the study nests. Temperature data were recorded at 15 minute intervals as described above and the data log- gers downloaded at 24 h intervals. Temperature records for a total of 19 colony-days were collected. Daily activity patterns of field colonies. Daily activity patterns of seven colonies of T. pityocampa oc- curring on different trees were monitored with in- frared activity monitors (Fitzgerald & Underwood 2000) from 18-26 February. Records for a total of 26 colony-days were collected. The monitors were placed on branches bearing the major trunk trails of the colonies, approximately 20 cm from each nest. Activity monitors were connected to event loggers (Onset computer Co.) which recorded the time of day when the caterpillars triggered the monitors. A reset delay of five seconds was programmed into the recorders to minimize the probability that a single passing caterpil- lar would trigger the monitor more than once. Data were off loaded with BoxCarPro software at 24 hour intervals. Colonies were also observed each evening with red-filtered light and again in the early morning to aid in the interpretation of the activity recordings. In reporting the time of onset and termination of daily activity periods, we ignored isolated early starters and stragglers by considering colony activity to have started when the number of caterpillars moving past the de- tector reached 10 or more per 15 minutes and to have ended when the number or returning caterpillars fell to fewer than that number. Statistical analyses. Statistical analyses as detailed below were conducted with SigmaStat and ProStat statistical software. Nest orientation was analyzed with Rayleigh’s test for circular distribution (Zar 1974). VOLUME 57, NUMBER 3 80 Number of nests 10 4 : 0 ae N S E Ww Orientation (quadrant) Fic. 1. Orientation of nests of T. pityocampa on pine trees at the study site. RESULTS Orientation of nests at the field study site. Ap- proximately 61% of the nests at the study site were lo- cated within the SE-SW (135-225°) quadrant on the crowns of trees. Nests were non-randomly distributed with a mean orientation of 204 + 6.8° (SE) (Rayleigh’s test of uniformity, p < 0.01, Fig. 1). Heat gain in nests under controlled conditions. Empty nests exposed to a radiant heat source in the 00, V Temperature °C Time lapse (15 min intervals) Fic. 2. Temperature within the irradiated (solid line) and shaded sides (dotted line) of four empty nests of T. pityocampa recorded under laboratory conditions. Ambient temperature range is indicated by the horizontal shaded bar. Arrows indicate points when artificial heat source was turned off, values indicate mass of nests. 163 40 February 20 February 19 February 18 Temperature (°C) 1200 2400 1200 2400 1200 2400 Time Fic. 3. Temperature inside the nest of a colony of T. pityocampa (gray) and ambient temperature (black) recorded over a three day period. laboratory showed a rapid increase in internal temper- ature on the irradiated side (Fi ig. 2). The larger nests showed virtually no heat gain on the shaded sides while the shaded sides of the smaller nests had modest gains relative to the irradiated sides of the structures (Fig. 2). Temperature differentials between the irradi- ated and shaded sides of the nests ranged from 29.3°C in the larger nests to 13.2°C in the smaller. All of the nests cooled precipitously to ambient temperature when the heat source was removed (Fig. 2) Diurnal fluctuation of nest temperature in the field. On 19-23 and 25 February, skies at the study site were largely cloud free. Four of our five study nests were in open areas and were directly irradiated by the sun for most of those days. The mean daily low temperature recorded in these nests was 3.8 + 0.25°C and the mean daily high 34.6 + 1.0°C (12 colony-days, range = 0-38°C). The mean difference between the daily high and low temperatures in the nests was 30.9 + 0. 9°C. The mean daily high ambient temperature recorded at the study site during this period was 16.9 + 1.5°C. One of our study nests was in a more shaded area and experienced direct radiation for only part of the day. For this nest, the mean low temperature was 3.8 + 1.0°C and the mean high 17.3 + 1.0°C (4 colony- days). The mean difference between the daily high and low temperatures in the nest was 13.5 + 0.9°C. For all nests, daily lows occurred in the morning just before dawn and daily highs between 1200 and 1500 h. Daily temperature fluctuation recorded in one nest over a three-day period is shown in Fig. 3. Our investi- gations in the study area were terminated at noon on 26 February and we obtained a temperature record for the period ‘from 2400 to 0900 h on that day. On the morning of 26 February, standing water in outdoor containers had iced over and our data loggers indi- cated the temperature dropped to -4°C by 0600 h, the lowest temperature recorded during the study period. Seasonal and daily activity patterns of field colonies. A seasonal temperature profile compiled for the study area for 2000 and 2001 shows that the cater- 164 Mean temperature (°C) E> (2) Jo, TS) >>320 0 7 <«t QHD 2001 Fic. 4. Monthly mean minimum (gray) and mean maximum (black) temperatures recorded near the study site over a two year interval. Horizontal bar indicates approximate periods when T. pity- ocampa actively forages (gray) and pupates underground (white). pillars feed and grow during the coldest part of the year and reside as pupa buried in the soil during the hot summer months (Fig. 4). Daily activity records ob- tained during the present study show that during the period of growth, caterpillar activity outside the nest is restricted to the coldest part of the day (Fig. 5). De- spite low early morning temperatures during the study period, colonies were active overnight on all of the study days. Colonies initiated foraging bouts an aver- age of 93.1 + 35.2 minutes after the end of civil twi- light in the evening (center of the sun 6° below hori- zon) and the last contingent of foragers returned to the nest 42.9 + 24.9 minutes before the onset of civil twi- light in the morning (N = 26 colony-days). The interval between the movement of the first contingent of cater- pillars from the nest in the evening and the return of the last contingent in the morning was 618.0 + 35.2 minutes. Thus, colonies were typically active on the tree throughout the evening and early morning hours. That colonies fed during these overnight forays was evidenced by the presence of fresh leaves in their guts after they returned to the nests. In 20 of the 26 forag- ing bouts recorded, colonies moved from the nest to feeding sites in the early evening and returned in the early morning, giving rise to bimodal activity patterns (Fig. 5). In the other six instances, activity between the nest and feeding sites occurred throughout the evening and early morning. Colony activity on the two coldest nights is illustrated in Fig. 6. Overnight on 17-18 February, colonies were continually active even though temperatures measured at nest sites fell to 0°C by 2400 h. The ambient temper- ature was slightly below freezing at 0900 h when the whole of colony 1 was observed to still be out of the nest moving about the tree in procession. It is not known when the caterpillars returned to the nest but all were back when the nest was next observed in late afternoon. Overnight on 25-26 February the temperature fell be- low 0°C at approximately 2200 h and reached an overnight low of -4°C at 0600 h. Inspection of the ac- tivity record for colony 3 (Fig. 6) shows that activity was JOURNAL OF THE LEPIDOPTERISTS SOCIETY Colony 1 100 4 February 19-24 Events Temperature (°C) | tu! ! Tl Li 2400 2400 2400 2400 2400 Time Fic.5. Bimodal daily activity bouts of a colony of T. pityocampa (vertical bars) and nest temperature (black line). Horizonal bar indi- cates daily photoperiods (white) and scotoperiods (black). Photo- periods begin at the onset of civil twilight and end at the termination of civil twilight. not initiated for the most part until the temperature fell below 0°C. Caterpillar activity largely ceased when the temperature reached —2°C but sporadic activity was recorded at —-4°C. When this colony was observed at 0900 h on 26 February, all the caterpillars were back in the nest. The extent to which the caterpillars fed at these sub-zero temperatures is not known. DISCUSSION Our study, providing the first empirical data set on the temporal foraging patterns of colonies of the pine processionary, supports the observation of Fabre (1916) that the caterpillars of the pine processionary feed throughout the winter on all but the most frigid nights. The pine processionary caterpillar is remark- able in its ability to locomote and feed at temperatures well below those at which the activity of most insects is curtailed. Only a few other invertebrates are active at such low temperatures. Some collemboleans (Aitchi- son 1983), several species of spiders (Aitchison 1984), the cricket Grylloblatta campodeiformis Walker (Prichard & Scholefield 1978), an amphipod (Dunbar 1957), and a copepod (Kigrboe et al. 1982) have been reported to move about and feed at temperatures at or slightly below zero. Laboratory studies of the sub- Antarctic caterpillar Pringleophaga marioni Viette (Tineidae) showed that the larvae are able to maintain motoric functions at temperatures as low as —1.6°C but there are no data to show that the caterpillars are ac- tive at temperatures this low in their natural environ- ment (Klok & Chown 1997). Locomotion and feeding in caterpillars has been recorded only rarely at temperatures below 5°C (Ke- van et al. 1982, Joos 1992, Kukal 1993, Klok & Chown VOLUME 57, NUMBER 3 Colony 1 ~ 2 February 17-18 |- 6) %| x 45 + 40 oe Lg zB 10- 2 § UG eI E =| [_ FE 0- 0 5 + —— —— - — T 71 0900 1300 1700 2100 0100 0500 0900 Time 305 Colony 3 r 80 25 | February 25-26 |- =~ + 60 O 204 Le ev 4 L4o 2 = 15 5 £ 104 lim ir) Q + 20 ena E = 04 anil 0 5 4 a a 1 pe 7 0900 1300 1700 2100 0100 0500 0900 Time Fic. 6. Temperature in the nest (black line) and activity outside the nest (vertical event bars) of two colonies of T. pityocampa. Hor- izontal bar indicates photoperiod (white), scotoperiod (black) and civil twilight (gray). 1997). A notable exception is the social caterpillar Eu- cheira socialis Westwood (Pieridae) which feeds throughout the winter in the mountains of Mexico (Fitzgerald & Underwood 2000). The daily tempera- ture profile in the winter in the montane forest where E. socialis occurs is similar to that of our study area in Catalonia in that the diurnal temperatures are moder- ate to warm while at night the temperatures plunge precipitously and are often near or below freezing. Like T. pityocampa, the caterpillars of E. socialis feed only at night and have been recorded outside their nests at subzero temperatures in midwinter (Fitzger- ald & Underwood 2000). Our study shows that the caterpillars of T. pityo- campa locate their nests preferentially in the SE-SW quadrant of host trees. Brueur et al. (1989) found ap- proximately 76% and Schwammer and Nemeschkal (1987) approximately 80% of the nests they surveyed in other areas of the Mediterranean to lie within this same quadrant. Thus, data from these studies show that colonies of the processionary locate their perma- nent nests on the side of the tree that is likely to re- ceive the most solar radiation. Although fewer nests were found within this quadrant in La Moixeta, this may have been the case because the trees occurred in young stands and were distantly spaced so that they ex- perienced little shading from nearby trees. Fitzgerald and Underwood (2000) found that the nests of E. so- cialis occurring on open grown and distantly spaced madrone trees were irradiated by the sun regardless of their position in the crown of the tree. 165 The ability of the inhabited nests of T. pityocampa to warm well above ambient temperature when irradi- ated was demonstrated in laboratory studies by Breuer and Devkota (1990), but our study shows that even when devoid of caterpillars the irradiated nests of T. pityocampa exhibit large heat gains. The overwinter- ing nests of the processionary are densely packed with silk, frass, and host material and the bodies of the rest- ing larvae are situated tightly within them, allowing the caterpillars to raise their T,,’s well above the tempera- ture of the outside air by conducting heat from the structures. Although the relationship between T, and the rate of digestion has not been determined for the processionary, nests situated on host trees to facilitate the absorption of solar radiation achieve thermal max- ima during sunlit days that are likely to be well in ex- cess of those required by the caterpillar for efficient food processing during midwinter. Data available for a few other species of caterpillars that feed at compara- bly low temperatures indicate that efficient food pro- cessing by the processionary is likely to require a T, that exceeds the winter time air temperatures typical of our study site. The Arctic caterpillar Gynaephora groenlandica (Wécke) (Lymanthriidae) has an assimi- lation efficiency of only 7% at 5°C compared to 40% at 15°C (Kukal 1993) but the assimilation threshold for Malacosoma americanum (Fabricius) (Lasiocampi- dae), which is reported to collect food at temperatures down to 7°C (Joos 1992), is not reached until the caterpillars warm to at least 15°C (Casey et al. 1988). Regardless of the thermal demands of the procession- ary, our study shows that the irradiated nests of the in- sect provide thermally heterogeneous environments (Fig. 2) within which the caterpillars might optimize their T,’s by varying their positions within the struc- tures during the daytime. In contrast, temperatures in shaded nests will be much cooler and may not allow optimization of T,, placing a premium on the correct siting of the permanent nest by the third instar cater- pillars. Although the caterpillars might also achieve T,,s conducive to food processing by basking outside the nest in midwinter, the cost would be greater expo- sure to predators. The mean daily temperature for the coldest months (November—February) recorded at the Font Rubi station from 1999-01 was 10.7 + 0.7°C and the mean daily maximum temperature for this same period was 14.7 + 0.5°C. Thus, if the caterpillars were to remain hidden outside the nest in protected loca- tions during these months their T,’s would be much lower than Those achievable in the nest during this same period and might be too low to allow the efficient processing of food. Despite the ability of the caterpillars to warm within 166 irradiated nests in midwinter, studies of caterpillars foraging under laboratory conditions (Fitzgerald in press) indicate that under the overall thermal regime the caterpillars experience in the field they grow at a rate well below that which they could achieve at sus- tained, higher temperatures. Caterpillars maintained in the laboratory at 22 + 2°C, under a photoperiod that simulated that experienced by field colonies of T. pity- ocampa, exhibited the same nocturnal pattern of ac- tivity as field colonies and did not feed during the day. These colonies, which eclosed from eggs in early Au- gust, completed their larval development and pupated by late October, at least 15 weeks sooner than field colonies require to complete their larval development. Halperin (1990) similiarly found that the social larvae of T. jordana (Stgr.), which also feed during the winter, required approximately 150 days to complete their lar- val development in the field but only 48 days when maintained in the laboratory at a constant 25°C. The likely reason for the difference in growth rate between field and laboratory colonies of both of these species is that when maintained at elevated temperatures cater- pillars are able to both collect and process food during their nocturnal forays. This allows them to assimilate more energy each day than caterpillars that experience temperatures too cold to permit food processing dur- ing the overnight foray. In addition, under field condi- tions, the caterpillars can be expected to assimilate little if any food on cloudy winter days. Electronic recordings of daily foraging activity are now available for five species of social caterpillars. All are central place foragers sharing overt features of their biology (Table 1, traits 1-6). Analysis of their activity records shows that the caterpillars fall into two distinct groups based on the temporal pattern of their foraging behavior. M. americanum (Fitzgerald 1980, Fitzgerald et al. 1989) and Eriogaster lanestris (L.) (Lasiocampi- dae) (Ruf & Fielder in press), feed both day and night and grow rapidly, achieving their full larval growth in approximately eight weeks. E. socialis (Fitzgerald & Underwood 1998a), Gloveria sp. (Lasiocampidae) (Fitzgerald & Underwood 1998b), and T. pityocampa (this study) feed only at night, grow slowly, and have ac- tive larval stages lasting 7-9 months. A major constraint on caterpillars foraging is predation pressure (Stamp & - Casey 1993) and for all of these caterpillars, defense against day active predators would best be served by foraging only at night and by sequestering themselves in the nest during the day. The eastern tent caterpillar, for example, has over 200 predators and parasitoids (Fitzgerald 1995), many of which might be avoided if the caterpillars hid in the nest during the daylight hours. Why then do the larvae of this species and those JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TABLE 1. Comparison of traits of five species of social caterpil- lars. Shaded box highlights differences between species 1-2 which forage in both daylight and darkness and species 3-5 which forage only in darkness. 1 = Malacosoma americanum, 2 = Eriogaster lanestris, 3 = Gloveria sp., 4 = Thaumetopoea pityocampa and 5 = Eucheira socialis. See text for references. Species Trait 1.2 Some aaes 1. Central place foraging Vas Ve Ay, 2. Univoltine y y V Va 2. Colonies average 200-300 siblings initially Yor Ya, SVR, eae 3. Colonies construct large, silken shelters y yW y Ve ways 4. Larvae have discrete, coordinated, bouts of en masse feeding interspersed with periods of rest y y y yo OY . Larvae experience warm days, cool/cold nights . Caterpillars distasteful/urticating . Larvae form aposematic aggregations on the nest in daylight y y n n on . Nest may overheat in sunlight causing evacuation Ot << 1D ie) 9. Rapid, progressive deterioration of host leaf quality y y n iy ia 10. Larvae grow rapidly y y n n on of E. lanestris commonly rest on the outside of the nest and feed away from the structure during the daylight hours, a foraging strategy distinctly different than that of T: pityopcampa? The answer may be sought in a suite of traits that distinguish these two species from the three nocturnal foragers (Table 1, traits 7-10). The caterpillars M. americanum and E. lanestris, both early spring feeders, are unable to feed on the aged leaves of their host trees. Their need to grow rapidly, in a race against declining food quality, may compromise safety for growth, favoring caterpillars that feed both day and night. Additionally, the nests of these species may easily overheat on hot and sunny days forcing the caterpillars to evacuate the structures and thus compromise their role as secure retreats (Joos et al. 1988, Ruf & Fiedler 2002). Both species are conspicuous against the white back ground of the nest during the day. Both are hairy and E. lanestris is reported to be urticating (Ruf & Fiedler in press). Both feed on species of Prunus whose cyanogenic glycosides may offer some defense against predators when regurgitated as cyanide (Peter- son 1987, Fitzgerald et al. 2002). Thus, aposematism and distastefulness may offset the risk of daytime expo- sure to some extent. More enigmatic is the fact that the colonies of E. so- cialis, Gloveria sp., and T. pityocampa feed only noc- turnally and do not take advantage of the warm day- light hours to collect additional food. Although it is not known if E. socialis is distasteful to predators, Gloveria VOLUME 57, NUMBER 3 sp. and particularly the older instars of T. pityocampa are urticating (Vega et al. 1999), and it would appear that they would be as well or better defended than M. americanum or E. lanestris were they to feed in the daytime. Perhaps most significant is the fact that these three nocturnally active species feed non-selectively on the leaves of their host trees and have no pressing need to accelerate their rates of feeding to keep pace with a seasonal decline in host quality. Furthermore, the nests of these species are denser and more opaque to radiation than those of M. americanum and E. lanestris and there is no evidence that the whole of these structures can become uninhabitable due to overheating. They therefore constitute dependable daytime retreats from predators. Thus, as may be the case for M. americanum and E. lanestris, none of these species needs to compromise safety for growth. ACKNOWLEDGMENTS We thank Joan Panades i Gallart, Merce Panades i Blas, and Rosa Maria Gallart i Pujol without whose kind support this project would not have been possible. M. Benincore-Posse assisted in the collec- tion of field data. Alfonso Pescador-Rubio provided the Spanish translation of the abstract. The Catalonia Meterology Service pro- vided the seasonal temperature data. LITERATURE CITED AITCHISON, C. W. 1983. Low temperature and preferred feeding by winter-active collembolans. Pedobiologia 25:27—36. . 1984. Low temperature feeding by winter-active spiders. J. Arachnol. 12:297-305. BREUER, M. & B. DEvKoTA. 1990. Studies on the importance of nest temperature of Thaumetopoea pityocampa (Den. & Schiff.) (Lep., Thaumetopoeidae). J. Appl. Entomol. 109:331—335. BREUER, M., B. DEVKOoTA, E. DOUMA-PETRIDOU, A. KOUTSAFTIKIS & G. H. Scumipr. 1989. Study on the exposition and tempera- ture of nests of Thaumetopoea pitycampa (Den. & Schiff.) (Lep., Thaumetopoeidae) in Greece. J. Appl. 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Englewood Cliffs, New Jersey. Journal of the Lepidopterists’ Society 57(3), 2003, 168-175 DEFENSE MECHANISMS IN PYRALIDAE AND CHOREUTIDAE: FECAL STALACTITES AND ESCAPE HOLES, WITH REMARKS ABOUT COCOONS, CAMOUFLAGE AND APOSEMATISM ANNETTE AIELLO Smithsonian Tropical Research Institute, Box 2072, Balboa, Ancon, Republic of Panama, email: aielloa@tivoli.si.edu AND M. ALMA SOLIS Systematic Entomology Laboratory, PSI, ARS, USDA, Smithsonian Institution, P.O. Box 37012, National Museum Natural History, E-517, MRC 168, Washington, D.C. 20013-7012, USA, email: asolis@sel.barc.usda.gov ABSTRACT. A novel behavior, linking fecal pellets with silk on the underside of leaves, to form what look like slender brown stalactites, is reported in two species of Pyralidae and two species of Choreutidae from the Republic of Panama. These fecal stalactites, constructed in asso- ciation with escape holes cut in leaves, may function as landmarks to locate those holes and/or as decoys or camouflage. We discuss fecal stalac- tites, camouflage and aposematism, and cocoons in these larvae as part of a multiple defense system. We provide the first larval description for Monoloxis flavicinctalis. Additional key words: Abaera, Brenthia, Monoloxis, Panama, host plants. Larvae of some lepidopterans living within enclosed spaces, e.g., leaf-eating species that roll leaves or cut and fold leaf shelters, remove fecal pellets, at times ex- plosively, from their habitations (e.g., Friedlander 1987, Rawlins 1984, M. Weiss pers. com., AA pers. obs.). These behaviors may have evolved to avoid pathogens (Rawlins 1984) or to eliminate olfactory cues that might attract predators or parasitoids (Stamp & Wilkens 1993). Contrary to this, some larvae use and live with fe- cal pellets in their habitations, e.g., many pyraloid lar- vae, including stored product pests (MAS pers. obs.). We document a novel use of fecal pellets by the lar- vae of four moth species. We use “fecula” and “fecal pellets” to refer to larval excrement, reserving “frass” for “The chips or particles cast aside by wood borers” (Frost 1959). The larvae of Monoloxis flavicinctalis (Sepp, [1852]) (Pyralidae: Chrysauginae) (Figs. 2-4), Abaera nactalis Walker, [1859] (Pyralidae: Chrysaugi- nae) (Figs. 6, 7), and two species of Brenthia Clemens, 1860 (Choreutidae: Brenthiinae) chew one or more escape holes near the blade midvein, then link their fe- cal pellets using silk to form what look like slender brown stalactites suspended from the underside of the leaf (Figs. 4, 7, 8). In addition, M. flavicinctalis and Brenthia sp. 1 are here reported to construct cocoons of fecula. We discuss the function of fecal stalactites and es- cape holes, briefly address the evolution of fecal sta- lactites in Brenthia species, in contrast to those species that do not construct them, and review other struc- tures that appear similar to fecal stalactites, with simi- lar or dissimilar functions. We also address the host plants of these four moth species and possible apose- matism of their larvae. MATERIALS AND METHODS The four species were collected as larvae (repre- senting various stadia), or pupae, on the dates and at the localities (all in the Republic of Panama) listed in Table 1. They were reared in petri dishes or in small cages fashioned from petri dishes and window screen- ing, and placed in Ziploc® bags with folded, moistened paper towel strips to regulate humidity. Their behavior was observed and recorded daily (with few excep- tions), and shed head capsules and pupal exuviae were collected and mounted. Larvae were preserved by bringing them to a boil in distilled water, then drop- ping them into 80% ethanol. The two Brenthia are not identified to species. Brenthia species-level identifications are possible only with genitalic dissection of males and only if the spec- imen belongs to a species described by Meyrick and il- lustrated by Clarke (1969) (V. Becker in litt.). Adult specimens and exuviae of Maracayia chlo- risalis Walker (Aiello lot 1978-45) and Monoloxis flavicinctalis (Aiello lot 1979-73) are deposited at the National Museum of Natural History (NMNH), Smithsonian Institution, Washington, D.C., U.S.A. All material relating to the remaining rearings, including other specimens of M. flavicinctalis, and plant vouch- ers, are at the Smithsonian Tropical Research Institute (STRI), Republic of Panama. In the accounts to follow, lot numbers are those of Aiello, and consist of the year plus a sequential number. When more than one individual is reared, an individual number (#) is appended. Thus “lot 1979-73 #2” refers to individual #2 of the 73rd lot for the year 1979. These numbers appear on the labels of all reared specimens VOLUME 57, NUMBER 3 169 Table 1. Collection and developmental data as of days in each stadium) and outcomes. Numbers include days spent preparing for molting or pupation, i.e., not eating. Final date is the date of eclosion, death, or preservation, and is not included in durations. If cocoon con- tents were not visible, only the duration in the cocoon is given. Eclosion dates are for the morning immediately following adult nocturnal emergence. Minimum durations (>) are given for the stages collected or for stages cut short by preservation or natural death. A = indicates a molt that may have occurred on a day when observations were not made. A lowercase “p” next to an individual indicates it was parasitized. Instar Instar Instar Instar Instar Cocoon/ Final Name and collection data Lot# Indiv# a b c d e pupa date Outcome PYRALIDAE (CHRYSAUGINAE): Monoloxis flavicinctalis, on Lacistema aggregatum Canal Area, Barro Colorado Island Snyder-Molino Trail-5.9 25 May 1979, A. Aiello 0n00002865 60 SHORE ACOOE ECE OECE PEE ee See 1979-73 1 >2 18 >12P 27 Jun Pupa died, discarded 1979-73 2 >I 12 15P 23 Jun Adult Canal Area, Barro Colorado ised Brokaw Ridge (off Balboa Trail-10) 25 August 1988, A. Aiello & E. Leigh RE eco oe oiciueaesuisans vavehabetbdd cadesseae 1988-19 1 >7 30C 1 Oct Adult 2000088000000 000d ODOBAS HEB ADS Ge TBE EEO RO BERe HOSE ener 1988-19 2 >7 — 1 Sep Larva preserved Panama Province, Arraijaén Loma del Rio 12 December 2001, A. Aiello RN re are ten ide ctscateatetaats 2001-44 25 39C+15P 9 Feb Pupadied, pointed nadonason0bbiCOO US BOEEEEC RIO SERE SRE cee BeeR EeePC eee 2001-44 2 Pal — 13 Dec Larva preserved Abaera nactalis, on Cordia panamensis Canal Area, Summit, Old Gamboa Road 27 June 1990, D. Windsor soneecisosbocedUsSAbaOSoe ROO SEERA BEER Eee ES ee 1990-7 1 22 =O) =6 8 9 37C 3 Sep Adult CHOREUTIDAE (BRENTHIINAE): Brenthia sp. 1, on Cojoba rufescens Panama Province, Arraijan Loma del Rio 12 January 1992, A. Aiello Bee ce aoc aiedne sre sctsscnsusunsw nen acsnntattye 1992-5 a — 12 Jan Empty cocoon cocobadbosddodasonseecee Soe Par eS uSCE ET CEN enc cee 1992-5 p2 27 Jan Parasitized, wasp pupa died 3 25 2C+11P 30 Jan Adult 4 215C 27 Jan Adult Brenthia sp. 2, on Calathea sp. Canal Area, Barro Colorado Island 26 August 1993, D. Windsor tadbea650aHS 000007 USaC ODIO SEE RECO ECP EBEDEELOR EEE EERE CEE 1993-70 Il 212C 7 Sep Adult ON ON ee eee t cece: estbue casts 1993-70 2 >12C 7 Sep Adult soca GOBRiAbe SAREE COS RAB STRAY SA a nO st 1993-70 3 >12C 7 Sep Adult Brenthia sp. 2?, on Calathea sp. Panama Province, Cerro Jefe Conservation Trail 10 September 1993, A. Aiello, D. Windsor & J. Miller cdncoacdeaacoeconnceen Me Seen erste center LOOO=No mpl 2] 1C+8P 20 Sep Parasitized, wasp adult scdoboab os punDOG EEG EoREE BE RBSEA DEE EERE EE DRAENOR BS aECeE 1993-73 2 Pal = 11 Sep Larva preserved 170 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 1-4. Monoloxis flavicinctalis (Sepp) (Pyralidae: Chrysauginae), reared on Lacistema aggregatum (P. J. Bergius) Rusby (Flacour- tiaceae), as Aiello lot 1979-73. 1, Pinned adult. 2, Mature larva, eating. 3, Larva (anterior portion), moving to upper surface of leaf, through es- cape hole. 4, Larva (posterior portion), returning to lower surface of leaf, by backing through escape hole. Photographs by A. Aiello. and their associated parts, and correspond to numbers on daily data forms maintained by Aiello at STRI. RESULTS Monoloxis flavicinctalis (Sepp, [1852]) (Pyralidae: Chrysauginae) (Figs. 1-4, 9) All six Monoloxis flavicinctalis larvae were found on mature leaves of Lacistema aggregatum (P. J. Bergius) Rusby (Flacourtiaceae). Table 1 gives collection and developmental data. Of these six individuals, two were preserved as larvae, two died as pupae, and two were reared to adults. In each larva the head was dark red and the body was purplish black, with white markings (dorsal and lateral on T1 and A7, and lateral on A2, A5, and A8—A10) (Fig. 2). The larvae lived on the under- surfaces of leaves, each within a wispy tangle of silk supported by a series of three or four, flexible, brown stalactites, several mm apart along one side of the mid- vein, and constructed by linking as many as 50 fecal pellets and securing them with silk. Each larva rested with its head near a small, neatly rounded hole located to one side of the midrib and near a fecal stalactite. When touched, the larva moved rapidly head first through the hole to the upper surface of the leaf, which was totally bare (Fig. 3). After a few seconds without further disturbance, it backed down through the hole into its silk tangle (Fig. 4). Except for these escape maneuvers, larvae remained within their silk tangles, extending out of them only to eat surrounding leaf tissue. In the only instance that new shelter con- struction was witnessed (lot 2001-44 #1), the larva pro- duced the escape hole before anything else. Molting took place within the silk tangles. Cocoons were constructed of fecula and silk on the leaf or on the container floor. The adults were brown, with or- ange forewing apices (Fig. 1), and rested with the forewings covering the hind wings, in a broad, flat, tri- angle. Plant vouchers for lot 2001-44 are Aiello 1582 and 1635. Though the setal pattern of M. flavicinctalis is typi- cal of the Chrysauginae, the larvae are distinctively patterned, an unusual trait within the Pyralidae. We provide the first larval description for the genus, to- gether with an illustration (Fig. 9). A pupa was not available for description. Larva (Fig. 9): Length: 18 mm (n = 1) (final instar). Head with reddish brown platelets; beige or white between platelets. Epicra- nial suture present. White between L1] and A3, medially across head and on frons dorsal to F1, on either side of epicranial suture, VOLUME 57, NUMBER 3 Fics.5-8. Abaera nactalis Walker (Pyralidae: Chrysauginae), reared on Cordia panamensis L. Riley (Boraginaceae), as Aiello lot 1990-7. 5, Pinned adult. 6, Penultimate larva within fecula and silk tangle beneath leaf; the largest dorsal, yellow marks are on Al. 7, Larva camouflaged within fecula and silk tangle; dark patches are feeding areas. 8, Larval shelter back-lighted, showing fecal stalactites, escape hole (arrow), and scraped feeding areas. Photographs by Carl Hansen. and ventroposterior to stemmata. Frontoclypeus area ventral to F1, including clypeus dark brown; anteclypeus white; labrum yel- lowish brown. Adfrontal area light brown. Mandibles yellowish brown with dark brown margins. T1-3 and Al—10 integument lightly rugose. With sclerotized rings at the bases of D2 on T3 and SD1 on AS. Prothoracic shield white, dark brown between D2 and SD2 along posterior margin and extending length of medial line. XD1, XD2, SD1 with small dark brown pinacula. T1 with lobes an- terior to thoracic legs and prothorax; 2 L setae below and anterior to spiracle on brown pinaculum. T1-3 legs with basal segments sclerotized dark brown; tarsus white. T2—3 with D1—D2, SD1- SD2, and L1—-L2 on same pinaculum; SV1 and L3 with one seta on separate pinacula. T1-3 and Al, light brown ventrally; A2-A10 white ventrally. A2, A5, and A7 with white areas anterior to spira- cle and SD1 and extending dorsal to SD1, but ventral to D setae. A7 also with a white area joining D2 on both sides. A8 with white between SD1 and D1 and between both D2 setae. A9 primarily white; pinaculum dark brown. A10 primarily white with brown mottling between SD2, D2, and D1. Al-8 with L1 and L2 present on separate pinacula ventral to spiracle; SD1 on a large, brown pinaculum dorsal to spiracle, except A2, A5, and A7 where the pinaculum is small, dark brown, triangular; D1 and D2 setae on separate small, round, dark brown pinacula. Al—-6 with three SV setae, A7—8 with two SV setae. Al—8 with one L3 seta. SD2 of Al anterior to SD1, SD2 of A2, A3, A4, A6, and AS anterodorsal to spiracle; SD2 not present on A2, A5, and A7. SD1 pinaculum on A8 protruding, seta at least 20 times the length of other setae. Spiracle on AS at least twice as large and slightly more dorsal than other abdominal spiracles. A9 with three L setae on same pinacula; D1, D2, and SD1 on separate pinaculum. Prolegs with crochets biordinal in a circle. Abaera nactalis Walker, [1859] (Pyralidae: Chrysauginae) (Figs. 5-8) The single Abaera nactalis larva (lot 1990-7) was found on a mature leaf of Cordia panamensis L. Riley (Boraginaceae). Table 1 gives collection and develop- mental data. The head was patterned with white and dark brown, and the body was checkered dark brown, pale brown, and bright yellow (Fig. 6). Like M. flavi- cinctalis, it lived within a loose, silk tangle supported by flexible fecal stalactites, and had an escape hole (Figs. 6-8) through which it scooted to the upper sur- face of the leaf when we disturbed it. Unlike M. flavi- cinctalis, this larva decorated the silk tangle with nu- merous individual fecal pellets, which, in conjunction with the complex markings of the larva, provided messy but highly effective camouflage (Fig. 7). Early instars ate only the tissue of the leaf undersurface, pro- ducing extensive, brown scraped patches bounded by the secondary veins (Fig. 7). The final instar ate areas of leaf, veins and all. Portions of three shelter-building efforts were ob- served. The first of these new shelter building events JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY prothorax mesothorax metathorax abdominal segment 6 abdominal segments 8-10 Fic. 9. Monoloxis flavicinctalis (Sepp) (Pyralidae: Chrysauginae), setal maps of larva body (above) and head (below). Illustrations by A. Solis. took place 5 July on the original leaf, and consisted of an escape hole near the midrib, one fecal stalactite, a silk tangle sprinkled with fecula, and two scraped feed- ing areas. When the leaf began to turn black, the larva was prodded onto a fresh leaf, thus precipitating the second building episode. Within 30 minutes the larva had moved to the underside of the leaf and chewed an escape hole next to the midrib. During the next 45 minutes, it constructed a fecal stalactite (4 mm long) near the hole. The next morning a second fecal stalac- tite (10 mm long), a silk tangle, and a small feeding area had been added. And the day after that a third stalactite (17 mm long) appeared. The third new shel- ter effort took place 20 July, after the newly molted fi- nal instar was transferred to a fresh leaf. The larva first chewed a new escape hole, then constructed several fe- cal stalactites and a silk tangle. The following day (21 July), it fashioned a cocoon-like shelter of fecula and silk instead of more stalactites, and began eating whole leaf rather than simply scraping the blade surface. The mature larva curled into an O next to the midrib at a leaf base and constructed a cocoon of silk and leaf material. The finished cocoon was cream color on the inside, dark brown on the outside, and was covered with tufts of leaf trichomes. The adult male, brown, with powder-blue forewing apices (Fig. 5), eclosed the night of 2-3 September. It held the wings in a slightly tented triangle. Brenthia Clemens, 1860, sp. 1 (Choreutidae: Brenthiinae) One larva and three fecula and silk cocoons (lot 1992-5) of a small species of Brenthia (6 mm wing span) were found on the undersurface of mature leaflets of Cojoba (=Pithecellobium) rufescens (Benth.) Britton & Rose (Fabaceae: Mimosoideae), in sec- ondary growth. Table 1 gives collection and develop- mental data. The three cocoons were suspended hori- zontally among stalactites, within fecula-sprinkled silk webbing. Cocoon #1 contained only a cast larval skin. Cocoon #2 held a pupa, which protruded from its housing and within which a wasp pupa could be seen clearly; the wasp pupa failed to develop, then molded and was discarded. The only larva (#3) was among fecula-sprinkled webbing and had made several neatly rounded holes in the leaflet, permitting rapid passage to the upper sur- face. Each hole had a flexible fecal stalactite next to it. The larva ate the undersurface leaf tissue only, pro- ducing small scraped patches. Cocoon #4 contained a healthy pupa. Individuals #3 and #4 yielded adults. Prior to eclosion, pupae protruded from their cocoons. VOLUME 57, NUMBER 3 Adults displayed in their petri dishes. This and the next species are among the microlepidopteran “pea- cock moths” that are seen frequently in the Canal wa- tershed area, performing a raised wing display (Aiello & Becker in prep.) on foliage, sometimes several indi- viduals per leaf. Brenthia Clemens, 1860, sp. 2 (Choreutidae: Brenthiinae) Table 1 gives collection and developmental data. Three cocoons (lot 1993-70) of a larger Brenthia species (1 cm wing span, versus 0.7 cm) were found on the undersurface of Calathea sp. (Marantaceae) leaves. Escape holes, short fecal stalactites, silk tan- gles, and scraped feeding areas on the leaf undersur- face indicated a larval life style similar to that of the preceding Brenthia species. However, this species was found on a monocotyledonous plant, and instead of constructing a cocoon of fecula and silk, it spun a white silk cocoon. The cocoon was composed of three parts. The main part, an elongate spindle-shaped structure that housed the pupa, rested suspended within a cloud of wispy silk between two silk sheets; the top sheet was flat and had numerous holes, espe- cially towards its margins, and was anchored all around to the leaf; the bottom sheet was creased lengthwise to form a V-shaped trough, shorter than the top sheet, and anchored to it along its sides. The final larval skin had been pushed out of one end of the spindle and into the silk cloud. Prior to eclosion the pupae pro- jected from their cocoons. All three cocoons yielded adult females. And all three moths dashed about their petri dishes displaying in the same manner as the smaller species. One of two larvae found on Cerro Jefe (Table 1, lot 1993-73), also on the leaves of Calathea sp., spun a white, three-layered cocoon like those of lot 1993-70, and al- most surely was Brenthia, very likely Brenthia sp. 2. Fol- lowing pupation, it pushed its final larval skin out of one end of the spindle-shaped cocoon. On the mis- taken conviction that an adult would be obtained from it, the other larva was preserved. Alas, a small braconid wasp emerged from the cocoon and no adults were ob- tained from this rearing. DISCUSSION The larval constructions, i.e., fecal stalactites and es- cape holes, and associated behavior described above are part of a complex defense system that includes camouflage and possibly aposematism. Fecal stalac- tites are always near an escape hole, and appear to act as landmarks to help larvae locate the holes and escape quickly. The escape hole and its accompanying stalac- tite were the first items to appear in the shelter con- struction sequences, underscoring their importance to larval survival. Unlike the larval behavior of the two Brenthia species described here, several other species of Brenthia fashion escape holes, but not stalactites. Specimen information and previous descriptions of Brenthia biology in two different parts of the World do not report the building of fecal stalactites. Brenthia pavonacella Clemens, reared by Busck in the United States (NMNH: specimens and leaf remains), lived among fecula-dotted silk tangles beneath leaves. In Japan, B. japonica Issiki (Arita 1971, Issiki et al. 1975) and B. pileae Arita (Arita 1971) have been reported to fashion escape holes and fecula-laden silk tangles; fig. 230 in Issiki et al. (1975) shows a larva that has “es- caped” through its hole. Similar behavior is found in an unrelated, more primitive, microlepidopteran species, Compsistis Meyrick (Elachistidae: Depres- sariinae), whose larvae cut escape holes but do not construct fecal stalactites, beneath mature leaflets of Pseudobombyx septenatum (Jacq.) Dugand (Bomba- caceae) (AA pers. obs. lot 1993-94). Because escape holes appeared before fecal stalactites in shelter con- struction and because some species of Brenthia create escape holes without fecal stalactites implies that es- cape holes may have come first in the evolutionary se- quence of this behavioral pattern. This idea could be tested by conducting a worldwide phylogenetic study of Brenthia. There are several structures in arthropods, includ- ing in other Lepidoptera, that are reminiscent of fecal stalactites. The sand pillars constructed by some fid- dler crabs help them locate their burrows rapidly and thus avoid predation (Christy 1991, 1995). In Mono- loxis and Brenthia, fecal stalactites may function sec- ondarily as decoy larvae, and they recall the “fake” lar- vae fabricated by early instar Adelpha_ basiloides (Bates) (Nymphalidae) (Aiello 1984). Fecal stalactites remind us of the horizontal and more rigid fecal rods produced by early instars of many Nymphalidae, ei- ther as continuations of leaf veins or as formations an- chored to the leaf margin. The techniques involved in construction of fecal rods and stalactites may be simi- lar, but their functions are quite different. Fecal rods are used as resting and molting perches by earliest in- star nymphalids and are thought to provide both cam- ouflage and safe haven from ants and other predators (Machado & Freitas 2001). Many nymphalids enhance those protections by barricading the base of the struc- ture with loosely attached leaf bits (Muyshondt & Muyshondt 1979) or, in Adelpha spp., clusters of fecal pellets (Aiello 1984). Though fecula cocoons are not common among Lepidoptera, examples are found in several families besides those reported here for Pyralidae and Chore- utidae, e.g., Synanthedon spp. (Sesiidae) (Barrett 1997), Mimallo amilia (Cramer) (Mimallonidae) (AA pers. obs. lots 1985-131, 1987-45, 1990-54, 1997-33, 2002-27). Fecula cocoons might help protect their oc- cupants from parasitoids and predators, but their ef- fectiveness has not been tested. Among known Bren- thia, Brenthia sp. 1 is unique, so far, in using fecal pellets to construct its cocoon. All other Brenthia species for which we have information spin white silk cocoons: B. coronigera Meyrick in India (Fletcher 1920, NMNH: Rangi specimen), B. japonica (Arita 1971, Issiki et al. 1975), B. pavonacella (NMNH: Busck specimens), B. pileae (Arita 1971), and Brenthia sp. 2 in Panama (this paper). As well, white silk co- coons among webbing are found in another member of the Choreutidae, Hemerophila albertiana (Stoll) (AA pers. obs. lot 2001-39). It would be a challenge for a predatory or parasitoid wasp to breach one of these multi-layered silk cocoons. Additionally, our observations indicate that though camouflaged, the larvae of M. flavicinctalis and A. nac- talis may also be aposematic, exhibiting both warning coloration and unpalatability (Bowers 1993). It is known that the degree of pigmentation in lepi- dopteran larvae tends to correlate positively with de- gree of exposure to visually hunting predators (Stamp & Wilkens 1993). Exposed feeders include mimetic, camouflaged, or cryptically patterned species as well as colorful aposematic ones (Bowers 1993, Stamp & Wilkens 1993). The larvae of hidden feeders tend to be colorless, or they may appear green or brown due to their gut contents, or white due to fat body, and in species that extend from their shelters to feed, or that reside in moveable cases, the head and prothorax are pigmented and the rest of the body is not (AA pers. obs.). In contrast to conventional notions that most camouflaged pyraloid larvae are watery-looking cater- pillars with pale or clear cuticles, M. flavicinctalis and A. nactalis are well-pigmented. The larva of A. nactalis is strikingly colored yellow and brown, and well camouflaged within its fecula- sprinkled webbing. As well, it may be protected from chance exposure to predators by chemicals obtained from its food plant, Cordia panamensis, a short-lived, second growth tree that also hosts six species of metal- lically colored tortoise beetles: two species of Omo- cerus Chevrolat and four of Discomorpha Chevrolat (Chrysomelidae: Cassidinae) (Windsor et al. 1992). The larva of M. flavicinctalis, less well camouflaged than that of A. nactalis, quite likely derives chemical protection from its food plant, Lacistema aggregatum, JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY a shrub or small tree that also is host to an as yet unidentified sexually dimorphic, wasp-mimicking diur- nal moth (Arctiidae: Ctenuchinae) (AA pers. obs. lot 1999-8). The Flacourtiaceae belong to the Violales, a cluster of families notable for supporting an array of aposematic lepidopterans, e.g., Heliconius spp. (Nymphalidae) on members of the Passifloraceae (Benson et al. 1976, Trigo 2000), Josia draconis Druce (Notodontidae: Dioptinae) on Turnera panamensis Urb. (Turneraceae) (AA pers. obs. lots 1994-37, 1994- 39, Miller 1996); Siderone marthesia (Cramer) (Nymphalidae) on Casearia guianensis (Aubl.) Urb. (Flacourtiaceae) (AA pers. obs. lots 1990-25, 1996-28, 2000-37); Zunacetha annulata Guérin (Notodontidae: Dioptinae) on Hybanthus prunifolius (Humb. & Bonpl.) Schulze-Menz (Violaceae) (Wolda & Foster 1978, AA pers. obs. lots 1977-25, 1979-26, 1997-11), to mention a few. Aposematism to protect camouflaged Pyraloidea lar- vae against chance exposure may be a more common defense mechanism than has been reported in the lit- erature. In addition to the two chrysaugine species dis- cussed above, the first author has reared the larva of Maracayia chlorisalis Walker (Crambidae: Spilomeli- nae), whose clear cuticle and large, black pinacula camouflage it beneath a silk and fecula tangle on the broad, succulent leaves of its foodplant, an epiphytic cactus, Epiphyllum phyllanthus (L.) Haw. (Cactaceae) (lot 1978-45); and the aposematic (white, ornamented with black pinacula and yellow supraspiracular blotches) larva of Palpita flegia (Cramer) (Crambidae: Spilomelinae) that eats the leaves of a toxic plant, Thevetia ahouai (L.) A. DC. (Apocynaceae) (lot 1984- 60). The adults of the latter two are white, the color most conspicuous and therefore most aposematic at night. The evolution of the ability to sequester defen- sive compounds as larvae and retain them into the adult stage has not been well studied (Bowers 1993). It is doubtful that Brenthia larvae, being small and inconspicuous, rely on chemical protection from their host plants. If any do so, the most promising host plants, as far as plant secondary compounds are con- cerned, would be the Sapindaceae, which is host to B. elongata Heppner in the West Indies and B. sapindella Busck in Cuba (Heppner 1985). Another group of sec- ondary compound candidates among known Brenthia host plants would be the Fabaceae, which are known to support a number of Lepidoptera aposematic as lar- vae and/or adults, i.e., Ormetica sicilia Druce (Arcti- idae) on Inga sp. (AA pers. obs. lot 1980-44), Utetheisa ornatrix L. (Arctiidae) on Crotalaria cajanifolia Kunth (Fabaceae: Papilionoideae) (Trigo 2000), Melanis pixe (Boisduval) (Riodinidae) on Albizia adinocephala VOLUME 57, NUMBER 3 (Donn. Sm.) Britton & Rose (AA pers. obs. lots 1988- 30, 1991-5). In addition to Brenthia sp. 1 in Panama, the Fabaceae are hosts to at least two other Brenthia species: B. albipunctata Arita on Spatholobus compar Craib in Thailand (Arita 1987) and B. pavonacella on Desmodium sp. in the U.S.A. (NMNH: Busck speci- mens from Great Falls, Virginia, 17 and 18 July 1913; Falls Church, Virginia, 5 August 1913) as well as on Inga vera Willd. at Lares, Puerto Rico, where “. . . in November 1931, Mr. Francisco Sein found them abundant, feeding on the underside of the leaves. . . (Wolcott 1948). In conclusion, fecal stalactites and escape holes are two mechanical constructions that may enhance larval survivorship in some species of Pyralidae and Chore- utidae, and may be just part of a multiple factor de- fense system (Bowers 1993) that includes camouflage and aposematism, against a variety of enemies, preda- tors or parasitoids, at different times of the night and day. ACKNOWLEDGMENTS Our thanks go to Jon Lewis (Systematic Entomology Laboratory, USDA) for his generous help with the literature and with locating specimens; to Vitor O. Becker (Research Associate, Universidade ae Brasilia, Brasil) for identifying the moths; to Donald Windsor (STRI) for collecting the A. natal and Compsistis larvae, and to Carl Hansen for photographing the former; to Walter Aiello (Duke University Medical Center), Naoki Takebayashi (Duke University Department of Biological Sciences), and K. T. Park (Kangweon Na- tional University, Chuncheon, Korea) for kindly translating the Japa- nese text; to Mare Epstein (Department of Systematic Biology, NMNH), David R. Smith and Steve Lingafelter ‘(Sy ystematic Ento- mology Laboratory, USDA), Bernard Landry (Muséum dhistoire naturelle, Geneva, Switzerland), Priya Davidar (Pondicherry Uni- versity, India), Jean-Philippe Puyravaud, and an anonymous re- viewer for divest helpful reviews of the manuscript; and to Martha Weiss, Georgetown University, for insights into the opposite larval behavior, the removal of fecal material. LITERATURE CITED AIELLO, A. 1984. Adelpha (Lepidoptera: Nymphalidae): deception on the wing. Psyche 91 (1—2):1-45. AIELLO, A. & V. BECKER. (submitted). Display of the “peacock moth.” [Submitted to J. Lepid. Soc.] Arita, Y. 1971. Genus Brenthia (Lepidoptera: Glyphipterygidae) of Japan. Kontyti 39 (4):346-352. . 1987. Brenthiinae (Lepidoptera, Choreutidae) of Thai- land, pp. 16 and 17. In Osaka Furitsu Daigaku (ed.), Microlepi- doptera of Thailand: scientific results of lepidopterological ex- peditions of the University of Osaka Prefecture to Thailand. Entomological Laboratory, University of Osaka Prefecture. Barretr, B. A. 1997. Insect borers of fruit trees. MU Guide G7190. 6 pp. University Extension, University of Missouri, Colombia. BENSON, W. W., K. S. BROWN, JR. & L. E. GILBERT. 1976. Coevolu- tion of plants and herbivores: passion flower butterflies. Evolu- tion 29:659-680. Bowers, M. D. 1993. Aposematic caterpillars: life-styles of the warningly colored and unpalatable, pp. 331-371. In Ste amp, N. E. & Casey, T. M. (eds.), Caterpillar: ecological and evolution- ary constraints on foraging. Chapman & Hall, New York. CHRISTY, eS ONE Comparative studies of reproductive behavior in mantis shrimps and fiddler crabs. Am. Zool. 31:329-337, . 1995. Mimicry, mate choice, and the sensory trap hypoth- esis. Am. Nat. 146 (2):171-181. CLARKE, J. F.G. 1969. Catalog of the type specimens of Microlepi- doptera in the British Niccum (Natural History) described by Edward Meyrick. Vol. VI, pp. 24-39. London: British Museum (Natural History). FLETCHER, T. B. 1920. Life-histories of Indian insects—Microlepi- doptera. Mem. Dept. Agr. India, Entomological Series 6:1-217. FRIEDLANDER, T. 1987(88). Taxonomy, phylogeny, and biogeogra- phy of Asterocampa Réber 1916 (Lepidoptera: Nymphalidae, Apaturinae). J. Res Lepid. 25 (4):215-338. Frost, S. W. 1959. Insect life and insect natural history. 2nd re- vised ed. Dover. New York, viii + 526 pp. HEPPNER, J. B. 1985. West Indies Brenthia (Lepidoptera: Chore- utidae). Insecta Mundi 1 (1):13—26. Isstki1, S., A. MuruuraA, Y. YAMAMOTO, I. Harrort, H. Kuroko, T. Kopama, T. Yasupa, S. Moriuti & T. Saito. 1975. Early stages of Japanese moths in colour. Vol. If. Hoikusha Publishing Co., Ltd., Osaka. 237 pp. Macuapo, G. & A. L. Freivas. 2001. Larval defence against ant predation in the butterfly Smyrna blomfildia. Ecol. Entomol. 26:436—-439. MILLER, J. S. 1996. Phylogeny of the Neotropical moth tribe Josi- ini (Notodontidae: Dioptinae): a hidden case of Miillerian mim- icry. Zool. J. Linn. Soc. 118:1-45. MuysHonpt, JR., A. & A. MuysHonpT. 1979. Notes on the life cy- cle and natural history of butterflies of El Salvador. Ic. His- toris odius and Coea acheronta (Nymphalidae: Coloburinae). J. Lepid. Soc. 33:112-123., RAWLINS, J. E. 1984. Mycophagy in Lepidoptera, pp. 382-423. In Wheeler, Quentin & Blackwell, Meredith (eds.), Fungus-insect relationships. Colombia University Press, New York. Stamp, N. E. & R. T. WILKENS. 1993. On the cryptic side of life: be- ing unapparent to enemies and the consequences for foraging and growth of Sita pp. 283-330. In Stamp, N. E. & Casey, T. M. (eds.), Caterpillar: ecological and evolutionary con- straints on foraging. Chapman & Hall, New York. Trico, J. R. 2000. The chemistry of antipredator defense by sec- ondary compounds in Neotropical Lepidoptera: facts, perspec- tives and caveats. J. Braz. Chem. Soc. 11 (6):551-561. WINDSOR, D. M., RILEY E. G. & STOCKWELL H. P. 1992. An intro- duction to the biology and systematics of Panamanian Tortoise Beetles. In Quintero A., D. & Aiello, A. (eds.), (1992). Insects of Panama and Mesoamerica: selected studies. Oxford Univer- sity Press. xxii + 692 pp. Wo cort, G. N. 1948. The insects of Puerto Rico. J. Agr. Univ. Puerto Rico 32:418—744. Wo tpa, H. & R. Foster. 1978. Zunacetha annulata (Lepidoptera; Dioptidae), an outbreak insect in a Neotropical forest. Geo- Eco-Trop. 2 (4):443-454. Received for publication 30 August 2002; revised and accepted 20 January 2003. Journal of the Lepidopterists’ Society 57(3), 2003, 176-192 HYBRIDIZATION OF CHECKERSPOT BUTTERFLIES IN THE GREAT BASIN GEORGE T. AUSTIN Nevada State Museum and Historical Society, 700 Twin Lakes Drive, Las Vegas, Nevada 89107, USA DENNIS D. MURPHY Department of Biology/314, University of Nevada, Reno, Nevada 89557, USA JOHN F. BAUGHMAN, ALAN E., LAUNER AND ERICA FLEISHMAN Center for Conservation Biology, Stanford University, Stanford, California 94305, USA ABSTRACT. Two putative species of Euphydryas butterflies, E. anicia and E. colon, may be hybridizing in the north-central Great Basin after an extended period of geographic separation. Surveys were conducted throughout northern Nevada to estimate the distribution of each species and of apparent hybrids. More detailed mark-recapture studies were made at one site in the Pequop Mountains in order to examine eco- logical interactions between the species. The two are largely allopatric and readily separated by wing color and genital morphology. Although in- terbreeding was apparent from the occurrence of intermediate phenotypes and known mating attempts, the taxa are largely temporally segre- gated and prefer different larval hostplants. There is also a suggestion of an unbalanced sex ratio of phenotypically intermediate individuals. These Euphydryas, although closely related and not strict biological species, are undoubted historical entities and seem to be best treated as phylogenetic species. Additional key words: distribution, Euphydryas, genitalia, hostplants, hybridization, Nevada, Nymphalidae, phenology, sex ratio. Many taxa exist as intergrading populations in which individuals from neighboring populations are morpho- logically and ecologically similar, whereas individuals from distant populations are quite distinct. In some in- stances, however, phylogenetically distinct portions of a series of intergrading populations, or of two closely related species, may be sympatric. Such cases of “ring species” have been considered manifestations of al- lopatric speciation providing evidence of the differen- tiation of populations along environmental gradients (Irwin et al. 2001, Irwin & Irwin 2002). Interactions between contiguous or sympatric populations not only augment the understanding of speciation phenomena, but also potentially provide important information on several aspects of population biology (e.g., Endler 1977, Harrison 1993, Bull 1991, Futuyma & Shapiro 1995, Jiggins et al. 1996). Hybrid zones have long intrigued biologists and an abundant literature speculating on the genetic, ecolog- ical, and evolutionary significance of interactions be- tween closely related taxa in such areas has developed (e.g., Sibley 1961, Mayr 1963, Moore 1977, Grant & Grant 1992, Harrison 1993) including one for Lepi- doptera (e.g., Remington 1968, Oliver 1979, Porter 1997, Sperling 1990, Scriber et al. 1995, Porter et al. 1995, 1997, Jiggins et al. 1996). More field research on the interactions between taxa in zones of overlap is needed; most investigations infer ecological interac- tions based on morphological and genetic data (but see Otte & Endler 1989, Lindroth et al. 1988a, b, Porter 1997, Mallet et al. 1998). Little attention has been paid to the extent of interbreeding in these zones (e.g., Blair 1950, Ficken & Ficken 1968, Collins 1984, John- son & Johnson 1985, Nichols & Hewitt 1988, Mallet et al. 1998, Benedict 1999). Ecological and demographic data from hybrid zones may, however, provide infor- mation crucial to reconstruction of paleoecological events, identification of biogeographic patterns, or prediction of future changes in closely related lineages (e.g., Hafner 1992, Scriber & Gage 1995, Benedict 1999). These data are particularly important since many interacting taxa do not readily fit into traditional taxonomic schemes (e.g., Cracraft 1989, Templeton 1989, Sperling 1990). This work focuses on butterflies of the Euphydryas chalcedona (Doubleday) complex (Lepidoptera: Nymphalidae) in the north-central Great Basin, an area where two morphologically distinct forms of the group appear to hybridize. The objectives were to de- termine their geographical overlap in northern Nevada and the present extent and nature of interaction. To address these, populations of Euphydryas were placed into a broad biogeographic context by conducting re- gional surveys of wing color patterns and male genital morphology. To examine ecological interactions be- tween forms in greater detail, concentrated investiga- tions were conducted on a site in the Pequop Moun- tains (Elko County, Nevada) at which both forms were present. At this site, data were collected on wing phe- notypes, genital morphology, and spatial and temporal distributions of the forms. Naturally occurring matings were quantified and oviposition hostplant preferences VOLUME 57, NUMBER 3 were tested as an indicator of recent evolutionary his- tory and potential overlap in hostplant utilization be- tween the forms. In addition, the apparent sex ratio of the Pequop population was compared with sex ratios of allopatric populations to determine if this may be unbalanced in the Pequop Mountains (Haldane 1922). These data were supplemented with information from other sites where more than one form occurs. STUDY SYSTEM The taxa of the Euphydryas chalcedona complex are distributed across much of western North America, from Alaska to Mexico and east to the Great Plains (Scott 1986). The group consists of three nominally distinct species, E. chalcedona, Euphydryas colon (W. H. Edwards), and Euphydryas anicia (Doubleday & Hewitson) (Miller & Brown 1981). These species were defined primarily by the shape of the male genitalia and by wing shape and coloration (Gunder 1929, Bauer in Ehrlich & Ehrlich 1961). Within each of the three putative species, there is considerable between- population phenotypic variation in both wing color (Austin & Murphy 1998b) and male genital morphol- ogy (Scott 1978). Largely in response to this phenotypic variation, nearly 80 names have been proposed for the various forms within the E. chalcedona group, many of which represent aberrations. At present, 11 subspecies are recognized for E. chalcedona, five for E. colon, and 22 for E. anicia (Miller & Brown 1981). Although Ferris (1989) synonymized E. colon with E. chalcedona, there is no consensus on their taxonomic status. Allozyme studies, however, suggested that variation in wing color and pattern was not accompanied by genetic differen- tiation to justify species-level characterization, and the three groups were tentatively lumped into one mor- phologically diverse species, E. chalcedona (Brussard et al. 1985, 1989, Scott 1986). The “messy” systematic situation is compounded by the predominantly allopatric geographic distributions of the forms. Although distribution maps suggest broad sympatry in some areas of the western United States (Stanford & Opler 1993), the seemingly geo- graphically sympatric taxa are usually ecologically al- lopatric, may have somewhat different flight seasons, and often have different larval hostplants. At most spe- cific locations, therefore, only a single member of the complex is present. There are a few locations, how- ever, at which two of the three named entities co- occur (Ehrlich & Murphy 1982, Austin & Murphy 1987, 1998b, Ferris 1988, Brussard et al. 1989). The phylogenetic history of each of the forms has not been fully clarified and it may be difficult to distinguish sec- 120°W 118°W 116°W 114°W i . = i ba { 2 = ey Bess 40°N i ikke Se IE ne rs (a) = E 116°W 114°W Fic. 1. Map of northern Nevada showing distributions of Eu- phydryas anicia (open circles), E. colon (closed triangles), and E. chalcedona (open triangles); closed circles indicate sites with more than one form. Irregular line passing through Elko and Winnemucca is the Humboldt River. Numbers refer to locations coded in Tables 1 and 3. ondary from primary contact and/or intergradation (Mayr 1942, Endler 1977, but see Hammond 1990). Based, however, on the known paleoclimate and paleo- vegetation of western North America (Mifflin & Wheat 1979, Wells 1983, van Devender et al. 1987, Benson & Thompson 1987, Grayson 1993), allozyme data (Brussard et al. 1989), and present distributions, it appears that E. chalcedona and E. colon were south- ern and northern Pacific isolates, respectively, and that E. anicia was isolated somewhere between the Rocky Mountains and Sierra Nevada. It is probable that pres- ent distributions reflect post-Pleistocene dispersal and rejunction and not the result of a single rampantly dif- ferentiating lineage. METHODS Data on the distribution of butterflies of the E. chal- cedona group in northern Nevada were collected as part of ongoing field studies throughout the region. These surveys, initiated in the mid-1970's, were ex- panded specifically to investigate the E. chalcedona complex during the late 1980's and early 1990's. In order to further clarify the distribution of the three E. chalcedona group species, the genitalia of more than 800 male butterflies from 32 sites within Nevada (including one reported by Scott 1978) were scored according to Scott (1978). At seven of these sites, two forms or intermediates of the Ewphydryas chalcedona complex have been recorded (Fig. 1). The remaining 25 sites supported only a single form. 178 TABLE 1. Number of individuals with each wing phenotype among museum specimens of the Euphydryas chalcedona complex from locations with sympatry or intermediates in Elko County, Nevada (location numbers refer to those in Fig. 1). R = red form (“pure” E. anicia), RI = intermediate phenotype (more similar to E. anicia), I = “true” intermediate, BI = intermediate phenotype (more similar to E. colon), B = black form (“pure” E. colon). Wing phenotype Location Sex B BI I RI R 1. Pequop Mountains male 35 3 3 8 Izy female 8 1 0 1 3 2. Windemere Hills male 42 0 Il 0 1 female 11 0 0 0 1 3. Snake Mountains male 11 0 IL 1 ] 4, Independence Mts. (Maggie Summit) male 16 il 0 9 0 female 1 1 0 0 0 5. Independence Mts. (Jack Creek) male 4 2 0 0 6. Owyhee River Valley male 32 2 0 0 0 female 2 O 0 0 0 At a site in the Pequop Mountains, south of I-80 about two miles east of Pequop Summit at a mean ele- vation of approximately 2200 m (41’05’N 114’33’W; see Table 1, Fig. 1), comparatively detailed determina- tions of wing color were made during mark-release- recapture studies (MRR, techniques according to Ehrlich & Davidson 1961) over two seasons. The MRR studies were conducted along a dirt road in the bottom of a predominantly east-west oriented canyon. This canyon is surrounded by mixed riparian and canyon wash habitats with pifion (Pinus monophylla) and ju- niper (Juniperus osteosperma) on the surrounding hill- sides. From 20 May to 12 June 1989 and from 1 to 12 June 1990, 387 and 194 individual butterflies, respec- tively, were marked along this road. Upon initial cap- ture, each butterfly was given a unique number and placed into one of five distinct categories based on wing coloration: red (R), red-intermediate (RI), inter- mediate (I), black-intermediate (BI), and black (B); voucher specimens from here and other sites are at the Nevada State Museum and Historical Society, Las Ve- gas. Categories R and B are here termed as “pure”, RI, I, and BI as “intermediate”, and I alone as “true inter- mediate.” All butterflies were released in the center of the area in which they had been captured. Similar de- terminations of phenotype were made on museum specimens collected from several sites in northeastern Nevada. Within field samples of butterflies, and accounting for developmental differences, males usually outnum- ber females, often by a broad margin (e.g., Ehrlich et al. 1984) due largely to behavioral differences between the sexes (e.g., Gall 1985). In general, with large sam- ples taken throughout the flight season, it may be as- JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY sumed that an apparent sex ratio within a taxon should be correlative across populations using comparable collection techniques (Boyd et al. 1999). Accordingly, the sex ratios of museum specimens and individuals captured as part of the MRR studies were tabulated. The MRR study in 1989 and 1990 also produced in- formation on relative spatial and temporal distribution of red and black forms. In 1989, the site was divided into 36 areas of similar size. Thirty sites straddled the road and the remaining six were located adjacent to the road in the broadest area of the canyon. In 1990, only the 10 subsites that had the highest butterfly den- sities in 1989 were again sampled. For the purposes of this spatial and temporal delineation, R and RI indi- viduals were considered “red”, B and BI were consid- ered “black”, and the few true intermediates were omitted. Probable differences in emergence timing between the two forms are indicated by differences in mean age of individuals. At each handling, all individuals were scored for wing wear, a common estimator of butterfly age (Orive & Baughman 1989). Butterflies were scored in 0.5 intervals from 0.5 (newly emerged) to 3.5 (worn, indicating extended flight). To test whether individuals of different color forms at least attempt to interbreed, the genitalia of each male captured during the MRR studies was dipped in a powdered fluorescent dye upon initial capture and on each subsequent recapture. Some of this dye is typ- ically transferred to a female during a subsequent mat- ing, and mated females were examined under ultravio- let light for evidence of dye (Wheye & Ehrlich 1985, Fleishman et al. 1993). Use of this technique to inves- tigate matings between members of different experi- mental classes assumes that dyed and undyed males are equally likely to achieve copulations, dyes of different colors are equally likely to be transferred during mating and retained by females after mating, and dye transfer occurs at consistent frequencies for successful and un- successful matings. The protocol does not assume that all matings are equally viable or are equivalent in an evolutionary sense. All R and RI males were dipped in pink dye, while B and BI males were dyed green. Some mated females were retained during the MRR study to determine oviposition preference using techniques developed by Singer and co-workers (e.g., Singer 1986). Each female was sequentially offered each of three locally available potential oviposition hostplant species, Castilleja angustifolia (Nutt.) G. Don, Penstemon speciosus Dougl. ex Lindl. (both Scrophulariaceae), and Symphoricarpos oreophilus Gray (Caprifoliaceae) at five-minute intervals or at in- tervals permitted by weather conditions. A plant was VOLUME 57, NUMBER 3 recorded as accepted if the female’s abdomen was fully curled and the ovipositor extruded for at least three seconds. Actual oviposition was not permitted. A fe- male was considered to have preferred plant species A over plant species B if a rejection of B was recorded af- ter an acceptance of A (Singer 1982). If females were captured before 11:00, their preference-testing com- menced on the day of capture. If they were captured after 11:00, testing commenced on the following day. In every case, plant species that were accepted on the first day of testing were recorded. In addition, ex- amples of these plants were searched on two dates at the Pequop Mountains study area to determine the presence of egg masses. Statistical significance (considered at p < 0.05 through- out) was determined using chi-square comparisons. RESULTS Distribution of Euphydryas in the northern Great Basin. Surveys in northern Nevada and adja- cent areas revealed a fairly clear distribution of popu- lations of the E. chalcedona group (Fig. 1). In the southern portion of the study area, the phenotypically red Euphydryas anicia wheeleri (Hy. Edwards) is widespread and often common. Across much of Nevada, this subspecies is associated principally with Castilleja angustifolia, but also with Castilleja linariae- folia Gray, Pedicularis centranthera Gray, and Penste- mon speciosus (all Scrophulariaceae) (Murphy & Ehrlich 1983, GTA unpublished data). The presently known northern distributional limit of E. anicia wheeleri in Nevada occurs in the Toana Range, Windemere Hills, Snake Mountains, and Independ- ence Mountains (all Elko County) westward generally south and east of the Humboldt River (Fig. 1). In the northeastern portion of the study area, the phenotypically black Euphydryas colon nevadensis Bauer predominates and is relatively widespread across sagebrush-dominated (Artemisia) slopes and along riparian corridors (Fig. 1). Its known larval host- plants are Syinnataovteera09 oreophilus and possibly Penstemon (Bauer in Howe 1975, GTA unpublished data). The southern distributional limit of E. colon nevadensis in eastern Nevada is in the Pequop Moun- tains. In the narrow geographic band of overlap between E. anicia wheeleri and E. colon nevadensis, three sites were found where black and red forms and intermedi- ates fly together (northern end of the Pequop Moun- tains, Windemere Hills in the Thurston Spring area, Tabor Creek in the Snake Mountains) and an addi- tional three sites with one form and intermediates (Wildhorse Crossing Campground in Owyhee River 179 TABLE 2. Wing phenotype of Ewphydryas chalcedona complex individuals marked during the mark-recapture-release study in the Pequop Mountains (wing phenotype as in Table 1). 1989 1990 Category Males Females Total Males Females Total R 61 27 88 29 9 38 RI 48 3 51 23 4 7 I 2 0 2 8 0 § BI 64 5 69 35 3 38 B 165 12 17 75 8 §3 Total 340 47 387 170 24 194 Valley, Jack Creek Campground and west of Maggie Summit in the Independence Mountains) (Fig. 1). The site in the Pequop Mountains supported large numbers of both red and black phenotypes during 1989 and 1990. Not surprisingly, the few locations of sympatry between red and black forms are topograph- ically complex; these sites are canyons where sharply defined warm and cool slope exposures supporting dis- tinctive plant communities are just meters apart. In northwestern Nevada (and also adjacent north- eastern California and southern Oregon), populations of E. colon are all far north of the Humboldt River (Fig. 1). These largely black Euphydryas colon walla- censis Gunder are sympatric in some locales on the Sheldon National Wildlife Refuge (Humboldt County), but apparently do not hybridize, with the also largely black Euphydryas anicia veazieae Fender & Jewitt. The two phenotypes have partially overlapping flight seasons, but the details of their microsympatry require definition. In this region, E. colon is associated with Symphoricarpos while E. anicia apparently uses both Penstemon and Castilleja (Bauer in Howe 1975, Austin & Murphy 1998b). Just south and east of this area E. anicia veazieae intergrades broadly with the redder Euphydryas anicia macyi Fender & Jewett. Outside these two regions, only one form of the E. chalcedona complex is present at any given location in the Great Basin of Nevada, although in some areas their distributions approach (Fig. 1). The affinity of each population is unequivocal and individuals are readily identifiable by superficial characters that are consilient with genital morphology (see below). Wing phenotypes. As noted, the broad scale sur- veys located sites in northeastern and northwestern Nevada where two E. chalcedona complex taxa were sympatric: the Pequop Mountains, the Windemere Hills, the Snake Mountains, and the Sheldon National Wildlife Refuge. The black form and intermediates were found in the Owyhee River Valley and at two sites in the Independence Mountains (Table 3, Fig ig. 1). 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Description. Male: Head: Vertex white, frons bulbous, ventral lip not produced, white. Labial palp white. Antenna filiform, scape and dorsal scales white. Thorax: Patagium, tegula, meso- and metathorax white. Venter white. Foreleg with femur darker medially, cream to gray, lighter laterally, cream to white; tibia longer than basitarsus, white, inner side with one large and 3 progressively smaller spines, outer side with 2 large and 2 smaller spines; tarsi white. Middle and hind legs white, some specimens can have pink on tibia and tarsi. Forewing: Male length 12-14.5 mm (N = 10). Basal patch pink to pink with a slight purplish cast; median area white with a broad light brown band; alienate band pink to pink with a slight purplish ne terminal area white tending to cream to light brown at margin; fringe cream to light brown. Hindwing: Ground color white; marginal band intensity ari color variable, absent to moderately developed and pink to a mixture of pink and light gray. Abdomen: Cream with small brown spots laterally. Canitrllte (Figs. 9-10): Uncus short (0.30 x valve length), robust. Vi alve elongate (length 8.3 x width), costal margin gen- tly curved; ampulla “atthe (0.015 x valve length); corona present; sac- culus well developed and greatly produced. Juxta quadrate, proximal margin slightly produced, heavily sclerotized, forming a distinct bar along margin. Remoereus slightly curved, dorsal patch of dense minute spicules; vesica with 2 and one- -half coils and minute spicules. Female. As in male except forewing length 13-16 mm (N = 12). Genitalia (Fig. 13): Papillae anales triangulate, apex pointed; dorsal margin concave. Eighth segment with fine spicules. Distal margins of seventh segment with a double row of elongate setae, distal row longer and more robust than proximal row. Ostium bursa lightly sclerotized, minute spicules present. Ductus bursa narrow, approxi- matly 0.2 x length. Appendix bursa coiled. Corpus bursa ovate; signa composed of two scobinate bars. Types. Holotype: ¢, in USNM, with the following labels: (1) Candian, Hemphill Co., Texas; 15 VIII 71; A. & M. Blanchard; (2) USNM ENT 00142656 [bar code label]; (3) Holotype ¢, Schinia regina Pogue and Harp. Paratypes. 47 4, 53 9, all in USNM unless atattedl COLORADO: No speciiic locality (1d 12), Bruce [collector] (USNM ENT 143145-6); (1 2), Oslar (USNM ENT 143147). Adams Co., E of Bennett, 39.74°N, 104.41°W, 21 Aug. 1999 (1 d) CLE. Harp (CEH). Arapahoe Co., S of Manilla, 39.74°N, 104.52°W, 21 Aug. 1999 (1 3), C.E. Harp (CEH). Baca Co., Picture Canyon, pic- nic area, Comanche National Grassland, sw of Campo, UV trap, 37°00.66’N, 102°44.64’W, 25 Aug. 2002 (3 36 1 2), M.G. Pogue & C.E. Harp, (USNM ENT 14410-3); Picture Canyon, n. of picnic area, Comanche National Grassland, sw of C ZB, at mv light, 37°01.4U/N, 102°44.65’W, 25 Aug. 2002 (7312), MG. Pogue & C.E. Harp, (USNM ENT 144459-66); Picture Canyon, Comanche National Grassland, sw of Campo, at mv light, 37°00.72’N, 102°44.60’W, 29 Aug. 2002 (2G) au Gabe Harp (CEH); Picture Canyon, Comanche National Grassland, sw of Campo, at UV light, 37°00.66’N, 102°44.60’W, 29 Aug. 2002 (1 3), CE. Harp (CEH); Springfield, s. End of town, along Hwy #385/287 at truckstop lights, 37°23.10’N, 102°36.92’W, 28 Aug. 2002 (3 4 12). (C18, Harp (CEH). Cheyenne Co., 2 mi e. of Aroya, Hwy. #94 at rd. T and rd. O, 38°50.98’N, 103°09.79’W, 28 Aug. 2002 (36 2), Cah, Harp (CEH). Fremont Co., Penrose, 38.42°N, 105.02°W, 17 Aug. 2001 (2 2) 24 Aug. 2001 (1 6 1 2), C.E. Harp (CEH). Jefferson Co., Morrison, June (2 d), Park, (USNM ENT 143165-6). Lincoln Co., Limon, 39.27°N, 103.71°W, 19 Aug. 1998 (1 9), C.E. Harp (CEH). Morgan Co., Wiggins, 40.23°N, 104.07°W, 9 Aug. 2000 (1 ¢ 2.9), C.E. Harp (CEH). Otero Co., Vogel Canyon Picnic Area, 15 mi S of La Junta, 4340 ft., 37°46’13”N, 103°30’46”W, 18 Aug. 1997 (1 3), D,J. Wright (DJW); Comanche NG, 15 mi S La Junta, 27 Aug. 2000 (1 2), D.J. Wright (DJW). Prowers Co., Holly, 38.05°N, 102.12°W, 25 Aug. 2000 (3 d 4 2), C.E. Harp (CEH). Pueblo Co., Pueblo West, 38.32°N, 104.74°W, 24 Aug. 2001 (262°) CE. Harp (CEH). Weld Co., Roggen, 40. 17°N, 104.37°W, 9 Aug. 1999 (1 3), 29 Aug. 1999 (1 3), C.E. Harp (CEH); E of Roggen, 40. 22°N, 104. 21°W. 22, Aug. 2001 (1 d), C.E. Harp (CEH); Keenesburg, 40.11°N, 104.52°W, 9 Aug. 2000 (15192), 16 Aug. 2000 (1 2), C.E. Harp (CEH). KANSAS: No specific locality (1 ¢), (USNM ENT 143148). Ellis Co., Hays, 6 Sep. 1935 (1 2), H.K. Walkden, (USNM ENT 143151). Finney Co., Garden City, 30 Aug. 1935 (1 9), H.K. Walkden, (USNM ENT 143150). Morton Co., Spee NG, 7.5 mi N of Elkhart, 25 Aug. 2000 (2 2), 26 Aug. 2000 (2 2), D.J. Wright (DJW). Riley Co., Man- hattan, 1 Sep. 1937 (1 A H. K. Walkden, (USNM ENT 143149). NEBRASKA: Dawes Co., Chadron, 42.83°N, 103.02°W, 26 July 1976 (1 3), H.R. Lawson (CSC). Scotts Bluff Co., Scottsbluff, 5 Aug. (1 3), 3 genitalia slide USNM 46786, 12 Aug. (1 9), 13 Aug. (2 9), Whelan, 14 Aug. (1 2), (USNM ENT 143152-6). NEW MEXICO: Eddy Co., Campsite, 31°21.4’N, 103°46.9’°W, 13 June 1979 (1 2), 19 June 1979 (1 2), D.R. Delorme & H. L. Carrola, (USNM ENT 143416-7) (TAMU); White[s] City, 18 Sep. 1963 (4 2), 22, Sep. 1962 (1 3), A. & E. Blanchard, (USNM ENT 143158, 143161-4). Luna Co., Deming, 1-7 Sep. (1 3 2 2), 2 genitalia slide USNM 46787, (USNM ENT 143157-60). OKLAHOMA: Cimarron Co., nw. of Black Mesa State Park, roadside along Gallinas Canyon, 36°57.72’N, 102°48.52’W, 29 Aug. 2002 (42), C.E. Harp (CEH). TEXAS: Brew- ster Co., 15-30 Aug. 1926 (1 3), O.C. Poling, (USNM ENT 142637). Cottle Co., Paducah, 19 Aug. 1971 (3 3), A. & E. Blanchard, (USNM ENT 142653-5). Hemphill Co., Canadian, 15 Aug. 1971 (1 1 2), A. & E. Blanchard, (USNM ENT 142656-8). La Salle Co., Artesia Wells, 28 Sep. 1971(1 ¢ 3 2), d genitalia slide USNM 46850, (USNM ENT 142646-9); Chaparral Wildlife Management Area, JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 29-30 Sep. 1959 (1 9), J. Schaffner, (USNM ENT 143413) (TAMU). Reeves Co., Pecos, 18 Sep. 1952 (5 2), R. Leuschner, (USNM ENT 142638-42). Ward Co., Monahans Sandhill State Park, Monahans, 20 Sep. 1999 (1 4), J.B. Lombardini, (USNM ENT 143415) (TAMU). me Co., ‘ered, 25 Aug. 1926 (1 6), (USNM ENT 142615). Additional material examined. COLORADO: Morgan Co., SSW of Ft. Morgan, 40.23°N, 103.80°W, 16 Aug. 1990, M.D. Bowers (JKA). Weld Cor No specific locality, 40.43°N, 104.72°W, P.A. Opler (CSU): KANSAS: Ellis Co., No specific locality, 38.88°N, 99. 33°W, 6 Sep. 1935, H.H. Walkden (SMEK). Finney Co., No specific locality, 37.95°N, 100.90°W, 4 Sep. 1935, H.H. Walkden (SMEK). Franklin Co., No specific locality, 38.58°N, 95.27°W, (FHSU). Morton Co., No specific locality, 37.02°N, 101.92°W, (FHSU). Seward Co., No specific locality, 37.05°N, 100.93°W, (FHSU). Sherman Co., E of Kanorado, 39.32°N, 102.05°W, 1 Sep. 1995, James K. Adams (JKA). Stanton Co., No specific locality, 37.53°N, 101.88°W, (FHSU). NEBRASKA: Deuel Co., No specific locality, 41.10°N, 102.48°W, R. Leuschner (RL). Lancaster Co., No specific locality, 40.82°N, 96.68°W, (UNL). Lincoln Co., No specific locality, 41.15°N, 100.75°W, (UNL). Scotts Bluff Co., Scotts Bluff, 41.87°N, 103.67°W, 30 Sep. 1935, H.H. Walk- den (FHSU). NEW MEXICO: Dona Ana Co., Las Cruces, 32.30°N, 106.78°W, 12 Sep. 1994, J.K. Adams (JKA). Socorro Co., No specific locality, 34.07°N, 106.92°W, (LACM). OKLAHOMA: Ellis Co., No specific locality, 36.27°N, 99.92°W, (ORU). TEXAS: Bailey Co., No specific locality, 34.23°N, 102.73°W, (AMNH). Briscoe Co., Caprock Canyon State Park, 34.47°N, 101.30°W, 29 Sep. 1994, E. Knudson (ECK). Culberson Co., No specific locality, 31.05°N, 104.85°W, (ECK). El Paso Co., Fabens, 31.43°N, 106.13°W, 7 Sep. 1997 (JKA). La Salle Co., No specific locality, 28.45°N, 99.25°W, (AMNH). Reeves Co., No specific locality, 31.42°N, 103.50°W, (RL). Sutton Co., No specific locality, 30.57°N, 100.65°W. Uvalde Co., Concan, 29.48°N, 99.44°W, (JKA). Larval foodplant. Palafoxia sphacelata (Nutt. ex Torr.) Cory (Asteraceae). Flight period. The main flight is from mid August to the end of September with a few specimens from mid June. Distribution (Fig. 14). From southern and western Texas north to the panhandle, northwestern Okla- homa, Kansas, and Nebraska and west to southern New Mexico and eastern Colorado. Discussion. Schinia regina is known to have a single brood, flying from early August through Sep- tember in the northern parts of its range and during September in the south. There is a record of a June specimen, indicating a partial second brood in some localities. The peak flight of S. regina is the end of Au- gust with a peak of early October for S. regia. Both species overlap in flight period during September. Although adults are readily taken at lights in proximity to their host plants, they may be seen resting across the tops of the flowerheads of Palafoxia during the early to mid-morning hours. Eggs are laid in pre-bloom flowers. Early instar larvae feed within the long, narrow flower- head. Their presence can be seen as the larvae continue to feed on the early seed parts. This feeding causes the maturing flowers to pull apart basally and start to squeeze up through the top of the flower. This unique appearance in still young flowerheads is indicative of the VOLUME 57, NUMBER 3 internal feeding larva. Only the latter instar larvae feed on the outside of the flowerheads at night and early morning, holding on to the stem just below the calyx and feeding outside through the of the bottom of the flower into the maturing seeds and flower parts. Forewing coloration can vary from pink to purplish pink, which is about the same color as in regia. Etymology. The species epithet is latin and refers to a queen. This reflects a relationship with the specific epithet of Schinia regia meaning royal. ACKNOWLEDGMENTS We thank Paul Z. Goldstein, Field Museum of Natural History, Chicago, Illinois for providing label data for the holotype of Schinia regia. For lending specimens for this study we thank Edward G. Ri- ley, Department of Entomology, Texas A & M Univ ersity, College Station, Texas and J. Donald Datontames Canadian National Collec- tion, Ottawa, Ontario, Canada. We thank Clifford D. Ferris, Laramie, Wyoming and Natalia J. Vandenberg and David R. Smith, Systematic Entomology Laboratory, Agricultural Research Service, U.S. Department of Agriculture, Washington, DC for helpful sug- gestions that greatly improved this paper. LITERATURE CITED Barnes, W. & J. McDuNNoucu. 1917. Check list of the Lepidoptera of boreal America. Herald Press, Decatur, Illinois. ix, 392 pp. Dyar, H. G. 1903 [1902]. A list of North American Lepidoptera and key to the literature of this Order of insects. Bulletin of the United States National Museum 52:i—xi, 1-723. . 1921. New American Noctuidae and notes. Insecutor In- scitiae Menstruus 9:40-45. 203 FRANCLEMONT, J. G. & E. L. Topp. 1983. Noctuidae, pp. 120-159. In Hodges, R. W. et al. (eds.), Check list of the Lepidoptera of America north of Mexico. University Press, Cambridge. xxiv, 284 pp. HArpwIcK, D. F. 1996. A monograph to the North American He- liothentinae (Lepidoptera: Noctuidae), David F. Hardwick, Ot- tawa, Ontario. 281 pp. McDunnoucu, J. 1938. Checklist of the Lepidoptera of Canada and the United States of America. Part 1. Macrolepidoptera. Memoirs of the Southern California Academy of Sciences. 275 pp POOLE, R. W. 1989. Fascicle 118, Noctuidae. Lepidopterorum Cat- alogus (New Series), E.J. Brill and Flora and Fauna Publica- tions, Leiden. xii, 1313 pp. POOLE, R. W. & P. GENTILI (EDS.). 1996. Nomina Insecta Nearc- tica. A check list of the insects of North America. Vol. 3. Diptera, Lepidoptera, Siphonaptera. Entomological Informa- tion Services, Rockville, Maryland. 1143 pp. SmiTH, J. B. 1891. List of Lepidoptera of boreal America. P.C. Stockhausen, Philadelphia. v, 124 pp. . 1893. A catalogue, bibliographical and synonymical, of the species of moths of the Lepidopterous superfamily Noctuidae, found in boreal America. Bull U.S. Nat. Mus. 44:1-424. . 1906. New Noctuidae for 1906. J. N.Y. Entomol. Soc. 14:9-30. STRECKER, F. H. H. 1876. Lepidoptera, Rhopaloceres and Hetero- ceres indigenous and exotic; with descriptions and colored il- lustrations. No. 13. Owen’s Steam Book and Job Printing Office, Reading, Pennsylvania, pp. 109-123. Topp, E. L. 1982. The noctuid type material of John B. Smith (Lepidoptera). U.S. Department of Agriculture, Technical Bul- letin 1645. 228 pp. Received for publication 9 September 2002; revised and accepted 3 April 2003. Journal of the Lepidopterists’ Society 57(3), 2003, 204-219 THE HISTORY AND TRUE IDENTITY OF MELITAEA ISMERIA (NYMPHALIDAE): A REMARKABLE TALE OF DUPLICATION, MISINTERPRETATION, AND PRESUMPTION JOHN V. CaLHOUN’ 977 Wicks Drive, Palm Harbor, Florida 34684-4656, USA ABSTRACT. John Abbot (1751—ca. 1840) supplied the watercolor drawing for the original description and accompanying engraved plate of Melitaea ismeria Boisduval and Le Conte. The plate was poorly executed, resulting in 170 years of debate regarding the identity of the fig- ured species. Most authors treated M. ismeria as synonymous with Chlosyne gorgone (Hiibner). More recently, a neotype of M. ismeria was des- ignated to reflect synonymy with Chlosyne nycteis (Doubleday), resulting in a proposed priority replacement of nycteis. During a study to eval- uate these findings, the original drawing of M. ismeria was discovered. John Abbot copied this drawing from an earlier painting of C. gorgone. Two other duplicates of this C. gorgone illustration were also located. The figured early stages and hostplant are consistent with C. gorgone. The proposed priority replacement of nycteis is therefore unwarranted. Also included are details about the drawings used by Boisduval and Le Conte and the discovery of a specimen of C. gorgone attributed to John Abbot. Additional key words: Chlosyne, Georgia, John Abbot, larva, pupa, South Carolina. Klots (1951) considered Melitaea ismeria Boisduval & Le Conte, [1833] to be “one of our greatest prob- lems.” Miller & Brown (1981) called it “a nomenclat- ural headache.” Due to a poorly engraved illustration that accompanied the original description, M. ismeria has remained enigmatic for 170 years. Since 1840, most authors have treated M. ismeria as synonymous with the insect now recognized as Chlosyne gorgone (Hiibner, [1810]), but enough uncertainty remained as to permit alternative interpretations. Attempts to re- solve this dilemma were as intriguing as the taxon itself and included a great deal of misleading information. The most recent was Gatrelle (1998) who considered M. ismeria to be synonymous with Chlosyne nycteis (Doubleday, [1847]). He designated a specimen of C. nycteis as the neotype of M. ismeria, resulting in the priority replacement of nycteis. I now submit new evi- dence that correctly defines the intended species and contradicts the findings of Gatrelle (1998). These re- sults finally bring resolution to this troublesome and persistent mystery. MATERIALS AND METHODS Historical literature pertaining to M. ismeria was ex- amined in detail and the conclusions of Brown (1974) and Gatrelle (1998) were evaluated. The publication history of Boisduval & Le Conte ([1833]) was investi- gated through the works of Oberthiir (1920), dos Pas- sos (1962), and Cowan (1969). Photocopies, microfilm, digital scans, and digital photographs of specimens, published figures, original illustrations, manuscript notes and other relevant data were obtained for analy- sis from many sources, including the Alexander Turn- bull Library (Wellington, New Zealand), Allyn Mu- seum of Entomology (Florida Museum of Natural History, Sarasota), Florida State Collection of Arthro- pods (FSCA, Division of Plant Industry, Florida Dept. ‘Research Associate, Florida State Collection of Arthropods, DPI, FDACS, Gainesville, Florida 32614, USA. of Agriculture and Consumer Services, Gainesville), Houghton Library (Harvard University), Macleay Mu- seum (University of Sydney, New South Wales, Aus- tralia ), The Natural History Museum, London (BMNH), Thomas Cooper Library (University of South Carolina), and Wittenberg University Library (Spring- field, Ohio). Specimens and photographs of early stages were obtained from several sources. Comparative stud- ies were conducted using original Abbot illustrations, species of Chlosyne, as well as engraved figures in Bois- duval & Le Conte ({1833]) and Smith & Abbot (1797) (authorship of this publication follows Wilkinson (1981)). Detailed biographies of John Abbot by Rogers- Price (1983) and Gilbert (1998) were consulted to more fully understand Abbot's life and artwork. RESULTS Original description. Melitaea ismeria was de- scribed and figured in Histoire Générale et Iconogra- phie des Lépidoptéres et des Chenilles de VAmérique Septentrionale by renowned French entomologist Jean Baptiste Alphonse Boisduval (or Déchauffour de Bois- duval) (1799-1879) and wealthy American naturalist John Eatton Le Conte, Jr. (1784-1860), whose sur- name is still in contention. His name was given as “Leconte” in this and other publications. Rehn (1954) believed that the family preferred “Leconte,” but Cowan (1969) considered this to represent the histori- cal version used by earlier Huguenot family members before they fled France to escape religious persecu- tion. I have employed the version used by J. E. Le Conte himself, who plainly signed his name as “Le Conte” (see Scudder 1889, vol. 2, frontispiece). His fa- mous nephew, Joseph (who referred to J. E. Le Conte as “Uncle Jack’), also signed his name as “Le Conte” and used only this version in his detailed autobiogra- phy (Armes 1903). Many documents from the Le Conte family are deposited in the library of the Ameri- can Philosophical Society, Philadelphia. Robert S. Cox, VOLUME 57, NUMBER 3 Curator of Manuscripts, confirmed (pers. com.) that “Le Conte” is the correct version for this family. The compressed variation, “LeConte,” is also frequently used (e.g., Bigley 2001). The cover page of most editions of Histoire Générale was dated 1833, but the plates and accompa- nying text were issued in 26 livraisons (fascicles) from 1829 to 1837 (dos Passos 1962, Cowan 1969). The publication included 78 hand-colored engraved plates, three issued per livraison. Melitaea ismeria was de- scribed on pages 168-169 and figured on Plate 46, which included depictions of a dorsal female, ventral female, mature larva, and pupa (Figs. 1, 7, 9, 12, 16). Boisduval probably based his species name on Ismeria, a beloved Sudanese woman who married the son of William II of France in the 13th century. The brief Latin description reads “Alis swbhdenticulatis, supra ni- gro fulvoque variis, anticis apice albo puntatis; posicis subtus fasciis albis fulvisque, serieque punctorum ni- grorum” (wings finely toothed, above variably colored with orange and black, white spots near the apex; hind- wings below with light orange bands and a series of black spots). Longer French descriptions of the adult, larva and pupa were also included. It was stated in French that, “This Melitaea is found in Carolina and Georgia. It is very rare in collections.” A printed notation at the bottom of Plate 46 of M. ismeria reads “Abbot pinx.” Most of the plates in Bois- duval & Le Conte ([1833]) were reproduced from original drawings by John Abbot (1751- ca.1840), an English artist and naturalist who resided in Georgia from 1776 until his death. Notations on other plates refer to French naturalist Emile (or Charles Emile) Blanchard, J. E. Le Conte, and artist/engraver Paul C. R. C. Duménil (misidentified by Gilbert (1998) as French naturalist André M. C. Duméril). dos Passos (1962) credited 62 plates to Abbot, while Gilbert (1998) listed 65. However, some of the plates attrib- uted to Abbot in Boisduval & Le Conte ([1833]) were undoubtedly not derived from his work. These include West Indian Eurytides celadon (Lucas) (as Papilio sinon, Plate 3), Battus devilliers (Godart) (as P. villier- sti, Plate 14), and Battus polydamus (L.) (as Papilio polydamus, Plate 15), as well as Asian Leptosia nina (Fabricius) (as Pieris chlorographa, Plate 17). The text confirms Abbot's unlikely involvement in these plates; the American distribution of the papilionids was given as “la Floride” and the inclusion of L. nina was based on two specimens of dubious North American origin. These illustrations are more consistent with the work of Duménil, who probably drew the originals from specimens in Boisduval’s extensive collection. Despite the collaboration of so many artists and en- gravers, many of the published plates were poorly exe- 205 cuted. In the preface to livraison ten, Boisduval an- nounced that, “certain of our subscribers have com- plained that, although our figures are accurately col- ored, they are not well drawn; most of the bodies are defective, with the wings and legs badly attached and the veins faulty. I am the first to recognize that one has the right to expect an amount of perfection, as this is acceptable, but these drawings were not totally cre- ated in France, but in North America by Mr. Abbot through my collaborator Mr. Leconte of New York, who has paid for his faithful drawing and coloring of wings, bodies and legs. I have attempted to change nothing among the original figures, but in the future, in order to avoid problems, and along with the pub- lisher who will not sacrifice perfection of the publica- tion, I will have the plates retouched to conform to the nature of Mr. Abbot's drawings and repair any inaccu- racies when present. Subscribers may be assured that from delivery 10 our figures will no longer show these faults” (translation from French). Plate 46 of M. isme- ria was published in livraison 19 during 1833-34 (dos Passos 1962), thus the engraving may have been al- tered prior to the prints hein issued. Abbot did not supervise the creation and alteration of the plates, thus many of his drawings lost their distinctiveness in the reproduction process (Rogers-Price 1983). Unsatisfac- tory efforts of engravers and colorists were a concern for many artists of the eighteenth and nineteenth cen- turies. This problem was especially acute for Abbot, whose renderings were very meticulous (Wilkinson 1984). The first 30 plates for Boisduval & Le Conte ({1833]) were engraved under the supervision of P. Duménil. Either because Duménil retired, or was dis- gusted by earlier criticisms of his work, another promi- nent engraver named Borromée completed the re- maining 48 plates, including that of M. ismeria (Oberthiir 1920, Cowan 1969). Although Boisduval made efforts to improve the plates, many remained unsatisfactory. To make matters worse, the precision of the plates varied from copy to copy, depending upon the vagaries of the colorists. Among numerous letters written between English lep- sdlopianst Edward Doubleday and American entomol- ogist Thaddeus W. Harris, Doubleday observed in 1840 that the plates were “poorly colored and not ex- act,” while Harris complained that some of the figures were “miserably represented” (Scudder 1869). In 1883, Albert C. L. G. Giinther (Keeper of Zoology, British Museum, 1875-95) remarked, “many of these productions are so unsatisfactory that many of them can only be determined by reference to the originals” (Gilbert 1998). In the years following the description of M. ismeria, lepidopterists failed to document speci- mens that clearly matched the published plate, sug- 206 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 1-21. Melitaea ismeria and Chlosyne gorgone. 1, Plate of M. ismeria in Boisduval & Le Conte ({1833]). 2, John Abbot drawing that in- cludes original figures of M. ismeria (left). 3, Abbot painting of C. gorgone for John Francillon*. 4, Ventral adult C. gorgone by Abbot for Francil- lon*. 5, Ventral adult C. gorgone by Abbot for William Swainson. 6, Original ventral figure of M. ismeria. 7, Ventral M. ismeria in Boisduval & Le Conte ([1833]). 8, Original dorsal figure of M. ismeria. 9, Dorsal M. ismeria in Boisduval. & Le Conte ({1833]). 10, C. gorgone larva by Abbot for Francillon*. 11, Original larva of M. ismeria. 12, Larva of M. ismeria in Boisduval & Le Conte ([1833]). 13, C. gorgone larva, fm. ‘bicolor.’ 14, C. gorgone pupa by Abbot for Francillon*. 15, Original pupa of M. ismeria. 16, Pupa of M. ismeria in Boisduval & Le Conte ([1833]). 17, C. gorgone pupa. 18, Enhanced sketch lines on original ventral figure of M. ismeria. 19, Body of ventral C. gorgone by Abbot for Francillon*. 20, Ventral fig- ure of Dryas gorgone by Jacob Hiibner. 21, Abbot painting of C. gorgone for William Swainson. (* © The Natural History Museum, London) VOLUME 57, NUMBER 3 gesting that it did not represent a valid species, or was poorly engraved and lacked the precision of the origi- nal drawing. A John Abbot painting. In a letter dated 27 May 1840, E. Doubleday told T. W. Harris that he had “cur- sorily examined Abbot's drawings in the British Mu- seum’ and that they included “a vast number of Abbot's manuscripts’ (Scudder 1869). The first illustration Doubleday mentioned was “Melitaea ismeria” and he transcribed the accompanying manuscript notes as “Feeds on crosswort. Frequents the oak woods of Burke County, but is not common. Caterpillar sus- pended itself May 16th, changed to chrysalid May 17th. Butterfly appeared May 26th.” Doubleday asked Harris, “Do you know this species? The name I think is Boisduval’s. The drawing has no name to it.” Dou- bleday obviously noted a resemblance between this painting and the published plate of M. ismeria. Nearly 30 years later, American entomologist Samuel H. Scudder read Doubleday’s 1840 letter while preparing to publish the correspondences of T. W. Har- ris (Scudder 1869). He received additional information on the Abbot illustrations in the British Museum from John. E. Gray (Keeper of Zoology, 1840-75; letter dated 1 October 1869, Houghton Library, Harvard University). In 1871, Scudder visited the British Mu- seum and personally examined these paintings (Scud- der 1872a). He sketched copies of at least 22 of the fig- ured larvae and pupae, which he later published (Scudder 1889). Scudder found the painting men- tioned by Doubleday and identified the depicted species as “Ismeria (carlota Reek. [sic.]).” He summa- rized Abbot's associated notes as “Feeds on cross wort (Helianthus trachelifolius?) and sunflower; frequents oak woods of Bruke [sic.] Co., but is not common; tied up May 15; chrysalis May 17, from which imago May 26.” Like Doubleday, Scudder perceived a similarity between this painting and the published plate of M. is- meria. He also believed that it represented the same species as Eresia carlota Reakirt, now generally con- sidered a subspecies of C. gorgone. Because ismeria had been described 33 years prior to carlota, Scudder (1872b, 1875, 1889) gave priority to ismeria. Strecker (1878) credited Scudder with resolving the identity of M. ismeria and wrote, “There has been some uncer- tainty as to what Bdl.-Lec.’s figures really represent. There can no longer be any doubt that they were in- tended to illustrate this species [C. gorgone].” Henry Edwards (1889) remained unconvinced about ismeria, stating, “there is still some doubt as the this species.” Nonetheless, most subsequent authors followed Scud- der’s arrangement and associated ismeria with the in- sect now recognized as C. gorgone. Holland (1898), Seitz ([1907]—1924), and Clark & Clark (1951) even identified their published figures of C. gorgone as Phy- ciodes ismeria and Melitaea ismeria. In 1950, Norman D. Riley (Keeper of Entomology, BMNH, 1933-55) contacted Georgia naturalist Lucian Harris, Jr. regarding the John Abbot painting in Lon- don. Harris (1972) attributed it to C. gorgone and noted, “Abbot's drawing is labeled Melitaca ismeria.” Abbot's associated notes were transcribed as “It fre- quents the oak woods of Burke County but is not com- mon. Caterpillar feeds on crosswort and sunflower. It tied itself up by the tail 16 May, changed into chrysalis 17, bred 26th.” This offers a slightly different version from that of Doubleday (Scudder 1869) and Scudder (1872a). Based on specimens collected by Harris, Klots (1951) mentioned that, “a few gorgone females taken recently in Georgia lean ward ismeria. Harris (1972) later figured three such C. gorgone specimens as “transition near ismeria.” After decades of research on the butterflies of Georgia, Harris agreed with the synonymy of dos Passos (1969) and concluded, “ismeria was named for a variant specimen of C. gorgone.” Neck (1975) referred to Harris’ figured specimens and also suggested, “a likely solution to the nomenclatural prob- lem is that ismeria is an extreme form of gorgone. The late F. Martin Brown did not readily accept the synonymy of M. ismeria and C. gorgone (dos Passos 1969). This conviction led Brown to more deeply ex- plore the subject and he remains the only author to ex- amine in detail the nomenclatural history of the names ismeria, gorgone and carlota (Brown 1974). To evalu- ate the synonymy of ismeria and gorgone, Brown asked entomologists at the British Museum (Natural History) to examine the Abbot artwork deposited there, including the painting mentioned by Scudder (1872a). Brown recorded Abbot's notes for this paint- ing as, “The caterpillar feeds on the Crop Wort, and Sun Flower. It tyed itself up by the tail, 16th May, changed into Chrysalis 17th, Bred 26th. It frequents the Oak Woods of Burke County, but is not common.” Once more, this offers a slightly different version than those given previously. Brown concluded that this painting did not serve as the model for the published plate of M. ismeria. He also unsuccessfully compared the published larva and pupa of M. ismeria with de- scriptions of immature C. gorgone, C. nycteis, and Chlosyne harrisii (Scudder). As a result, he still could not comfortably assign the published figures to any known species of Chlosyne. He proposed that ismeria be considered “nomen incognitum” (=nomen dubium), as had Higgins (1960). Un onturnaiall Brown did not reproduce or discuss details of the Abbot painting in London. 22 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 22-25. John Abbot specimen and illustrations of C. gorgone. 22, Dorsal male painting for Francillon*. 23, Dorsal (left) and ventral of male specimen in The Natural History Museum, identified by E. Doubleday as M. ismeria. 24, Ventral painting for Francillon*. 25, Labels for specimen in The Natural History Museum and (bottom) original Abbot label from beetle in Macleay Museum. (* © The Natural History Museum, London). The John Abbot painting first mentioned by Dou- bleday (Scudder 1869) was transferred in 1883 from the British Museum, Bloomsbury, to the newly com- pleted South Kensington location (Giinther 1912). Originally called the British Museum (Natural His- tory) (or BMNH), this institution is now known as The Natural History Museum, London. The Abbot water- color measures 23 cm X 30 cm and was among those completed between 1790 and ca. 1816 for John Fran- cillon (1744-1816), a London jeweler who collected Abbot's drawings and specimens and acted as his agent, selling duplicates to the naturalists of Europe (Rogers-Price 1983) (he is also famous for having sold the Hope Diamond in 1812). Francillon had divided his Abbot illustrations into 17 bound quarto volumes. This painting is Plate 7 of Folio 34, Volume 16. Vol- ume 16 contains 130 paintings and is dated ca. 1816 (V. Veness pers. com.), which is consistent with Abbot's manuscript reference to Burke County (he departed Burke County in 1806 to reside in Savannah, Chatham County, Georgia). The painting depicts life-sized dor- sal aspects of male and female adults, ventral female, mature larva, pupa, and hostplant (Fig. 3). Inscribed in ink on the previous page are the following notes writ- ten in Abbot's hand (confirmed from digital scan); “Tab. [Plate] 7. Papilio. Cross wort Frittilary [sic.] But- terfly. The Caterpillar feeds on the Cross Wort, and Sun Flower. It tyed itself up by tail 16th May, changed into Chrysalis 17th, Bred 26th. It frequents the Oak Woods of Burke County, but is not common.” Scud- der’s (1872a) date of May 15th was in error, as was the reference by Brown (1974) to “Crop Wort.” Although Harris (1972) stated that the illustration was labeled as M. ismeria, there is no such inscription associated with the painting or notes. Harris (1972) and Brown (1974) also mistakenly believed the notes were written on the painting itself. Oddly, none of the previous authors mentioned Abbot’s common name for the butterfly. The notation “Helianthus trachelifolius” is inscribed faintly in pencil on the notes page, not on the painting as stated by Harris (1972) and Brown (1974). It is not written in Abbot’s hand. Doubleday did not mention this notation in his 1840 letter, but Scudder saw it dur- ing his visit to the British Museum in 1871. Although Brown (1974) could not determine the origin of this entry, Scudder (1872a) postulated that botanical iden- tifications had “in most cases, been inserted . . . by some subsequent student.” Helianthus tracheliifolius (or trachelifolius) Miller (Asteraceae) is now generally considered a junior synonym of Helianthus de- capetalus L. (Asteraceae) (Heiser et al. 1969, Cron- quist 1980, Kartesz 1994, USDA 2003). Charles B. Heiser, authority on the genus Helianthus, and promi- nent Florida botanists Richard P. Wunderlin and Mark A. Garland examined a digital photograph of the paint- ing and agreed the plant actually represents He- lianthus divaricatus L. (Asteraceae), a widespread species in Georgia. Helianthus decapetalus (=tracheli- ifolius) is restricted in Georgia to the mountainous Blue Ridge and Piedmont regions (Duncan & Kartesz 1981, Jones & Coile 1988, USDA 2003). Although I was unable to locate herbarium specimens of H. divar- icatus from Burke County, Georgia, accurate botanical illustrations by Abbot could be considered as valid records (Ewan 1985). The illustrated adult butterflies clearly represent C. gorgone (Figs. 3, 4, 22, 24), which Scudder (1872a) identified as carlota. The associated sunflower, H. divaricatus, is the only known hostplant of C. gorgone within the coastal plain region of eastern Georgia and adjacent portions of South Carolina (Gatrelle 1993, 1998). Various authors (e.g., Harris 1972, Opler and Krizek 1984) have assumed Abbot's “cross wort” hostplant of C. gorgone referred to a species of Lysimachia L. (Primulaceae), but Abbot apparently used this com- mon name for H. divaricatus. In Smith & Abbot (1797), Abbot gave “cross-wort” as the primary host- plant for Phalaena phyllira Drary (=Grammia phyl- VOLUME 57, NUMBER 3 lira) (Arctiidae) and his associated Plate LXIV portrays the same species of sunflower as in his C. gorgone painting. In the text, J. E. Smith (a competent botanist) correctly identified Abbot's figured plant as H. divaricatus. Furthermore, Abbot’s common name for C. gorgone was the “Cross Wort Frittilary,” and he illustrated the species with H. divaricatus. An alternative theory. Gatrelle (1998) did not lo- cate John Abbot's original drawing of M. ismeria, but announced, “enough evidence now exists to resurrect ismeria and define it correctly as that insect long known as C. nycteis.” He collected three male C. nycteis on 20 August 1989 at mud along the Savannah River in Burke County, Georgia, and designated one of these specimens as the neotype of M. ismeria. Because ismeria was described 14 years earlier, he proposed the priority replacement of nycteis. Despite his state- ment that C. nycteis specimens from Burke County “possess all the major phenotypic characters of the original painting of ismeria,” he did not examine Ab- bot'’s original drawing and based his comparisons strictly on the published plate. He considered popula- tions of C. nycteis distributed from eastern Georgia, across northern Florida to southern Louisiana as C. is- meria ismeria and other eastern populations as C. is- meria nycteis. Western North American populations would be referable to C. ismeria drusius (W. H. Ed- wards) and C. ismeria reversa (F. & R. Chermock) (Gatrelle 1998, 2000b). Gatrelle also collected speci- mens of C. gorgone in eastern Georgia and designated the neotype of Dryas reticulata gorgone Hiibner, ap- parently unaware that Hiibner’s intermediate name, “reticulata, is comparable to a subgeneric category and was not intended as part of the name of the insect (Hemming 1937). Both of Gatrelle’s neotypes are de- posited in the Allyn Museum of Entomology, Florida Museum of Natural History, Sarasota, Florida. Gatrelle primarily based his arguments on the con- clusions of Brown (1974) and John Abbot's life history notes, but he committed critical errors with this ap- proach (see Discussion). Doubts about the validity of his neotype designations prompted Gatrelle (2000a) to defend his publication format as compliant with ICZN (1999). Kons (2000) disagreed with Gatrelle’s findings about the identity of M. ismeria and hesitantly sug- gested that C. harrisii was the intended species. It was obvious that additional proof was still necessary to con- firm the identity of M. ismeria. As Brown (1974) sur- mised, John Abbot’s original drawings would “provide the proper measure of accuracy.” Original drawings for Boisduval & Le Conte ([1833]). Oberthiir (1920) and Cowan (1969) summa- rized the early history of an original set of drawings 209 used for the published plates in Boisduval & Le Conte ({1833]). Cowan lost track of them after 1963. Art his- torian Vivian Rogers-Price (1983) relocated these drawings and offered a brief historical overview up to that time. Her treatise was published as an exhibition catalog and was overlooked by lepidopterists. Based on an exhaustive review of historical and contemporary evidence, I now offer a detailed account that connects these original drawings with the published plates of Boisduval and Le Conte. Discovered in this set of wa- tercolors is the original drawing of M. ismeria. In the front of a copy of Boisduval & Le Conte ([1833]), shelved in the Entomology Library, The Nat- ural History Museum, London, is a brief inscription that reads, “The originals of these plates passed into the possession of M. Chas. Oberthiir from the library of Dr. Boisduval. Seen by F. A. Heron, 11 x 1904” (P. Ackery pers. com.). Francis A. Heron served as Assistant-in-Charge of Butterflies for the British Mu- seum (Natural History) from 1901-10 (Harvey et al. 1996). At least 25 years earlier (probably in 1871), S. H. Scudder had visited Boisduval in Paris who showed him drawings by John Abbot that were “contained in a little oblong folio volume, on sheets broader than high (27 x 16.5 cm), instead of on ordinary large folio sheets” (Scudder 1888). Scudder obtained permission from Boisduval to draw at least 23 of the figured but- terfly larvae and pupae. Scudder later published these copies and confirmed that the original figures were “formerly used in Boisduval and Leconte’s Iconogra- phy” (Scudder 1889). Holland (1898) and Klots (1951) also reproduced some of the figures copied by Scud- der. I discovered Scudder’s loosely written notes about these drawings in the Houghton Library, Harvard Uni- versity. Under the heading “Abbot’s Drawings in Bois- duval’s Possession,” Scudder identified the butterfly species depicted in the drawings, listed the illustrated early stages, and indicated the figures he desired to copy. At a later date, Scudder haphazardly inserted J. E. Le Conte’s name into the title of the notes because he suspected that some of the drawings in this set were actually by Le Conte (Scudder 1888). Among the many drawings Scudder identified in this set was “Ismeria.” The John Abbot drawings in this set were commis- sioned in 1813 by J. E. Le Conte, who asked Abbot to illustrate Georgia Lepidoptera, including adults and early stages, but not hostplants (Rogers-Price 1984). Three years earlier, Le Conte’s brother, Louis, had in- herited the family’s immense rice plantation (over 1250 hectares) near Riceboro, Liberty County, Geor- gia. Called “Woodmanston,” this plantation was lo- cated 40 km (25 mi) southwest of Savannah, where John Abbot resided during most of the years from 210 1806 to 1816. A small portion of this plantation re- mains as a botanical garden on the National Register of Historic Places (Armes 1903, Bigley, 2001). J. E. Le Conte resided in New York, but regularly visited his brother at the plantation during the winter months (Scudder 1889, Barnhart 1917). The proximity of Woodmanston to Savannah surely enhanced Le Conte’s relationship with Abbot, who may even have visited the plantation (Bigley 2001). Between the years 1813 and 1834, Abbot completed as many as 3000 il- lustrations for Le Conte (Gilbert 1998). The drawings commissioned in 1813 changed hands at least eleven times, were taken from Georgia to New York, then to France and England aboard ship. 135 years after their journey to Europe, they were returned to New York and ultimately found a home in South Carolina within 215 km (135 mi) of their origin. In 1828, Le Conte took these Abbot drawings (and probably others) to Paris where he met with Boisduval to discuss the book they would eventually coauthor (Sallé 1883, Cowan 1969). After some were duplicated for engravings in Boisduval & Le Conte ([1833]), Bois- duval apparently kept them for many years with the other illustrations he had assembled. Probably around 1850, Boisduval temporarily loaned the entire set to French lepidopterist Achille Guenée for his multi- volume publication on moths (Oberthiir 1920, Cowan 1969). A number of moth species were described and figured by Guenée [1852-58]) based on Abbot draw- ings, but the disposition of these illustrations was un- known (Gall & Hawks 2002). In 1876, three years prior to his death, Boisduval presented his library, os- tensibly including these drawings, to good friend and fellow Parisian lepidopterist Louis M. A. Depuiset (Oberthiir 1$80). Depuiset organized all of Boisduval’s assorted illustrations sometime before his death in 1886 (Oberthiir 1920). Depuiset had also helped maintain Boisduval’s enormous insect collection that was bequeathed in 1876 to lepidopterist Charles M. Oberthiir of Rennes, France (Oberthiir 1880, Clément 1887). Either before or after the death of Depuiset, Oberthiir also acquired the set of original drawings (Oberthiir 1920). In 1928, four years after Oberthiir died, a book dealer named La Chavalier purchased his library (Brown 1974). During the next four decades, the drawings remained in private hands. They resur- faced on 4 November 1963 when Sotheby and Com- pany auction house of London offered them for sale on behalf of “a lady” (Lot 1). They were then mounted in two half-morocco albums (Sotheby & Co. 1963). The Sotheby catalog included a full-page black and white reproduction of Abbot's drawing of Citheronia regalis (Fab.). Rare book firm H. P. Kraus of New York JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY City purchased the set from the Sotheby auction for a meager $1456 U.S. (post-auction edition of Sotheby & Co. 1963). In 1964, H. P. Kraus again offered these drawings for sale, incorrectly describing them in the sales catalog (Kraus [1964]) as the original paintings for Smith & Abbot (1797). This catalog featured a full- page color reproduction of Abbot’s drawing of Nymphalis antiopa (L.). H. P. Kraus had matted and repackaged the drawings in six blue half-morocco portfolio cases with gilt-lettered backs. They were of- fered for sale with a matching boxed copy of Smith & Abbot (1797) at a total price of $12,500 U.S. (Kraus [1964]). Thankfully, the University of South Carolina obtained the drawings from this sale (Ridge 1966) and they are now safely deposited in the Department of Rare Books and Special Collections, Thomas Cooper Library, Columbia. Rogers-Price (1983) and Gilbert (1998) followed Cowan (1969), who claimed this set included 148 drawings, all rendered by Abbot. However, it actually includes 149, and only 105 are consistent with the work of Abbot. These Abbot drawings were prepared in a horizontal format and depict life-sized figures, with early stages placed above the adults. Many have names and other pencil notations written by Abbot, Boisduval, and Le Conte (compared with known writ- ing samples). Boisduval combined these drawings with others for use in Boisduval & Le Conte ({1833]) and perhaps other publications. All the drawings in the current set are rendered in watercolor and graphite, mostly on cream-colored wove paper, and mounted on stiff paper backing. The sheets measure approximately 26 cm X 16.5 cm, which is consistent with Scudder’s (1888) description. The margins appear to have been trimmed, perhaps for their arrangement into volumes. They are numbered in pencil and the numbers match the butterfly drawings listed in Scudder’s notes. Only 34 of the 55 butterfly drawings in this set are by Abbot. Oberthiir (1920) attributed 17 watercolors to Emile Blanchard; nos. 13-15, 17, 20, 23, 25, 32, 34, 40, 48-54. Undoubtedly ignorant of Oberthiir’s assess- ment, an unpublished inventory list of these drawings compiled by H. P. Kraus also credited 17 of them to Blanchard, matching those listed by Oberthiir with two exceptions; no. 13 (as by Abbot) and no. 45 (as by Blanchard). Blanchard’s drawings are quite distinctive, most being signed in ink “E. Blanchard, pit.” They are rendered in a vertical format, do not include early stages, and depict only one side of dorsal adult figures. Until recently, one of these drawings (34) hung in the President’s office at the University of South Carolina. Based on my own evaluation, the Blanchard drawings are 14, 15, 17, 20, 23, 25, 32, 34, 40, 45, 48-54. Num- VOLUME 57, NUMBER 3 ber 37 is by P. Duménil. Numbers 4 and 13 may also be by Duménil. Number 44, depicting only the mature larvae of Megathymus yuccae Boisduval and Le Conte, is formatted similar to larval moth drawings in this set and was probably drawn by J. E. Le Conte. Figures from this set of drawings were copied for 43 of the butterfly plates in Boisduval & Le Conte ([1833]). Many of the drawings include old pencil no- tations that refer to the corresponding published plates (e.g., “Pl. 1”), as well as numbers that were used to identify individual figures. All of Abbot’ illustrations were rearranged for the published plates, but ten of Blanchard’s multi-species drawings were reproduced in their original layouts. Several published plates in Boisduval & Le Conte ([1833]) lack similarly format- ted original drawings in this set, explaining Oberthiir’s (1920) fear that some watercolors had been lost. Plant leaves and stems were inserted by the engravers into several published plates derived from Abbot's draw- ings in this set. 15 published plates included large hostplants and were evidently copied from other sets of Abbot illustrations. The whereabouts of these paint- ings is unknown, but S. H. Scudder obtained three sets of Abbot's “Notes to the Drawings of Insects” from Boisduval during his trip to Paris (Scudder 1888) (in Harvard University). They pertain to 191 paintings of insects with hostplants, including 172 Lepidoptera. The moth drawings at the University of South Car- olina are rendered in several formats and represent the work of at least one other artist in addition to Ab- bot. Seventy-one are consistent with Abbot's butterfly drawings in this set and some include Abbot's hand- written names. Many of the moth drawings, including 13 depicting only larvae, were prepared on smaller pieces of paper that were then pasted onto sheets matching the size of the larger Abbot drawings. One of these (90) includes an inscription by Boisduval that ap- pears to credit the drawing to J. E. Le Conte, suggest- ing that at least some of these smaller drawings are by Le Conte. Oberthiir (1920) noted that Boisduval sepa- rately kept 452 drawings by Le Conte measuring 13.8 cm x 8.8 cm, a size very similar to the small drawings in this set. This further explains Scudder’s (1888) sus- picion that some of the drawings in this set were actu- ally rendered by Le Conte. Similarly formatted draw- ings attributed to Le Conte are deposited in the library of the American Philosophical Society (Rehn 1954). I examined digital scans of two such drawings and the style can be considered comparable to the smaller drawings in South Carolina. Boisduval planned, but never executed, a companion moth volume to Histoire Générale (Cowan 1969) and the plates for this install- ment would surely have been derived from this set of drawings. This is implied by the presence of many un- published names on the illustrations that were written by Boisduval and include the Latin suffix “nob.” or “nobis”, meaning “of us.” Two small drawings in this set are identified as Sphinx ulmi (=Ceratomia amyntor Geyer) (90, 91), which Boisduval did not describe un- til 1875. Several of Boisduval’s inscribed names were apparently “borrowed” by Guenée ({1852—-58]), who used them for his own descriptions. Lawrence F. Gall recently examined Abbot's original drawings in this set and confirmed (pers. com.) that they were likely among those that Guenée consulted for his publica- tion. I am assisting Patrick G. Scott (Associate Univer- sity Librarian for Special Collections, Thomas Cooper Library) to identify the species depicted in this set of drawings, which will be made available for viewing on the Internet. The original drawing of Melitaea ismeria. The original illustration used by Boisduval and Le Conte for their description of M. ismeria (Fig. 2) is contained in the first of six portfolio cases as packaged by H. P. Kraus. It is included on drawing 24; the number “24” being written in graphite in two different hands across the top margin. The numbers “5” and “6” are also writ- ten in graphite at the top right and extreme lower left, respectively, but their meaning is unknown. The fig- ures of M. ismeria were drawn on the left half of the sheet. They are positioned under Boisduval’s small handwritten pencil heading of “Diurn. [Diurnes] 277 and consist of a dorsal female, ventral female, mature larva, and pupa that match the figures on Plate 46 in Boisduval & Le Conte ({1833]). There are visible cor- rections to the heads, legs and abdomens of the adult figures. The right half of the sheet, “Diurn. 26,” ex- hibits a dorsal female, ventral female, mature larva, and pupa of Euptoieta claudia (Cramer), matching the figures on published Plate 44 in Boisduval & Le Conte ({1833]). The left wings of both dorsal adult figures are unfinished, undoubtedly because engravers required only one completed side from which to extrapolate an entire illustration (probably the same reason E. Blan- chard rendered only one half of his dorsal adult figures). There are several inscriptions on the sheet in Bois- duval’s hand. Faint pencil notations are present below the figures of M. ismeria, reading “myrina Cr” and “myrissa God.” and probably represent Boisduval's ini- tial attempt to compare the figures with Brenthis my- rina Cramer (=Boloria selene myrina) and Argynnis myrissa Godart (a proposed replacement name for B. myrina). Written below the figures of E. claudia are “claudia Cr’ and “columbina F” (a synonym of the closely related Euptoieta hegesia (Cramer) and the name used by Boisduval & Le Conte ({1833]) for their Plate 44). Inscribed in ink at the bottom left, also in Boisduval’s hand, is “M. pyone Bd.” This name does not conform to any butterfly taxa of the era, including those described by Boisduval (Kirby 1871). Boisduval perhaps proposed this name (i.e., Melitaea pyone), but later abandoned it in favor of M. ismeria. The original figures of M. ismeria are clearly copies of those in Abbot's earlier painting of C. gorgone in The Natural History Museum, London (Figs. 6, 8, 11, 15). Abbot probably provided notes with these draw- ings, but they were undoubtedly lost during the nu- merous transfers of ownership. Abbot may have col- lected natural history specimens in South Carolina (Sanders & Anderson 1999), but the reference to ‘Carolina” in the original description of M. ismeria likely came from J. E. Le Conte, who traveled more widely in the southeastern United States. Analysis of immatures. Brown (1974) discussed at length his inability to match the larva and pupa in the published plate of M. ismeria with known species of Chlosyne. However, he relied primarily upon 19th century larval descriptions and did not fully under- stand the polymorphic nature of C. gorgone larvae. Gatrelle (1998) attempted to rear a large number of C. gorgone larvae, but few reached maturity and he did not discuss their coloration. Several of these larvae were forwarded to Thomas J. Allen to be pho- tographed, but they also failed to reach maturity (T. J. Allen pers. com.). To settle this issue, I contacted lepidopterists famil- iar with the immatures of eastern Chlosyne species. Nick V. Grishin has reared C. gorgone and C. nycteis from Texas, Paul M. Catling has reared C. gorgone from Ontario, Canada (Catling & Layberry 1998). sane Richard F. Boscoe has reared C. gorgone from South Carolina, as well as C. nycteis and C. harrisii from populations in the northeastern United States. Boscoe reared C. gorgone from eggs obtained in Orangeburg County, South Carolina, only 96 km (60 mi) northeast of Burke County, Georgia, where Abbot obtained his figured specimens. Gatrelle (1998) applied both popu- lations to the nominate subspecies. Grishin, Catling and Boscoe compared the mature larva in the original drawing of M. ismeria (Fig. 11) with mature larvae of all three eastern Chlosyne species. Grishin and Boscoe observed that mature lar- vae of C. nycteis are black with broad yellow or orange lateral bands (see Allen 1997, Plate 36, row 4). Boscoe noted that mature larvae of C. harrisii are orange with transverse black stripes on each segment (see Allen 1997, Plate 37, row 1). Grishin and Boscoe confirmed that mature larvae of C. gorgone are highly variable, possessing three primary color forms; all black (ni- JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY gra’), black with orange or fulvous longitudinal band- ing (‘bicolor’), and nearly all orange (‘rubra’). Interme- diates are common. Catling found young instars of C. gorgone to always be pure black, but mature larvae are either totally black (fm. ‘nigra’) or black with brownish- orange banding (fm. ‘bicolor’). Although mature larvae of C. nycteis and C. gorgone can be similar, those of C. nycteis consistently lack orange or fulvous dorsal band- ing often present in C. gorgone fm. ‘bicolor.’ Abbot's larval figure displays yellowish-orange dorsal banding and as such most closely matches C. gorgone. Grishin provided a color photograph of a mature larva of C. gorgone fm. ‘bicolor’ approaching the pattern figured by Abbot (Fig. 13). The pupa of C. nycteis is white with extensive black mottling (see Allen 1997, Plate 47, row 1). The pupa of C. harrisii is similarly white with irregular black, or- ange, and brown spotting (see exuvia photo in Williams 2002). Grishin and Boscoe described the pupa of C. gorgone as more uniform in color, brownish or grayish. I examined pupal exuvia of C. gorgone from three males and three females reared in 2000 and 2002 by Grishin from the vicinity of Dallas, Texas, and three males and one female reared in 1995 by Boscoe from ova obtained near the town of North, South Car- olina (FSCA collection). Grishin also provided two color photographs of living C. gorgone pupae from Texas. These examples all possess an extremely intri- cate pattern of brown, gray and white maculation, re- sulting in an overall brownish-gray or reddish-brown coloration. There are pale dorsal highlights on many abdominal and thoracic segments, as well as an undu- lating series of small white spots across each wing en- casement (Fig. 17). Abbot's painting of C. gorgone, as well as the original drawing and published plate of M. ismeria include the same depiction of a pupa that is unmistakably consistent with C. gorgone. These fig- ures even include the pale segmental highlights and row of white forewing spots (as an unbroken line) (Figs. 14-17). Written descriptions of larval C. gorgone by several authors, including Klots (1951), and Brown et al. (1955), obviously repeated the description given by Holland (1898), who considered gorgone and ismeria synonymous and derived his information from the published plate of M. ismeria. Consequently, these later authors unwittingly associated M. ismeria with C. gorgone, including F. M. Brown who fundamentally disagreed with this synonymy(!). A search for John Abbot specimens. Surviving John Abbot specimens of Chlosyne would reveal much about the species he encountered in Georgia. Brown (1974) and Gatrelle (1998) could not locate any such VOLUME 57, NUMBER 3 specimens. I searched additional sources for evidence of their existence. Jacob Hiibner (1806-[1838]) figured at least four species based on specimens from “Georgia” and “Georgien.” Authors, such as Clark & Clark (1941) and Brown (1974), have speculated that such specimens came from John Abbot, but their actual source remains obscure. Unfortunately, no text accompanied Hiibner’s plate of Dryas gorgone and Hiibner’s manuscripts do not provide additional insight into their origin (Hem- ming 1937). According to notations in the publication, as well as Hiibner’s manuscripts, North American spec- imens used for his plates came from Georgia, New York, Pennsylvania, Virginia, and “America” (Hemming 1937). Although the specimens of C. gorgone were most likely from Abbot, we may never be certain. Dur- ing the early 19th century, Hiibner’s Lepidoptera type specimens were obtained by Vincenz Abbate Edler von Mazzola. In 1823, Mazzola’s European Lepidoptera collection was deposited in the Emperer’s “Naturalien- Kabinett” in Vienna, Austria. It is believed that many of these specimens bummed in a fire in 1848 (probably dur- ing the revolution that year). The few surviving Hiibner specimens are now deposited in the Naturhistorisches Museum in Vienna (Horn et al. 1990). Regrettably, Mazzola removed all of Hiibner’s original labels, com- plicating positive identification of Hiibner material. The fate of Hiibner’s North American specimens is un- known and no C. gorgone, C. nycteis, or C. harrisii are now deposited in the Naturhistorisches Museum (M. Lédl pers. com.). Edward Doubleday wrote in 1840 that, “In all old collections are many specimens collected by Abbot; at Francillon’s, Donovan’s and other sales, some of these have been dispersed, and have crept into collections nominally British only” (Scudder 1869). After John Francillon’s death in 1816, his collection of insects was sold in London in four separate auctions in May and July 1817 and June 1818. The sales catalog from July 1817 (King 1817) contained numerous listings for “beautiful Georgian Lepidoptera” and other insects. Unfortunately, there were no specific listings that could suggest Chlosyne. The bulk of Francillon’s col- lection, including 72 drawers of foreign Lepidoptera, was sold 11-19 June 1818. The Lepidoptera portion of the 1818 auction catalog (King 1818) listed dozens of specimens from Georgia, especially moths. Among the contents of Drawer 32, Lot 6, were three Georgia specimens of an unidentified “Argynnis.” This lot con- tained similar small species, including North American B. s. myrina (as “Myrina”) and Phyciodes tharos (Drury) (as “Tharos”). Alexander Macleay, an English naturalist and honorary Secretary of the Linnaean So- ciety of London, acquired a large portion of this col- lection. In 1825, Macleay moved from England to Australia to serve as Colonial Secretary of New South Wales. His insect collection now serves as the core of the Macleay Museum, University of Sydney (Barker 1999). Among numerous North American insect spec- imens in the Macleay Museum are many labeled sim- ply “Georgia” that undoubtedly originated with Abbot. A digital scan of a label taken from a beetle specimen from Georgia shows it was written in Abbot's hand (Fig. 25), confirming that Abbot personally labeled at least some of his own specimens. A late 19th century curator foolishly discarded many of the original labels in favor of more carefully written substitutes (M. Humphrey pers. com.). Macleay incorporated Francil- lon’s specimens into his own collection and the original organization was lost. Unfortunately, no C. gorgone, C. nycteis, or C. harrisii were found in the Macleay Mu- seum collections (M. Humphrey, K. Fairey pers. com.). In an astonishing letter to T. W. Harris dated 30 April 1842, E. Doubleday wrote that he had found in the British Museum “some specimens of Melitaea Is- meria, collected by Abbot,” adding, “It is nearer M. tharos than Boisduval’s plate would lead you to imag- ine” (Scudder 1869). In 1847, Doubleday again re- ferred to M. ismeria in the British Museum (Double- day & Hewitson 1846-50). Probably between 1906 and 1908, when he taught classes in Europe, W. T. M. Forbes saw Georgia specimens of C. gorgone in The British Museum (NH) that he later reported as “from Abbot” (Forbes 1960). Ironically, Forbes (1960) mir- rored the earlier observations of Doubleday, stating that these specimens looked “at first glance much more like tharos than carlota.” Gatrelle (1998) at- tempted to locate potential Abbot Chlosyne specimens in The Natural History Museum (“BMNH_’), but was unsuccessful. Nonetheless, a single male C. gorgone, labeled simply “Georgia,” was discovered among spec- imens pulled from the main collection by Lionel G. Higgins during his work on Chlosyne (P. Ackery pers. com.). The specimen (Fig. 23) has a damaged right hindwing and lacks a left antenna, but is otherwise in good condition. In addition to the locality label, this specimen of C. gorgone bears a small round label reading “520” with another character nearly obliterated by pinholes. A third label, probably placed during the late nineteenth century, reads, “carlota Reak.” (Fig. 25). Phillip R. Ackery (Collections Manager) confirmed my suspicion that “520” corresponded to a species listing in E. Dou- bleday’s manuscript catalog of Lepidoptera specimens in the British Museum (Entomology Library, The Nat- ural History Museum) (see Harvey et al. 1996). The entry for species 520 was given as, “Argynnis Ismeria Boisduval” and listed specimens “a, b, Georgia; c, Ohio.” The published version of this catalog (Double- day 184448) was not numerical and these specimens were identified as “Melitaea Ismeria, Boisd. et Leconte.” Doubleday undoubtedly affixed the numeric label dur- ing the preparation of his manuscript catalog and the obscure character on this label is likely a “b,’ matching the specimen he listed. Doubleday’s association of M. ismeria with C. gorgone is consistent with his 1840 identification of Abbot's painting in London. The Ohio specimen listed by Doubleday is also extant and repre- sents C. gorgone. The locality labels on both the Ohio and Georgia specimens are similar, being less discol- ored with a characteristic double black line drawn across the lower edge (Fig. 25). Although Doubleday’s published catalog (Double- day 1844-48) did not indicate the origin of these C. gorgone specimens in the British Museum, his original manuscript gave “Dyson” as the source of the Ohio specimen. English naturalist David Dyson (1823-56) spent nearly the entire year of 1843 in America where he collected insects, birds, shells and plants, “across the Allegheny Mountains, and as far as St. Louis” (Anonymous 1856). Other old butterfly specimens in the collection from the United States bear locality la- bels with the same characteristic double black line. The similarity of the labels suggests that Dyson col- lected them all. However, Ives (1900-01) claimed Dyson was unable to read or write and utilized “a kind of hieroglyphic marking understood only by himself.” If this is true, Dyson may have verbally communicated his collecting localities to Doubleday, who recorded the data only in his manuscript catalog. Comments by Doubleday (1844) show that they were personally ac- quainted at the time. The locality labels actually look to be of more recent provenance and were probably affixed by a later museum worker in an attempt to standardize the data on these old specimens. In addi- tion, Dyson’s route in America implies that he followed the Ohio and Mississippi River Valleys and did not reach as far south as Georgia. Doubleday (1844-48) listed Dyson as the source of other Ohio specimens, but none from Georgia. Finally, Doubleday’s 1842 dis- covery of M. ismeria in the British Museum predated Dyson’s trip to America and there is no indication that Doubleday ever applied the name ismeria to any species other than C. gorgone. Based on available evi- dence, there is little doubt that the surviving C. gorgone from Georgia is one of the specimens that Doubleday identified as M. ismeria from John Abbot. Purported Abbot specimens were acquired by the British Museum from many sources. Two years before JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Doubleday discovered the M. ismeria specimens in the British Museum, he wrote that “many Lepidoptera of Abbot's collecting” were bought by the museum from “the late Mr. Milne’s collection” (Scudder 1869). The George Milne (or Mylne) collection of 1749 spec- imens, mostly Lepidoptera and Coleoptera, was pur- chased by the museum in June 1839 (Stevens 1839, Stearn 1981). Auction lots 195 and 196 of the Lepi- doptera portion of the Milne sales catalog listed “sev- eral rare species of Melitaea” and “various species of Melitaea,” respectively (Stevens 1839). The surviving Georgia specimen of C. gorgone may have been ob- tained from this collection. The fate of the remaining Georgia specimen of M. ismeria that Doubleday listed as “520a” is unknown. No Georgia specimens of C. nycteis or C. harrisii are currently deposited in The Natural History Mu- seum (P. Ackery pers. com.). Doubleday’s original de- scription of nycteis did not include Georgia within the general distribution of “Middle States” (M. ismeria was listed separately from “Southern States”). Double- day’s catalog predated the original description of har- risii by nearly 20 years. Most assuredly, if Doubleday had found this insect in the British Museum, he would have recognized it as new and promptly described it with nycteis in Doubleday & Hewitson (1846-50). DISCUSSION The true identity of Melitaea ismeria. The plate of M. ismeria in Boisduval & Le Conte ([1833]) was engraved from the original John Abbot drawing now deposited in the Thomas Cooper Library, University of South Carolina. The figures in this drawing (ca. 1815) (Fig. 2) are analogous to those in Abbot's earlier paint- ing of C. gorgone (ca. 1804) (Fig. 3) deposited in The Natural History Museum, London. Therefore, M. is- meria is synonymous with C. gorgone (Figs. 4, 6-9, 10-12, 14-16). The figured adults and early stages were simplified with each successive copy, resulting in a published plate that held little resemblance to the initial painting. This imprecision contributed to nearly two centuries of nomenclatural confusion. John Abbot was approximately 64 years of age when the original drawing for M. ismeria was completed. Al- though he apparently collected and painted natural history specimens into his eighties, he became less ca- pable of travel in his later years, spending more time painting than exploring the countryside for new dis- coveries. In an 1834 letter to T. W. Harris from Abbot's long-time friend, Augustus G. Oemler, Abbot was de- scribed as “very corpulent, but still exercises his pur- suit of hunting birds and drawing—but engaging boys to run after butterflies” (Dow 1914). J. E. Le Conte re- VOLUME 57, NUMBER 3 quested that Abbot include both adults and immatures in his paintings. Surely, it would have been a daunting task for Abbot to collect all new specimens and repeat his laborious life history studies. This is especially true for species he considered rare or uncommon, such as C. gorgone. Throughout his career, Abbot was known to main- tain a master set of template drawings with accompa- nying life history notes from which to create additional renderings of the same species. Paintings in John Francillon’s volumes were numbered so additional copies could be ordered for other buyers (Rogers- Price 1983, Gilbert 1998). Abbot completed duplicate paintings for many individuals, including J. Francillon, A. G. Oemler, and English naturalist William Swain- son. Ten out of 30 surviving Abbot paintings of Cato- cala Schrank moths for Francillon and Oemler are ex- act duplicates (Gall & Hawks 2002). One of the illustrations that Abbot duplicated was the “Cross wort Frittilary Butterfly” (C. gorgone). From about 1813 to 1818, Abbot provided to A. G. Oemler at least 193 paintings that are now deposited in the Houghton Library, Harvard University. Plate 11 of this set, measuring 34 cm x 24 cm, is a duplicate of the earlier C. gorgone painting in The Natural History Museum, London. In the accompanying “Notes to the Drawings of Insects,’ Abbot identified it as the “Cross Wort Frittilary” and added, “Feeds on Cross Wort, and Sun flower, changed 17th May—bred 26th. Frequents the Oak woods of Burke County, but is not common” (S. Halpert pers. com.). Between 1816 and 1818, Ab- bot also completed 103 illustrations of insects for W. Swainson, mostly Lepidoptera not figured in Smith & Abbot (1797). Swainson emigrated to New Zealand in 1840 and the paintings were acquired in 1927 by the Alexander Turnbull Library, Wellington (Parkinson 1978, 1983). Plate 17 in this set is another duplicate of Abbot's C. gorgone painting in London (Figs. 3, 5, 21). It measures 34.2 cm x 24.8 cm and was figured in color by Parkinson and Rogers-Price (1984). Again, Abbot's entry in his accompanying “Notes to the Drawing of Insects” is the same: “Cross wort Frittilary Butterfly. Feeds on Cross wort, and sunflower, Tyed itself up by the tail 16th May, changed 17th bred 26th. Frequents the Oak Woods of Burke County, but is not common.” To fulfill Le Conte’s commission, Abbot likely relied on his template drawings as often as possible. A com- parison of engraved plates in Smith & Abbot (1797) and Boisduval & Le Conte (|1833]) shows that many contained duplicate figures. Ten of the 23 species treated in both publications included identical depic- tions of larva and/or pupa. Many of Abbot's other paintings also share figures with the drawings used by Boisduval & Le Conte ([1833]). It is obvious that the drawing used for the description of M. ismeria is no more than an abbreviated version (no male butterfly or hostplant) of the same C. gorgone illustration that Ab- bot provided to Francillon, Oemler and Swainson. Traces of corrected graphite sketch lines are visible around the adult figures in the original drawing of M. ismeria. These lines correspond to the outlines of the counterpart figures in Abbot's duplicate paintings of C. gorgone (Figs. 18-19) and offer convincing evidence that Abbot indeed copied this drawing from his tem- plate of C. gorgone. Abbot's later copies were more carelessly rendered than the earlier paintings for Fran- cillon (Figs. 4-6). In 1819, Swainson even complained to Abbot that the drawings he received were “not so highly finished” as those published in Smith & Abbot (1797) (Parkinson 1978). The three known copies of the C. gorgone illustration, including the original draw- ing of M. ismeria, were probably completed within a five-year period (1813-18) during Abbot's 64-year resi- dency in Georgia. Artwork of John Abbot is deposited at many locations and there may be additional surviv- ing copies of this rendering. Assessment of the current neotype. Article 75.3.5 of ICZN (1999) states that a neotype is validly designated only if it is “consistent with what is known of the former name-bearing type from the original de- scription and from other sources.” Although the origi- nal description of M. ismeria did not include a name- bearing type specimen, the neotype of Gatrelle (1998) is not consistent with the identity of the intended species. To promote nomenclatural stability, the neo- type Melitaea ismeria Boisduval & Le Conte, [1833] should be set aside and another designated to reflect synonymy with C. gorgone. An ICZN application has been prepared to achieve this objective (Calhoun et al. under consideration). Opler and Warren (2002) re- ferred to the preparation of another petition to sup- press the use of ismeria as “a possible senior synonym of nycteis,” but it was not submitted in deference to the present study. Commentary on Brown (1974) and Gatrelle (1998). Despite his thorough treatment, Brown (1974) provided misleading information. He main- tained that, “Scudder (1872) stated that he had found the original of Abbot’s plate of ismeria in the British Museum (N. H.) and that it represented the male of Huebner’s gorgone.” In actuality, Scudder (1872a) made no such allusions and simply listed ismeria among the John Abbot paintings in the British Mu- seum. Scudder did not elaborate. Strecker (1878) im- plied this claim when he credited Scudder with reveal- ing the published figures of M. ismeria “were copied from Abbot's unpublished drawings and poorly enough copied at that.” It is possible that Scudder wrote to Strecker about the original drawing he found in Bois- duval'’s library. Nonetheless, Scudder recognized the resemblance between Abbot’ earlier painting and the published plate of M. ismeria. He further associated the adult figures with Eresia carlota (=C. gorgone), which was loosely described five years prior to his visit to London. Brown inexplicably disregarded the obvi- ous similarity of Abbot’s painting to the published fig- ures of M. ismeria. He apparently intended to repro- duce the painting, but there is no figure associated with his reference to “(our figure 5).” Furthermore, he never actually reported the identity of the species de- picted, undoubtedly contributing to the misconcep- tions of Gatrelle (1998). Gatrelle (1998) misinterpreted crucial information. He alleged Brown (1974) “established ismeria as a valid (but unidentified) species separate from gorgone and postulated that it could well be C. nycteis.” In fact, Brown could not positively identify M. ismeria and recommended that the name be ascribed only to the published plate and not to any existing species. Brown finally suggested the published plate was a fictitious representation. Most importantly, Gatrelle misunderstood the status of Abbot’s painting and accompanying notes in The Natural History Museum (Fig. 3) and did not confirm the identity of the depicted butterfly or hostplant. He wrongly assumed Helianthus trachelifolius, as in- scribed on Abbot's notes page, was the identity of the figured plant and mistakenly associated it with He- lianthus strumosus L. (Asteraceae). Gatrelle ultimately disregarded the painting (as unidentifiable?) and erro- neously applied the life history notes to support his proposed synonymy with C. nycteis. Not only are the notes referable to C. gorgone, their forced application to C. nycteis is tenuous. Abbot’s reference to “oak woods” is consistent with Gatrelle’s “oak sandhill” habitat of C. gorgone, but not the riparian habitats as- sociated with C. nycteis in Georgia (Harris 1972), in- cluding the three specimens Gatrelle personally col- lected along the Savannah River. In the notes for his various illustrations, Abbot plainly differentiated up- land “oak woods” from bottomland “swamps.” Gatrelle also argued that the dates given in Abbot's notes more accurately coincide with C. nycteis, which emerges a month later than C. gorgone in Burke County, Geor- gia. These dates cannot be directly compared, as Ab- bot’s data was not from a wild-collected adult and his rearing conditions could have resulted in abnormal de- velopment. According to Gatrelle (1998), populations of C. gorgone in coastal Georgia and South Carolina JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY are univoltine with diapausing third instar larvae, but Gatrelle (1993) reared two adults from ova obtained earlier the same year. The life cycle for these individu- als was 42 days, showing Abbot could also have reared his specimen from an ovum he obtained at the onset of the normal adult flight period in mid-April, which pro- duced an adult on 26 May of the same season. Finally, Gatrelle (1998) agreed with Harris (1972) who believed the original plate of Dryas gorgone in Hiibner (1806-[1838]) was engraved from a painting by John Abbot. While the figured specimens probably came from Abbot, the original illustration and engrav- ing were undoubtedly the work of Jacob Hiibner him- self. Representations of C. gorgone by Hiibner and Abbot reveal very different portrayals and artistic styles (Figs. 4-6, 20). Hemming (1937) described Hiibner as a “draughtsman and illustrator of excep- tional skill” whose propensity for drawing was noted at an early age. Unpublished, typewritten research of Cyril F. dos Passos, dated 14 October 1955, was found inserted into his personal copy of Hiibner (1806-[1838]) (Wittenberg University), in which he determined, “the plates of volumes 1 and 2 are by Hiibner with the exception of four (4) plates by Geyer, numbers [85], [119], [186], and [209], and the plates of volume 3 are by Geyer” (brackets of dos Passos). Many of the original paintings for this publication by Carl Geyer (Hiibner’s assistant) were acquired in 1949 by The Natural History Museum, London, as part of the Baron von Rosen manuscript collection (Harvey et al. 1996). Hiibner’s original illustration of Dryas gorgone remains elusive. Hemming (1937) did not find it in any institutions known to contain artwork used for this publication. The Natural History Museum acquired additional Hiibner manuscripts with the von Rosen documents (Harvey et al. 1996), but a search of this material was also unsuccessful (V. Veness pers. com.). John Abbot may have encountered more than one species of Chlosyne in Georgia, but available evidence precludes all but C. gorgone. When Edward Double- day described Melitaea nycteis in 1847, he failed to see any resemblance with Melitaea ismeria. Within seven years of the original description of M. ismeria, Dou- bleday had correctly determined the intended species as the insect now known as C. gorgone. Samuel H. Scudder corroborated Doubleday, but his findings were disregarded. It took 160 years to prove they were both correct. To quote Herman Strecker (1878), “Time at last sets all things even.” ACKNOWLEDGEMENTS My appreciation is boundless for the many individuals who assisted in this study. Patrick G. Scott (Thomas Cooper Library, Univ. of South VOLUME 57, NUMBER 3 Carolina) provided access to the set of original John Abbot drawings. Without his help, this study would not have been possible. Mark Maier digitized these drawings for examination. For providing digital photos of specimens, artwork and published plates, as well as litera- ture and other information, I am very grateful to Phillip R. Ackery, Kim Goodger, Dawn Hathaway, Peter B. Mordan, Robert P. Prys- Jones, Richard I. Vane-Wright, and Vicki Veness (all The Natural His- tory Museum, London), John M. Burns and Donald J. Harvey (Na- tional Museum of Natural History, Smithsonian Institution, Washington, D.C.), Margaret Calder and Philip G. Parkinson (Na- tional Library of New Zealand, Alexander Turnbull Library, Welling- ton), Robert S. Cox (Library of the American Philosophical Society, Philadelphia), Mary Ann Folter (H. P. Kraus, New York), Susan Halpert (Houghton Library, Harvard Univ.), Margaret A. Humphrey and Kenneth Fairey (Macleay Museum, Univ. of Sydney), Martin Lédl (Naturhistorisches Museum, Vienna), ), Jacqueline Y. Miller and Lee D. Miller (Allyn Museum of Entomology, Florida Museum of Natural History, Sarasota, Florida), Beverly L. Pope and Alice R. Saunders (Division of Plant Industry Library, F lorida Dept. of Agri- culture & Consumer Services, Gainesville), Linda Seckelson (Thomas J. Watson Library, The Metropolitan Museum of Art, New York), and Suzanne Smailes (Wittenberg Univ. Library, Springfield, Ohio). Thomas J. Allen, Erika Bandera, Richard F. Boscoe, Paul M. Catling, Nick V. Grishin, John B. Heppner (Florida State Collection of Arthro- pods, Division of Plant Industry, Florida Dept. of Agriculture & Con- sumer Services, Gainesville), and Michael A. Quinn supplied speci- mens and additional assistance. Richard Carter (Valdosta State Univ.), Mark A. Garland (Univ. of Florida), Charles B. Heiser, Jr. (Indiana Univ.), Richard P. Wunderlin (Univ. of South Florida), and Wendy B Zomlefer (Univ. of Georgia) provided botanical expertise and searched for herbarium specimens. I am very grateful to Jonathan P. Pelham, John A. Shuey, Niklas Wahlberg (Univ. of Stockholm), An- drew D. Warrren (Oregon State Univ.), and especially Jacqueline Y. Miller and Lee D. Miller for patiently reading and/or critically re- viewing various drafts of the manuscript. Upon reading an earlier ver- sion of this manuscript, Jackie Miller and Lee Miller offered their full support of this project, even coauthoring my application to the ICZN. I cannot thank them enough. 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The American rhopalocera (Macrolepidoptera of the American fau- nistic region). Vol. 5. Alfred Kemen Verlag, Stuttgart, Germany. 1139 pp., 194 pl. SmitH, J. E. & J. Abbot. 1797. The natural history of the rarer lep- idopterous insects of Georgia, including their systematic char- acters, the particulars of their several metamorphoses, and the plants on which they feed, collected from the observations of Mr. John Abbot, many years resident in that country. 2 vols. T. Bensley for J. Edwards, Cadell and Davies, and J. White, Lon- don. 214 pp., 104 pl. SOTHEBY & Co. (FIRM). 1963. Catalogue of a fine collection of rare books and drawings on natural history and travel. Lon- don. 85 pp. STEARN, W. T. 1981. The natural history museum at South Kens- ington. A history of the British Museum (Natural History) 1753-1980. Heinemann, London. 414 pp. STEVENS, J. C. 1839. Collection of foreign and British insects formed by the late George Mylne, Esq. [sales catalog]. London. 23 pp. STRECKER, H. 1878. Butterflies and moths of North America with full instructions for collecting, breeding, preparing, clas- sifying, packing for shipment, etc., a complete synonymical catalogue of macrolepidoptera with a full bibliography to which is added a glossary of terms and an alphabetical and descriptive list of localities. H. F. Owen, Reading, Pennsylva- nia. 283 pp. USDA (UNITED STATES DEPARTMENT OF AGRICULTURE). 2003. The PLANTS database (version 3.5, 3 February 2003). Nat. Res. Cons. Service. http://plants.usda.gov. WILKINSON, R. S. 1981. Smith and Abbot, the natural history of the rarer lepidopterous insects of Georgia (1797): its authorship and later history. Entomol. Rec. 93:213-218. . 1984. John Abbot's London Years. Entomol. Rec. & J. Var. 96:110-123, 165-176, 222-229, 273-285. WILLIAMS, E. H. 2002. Harris’ checkerspot: a very particular but- terfly. Amer. Butterflies 10:18-25. VOLUME 57, NUMBER 3 Received for publication 31 December 2002; revised and accepted 7 April 2003. NOTE ADDED IN PRESS: On 26 April 2003, after this paper had gone to press, I visited the sites in Burke County, Georgia where R. Gatrelle had found C. gorgone and C. nycteis (locality data obtained from his neotypes in the Allyn Museum of Entomology). I obtained one male and one female C. gorgone that are very consistent with 219 Abbot’s illustrations and purported specimen in London. Gatrelle designated the type locality of Dryas gorgone as “Burke County, Georgia,” but this county is 2,155 sq. km (832 sq. mi) in size. Gatrelle (1998) did not publish all the information that appears on the labels of his neotype specimen. The collection location was given as “River Rd at Hancock Landing Rd.” The type locality should be further restricted to the town of Hancock, Burke County, Georgia. Hancock is located only 11 km (7 mi) northeast of Abbot's former residence in Burke County. Journal of the Lepidopterists’ Society 57(3), 2003, 220-229 LATE-INSTAR SHIFT IN FORAGING STRATEGY AND TRAIL PHEROMONE USE BY CATERPILLARS OF THE NEOTROPICAL MOTH ARSENURA ARMIDA (CRAMER) (SATURNIIDAE: ARSENURINAE) James T. Costa', DIETRICH A. GOTZEK° Department of Biology, Western Carolina University, Cullowhee, North Carolina 28723, USA AND DANIEL H. JANZEN Department of Biology, University of Pennsylvania, Philadelphia, Pennsylvania 19104, USA ABSTRACT. Caterpillars of Arsenura armida (Cramer) (Saturniidae: Arsenurinae) are diurnal nomadic foragers in early instars, maintain- ing aggregations within the host tree crown through the use of a trail pheromone. In the fourth instar, larvae switch foraging strategies to be- come nocturnal central place foragers. In central place foraging mode, the caterpillars rest by day on the trunk of the food plant, ascend to the canopy at nightfall to feed, and then return to the lower trunk by dawn, often at the same resting (bivouac) sites as used previously. Peak activity ascending to and descending from the canopy occurs in twilight. Central place foraging A. armida caterpillars do not maintain colony structure at night, but disperse in the canopy to feed singly. The caterpillars appear to use tree architecture and their trail pheromone to relocate con- specifics (which are generally confamilials) upon descending. While bivouac sites are often reused, individual caterpillars do not exhibit strict site fidelity and may go to a bivouac site different from whence they came. This shift in foraging behavior entails a concomitant change in reac- tion to the information content of A. armida’s trail pheromone, from maintaining groups as the caterpillars move from patch to patch, to relo- cating distant resting sites. Diurnal resting bivouacs are probably warning displays, and we discuss this behavior in the context of A. armida’s de- fensive ecology. Additional key words: _ trail-following, group foraging, social caterpillars, chemical communication, tropical dry forest. Larval sociality is widespread in the Lepidoptera, (Pieridae) (see additional examples in Fitzgerald occurring in an estimated 27 families in 19 superfami- 1993a, Costa & Pierce 1997). lies (Costa & Pierce 1997). Expressions of sociality in Although the behavior of relatively few social Lepi- this order vary considerably, and may include group doptera have been studied to date, species represent- defense, group nest or shelter construction, and/or ing each of these foraging strategies have been shown group foraging (e.g., Fitzgerald 1993a, Costa & Pierce to communicate via trail pheromones. A. cerasivo- 1997). Fitzgerald and Peterson (1988) suggested that ranus, for example, uses a pheromone to promote communication complexity of caterpillar societies can within-patch aggregation (Fitzgerald 1993b). Fitzger- be categorized by foraging strategy, identifying three ald and Costa (1986) studied the nomadic foraging la- basic modes of group foraging (patch-restricted, no- siocampid Malacosoma disstria Hiibner, the forest madic, and central place foraging) that in part reflect tent caterpillar, which also uses a trail marker for the extent of communication and cooperation by group cohesion. Its congener, the central place forag- group members. Patch-restricted foragers remain ing M. americanum, is the only social caterpillar thus more or less in a single patch or feeding site for the far shown to engage in elective recruitment to food. duration of the larval stage, typically constructing nests This species employs a two-part trail system consisting by tying leaves with silk or wholly enveloping leaves in of exploratory and recruitment trails used to direct masses of silk and extending the bounds of the nest as tentmates to high quality food patches (Fitzgerald & food becomes locally exhausted (e.g., the ugly nest Peterson 1983). caterpillar Archips cerasivoranus (Fitch), Tortricidae). The foraging strategy of some social caterpillars is Nomadic foragers, in contrast, move en masse among intermediate between the three basic strategies delin- patches, constructing no shelters and often exhibiting eated by Fitzgerald and Peterson (1988), and some aposematic coloration (e.g., notodontids like Anisota species switch strategies as they grow. It is very com- or Datana spp.). Finally, central place foragers nest or mon among social Lepidoptera and Symphyta for rest in a fixed location, periodically leaving the site to group fidelity to wane over time such that the penulti- feed; for example the eastern tent caterpillar Malaco- mate or ultimate instars forage solitarily, and many be- soma americanum Fab. (Lasiocampidae) and the gin their lives feeding as a group at the site of larval madrone caterpillar Eucheira socialis Westwood eclosion through at least the first instar, but soon shift to nomadic or central place foraging. These cases ‘To whom correspondence should be addressed. Email: might represent, respectively, the beginning of the dis- costa@wcu.edu all aly delaved t of active fe > Current address: Department of Genetics, University of Geor- persal phase or delayed commencement Or active tor- gia, Athens, Georgia 30602 USA. aging, and so may not be good examples of strategy VOLUME 57, NUMBER 3 switching per se. Only a few species are known to ex- hibit behavioral changes in later instars, but the list is growing. For example, in the penultimate or ultimate instar the arctiid Hyphantria cunea Drury switches to central place foraging, as does the European pine pro- cessionary Thaumetopoea processionea (Linnaeus) (Notodontidae) (T. D. Fitzgerald pers. obs.). The sat- urniids Hylesia lineata Druce and Automeris zugana Druce feed diurnally as a group of sibs in nomadic fashion through the second instar, when they com- mence nest building and switch to nocturnal central place foraging (DH] pers. obs.). Insofar as shifts in for- aging strategy by social caterpillars reflect an evolved response to temporal changes in host quality or preda- tion pressure, and are likely to be accompanied by shifts in communication mode, these species may be informative model systems with which to study factors shaping caterpillar social behavior. Few studies to date have focused on developmental shifts in social caterpillar foraging strategy, and no so- cial caterpillar has yet been shown to use a trail marker for different uses over time (though this is likely to be common). Here we report a change in foraging strategy and trail marker use by larvae of the Neotrop- ical arsenurine saturniid Arsenwra armida (Cramer) in northwestern Costa Rica. Costa et al. (2003) showed that the aposematic early instars of A. armida forage nomadically and use a trail pheromone to promote grouping. One of us (DH]J) previously observed that this species switches its foraging behavior in later in- stars, becoming a central place forager in that the lar- vae form sizable groups on the trunk of their host tree, leaving these sites to forage at night and reassembling at dawn or earlier. The present study documents this foraging mode by late instar A. armida with observa- tional and experimental data. In particular, we address the following questions: (1) Do A. armida larvae con- tinue to forage socially (in groups) in late instars? (2) Do larvae exhibit fidelity to a resting site, returning daily to the same location on the host tree? And (3) if group foraging and/or reusing resting sites, how might larvae use their trail marker to do so? Arsenura armida belongs to subfamily Arsenurinae, consisting of approximately 57 species of Neotropical saturniids found from tropical Mexico to northern Ar- gentina (Lemaire 1980, Hogue 1993). This species oc- curs from tropical Mexico to Bolivia and southeastern Brazil (Lemaire 1980, Balcézar & Beutelspacher 2000), and is found throughout lowland Costa Rica from dry forest to rainforest (for label data from spec- imens in INBio, see http:/Avww.inbio.ac.cr). In the tropical dry forest of northwestern Costa Rica, oviposi- tional hosts include guacimo (Guazuma_ ulmifolia bo bo — Lam., Sterculiaceae), on which it is an occasional pest (Hilje et al. 1991), Rollinia membranacea Triana & Planch (Annonaceae), and pochote (Bombacopsis quinatum (Jacq.), Bombacaceae) (Janzen & Hallwachs 2003). The following life history overview is based on rear- ing and observation records available in Janzen and Hallwachs (2003) and Janzen’s personal observations between 1978 and 2000. A more detailed overview with photographs is provided in Costa et al. (2003). The peak of A. armida adult eclosion occurs between the first and last week of June in the Area de Conser- vacion Guanacaste (ACG) dry forest, approximately three weeks after the rains begin (Janzen 1993), having passed the dry season as a solitary pupa in a chamber excavated in the soil. Part of the first generation enters dormancy as pupae and part ecloses about 5-8 weeks after pupation to create a second generation (November-December). All of the second generation pupae become dormant until the start of the following rainy season. After mating, female A. armida usually lay their entire egg load (350-500 eggs) in a single mass on the underside of a leaf, although split clutches are sometimes observed. The eggs hatch after about 2 weeks and the larvae remain brightly aposematic through their first three instars. Colonies of young in- stars forage nomadically using trail pheromones to promote colony cohesiveness, and silk plays no role in trail following (Costa et al. 2003). In the fourth (penul- timate) instar, the larvae switch their foraging strategy and begin to rest diurnally in large bivouacs on the lower trunk and underside of larger branches. Larvae remain together in this manner until late in the termi- nal stadium, when they abandon the tree as prepupae. MATERIALS AND METHODS Insect collection. One hundred twenty five penul- timate (fourth) instar A. armida caterpillars, 4-5 cm in length, were collected from a large (ca. 15 m tall) B. quinatum tree in the Cafetal area of Sector Santa Rosa, ACG, Guanacaste Province, Costa Rica. At least two large aggregations were found on this tree, each consisting of >200 larvae, with smaller groups nearby. Based on an average clutch size of about 450 eggs (Costa et al. 2003, Janzen & Hallwachs 2003), these aggregations may represent one or two family groups that have fragmented. Approximately half the individ- uals of an accessible lower aggregation were collected, leaving one large upper group and approximately half the second lower group (collectively numbering three to four hundred larvae) on the tree for later field ob- servations. The larvae were transported to the re- search center at the Administration Area of the ACG, bo i) bo where they were divided into three groups, each of which was transplanted onto a young host tree so they would be accessible for observation and experimenta- tion. One group was transplanted onto a 5 m tall B. quinatum sapling and two groups were transplanted onto adults of an alternate host, Guazuma ulmifolia. The first transplant tree (G1) was a G. ulmifolia, ca. 4 m in height and split into two main trunks about 30 cm above the base, DBH = 17 cm and 18 cm, respec- tively. A bag containing 40 larvae on G. ulmifolia fo- liage was attached to the lower trunk of the tree, and the larvae were permitted to exit the bag ad libitum. Within two hours nearly half of the caterpillars had ex- ited and formed a group on the trunk about 40 cm above the ground, and within 24 h, 37 were accounted for, occurring singly or in one of several groups or bivouacs. Each individual was then given a bivouac- specific mark on the dorsum of the anal segment (anal plate) or on one or both anal prolegs using a fine tip Sharpie® marker, after observations of test larvae showed that marking did not have an adverse effect. Singletons were also identified with a unique mark, and bivouac sites were marked with labeled pins to test for bivouac fidelity. The second G. ulmifolia (G2) was ca. 2 m in height, DBH = 4 cm, small enough to document the position of each larva in the canopy. Prior to release, 15 cater- pillars were marked on the anal plate using white Liq- uid Paper® correction fluid and given a unique num- ber with a fine-tip Sharpie® marker. Prior to release on the tree, larvae were observed for a 24 h period fol- lowing marking to ascertain they were not adversely affected by the marking procedure. There was no mor- tality and all larvae seemed to feed normally. Larvae were transplanted onto the tree using the same method as for G1. At the end of the first day, eleven of the larvae were relocated. Finally, the B. quinatum (G3) was about 5 m in height, DBH = 16 cm. Forty larvae were permitted to move onto the new host in the same manner as the transplanted G1 caterpillars. Observational and experimental data were collected from these groups over a 10 day period and used to ad- dress (1) larval foraging periodicity, (2) mode of larval foraging (ie., group vs. solitary foraging) day and night, and (3) the use of trail markers in foraging and bivouac formation. Group foraging. The group foraging dynamics of A. armida was assessed in three interrelated sets of ob- servations. First, mobilization of the caterpillars from their daytime resting bivouacs to nocturnal foraging mode, and their subsequent return and reassembly into bivouacs at dawn, was observed for seven consec- utive days. Bivouac fidelity and group composition was JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY documented by noting the position of marked G2 lar- vae each morning after all larvae had returned and se- lected a resting site. Because resting groups can be loose, often broken into two or more closely situated subgroups, initial bivouac sites were arbitrarily defined as a roughly circular area 15 cm in diameter centered on resting groups. Any larvae subsequently found within that area were scored as resting at that bivouac. Each bivouac site, including any new sites, was checked daily between 10:00 and 11:00 h for presence of larvae, and each larva’s bivouac of origin was noted daily. In addition, the canopies of each tree were searched to account for larvae that had not returned to a bivouac by the daily census time. The location of each larva of the G2 colony was doc- umented in pre-dawn counts prior to the return of the larvae from the canopy on two separate days. Counts were made with brief use of indirect lighting so as to minimize disturbance to the larvae. To determine if colonies mobilize and depart from their bivouac en masse more rapidly than they recoalesce at dawn, we timed the rate of departure from and arrival to bivouac sites for all three experimental groups. Finally, we ob- served mobilization of two naturally-occurring field colonies at dusk to ensure the behavior we observed in the manipulated groups was consistent with larval be- havior on larger food plants. Late-instar trail following. To supplement obser- vations of putative trail-following behavior, we pre- pared a pheromone extract with hexanes. Five 4th- instar larvae were soaked whole in 10 ml pure hexanes for 24 h. Larvae were then removed and the hexane extract concentrated by evaporation to 2 ml. Extract activity was tested using a Y-maze procedure with young 3rd-instar A. armida (Costa et al. 2003). 50 ul of extract was micropipetted onto the stem and one arm of a Y-maze on an index card, each of which was 4 cm in length; the same quantity of pure hexane was mi- cropipetted onto the alternate arm as a control. Test larvae were allowed to walk up the main stem of the Y-maze and choose an arm in each of 10 trials, each of which used a fresh Y-maze and a new test larva. Larval choices were statistically evaluated using a Chi-square test corrected for continuity (Zar 1999). Costa et al. (2003) suggested the cuticle of A. armida may be impregnated or coated with a trail pheromone, based on the observation that cuticular wipes from the dorsum and venter elicited trail follow- ing. At the end of one week the experimental groups in the present study molted to the final instar, affording an opportunity to test for activity of hexane extracts of the exuviae. Fresh exuviae of 20 caterpillars were col- lected and soaked in about 2 ml hexane for 36 h. We VOLUME 57, NUMBER 3 tested the extract for trail-following activity using the method described above using early 3rd instar A. armida (15 replicates), and analyzed arm choice data with a Chi-square test as above. Testing if late instar larvae employ trail pheromones when foraging proved difficult, as late instars become agitated when handled. To get around this problem we conducted the following in situ trail-following experi- ment: we presented randomly chosen larvae walking ad libitum on their host tree during their mobilization or reaggregation periods at dusk or dawn, respectively, with either 50 ul of pheromone extract or 50 ul of pure hexane, applied by micropipette directly to the tree sur- face at an approximate 45° angle leading away from the larva’s direction of movement. The response of each larva was scored as positive (stopping to sweep and in- vestigate the trail and/or deviating to follow the trail) or negative (no discernable response). We conducted a to- tal of 28 trials over two days (18 treatments and 10 con- trols), and statistically evaluated responses as a 2 x 2 contingency table using Fisher's Exact Test (Zar 1999). RESULTS Group foraging and bivouac use. The groups transplanted on G. ulmifolia and B. quinatum readily assembled into one or more diurnal bivouac. After 24 h a total of 36 of the 40 G1 larvae were recovered; these established 5 initial bivouacs with 3, 9, 6, 7, and 5 larvae, respectively, plus 6 singleton larvae. All 11 G2 larvae were recovered after 24 h, in a single bivouac, but only 22 of 40 initial G3 larvae were recovered after this period, and these occurred in two closely situated bivouacs near the base of the trunk. Many of the larvae not recovered after this initial 24 h period were subse- quently observed to join bivouacs, indicating that they had been undetected, probably in the canopy, and had not left the food plant or succumbed to predation. Daily observations of groups on all three trees re- vealed that larvae remain largely quiescent during the day and do nearly all of their foraging at night. A vari- able number of larvae were found bivouacked each day, ranging from all to about half of accountable lar- vae in a given colony (G1: 94-53%, G2: 100%, G3: 81-67%; Table 1). Most of the remainder were relo- cated resting as singletons or in pairs. In addition, we found that diurnal quiescence may be punctuated with brief periods of activity in which larvae either change resting position or temporarily ascend the trunk to feed. In several cases larvae that became active were observed to return to a resting site within about 30 min, but often not the same site from which they de- parted. These mid-day movement events are evident in a comparison of bivouac censuses taken in the bo bo ie) TABLE 1. Proportion of Arsenura armida caterpillars in study groups occurring in diurnal bivouacs. Number of larvae in a bivouac (%) Day of Total number of observation larvae observed A. G1 on Guazuma ulmifolia 1 35 33 (94) 2 32 29 (91) 3 32 17 (53) 4 24 18 (75) 5 27 14 (52) 6 23 13 (56) B. G2 on Guazuma ulmifolia 1 11 11 (100) 2 11 11 (100) 3 11 11 (100) 4 11 11 (100) 5 JUL 11 (100) C. G3 on Bombacopsis quinatum It 23 18 (78) 2 23 18 (78) 3 16 13 (81) 4 16 13 (81) 5 15 10 (67) morning and afternoon for colony G1 (Table 2), show- ing low levels of diurnal larval movement both be- tween different bivouacs and between bivouacs and the canopy over four consecutive days. On a daily basis we found that most resting larvae occurred in a group, but we consistently observed a subset of caterpillars that rested apart from con- specifics as singletons or doubletons on the host trunk, under a branch, or on foliage. These caterpillars would cycle in and out of groups seemingly at random. For example, we marked with correction fluid four single- ton caterpillars found in the G3 tree canopy, and on four successive days of observation found one, two, or none of these caterpillars had joined the group in the bivouac at the base of the trunk. Similarly, in several instances certain marked G1 and G3 individuals would disappear (presumably remaining high in the canopy where they were missed in our searches), reappearing after one or more days either solitarily or with a group. The number of caterpillars in bivouacs accordingly fluctuates from day to day in our study due to this semi-independence of larvae (see below). As darkness falls, all larvae ascend to the canopy to feed. Grouped larvae do not depart their bivouacs si- multaneously, but mobilize over a period of up to two hours. The first larvae can become active as early as an hour or more prior to sunset, departing their group and ascending to the canopy. Most, however, mobilize dur- ing twilight. In one week of observations we found that most larvae ascend by the end of astronomical twilight (approx. 18:30 h local time through the first half of July at the ACG), with peak departure occurring during twi- light (Fig. 1A). As twilight progresses larvae become in- A. Evening departure from bivouacs JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 17.00 18.00 19.00 20.00 SEE ee (10) Fic. 1. Periods of daily departure from (A) and reassembly at (B) diurnal bivouacs for 5 and 3 groups of larvae, respectively. Each bar cor- responds to a single group of caterpillars at one bivouac site (number of larvae initially or ultimately occupying each bivouac is given in paren- theses), and begins with the departure or arrival of the first caterpillar and ends with the departure or arrival of the last caterpillar at that bivouac. Shading in each graph denotes solar position: stippled = astronomical twilight, black = night, open = day. Observations were made 16-19 July, but times of sunrise and sunset change only slightly from day to day in circumsolsticial weeks. Note that larvae departing bivouacs in the evening become mobilized largely toward the end of twilight, but must begin to leave the canopy prior to morning twilight in order to have been ob- served arriving to bivouacs during twilight. The time taken to return to bivouacs in the morning is far longer than that of departing bivouacs in the evening, presumably because the Trae are returning from widely varying distances in the canopy. creasingly active, and small columns indicative of trail following were often observed when pairs or small groups of larvae chanced to become active at the same time. Night observations of feeding and resting G1 and G2 caterpillars revealed that the caterpillars do not for- age in groups in the canopy. The small size of the G2 host tree made it possible to map the position of each larva, and in three pre-dawn position plots all caterpil- lars were observed actively feeding or resting solitarily. This was consistent with positional plots of larvae we could relocate on the other trees: larvae appear to feed and rest individually in the canopy at night and less fre- quently in loosely associated pairs or small groups. Each morning the foraging caterpillars returned from the canopy and reassembled into bivouacs. Reassembly also takes place largely during twilight, and takes consid- erably longer than the evening departure (Fig. 1B). Our data reveal a stochastic element to larval reassembly de- spite evidence for trail following (discussed below): some bivouac sites were reused on several consecutive days, while others were abandoned after a single use (Table 2). Nonetheless, in our study larvae were more likely to re- assemble at bivouac sites used the previous day (recently occupied bivouacs were reused in 20 of 29 bivouac re- assembly observations of groups G1 and G3; x” = 4.17, p = 0.041). Significantly, although bivouac sites were often reused (Table 3), individual larvae did not exhibit consis- tent bivouac fidelity, but often regrouped each moming with at least some different conspecifics (Fig. 2). TABLE 2. Representative daytime movement observations of bivouacked Arsenura armida caterpillars over 4 consecutive days of observation. 10 July 11 July 12 July 13 July Bivouac! AM? PM AM PM AM PM AM _- PM B 0 0 7 0 9) 2 9 7 C 19 19° 0 0 0) 0 0 0 D 8 8 7 6 9 3 3 3 E 1 0 8 8! 0) 0 3 2 F 6 6 0 0 0) 0 0 0 G "/ Ti he 08s wi5h SOMO H "/ "/ " EM abe. O a a i 1 Subset of total bivouac sites tracked (see Table 3). *Number of caterpillars observed at each bivouac was noted at approximately 10:00 and 14:00 h Local Time each day; instances of change in larval makeup between AM and PM observations are noted i in bold. "/, indicates bivouac was not yet established. 5° Some change i in larval position within bivouac. *One larva departed and one joined. VOLUME 57, NUMBER 3 Bivouac B (9) (0) Bivouac Membership Bivouac D (7) (8) Bivouac Membership Bivouac of Cs (9) (10) (8) DAN RSHCE NA Is} [St Yy, Uj j Z S LS Is IS 1 GZ ' L ] L Y j Day of Observation Fic. 2. Daily composition record for two representative bivouac sites monitored on one tree. All larvae were initially (Day 1) given bivouac- specific markers and the bivouac position was marked with a pin (see methods for details). Note that the subsequent number of caterpillars and their initial bivouac of origin vary greatly over successive days, though bivouac B illustrates repeated reuse by many of the same larvae. Trail following by late-instar foragers. The tendency for bivouac sites to be attractive to larvae on successive days does not necessitate being relo- cated pheromonally, but our observational and exper- imental data confirm that trail markers are used by A. armida when moving between resting and feeding sites. Y-maze bioassays of cuticular hexane extracts made from late instars proved attractive to early in- sams GC = 648 7) = OOD), ancl tin stm Wess OF pheromone extracts indicated that late instars find the cuticular extracts attractive: test larvae responded positively in 16 of 18 treatment (hexane extract) in situ tests, but showed no response in 7 of 10 control (pure hexane) tests, a highly significant result (Fisher's Exact Test, p = 0.0028). Further evidence supporting the idea that the pheromone is in or on the cuticle is provided by the exuvium extract bioas- say results, in which test larvae selected the exuvium extract-treated Y-maze arm in 13 of 15 trials (y? = 8.07; p = 0.005). DISCUSSION Arsenura armida is one of a few social Lepidoptera or Symphyta known to exhibit a late-instar shift in for- aging strategy. Through the 3rd instar A. armida larvae forage as nomadic groups, feeding and resting on the leaves of the food plant at each patch. Our observations and experimental data confirm that this species switches to central place foraging in the 4th (penulti- mate) instar. In switching from nomadic to central place foraging, A. armida also shifts the manner in which its trail pheromone is used. As nomads, the trail pheromone is likely used to promote group cohesion, and as such would be used to mark trails between feed- ing sites. As central place foragers, in contrast, the pheromone conveys information for relocation of a pre- viously-occupied site (or at least return to the vicinity of that site) when forming dawn bivouacs. The dawn reaggregation dynamic has an element of stochasticity stemming from several sources. First, reaggregation is influenced by tree shape. Trees with a single trunk funnel or channel scattered nocturnally foraging larvae to a common location more readily than do trees divided low into multiple trunks (G. ul- mifolia often exhibits a coppice-like multi-trunked growth form in the ACG, while B. quinatum does not). We are confident that the caterpillars do not rely on tree architecture alone to assemble into bivouacs, however, since they often make directed, non-random movements to previously-used sites, many of which are reached following a circuitous path around the trunk. We are also confident that silk plays no role in relocation: as previously documented for early instars, late instar A. armida produce no silk when walking. Larvae use their pheromone to relocate the vicinity of the bivouac at which they rested the day before, but often end up resting at alternative sites. It is likely that tactile contact with conspecifics pre- sents a strong proximate cue to join bivouacs. Return- ing caterpillars encountering groups of bivouacked conspecifics often stopped searching and joined the group even if the bivouac was not the same one these caterpillars occupied previously, though we observed restlessness in some bivouacked larvae, which eventu- ally moved on and joined other bivouacs. Accordingly, bivouac sites or locations were often reused in our study, but the individuals making up the groups as- sembling at those bivouac sites changed to some de- gree each day. Our observations suggest, then, that A. armida’s use of trail pheromone in their daily descent from the canopy is not highly precise, a condition that may arise from the rough texture of the bark substrate JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY of hosts like G. ulmifolia. The combined effect of trail pheromone and tree shape makes relocation of con- specifics likely, however. It should be pointed out that the relatively small size of the groups observed in our study is not unusual: field observations of large intact colonies showed that small “satellite” subgroups of lar- vae regularly form separately from the main colony. We could not mark and track individuals in these colonies due to the size of the tree they occupied, but it is likely that the makeup of the satellite subgroups similarly changed over time judging from observed changes in group size and location. Beyond the shift to nocturnal foraging, A. armida’s temporal shift in foraging behavior is of further interest in that the larvae feed solitarily at night. In early instars the caterpillars are aggregated at all times. Late-instar bivouacked larvae mobilize and ascend to the canopy at roughly the same time and clearly follow chemical trails as they do so. The near simultaneity of mobilization and use of chemical trail markers might suggest that the group remains more or less together in the canopy, but we found no evidence of this. After ascending to a certain point, many larvae appeared to go their own way, and pre-dawn position checks of larvae in one of our study groups showed only solitary feeding. This, too, is likely a stochastic dynamic, and as large colonies move to the canopy to feed it is probable that loose groupings occur, but over the course of feeding through the night the larvae increasingly spread out. A. armida is convergent with Australian Perga sawflies in this general foraging pattern. Perga spp. (Pergidae) rest in diurnal aggregations on the branches or trunk of their Eucalyptus host trees but forage solitarily at night (Evans 1934, Carne 1962). Unlike A. armida, Perga lar- vae are thought to relocate resting groups with acoustic cues generated by tapping the substrate with their scle- rotized anal plate. It remains to be determined if other TABLE3. Reuse of diurnal bivouac sites on Guazuma ulmifolia by Arsenura armida caterpillars on 7 consecutive days of observation. Bivouac site Day. A BY © Di Be Ol) aC ame 1 e e ° e e ay yf ” yf, 2 fest ae e e e GC nf yf, n i. 3 py e Fe e poe’ O ap n i 4 Ss e nite e aS = e a) n ; 5 = e == CO) e = = e n j 6 = e = ck e a = e ae it * e = = e e = site occupied by 22 larvae. site occupied by singleton. site not occupied. »/ = site not yet established. VOLUME 57, NUMBER 3 Neotropical Lepidoptera that rest in aggregations diur- nally and forage nocturnally (for example, the saturniid Dirphia avia (Cramer) or the papilionid Papilio an- chisiades Esper) remain cohesive in the canopy at night or forage solitarily. Why does A. armida change foraging strategy so dramatically in the 4th instar? Caterpillar foraging strategy—encompassing among other things food plant specificity, shelter building, sociality, feeding po- sition and periodicity—is shaped by the joint effects of phylogenetic history, larval nutritional ecology, size or apparency, and defensive ecology (see reviews in Stamp & Casey 1993). Lepidopteran larvae may be ex- pected to experience a shifting milieu of selective pressures associated with these factors as they age, particularly when growing significantly in size and bio- mass, and hence apparency. Accordingly, some species shift defensive strategy over time, and such shifts may be manifested in both coloration and/or behavior (Booth 1990, Montllor & Bernays 1993). For example, larvae of many swallowtails (Papilio and Pterourus spp.) are described as cryptic mimics of bird- droppings in early instars and switch to aposematism or aggressive mimicry in later instars, a coloration change that is not accompanied by a change in forag- ing behavior, while species like Uresiphita reversalis (Guenee) (Pyralidae) and Chlosyne lacinia (Geyer) (Nymphalidae) experience changes in foraging behav- ior as well as coloration over time (Stamp 1977, Bernays & Montllor 1989). Similarly, Cornell et al. (1987, 1988) found that caterpillars of the buckmoth Hemileuca lucina Henry Edwards, another social sat- urniid, exhibit behavioral changes that appear to be driven by predation: buckmoth caterpillars become less aggregative with age as they shift from predomi- nantly defensive to escape behaviors, apparently in re- sponse to changes in predator milieu (biting predators vs. parasitoids). Predation and/or parasitism have presumably played a role in the striking ontogenetic coloration and behav- ioral changes of A. armida caterpillars. This species is avoided by most caterpillar-hunting visual predators in the ACG dry forest, including birds and monkeys, and one of us observed that late instar larvae are lethally toxic to trogon (Trogon elegans Gould) nestlings when swallowed (DHJ unpublished obs.). Moreover, A. armida is attacked by few parasitoids. In several years of mass rearing at the ACG, by far the most abundant parasitoids are the tachinid Winthemia subpicea and, less commonly, the ichneumonid Barylypa broweri (Heinrich) (see Janzen & Hallwachs 2003 for para- sitoid rearing data). The tachinid is known to hunt di- bo iN) ~l urnally, and the ichneumonid is also likely to be diur- nal. Elucidating the present or historical selective pressures favoring A. armida’s foraging strategy is con- tingent on correctly interpreting elements of that strategy. Is, for example, this species hiding, displaying or both when resting in large aggregations on the host trunk? While not cryptic per se, late instars are not as bril- liantly aposematic as they are in early instars (green- yellow soma ringed with black; see Janzen & Hallwachs 2003 for pictures of early instar larvae). Older caterpillars are duskier than early instars, but the intersegmental membrane is colored, giving the appearance of a dark body with narrow orange-yellow rings. In addition, late instars have a chestnut-brown head, a soma covered with fine short setae, and (until the ultimate instar) black tentacle-like protuberances on the dorsum of the thoracic segments. Given their coloration, toxicity, and grouping behavior, it seems most reasonable to conclude that late instar A. armida are making a group display rather than hiding (see Vu- linec 1990 and Bowers 1993 for discussions of apose- matic signaling strategies). This suite of traits may have different effects for different classes of predators, how- ever. Current or past visual predators of A. armida may learn to avoid groups of larvae sporting colored rings, and such predators would be lacking at night when the larvae are active. Coloration is probably less important for diurnal parasitoids than other aspects of grouping. Parasitoids might be better rebuffed by caterpillars in aggregations, and even if parasitoids are unaffected by coordinated group defense (which has not been ob- served in A. armida) or larval toxicity, the caterpillars might benefit from reduced per capita parasitism rates through group dilution effects (Hamilton 1971). It may be impossible to establish whether extant or past predators and parasitoids have selected for the foraging strategy displayed by A. armida. In the con- temporary ecological context these caterpillars are avoided by vertebrate predators and are attacked by two diurnal parasitoids, at least in the Area de Conser- vacion Guanacaste. One approach to test current de- fensive benefits of grouping would be to manipulate groups to document survivorship and parasitism rates as a function of group size. In another type of manipu- lation, larvae in groups of varying size could be con- fined to branches with a barrier that prevents them from moving to aggregation sites on the trunk. In- creased rates of mortality in such groups relative to control groups moving ad libitum would suggest pre- dation pressures currently in existence might help maintain the diurnal trunk-aggregation strategy. It may also be informative to take a phylogenetic view. The genus Arsenura includes about 23 species, and A. armida is the only member of the genus with social larvae for all or nearly all larval instars. Some, like the Brazilian species A. orbignyana (Guérin- Méneville), remain gregarious in early instars and dis- perse afterwards (Furtado 2001a), while most others are solitary in all instars. Solitariness is almost certainly an ancestral behavioral trait in the genus judging by its widespread occurrence in other Arsenurinae, includ- ing Titaea, Loxolomia, Coniopteryx, and Caio (Wolfe & Pescador 1994, Wolfe & Bénéluz 1997, Furtado 1998, 1999, 2001b). Costa et al. (2003) speculated that the pheromone-based trail following behavior demon- strated for A. armida may have its origins in individual trail marking by solitary progenitors. This hypothesis also has relevance for the late-instar foraging shift re- ported here for A. armida, as at least some solitary congeners show parallel ontogenetic changes in forag- ing strategy. For example, A. batesii Druce, which is solitary in all instars, forages diurnally in the canopy in early instars, remaining on foliage, but in the penulti- mate instar reportedly switches to resting on the trunk by day and moving to the canopy at night. The behav- ioral change is accompanied by a color change from mottled brown and green to cryptic brown. If the caterpillars return to the same resting site, they may use a trail pheromone to find their way much like that described for solitary caterpillars of the charaxine but- terfly Polyura pyrrus (Fabricius) (Tsubaki & Kitching 1986). In general this foraging switch is not uncom- mon in Neotropical Lepidoptera (DH] pers. obs.), and may mean A. armida’s shift is an ancestral trait, albeit one that is expressed in a new, social, context. ACKNOWLEDGEMENTS We thank José Manuel Pereira, Robert Pringle, and Ruth Franco for assistance locating and collecting Arsenura colonies. Roger Blanco and Maria Marta Chavarria olhined our work at the re- search station in many ways, and their considerable helpfulness is gratefully acknowledged. Finally, we thank Carla Penz and two anonymous reviewers for their critical comments and suggestions on an earlier draft of the paper. This project was made possible by NSF grants DEB-9024770, DEB-9400829, DEB-9705072 and DEB- 0072730 to DHJ, and an NSF-ROA grant to JTC. LITERATURE CITED BacAzar, M. A. & C. R. BEUTELSPACHER. 2000. Saturniidae (Lep- dioptera), pp. 901— 513. In Llorente Bousquets, J., Gonzalez So- riano, E. & Papavero, N. (eds.), Biodiversidad, Taxonomia y Biogeographia de Artrépodos de México. Hacia una Sintesis de su Conocimiento. Vol. II. Facultad de Ciencias, UNAM, CONABIO & BAYER. Bernays, E. A. & C. B. MONTLLOR. 1989. Aposematism of Ure- siphita reversalis larvae (Pyralidae). J. Lep. Soc. 43:261-273. Boot, C. L. 1990. Evolutionary significance of ontogenetic colour change in animals. Biol. J. Linn. 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Lemaire, Neuilly, France. 199 pp. MOonzrLLor, C. B. & E. A. BERNAyS. 1993. Invertebrate predators and caterpillar foraging, pp. 170-202. In Stamp, N. E. & Casey, T. M. (eds.), Caterpillars: ecological and evolutionary con- straints on foraging. Chapman and Hall, New York. Stamp, N. E. 1977. Aggregation behavior of Chlosyne lacinia larvae (Nymphalidae). J. Lep. Soc. 31:35-40, STamp, N. E. & T. M. Caszy (EDs.). 1993. Caterpillars: ecological and evolutionary constraints on foraging. Chapman and Hall, New York. TsuBAKI, Y. & R. KiTcHING. 1986. Central-place foraging in larvae of the charaxine butterfly, Polyura pyrrhus (L.): a case study in a herbivore. J. Ethol. 4:59-68. bo ts) Ke) VULINEC, K. 1990. Collective security: aggregation by insects as a defense, pp. 251-288. In Evans, D. L. & Schmidt, J. O. (eds.), Insect defenses. State University of New York Press, Albany, New York. Wo FE, K. L. & F. BENELUZ, 1997. Coniopteryx jehovah and its immature stages (Lepidoptera: Satumiidae: Arsenurinae). Trop. Lepid. 8:67—70. WOLFE, K. L. & A. PEscapor. 1994. Caio richardsoni: its imma- ture stages and natural history (Lepidoptera: Satumiidae: Ar- senurinae). Trop. Lepid. 5:29-32. Zar, J. H. 1999. Biostatistical analysis. 4th ed. Prentice-Hall, Up- per Saddle River, New Jersey. Received for publication 29 December 2002; revised and accepted April 13 2008. Journal of the Lepidopterists’ Society 57(3), 2003, 230-234 A NEW SPECIES OF EPIBLEMA (TORTRICIDAE) FORMERLY MISIDENTIFIED AS E. WALSINGHAMI (KEARFOTT) AND E. INFELIX HEINRICH DONALD J. WRIGHT 3349 Morrison Avenue, Cincinnati, Ohio 45220-1430, USA AND CHARLES V. COVELL JR. Department of Biology, University of Louisville, Louisville, Kentucky 40292-0001, USA ABSTRACT. Epiblema gibsoni, new species, is described from 84 adult specimens (69 4, 15 2). This moth is commonly encountered at black light in Ohio and Kentucky during July, especially in habitat supporting prairie vegetation. Its range extends from northwest Arkansas to central Mississippi and western South Carolina, and north to southern Michigan. Epiblema gibsoni is distinguishable from other members of the genus on the basis of forewing maculation, but its similarity to E. walsinghami (Kearfott) and E. infelix Heinrich has caused it to be misiden- tified in the past. Genitalic characters suggest that its closest congener is E. infelix. Additional key words: Olethreutinae, Eucosmini, prairie. Changes since publication of the most recent check list (Powell 1983) bring the current number of North American species of the genus Epiblema (Hiibner) to 41 (Blanchard 1979, 1984, Brown & Powell 1991, Miller 1983, 1985, 1995, Miller & Pogue 1984, Wright 2002). For about half of these species the larvae are known to be late instar stem and root borers in Aster- aceae, usually inducing a conspicuous gall. Recent survey activity in Kentucky (Covell 1999), Ohio, and Illinois generated numerous specimens of a previously unrecognized species of Epiblema, de- scribed below as E. gibsoni, new species. Efforts to identify these specimens led to the discovery of a rather extensive history of misidentification involving E. gibsoni, E. walsinghami (Kearfott), and E. infelix Heinrich. The purpose of this paper is to eliminate the confusion surrounding these taxa and make a name available for the new species. To our knowledge, the earliest literature reference to a specimen of E. gibsoni occurs in the description of Enarmonia walsinghami Kearfott. Kearfott (1907:57) mentioned a series of seven syntypes. Heinrich (1923:150) pointed out that two of those specimens, a male and female from Tryon, North Carolina, were not conspecific with the other five. He designated the fe- male as a paratype of his new species, E. infelix (Hein- rich 1923:151). He also identified the male as infelix but considered it somewhat aberrant and declined to include it among his paratypes. Our examination of the latter specimen revealed it to be E. gibsoni. For reasons explained by Klots (1942:392), it can be difficult to locate Kearfott’s syntypes. Klots (1942:412) listed four specimens in the American Museum of Natural History (AMNH) as belonging to the type se- ries for walsinghami, including a specimen labeled LECTOTYPE, which Klots interpreted as having been designated by Heinrich (1923:151). We examined the walsinghami material at AMNH and the United States National Museum of Natural History (USNM). Based on Klots’ (1942) remarks and the scant data provided by Kearfott (1907), we believe we found six of the seven syntypes. As mentioned above, one is infelix, and one is gibsoni. The other four are listed below under lectotype and paralectotypes for walsinghami. Five of the six specimens bear Kearfott’s handwritten cotype labels. We were unable to locate a syntype mentioned by Kearfott (1907:58) from Essex Co., N. J., dated 4 May. Kearfott is known to have published incorrect dates for some of his syntypes (Klots 1942:392), and this could be one such instance. Otherwise, that spec- imen is probably lost. We also examined the holotype and both paratypes of E. infelix. During this study we frequently encountered speci- mens of infelix and gibsoni that had been misidentified as walsinghami. In particular, the photograph and gen- italia drawings in Miller (1987:58) of walsinghami are actually illustrations of gibsoni. MATERIALS AND METHODS We examined material from the following institu- tional and private collections: AMNH, Canadian Na- tional Collection (CNC), Field Museum of Natural History (FMNH), Loran D. Gibson (LDG), Todd M. Gilligan (TMG), Illinois Natural History Survey (INHS), University of Louisville (UL), Mississippi En- tomological Museum (MEM), Mogens C. Nielsen (MCN), Ohio Lepidopterists (OL), Ron Panzer (RP), USNM, and Donald J. Wright (DJW). Other cited col- lectors are abbreviated as follows: Richard L. Brown (RLB), C. V. Covell Jr. (CVC), John G. Franclemont VOLUME 57, NUMBER 3 Bis sa fp Nias | eRe een EOE Fics. 1-6. 1, E. gibsoni, holotype male, Rowan Co., Kentucky. 2, E. walsinghami, lectotype female, Essex Co., New Jersey. 3, E. gibsoni, female, Adams Co., Ohio. 4, E. gibsoni, male, Adams Co., Ohio. 5, E. gibsoni, male, Cook Co., Illinois. 6, E. infelix, male, Laurel Co., Kentucky. (JGF), J. Richard Heitzman (JRH), Ronald W. Hodges (RWH), Eric H. Metzler (EHM), and Alex K. Wyatt (AKW). Line drawings were made with the aid of a Ken-A-Vision microprojector (Model X1000-1). Forewing length indicates the distance from base to apex, including fringe, and the number of specimens supporting a particular statistic is denoted by (n). Wing pattern terminology follows Brown and Powell (1991). SYSTEMATICS Epiblema walsinghami (Kearfott) (Figs. 2; 7, 8) Enarmonia walsinghami Kearfott 1907:57. Laspeyresia walsinghami; Barnes & McDunnough 1917:174. Epiblema walsinghami; Heinrich 1923:150; McDun- nough 1939:48; Powell 1983:35. Lectotype. °: Essex Co., New Jersey, 30 April 1899, W. D. Kear- fott, AMNH, designated by Heinrich (1923:151). Paralectotypes. NEW JERSEY: Watchung Mts., G. N. [Great Notch], 4 May 1902, W. D. Kearfott (1 3), AMNH; Hmlck Fls [Hemlock Falls], 29 April (1 d, genitalia slide RLB 98), USNM; Hmlck Fls, 29 April (1 2), AMNH. [These three specimens and the lectotype bear Kearfott’s handwritten label “Enarmonia walsing- hami Cotype Kearf.” and his red printed label “TYPE, Collection of W. D. Kearfott”. | Additional material examined. CANADA: Ottawa, 11 June 1907, Arthur Gibson (1 2), CNC. ILLINOIS: Putnam Co., 5 May 1965, M. O. Glenn (1 3), USNM. KENTUCKY: Bullitt Co., 13-17 April 1976, C. V. Covell (1 2; genitalia slide LDG 192), UL. NEW JERSEY: Gt. Notch, 10 May 1914 (1 8, genitalia slide DJW 869), USNM:; Hmlck Fs, 29 April (1 2), USNM; Hmlck Fls, 29 April (1 ¢, genitalia slide DJW 750), FMNH; Hemlock Falls, So. Orange, 19 April 1903, F. E. Watson (1 ¢), AMNH:; Newfndland, 17 May, G. P. Engelhardt (1 2), USNM; Palisades, 25 April 1915 (1 2; genitalia slide RLB 68), USNM; Plainfield, 9 May (1 8, genitalia slide CH 14), USNM. OHIO: Clermont Co., 13 May 1931, Annette Braun (1 °), CNC; Montgomery Co., 21 April 1987, Val Albu (1 ; genitalia slide LDG 184), LDG. PENNSYLVANIA: Oak Station, Alleg. Co., 18 May 1916 (1 4, genitalia slide USNM 70798), USNM. Remarks. Judging from these 17 specimens, forewing macula- tion of E. walsinghami exhibits very little variation. We found the following features most useful for diagnostic purposes: white inter- fascial spot on forewing roughly triangular, its base occupying mid- dle third of dorsum and marked by three to five small, variably ex- pressed, black dashes, its anterior vertex extending toward costa to two-thirds distance from dorsum to costa; basal and subbasal fasciae confluent, forming blackish basal patch, sometimes tinted with dull gray; median fascia represented at costa by blackish transverse bar extending to middle of discal cell and disintegrating into various blackish spots from there to pretornal portion of dorsum; post- median fascia represented by black mark on costa and three to four longitudinal black marks in ocellus; subterminal fascia a narrow black line arising on costa and following terminal margin of ocellus, sometimes broken below costa, often joined to postmedian fascia by variously expressed black mark anterior to ocellus; terminal fascia a short black apical streak; lateral margins of ocellus formed by dull gray transverse bars; another gray bar arising on costa between sub- basal and median fasciae, extending through discal cell along distal margin of interfascial spot. Forewing length: ¢ 6.7-7 mm (mean = 6.9, n = 6), 2 6-7.8 mm (mean = 7.3, n = 11). Male costal fold ex- JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY K — j d Fics. 7-10. Genitalia. 7, Male, E. walsinghami, slide DJW 750 (FMNH). 8, Female, E. walsinghami, slide DJW 869 (USNM). 9, Male, E. gibsoni, slide DJW 636 (DJW). 10, Female, E. gibsoni, right apophysis posterioris omitted for clarity, slide DJW 635 (DJW). Scale bars 0.5 mm. tending from base to 0.5 x length of forewing. Hindwing uniformly blackish brown. Male genitalia (Fig. 7): Uncus a rounded lobe, supported by moderately developed shoulders; socii short and mildly setose; costal margin of valve strongly concave basally, becoming nearly straight distally, ventral margin weakly convex with only slight invagination at neck, apex of cucullus rectangular with rounded corners, ventral an- gle V-shaped, basal margin of cucullus narrowly overlapping neck; clasper situated on upper third of inner margin of saccular opening. Female genitalia (Fig. 8): Papillae annales laterally facing and moderately setose; anterior margin of sterigma circular and collar- like; posterior margin of lamella postvaginalis convex and circular, VOLUME 57, NUMBER 3 with posteriorly directed lateral projections; posterior margin of sternum VII concavely invaginated to 0.3 x length of sterigma, ap- proximate to sterigma medially; ductus bursae constricted below os- tium, widening gradually toward corpus bursae; corpus bursae with two signa arising opposite one another posterior to mid-bursa, one narrow and awlshaped, the other much larger and spadelike. Distribution and biology. The study specimens indicate a geographic distribution from central Illinois east to New Jersey and north to southern Canada. The specimen from Bullitt Co., Kentucky, was taken in a malaise trap; the one from Montgomery Co., Ohio, was captured during the day. The mode of collection of the remaining specimens is not known to us. How- ever, the scarcity of specimens in major collections and the fact that no specimens taken at black light have come to our attention in more than twenty years of field work in Ohio and Kentucky lead us to suspect that walsinghami is either diurnal or not attracted to ultraviolet light. No larval host has been reported. Epiblema infelix Heinrich (Fig. 6) Epiblema infelix Heinrich 1923:151, Fig. 276 (genitalia of d holotype); McDunnough 1939:48; Powell 1983:35. Remarks. This species recently was reviewed by Wright (2002). A photograph of the adult is included here for comparison; illustrations of the genitalia can be found in Wright (2002: Figs. 13, 17) and Heinrich (1923: Fig, 276). Epiblema gibsoni Wright and Covell, new species (Figs. 1,3, 4,5, 9, 10) Epiblema walsinghami (not Kearfott) Miller 1987:58. Diagnosis. The three species under consideration here differ in forewing pattern and coloration. Moder- ately fresh specimens of gibsoni have a lavender cast, whereas infelix and walsinghami have a dull gray to blackish gray appearance. Of the latter two, walsing- hami is the more mottled, and its median fascia is more strongly expressed, particularly at the costa where it is represented by a distinct blackish mark. The shape of the interfascial spot in gibsoni (Figs. 1, 3, 4) is usually diagnostic (see description below). How- ever, our gibsoni sample did include four specimens from the Chicago area in which this feature was greatly reduced (Fig. 5). With regard to genitalic char- acters, gibsoni is distinct from walsinghami in the shape of the valva (Figs. 7, 9), and there are subtle but consistent differences in the shape of the cucullus be- tween gibsoni and infelix. In infelix (Wright 2002: Fig. 13) the inner and outer margins are nearly “parallel”, being concave and convex, respectively, producing a cucullus of roughly constant width. In gibsoni the outer margin is nearly straight and the inner margin is weakly convex toward the apex, causing a gentle taper- ing of the cucullus from the ventral angle to the rounded apex. The female genitalia of gibsoni and in- felix are similar, but the papillae anales of gibsoni tend to be larger and more nodulous than those of infelix (Wright 2002: Fig. 17). The marked difference in size between the two signa in walsinghami easily distin- guishes that species from the other two. The flight pe- riods of walsinghami and gibsoni are essentially non- overlapping: adults of the former species emerge in April and May, those of the latter largely in July. Epi- blema infelix flies from late April to early July. Description. Head: Scales of lower frons white, short and closely appressed, those of upper frons and vertex moderately long, yellowish brown basally, darker brown distally, often with pale laven- der hues; outer surface of labial palpus brown, inner surface cream white to light tan, third segment light tan to brown, often with tan apex; dorsal surface of antenna concolorous with head or slightly darker, ventral surface tan; ventral surface of scape tan. Thorax: Dorsal surface concolorous with head, ventral surface light tan; legs brown outwardly, light tan inwardly, with light tarsal annulations. Forewing (Figs. 1, 3, 4, 5): d length 6-9 mm Gaean = = 7.5,n=77),°@ length 7-9.5 mm (mean = 8.7, n = 13); costa weakly convex, termen straight to weakly concave and perpendicular to costa, tommus gently rounded, male costal fold extending from base to 0.5 x length of forewing. Dorsal surface lavender brown with brown to black mark- ings and an immaculate white dorsal spot between subbasal and me- dian fasciae; basal and subbasal fasciae confluent, forming brown to lavender brown basal patch; median fascia brown, weakly contrast- ing with adjacent lavender brown scaling, narrow and sometimes in- complete near costa, broader and more sharply defined toward dor- sal margin, overlaid with varying amounts of black scaling between interfascial spot and ocellus; postmedian fascia reduced to two to four longitudinal black dashes in ocellus and a brown spot, variably overlaid with black scaling, anterior to ocellus; subterminal and ter- minal fasciae expressed as a narrow brown dash and brown apical spot, respectively; interfascial spot bright white, extending from dor- sal margin to two-thirds distance to costa, sharply defined but vari- able in shape, width increasing gently from dorsum to fold and nar- rowing anteriorly to form a rounded nipple-shaped apex (Fig. 4), or lateral margins parallel and apex rounded (Figs. 1, 3), or, in a few in- stances, spot weakly expressed to nearly obsolescent (Fig. 5); central field of ocellus brown, bordered on lateral and tornal margins with lavender gray; distal half of costa with four, white, paired strigulae: ocellus separated from termen by narrow band of brown scales; scales along terminal edge of membrane gray with pale white apices; fringe gray to lavender oun Hindwing: Uniformly gray brown, fringe scales with pale white apices. Male genitalia (Fig. 9): Uncus a well developed, dorsall y setose lobe, supported laterally by well de- veloped shoulders; socii long, fingerlike, and densely setose; gnathos narrow and bandlike laterally, considerably expanded medially; juxta triangular, caulis short; costal mar gin of valva strongly concave basally, weakly convex toward apex, distal margin weakly convex, ventral invagination narrow and shallow, sealirall angle gently rounded, apex evenly rounded; cucullus narrowing gradually neorrerel apex, its inner surface densely setose; clasper centr ally located on in- ner margin of saccular opening, its basal surfaces supporting nu- merous, short, stout setae; sacculus moderately setose on ventral half of inner surface. Female genitalia (Fig. 10): Papillae anales ventro-laterally facing, nodulous, and strongly setose; anterior mar- gin of sterigma rounded and collarlike; posterior margin of lamella postvaginalis convexly rounded medially and flaring into posteriorly directed projections at the lateral margins; posterior margin of ster- num VII concavely invaginated to 0.5 x length of sterigma, closely approximate to sterigma medially, diverging therefrom laterally; ductus bursae long, constricted below ostium, and mildly sclerotized at juncture with ductus seminalis; corpus bursae with two finlike signa of nearly equal size arising roughly opposite one another and slightly posterior to mid-bursa. Holotype. d: KY: Rowan Co., Rt. 1274, 2 mi. W. Rt. 519, 16 July 1994, leg. L. Gibson; deposited in USNM. Type Locality: 38°06’47’N, 83°25/15”W. Paratypes (n = 83). ARKANSAS: Washington Co., Devil’s Den St. Pk., 29 June 1966, RWH (1 3). ILLINOIS: Cook Co., Gens. Markham Pr., 7 June 1999, RP (1 ¢, genitalia slide DJW 606). IN- DIANA: Hessville, 4 July 1914, AKW (2d); Lake Co., Du Pont Sav., 30 July 2000, RP (1 2), Ivanhoe D & S, 17 June 2000, RP (1 d, 12; 4 genitalia DJW 738, ° genitalia DJW 740). KENTUCKY: Barren Co., Mammoth Cave N. P., Wondering Woods, 26 June 1998, CVC (3 4); Bullitt Co., Pine Creek Forest, 0.5 mi. N of Rt. 480, 4.5 mi. E of 165, 22 July 1989, CVC (14), DJW (14), LDG (34, 19, d genitalia slides LDG 91, 185, 2 genitalia slide LDG 183); Menifee Co., Leather- wood Fork, Indian Creek Rd. 9A, 6 July 1991, LDG (1 4, genitalia slide LDG 194); Owsley Co., 3 mi. NE of Booneville, 20 July 1991, LDG (1 2), 24 July 1982, LDG (2 3, genitalia slide CVC 1197); Rowan Co., E side Rt. 1274, 2 mi. Ww Rt 519, 1 July 1995, LDG (2 3,1), 16 July 1994, LDG (1 4, genitalia slide LDG 195), MICHI- GAN: Monroe Co., T7S R6E Sec 15, 22 July 1988, MCN (1 3). MISSISSIPPI: Oktibbeha Co., 6 mi. SW Starkville, 15 August 1985, RLB (1 2); Winston Co., Noxubee N. W. Refuge, 14 June 1992, T. L. Schiefer (1 ¢). MISSOURI: Randolf Co., Rudolf Bennitt Wildlife Area, 24 July 1971, JRH (2 ¢). NORTH CAROLINA: Macon Co., Highlands, 3865’, 19 July 1958, RWH (1 2), 22 July ee JGF (1d genitalia slide CVC 1196), 23 August 1958. te 3). OHIO: Adams Co., Lynx Prairie Pr., 8 June 1989, DJW (2 4), 1 mi. S.E. of Lynx, 18 June 2002, DJW (5 ¢), 5 Ee 1996, are ae slide DJW 636), 5 July 2002, DIW (7 2), 16 July 1990, DJW (14, 1 Q) fea 1997, DJW (4 3), 29 July ee DJW (12), 3 August 1998, DyW (2 2), 3 August 2000, DIW (4d, 29; d genitalia slide DJW 810, 2 genitalia slide DJW 811), 19 August 1998, DJW (1 8, genitalia slide DI 635); Erie Co., Resthaven Wildlife Area, 13 July oh LDG (2 3, 2 2, d genitalia slide LDG 186), 16 July 1996, DJW (5 ¢, 2 2), 20 July 1990. DJW (2 2), 21 July 1990, TMG (2 2); Lucas cor Kitty Todd Preserve, 8 ae 1996, EHM (2 6). SOUTH CAROLINA: Oconee Co., ae Hill Recreation Area, Rte. 107, 2000’, 7 August 1958, JGF (2 4). Paratype depositories: AMNH, CNC, FMNH, LDG, TMG, INHS, UL, MEM, MCN, OL, USNM, DJW. Etymology. We are pleased to name this species af- ter Loran D. Gibson in recognition of his many contri- butions to the knowledge of Kentucky Lepidoptera. Distribution and biology. Our study sample con- sisted of 151 specimens from Arkansas, Illinois, Indi- ana, Kentucky, Michigan, Mississippi, Missouri, Ohio, North Carolina, and South Carolina. They document a flight period extending from the first week of June to the third week of August, but 70% of the records are from July. No larval host has been recorded. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY ACKNOWLEDGMENTS We thank J. W. Brown, R. L. Brown, P. T. Dang, P. Z. Goldstein, K. R. Methven, and E. L. Quinter for the loan of specimens under their care, G. Derkovitz and K. Gnaedinger for providing specimens from remnant prairie habitat near Chicago, E. H. Metzler for help with the adult photographs, and J. W. Brown and G. J. Balogh for helpful reviews of the manuscript. LITERATURE CITED BarNEs, W. & J. McDuNNouGH. 1917. Checklist of the Lepi- doptera of Boreal America. Herald Press, Decatur, Illinois. 392 Pp: Ces A. 1979. New status for Epiblema minutana (Kear- fott) and new species of Epiblema Hiibner and Sonia Heinrich (Tortricidae). J. Lepid. Soc. 33:179-188. . 1984. Epiblema luctuosana A. Blanchard, a homonym, is changed to Epiblema luctuosissima, new name. J. Lepid. Soc. 38:245. Brown, R. L. & J. A. POWELL. 1991. Description of a new species of Epiblema (Lepidoptera: Tortricidae: Olethreutinae) from coastal redwood forests in California with an analysis of forewing pattern. Pan-Pacif. Entomol. 67:107-114. COVELL, C. V., JR. 1999. The butterflies and moths (Lepidoptera) of Kentucky: an annotated checklist. Kentucky State Nat. Pre- serves Comm. Tech. Ser. 6:1—220. HEINRICH, C. 1923. Revision of the North American moths of the subfamily Eucosminae of the family Olethreutidae. U.S. Nat. Mus. Bull. 123:1—298. KeEARFOTT, W. D. 1907. New North American Tortricidae. Trans. Am. Entomol. Soc. 33:1-98. Kiots, A. B. 1942. Type material of North American microlepi- doptera other than Aegeriidae in The American Museum of Natural History. Bull. Amer. Mus. of Nat. Hist. 79:391-424. McDuNNoucH, J. 1939. Check list of the Lepidoptera of Canada and the United States of America. Part II. Microlepidoptera. Mem. South. Calif. Acad. Sci. 2:3-171. MILLER, W. E. 1983. Genus Phaneta: new synonymies and a new species (Lepidoptera: Tortricidae). Ann. Am. Entomol. Soc. 76:98-103. . 1985. Nearctic Epiblema: a new synonymy, a revised iden- tity, and two new species (Lepidoptera: Tortricidae). Great Lakes Entomol. 18:33-38. . 1987. Guide to the olethreutine moths of midland North America (Tortricidae). U.S.D.A. For. Serv. Agric. Handbook 660:1—104. 1995. Beringian Tortricidae: a new synonymy in Epiblema xn a recount of holarctic species. J. Lepid. Soc. 49:250-252. MILLER, W. E. & M. G. PocuE. 1984. Ragweed borer (Lepi- doptera: Tortricidae: Eucosmini): taxonomic implications of an allometric analysis of adult characters. Ann. Am. Entomol. Soc. Heo —oole POWELL, J. A. 1983. Tortricidae, pp. 31-41. In Hodges, R. W. et al. (eds.), Check list of the Lepidoptera of America north of Mex- ico. E. W. Classey & Wedge Entomol. Res. Foundation, Lon= don. Waricut, D. J. 2002. A new species of Epiblema previously con- fused with E. tripartitana (Zeller) and E. infelix Heinrich (Tor- tricidae). J. Lepid. Soc. 56:277-285. Journal of the Lepidopterists’ Society 57(3), 2003, 235-238 TOUGH AFRICAN MODELS AND WEAK MIMICS: NEW HORIZONS IN THE EVOLUTION OF BAD TASTE P. J. DEVRIES Center for Biodiversity Studies, Milwaukee Public Museum, Milwaukee, Wisconsin 53233, USA ABSTRACT. Mean hindwing toughness was measured experimentally and compared among three sympatric African nymphalid butterflies comprising an aposematic model, its Batesian mimic, and a palatable, non-mimetic relative of the mimic. The unpalatable model species had the toughest wings and palatable species had the weakest. Implications for assessing butterfly palatability and mimicry are discussed in light of previous work, aatale a wing toughness spectrum is proposed as a potential correlate Gi the pe alatability spectrum. Additional key words: butterfly mimicry, Amauris albimaculata, Pseudacraea lucretia, Cymothoe herminia. Insectivorous birds have likely influenced the evolu- tion of butterfly coloration and behaviors by attacking and eating adult butterflies (Poulton 1902, 1908, Car- penter 1932, 1937, 1938, Wourms & Wasserman 1985). Depending on where they fall on the theoreti- cal palatability spectrum, some butterfly species are eaten by birds, while other species are avoided (e.g., Brower 1958a, b, Turner 1984, Turner & Speed 1999). Generally distasteful butterflies minimize predation by advertising noxious qualities with conspicuous color patterns and a slow flight, while palatable ones use cryptic coloration and rapid flight to evade predators (Fisher 1958, Chai 1986, 1996, Chai & Srygley 1990, Pinheiro 1996). Still other palatable butterflies dimin- ish predation by mimicking distasteful species. The el- egance of mimicry stems from the fact that mimics may show strong phenotypic and behavioral resem- blance to their rely: regardless of taxonomic relat- edness among the species involved (Fisher 1958, Turner 1987, Srygley 1994, Joron et al. 2001). The evolution of warning coloration and mimicry re- quires differential survival of some individual butter- flies following attacks and tasting by predators, and that the experience be memorable to predators (Fisher 1958). For example, the bodies of aposematic and unpalatable Danainae are well known to be more resilient to damage from bird attacks than cryptic and palatable Satyrinae (Poulton 1908, Carpenter 1942, Chai 1996, Pinheiro 1996). Here natural selection seems to have favored aposematic phenotypes that are resistant to handling by predators, and at the same time allowed for continued advertising of the unpalat- able phenotypes. In sum, body toughness in butterflies appears to be correlated with unpalatability. Recent experimental work extends our understand- ing of unpalatable traits in butterflies by showing that wings of aposematic African danaine and acraeine species are significantly tougher than those of cryptic, palatable nymphalines and satyrines (DeVries 2002). The study suggested that, in addition to body re- silience, relative wing toughness may be correlated with palatability, and that the spectrum of butterfly wing toughness needs to be documented more broadly, Ac cordingly this report explores palatability and tough- ness in a different light by asking whether African mod- els are tougher than their mimics. To do so differential wing toughness was estimated among three sympatric nymphalid butterflies that represent an unpalatable model, a Batesian mimic, and a palatable, non-mimic. MATERIALS AND METHODS The study was conducted from 12—25 August 2001 in western Uganda at the Kibale Forest field station that forms part of the 766 km? Kibale National Park (0°13" to 0°41’N; 30°19’ to 30°32’E) adjacent to the western arm of Africa’s Rift Valley. The park lies be- tween altitudes 1110 m in the south and 1590 m in the north. Classified as a moist evergreen forest, Kibale Forest has affinities with both montane forest and mixed tropical deciduous forest. The area around the preserve is a matrix of second growth forest, small agricultural plots, associated riparian edges, and has a lone history of various human activities, including long-term studies of forest primates (summarized in Struhsaker 1997). Based on their relative abundance during the study three butterflies were selected to represent palatable or unpalatable species. The trio was formed by a model species, its Batesian mimic, and a cryptic, non- mimetic species that is closely related to the mimic. Palatability and mimetic resemblance were assessed by direct field observations on their color pattern, flight behavior, sympatry, and inference from a de- tailed literature (Marshall 1902, Swynnerton 1915a, b, Carpenter 1941, Brower 1984, Ackery & Vane-Wright 1984, Turner 1984, Ackery 1988, Larsen 1991). These criteria strongly suggested that Amauris albimaculata Butler (Danainae) is an unpalatable model for the pu- tatively palatable Batesian mimic Pseudacraea lucretia Neave (Nymphalinae), and that Cymothoe herminia Grosse-Smith (Nymphalinae) is a palatable, non- mimetic species closely related to P. lucretia. Wing Tear (gms) Amauris albimaculata Cymothoe herminia Pseudacraea lucretia Wing Tear (gms) unpalatable palatable Fic. 1. Box plot comparisons of wing tear weights. Each box spans the first to third quartile and the vertical bars extend to the maximum and minimum values of the sample. Within each box the median is shown by the dashed line, and the mean by the solid line. A, Comparison of wing tear weights for species. Sample sizes are as follows: Amauris albimaculata (N = 10), Pseudacraea lucretia (N = 23) and Cymothoe herminia (N = 14). B, Comparison of wing tear weights of all species grouped by palatable and unpalatable cate- gories. As done in DeVries (2002) an experimental bird-bill was fashioned using a small metal electrical clip with a small plastic weighing dish tied with thread opposite the clip’s jaws (hereafter, the clip assembly). A butterfly was killed by a pinch to the thorax, then immediately secured in the jaws of a wooden clothes peg attached to a rigid wire suspended from the center post below the legs of a photographic tripod. All individuals were se- cured with the wings closed in a natural resting position such that the clothes peg gripped all four wings. The clip assembly was then carefully attached to the hind- wing distal margins of the butterfly such that the jaws gripped the wings between veins Cu, and 2A. This po- sition closely approximates that of beak marks made by birds attacking resting butterflies (e.g., Carpenter 1932, 1937, 1938, 1941, Collenette & Talbot, 1928, PJD pers. obs.). The tripod center post was then raised slowly un- til the weighing dish was freely suspended about 20 mm above a receptacle. Once suspended, tiny ball bearings were slowly added to the dish until the clip as- JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY sembly tore free of the wing, falling into the receptacle below. The tear in the wing closely simulated wing damage inflicted by birds in the wild (DeVries 2002). The clip assembly and ball bearing weights were then weighed to the nearest 0.001 g on a model PB53 Met- tler-Toledo™ electronic balance. This weight estab- lished the force necessary to tear the clip assembly free of the hindwings, and provided a measure of relative wing toughness for each individual specimen. Individual butterflies that had any wing damage or faded wing-patterns due to old age were not used. This avoided potential effects of wing condition on measures of wing-length or relative wing toughness. To estimate body size by species the distance from base to apex of one wing was measured with dial calipers to the nearest 0.1 mm for all individual spec- imens. Differences in wing tear weights and forewing lengths among species were evaluated using a one-way ANOVA. The potential relationship between tear weight and wing length was tested for each species us- ing linear regression. Significance levels for mean wing tear-weight and length in paired comparisons were ad- justed for non-independence using the sequential Bonferroni-Dunn method (Rice 1989). Wing tear weights were evaluated using a one-way ANOVA for model, mimic and non-mimetic species, and for pooled palatable and unpalatable species. RESULTS Mean wing tear weights differed significantly among the individual species (F = 35.523, p < 0.001, df = 2), where A. albimaculata had the toughest wings, P. lu- cretia less tough wings, and C. herminia had the weak- est wings (Fig. 1A). Comparison of species pairs showed significant wing tear weight differences be- tween species (Table 1A). As a group, unpalatable but- terflies had significantly higher wing tear weights than palatable ones (Fig. 1A, B) (F = 51.135, p < 0.0001, df = 1). Tear-weights also differed among species pairs representing model, mimic and non-mimetic butter- flies (Table 1A). Wing lengths differed among species (F = 5.562, p = 0.007, df = 2), between species (Table 1B), and un- palatable butterflies had greater mean wing lengths than palatable ones (F = 5.084, p = 0.029, df = 1). Al- though the largest species, A. albimaculata, had the highest tear weight (Fig. 1A, Table 1), linear regression showed no significant relationship between wing- length and tear weight among species; all probability values were between 0.8580 and 0.4599, and all R? val- ues were between 0.004 and 0.044. VOLUME 57, NUMBER 3 DISCUSSION Butterflies are not discretely palatable or noxious to predators, but rather they encompass a theoretical palatability spectrum (reviewed in Turmer 1984, 1987). The palatability spectrum refers to the relative tasti- ness of potential prey that, depending on the species, is potentially distributed from delicious to positively noxious for particular predators. For example, groups of closely related butterflies (e.g., Danainae, Heliconi- inae) may include species that range from those eaten by birds to those that are always rejected because they possess a nasty taste (Turner 1984, Ritland 1991, Chai 1996, Srygley 1994, Pinheiro 1996). The concept of a palatability spectrum has challenged the traditional separation of Batesian and Miillerian mimicry in but- terflies, and forces us to consider these discrete mimetic categories in a new light (Rothschild 1971, 1981, Huheey 1988, Turner 1984, 1987, Speed & Turner 1999, Turner & Speed 2001, Joron et al. 2001, Mallet 2001). Empirical and theoretical work suggests that un- palatable butterflies should evolve physical attributes making them resistant to handling by predators (e.g., Poulton 1908, Carpenter 1938, 1941, 1942, Fisher 1958). By estimating the force necessary to tear wings this report corroborates the hypothesis that wing toughness may be a correlate of unpalatability in but- terflies (DeVries 2002). Here the aposematic model (A. albimaculata) had significantly tougher wings than its putative Batesian mimic (P. lucretia) and a palatable non-mimic (C. herminia), and that the mimic had sig- nificantly tougher wings than its non-mimetic relative (Fig. 1, Table 1). If predators use wing toughness to help assess butterfly palatability, these observations support the idea that, in addition to sharing behaviors and color patterns with their models, some Batesian mimics may be to some degree unpalatable (e.g., Car- penter & Ford 1933, Rothschild 1971, 1981, Turner 1984, Ritland 1991). Using wing toughness as a metric, the cryptic species, C. herminia, would be the most palatable of the trio examined here. Obviously a larger study comparing many aposematic, mimetic and cryp- tic butterfly species is needed to help reveal evolution- ary correlates and phylogenetic patterns of wing toughness. Nevertheless, in concert with other work (Carpenter 1941, DeVries 2002), the present investi- gation supports the concept of a wing toughness spec- trum that has evolved in parallel with the palatability spectrum. It seems likely that differential wing toughness is correlated with the category and location of damage marks left by predators on the wings of palatable and unpalatable nymphalid butterflies. Because their wings are tougher, beak marks (impressions on the wings) should be observed more frequently among un- palatable species whereas wing tears (areas removed from the wing) should be observed with a higher fre- quency among palatable species than unpalatable ones. This indeed seems to be the case in specimens recovered from nature (e.g., Carpenter 1932, 1937, 1938, 1941, Collenette & Talbot, 1928), and it would be useful to compare predator damage among species that fall along a wing toughness spectrum. Bird attacks are most frequently directed to the hindwing in resting butterflies (Carpenter 1944), and in palatable species distinct patterns at the hindwing margin may function as targets that divert predator attacks away from vital body areas (Blest 1957; Wourms & Wasserman 1985): the attacked butterfly may escape leaving the predator with only a piece of wing. Thus, we might expect to find the location of wing tears to be biased toward the target areas (e.g., eyespots of Satyrinae) in palatable species, and greater variance in location of beak marks in unpalatable species without target areas. As pointed to previously (DeVries 2002), differential wing tough- ness raises the question as to whether hindwing target areas in palatable species are weaker than the wing ar- eas surrounding them. Our understanding of butterfly mimicry has de- pended on continued reassessment of theory in light of empirical observation (e.g., Carpenter & Ford 1933, Fisher 1958, Rothschild 1971, 1981, Benson 1977, Owen 1971, Cuthill & Bennett 1993, DeVries et al. 1999, Joron et al. 1999, Speed & Turner 1999, Turner & Speed 2001). This and a previous study (DeVries 2002) establish a motive for a comparative study on differential wing toughness as an evolutionary corre- TaBLE 1. A, Wing tear differences among species pairs. B, Wing length differences among species pairs. Bonferroni/Dunn compar- isons are significant at p < 0.0167. Abbreviations: * = significant, n.s. = not significant. A Mean wing Critical Comparison tear difference p Significance albimaculata x herminia 24.433 7.299 <0.0001 * albimaculata x lucretia 16.897 6.678 <0.0001 = herminia x lucretia —~7.536 5.976 <0.0030 * B Mean wing Critical Comparison length difference p Significance albimaculata x herminia 3.327 2.562 0.0023 albimaculata x lucretia 1.346 2.343 0.1599 ns. herminia x lucretia —1.981 2.097 0.233 n.S. late among many palatable and distasteful butterflies. They also suggest new ways of assessing the palatabil- ity spectrum among butterflies that have been tradi- tionally considered palatable mimics. Finally, the methods used here provide a means for asking whether model butterflies are tougher than mimics, and if non-mimic butterflies are the weakest of all. By exploring the parallel between the palatability spec- trum and wing toughness we may potentially open new horizons in the evolution of bad taste. ACKNOWLEDGMENTS Iam grateful for the student and faculty interactions on the 2001 Tropical Biology Association course at Kibale, and in particular P. Brakefield, T. Glutton Brock, F. Molleman for field assistance, dis- cussions and enthusiasm. Special thanks are due to Rosie Trevelyan for facilitating my work in Kibale and introducing me to Sparky's place. I thank A. L. Freitas, R. Hill, N. Luebke, R. Lande, N. Mar- tin, C. M. Penz, and J. R. G. Tumer for discussions and comments relevant to this manuscript. This study was supported in part by the National Science Foundation (DEB 980679), and is dedicated to the magnificent body of work left behind by the late Tommy Flanagan. LITERATURE CITED Ackery, P. R. 1988. Hostplants and classification: a review of nymphalid butterflies. Biol. J. Linn. Soc. 33:95-203. ACKERY, P. R. & R. I. VANE-WricuT. 1984. Milkweed butterflies: their cladistics and biology. British Museum (Nat. Hist.), London, BENSON, W. W. 1977. 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GENERAL NOTES Journal of the Lepidopterists’ Society 57(3), 2003, 239-243 NOTES ON LARVAL MANDIBLE MORPHOLOGY OF HYLEPHILA PHYLEUS PHYLEUS (DRURY) (HESPERIIDAE, HESPERIINAE) Additional key words: fiery skipper, grass-specializing feeders, scanning electron microscopy, caterpillars. The present paper is part of a project describing the mandibular morphology of butterfly caterpillars and how it changes among larval instars. The goal is to rec- ognize and better understand the behavior patterns within the four largest butterfly families in the Neotropics—Hesperiidae, Nymphalidae, Lycaenidae and Riodinidae (Heppner 1991, Brown 1996, Robbins & Opler 1997). For butterflies, little is yet known about larval feed- ing strategies and how they evolved, especially in rela- tion to mouthpart morphology and the characteristics of their foods. DeVries et al. (1985) examined variation in the mandibular morphology of some Nymphalidae caterpillars. With regard to the structure of the cutting edge, they recognized two morphological patterns— toothed vs. smooth mandibles. Smooth mandibles were coded as apomorphic, occurring in the subfami- lies Satyrinae, Morphinae, Charaxinae, and Apaturi- nae. Recently, Ackery et al. (1999) suggested that the reduction and loss of larval mandibular teeth could be used to separate the grouping Heteropterinae + Trapezitinae + Hesperiinae from the other Hesperi- idae subfamilies. For moths there is more information available. A number of species of the families Saturniidae, Sphin- gidae, Noctuidae and Notodontidae have been investi- gated (Bernays 1986, Bernays & Janzen 1988, Godfrey et al. 1989, Miller 1991, Dockter 1993, Dewhurst 1999, Passoa & Passoa 2000), and some patterns are hypothesized. Bernays & Janzen (1988), for example, showed two larval feeding strategies in Saturniidae and Sphingidae (snipping vs. chewing, respectively), con- sidering these strategies as adaptive processes corre- lated with both the morphology of the mandibles as well as the physical and chemical features of the larval food plants. As in Nymphalidae (DeVries et al. 1985), smooth mandibles were found to be uncommon and an apomorphic feature in Notodontidae (as mentioned by Miller 1991). From the published information on the mandibular morphology of lepidopterous larvae, smooth mandibles seem to have had independent ori- gins in the evolutionary history of Lepidoptera, as also occurred in Orthoptera (see Tables 2 and 3 in Bernays 1991). The fiery skipper Hylephila phyleus phyleus (Drury, 1773) is acommon species of open areas (Scott 1986). It occurs from Canada to Rio Negro in southern Ar- gentina, and throughout the Greater and Lesser An- tilles (Evans 1955, Hayward 1973, Smith et al. 1994, MacNeill & Herrera 1999). The biology of the imma- ture stages of H. p. phyleus has been described several times since it was recorded as one of the most serious lepidopterous pests of lawn grasses in Hawaii (Kawa- mura & Funasaki 1971, Tashiro & Mitchell 1985, Tashiro 1987, Toliver 1987). However, with regard to the immature morphology of this species, the descrip- tions are not very detailed. The purpose of this paper is to describe the mor- phology of the mandibles and feeding habits of the five larval instars of H. p. phyleus. Ontogenetic changes in the mandibular morphology are documented with the aid of scanning electron microscopy (SEM). Mandibles of grass-feeding specialists (Isely 1944, Godfrey 1972, Brown & Dewhurst 1975, Bernays 1986) as well as of species feeding on other monocotyledonous plants (Peterson 1962, Casagrande 1979, DeVries et al. 1985, Ackery et al. 1999) have been characterized as having chisel-like edges (this is the terminology used by Bernays 1986; other names can be found elsewhere). Specimens used in this study were obtained from eggs (n = 26) laid by a single female collected on 28 March 1999, at noon, in an urban lawn next to the rail- road in the neighborhood of Cristo Rei, Curitiba, Parana State, Brazil (49°16’15”W and 25°25'48"S, ele- vation 900 m). Before netting we observed oviposition behavior of H. p. phyleus for approximately 10 min- utes. Our field observations corroborated the results of Tashiro & Mitchell (1985) who stated that “females [of a Hawaiian population of H. phyleus| alight on the turf for a few seconds for oviposition before flying a short distance to repeat the process”. In the laboratory, the female was confined in a 30 x 30 x 30 cm screen cage, fed 10% honey: water solution, and given fresh grass leaves daily for oviposition. After hatching, larvae were reared individually in plastic containers under green- house conditions with daylight temperatures that fluc- tuated from about 14 to 28°C and relative humidity of 63-88%. As larvae molted head capsules were pre- served in 70% ethanol for future measurements and analyses. The mandibles were dissected following a specific methodology so that the other mouthparts and the head itself were not damaged (Godfrey 1987:551). Left mandible width, here considered the lower edge of the mandible, and head capsule greatest width were measured with an ocular micrometer. These measure- ments are summarized in Table 1. Preparations for SEM analysis followed techniques in Bonatto & Car- valho (1996). Voucher specimens are deposited in the Colecao de Entomologia Padre Jesus Santiago Moure, Departamento de Zoologia, Universidade Federal do Parana, Parana, Brazil. Few conspicuous changes in mandibular morphol- ogy were observed. Mandibles of all instars of H. p. phyleus are relatively short (ca. one-fourth of the head capsule greatest width, Table 1) with a broad base. This overall mandible shape is shared with other taxo- nomically unrelated lepidopterous species that eat ei- ther monocotyledons or dicotyledons with hard or tough leaves (Godfrey 1972, Brown & Dewhurst 1975, Casagrande 1979, Bernays & Janzen 1988, Bernays 1991). The cutting edge is flat and smooth with dis- tinct notches that resemble inter-tooth depressions (Figs. 1-6). In worn mandibles these notches may be blurred or absent as a consequence of the abrasive agent (amorphous silica) deposited in the cell wall and cell lumen of grass leaves (Schoonhoven et al. 1998). Mandibles of the first two larval instars of H. p. phyleus differ from the other instars by the number of setae and absence of a transverse ridge in the oral sur- face (Fig. 2, 3). Mandibular setae are present in all in- stars. In the first and second instars there are only two widely separated setae, with the one closer to the cut- ting edge slightly longer (Fig. 1, 3). Third instar larvae have three mandibular setae, with the longest seta about four times the length of the shortest (Fig. 4). In the last two instars the number and size of mandibular setae varies intraspecifically, but usually with two long setae and 4-6 short setae. In addition, the oral surface is deeply concave in the last two instars, and the trans- verse ridge is well developed (Figs. 5, 6) dividing the oral surface in two portions, the distal portion wider and shorter than the basal one where some pores (pos- sibly glandular openings, see Snodgrass 1935:153-154) occur near the inner margin (Fig. 6). The similarity of mandibular morphology among H. p. phyleus larval instars seems to be associated with the larval feeding strategy that is very similar in all in- stars (Fig. 7). Larvae of H. p. phyleus process the food plant by snipping off pieces of the plant tissue, which are swallowed after a quick mechanical processing by the oral surface of the mandibles. However, we predict that the leaf tissues are not mechanically processed by the first two larval instars due to the simplicity of the mandibular morphology, i.e., there is no transverse ridge nor any undulated area in the oral surface that JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TaBLE 1. Head capsule and left mandible widths (in mm) of fiery skipper larvae from Curitiba, Parana State, Brazil. SD = standard deviation, N = number of specimens. Larval instars 1 } 3 4 5 Head capsule width Mean 0.48 0.68 1.00 1.57 2.24 SD 0.02 0.04 0.09 0.13 0.05 N 4 5 U 4 2) Left mandible width Mean 0.10 0.15 0.23 0.33 0.58 SD 0 0.02 0.03 0.03 0.01 N 4 5 7 4 2 may have a mashing or crushing function, as noticed in some notodontid species by Godfrey et al. (1989). Early instar larvae generally began consuming the edge of softer leaves. Third and subsequent instars readily accepted both young and old grasses. It is pos- sible that younger larvae (first and second instars) of H. p. phyleus may have trouble in processing tougher grass leaves than mature larvae because of the high levels of silica and the arrangement of the lignified veins (Bemays 1986, 1991). The toughness or hardiness in some grasses can be very high. For example, C4 grasses, i.e., species with a photosynthetic pathway pro- ducing a four-carbon acid, are about six times tougher than an average herbaceous plant (Bernays 1991). Whether the presence of smooth mandibles in some lepidopterous larvae, at least during the late instars, is primarily associated with feeding on specific plant taxa, still awaits a thorough examination. DeVries et al. (1985:26) cited cases where the evidence does not sup- port this hypothesis. A modified version of this hy- pothesis is that mandibular adaptations (toothed vs. smooth mandibles) of forb and grass feeders are asso- ciated with plant hardiness or toughness (Bernays & Janzen 1988, Bernays 1991). Curiously, Ackery et al. (1999) have brought this topic for discussion again. While discussing on the monophyly of the grouping Heteropterinae + Trapezitinae + Hesperiinae they suggested that 1) “a reduction and eventual loss of mandibular teeth” in Hesperiidae could 2) “possibly be related to a diet of grasses and other tough mono- cotyledons” (in the original text of Ackery et al. sen- tences | and 2 are reversed). We hope that the present contribution stimulates other researchers to begin ac- cumulating and reviewing as much information as pos- sible on lepidopterous larval morphology, ecology and behavior. We are grateful to George. L. Godfrey and C. F. Dewhurst for providing reprints of their papers on mouthparts of lepidopterous larvae, Daura R. Eiras-Stofella, Matilde Machado and Vera Regina F. Pionteke for helping with the SEM techniques. We also thank VOLUME 57, NUMBER 3 24] Fics. 1-6. Hylephila phyleus phyleus (Drury). 1, anterior view of the head capsule of first instar larva, arrow indicating an inter-tooth like notch; 2, second larval instar, left oral surface; 3, idem, lower outer surface with two mandibular setae; 4, third larval instar, outer surface with three mandibular setae; 5, fifth larval instar, left mandible (oral view), arrow indicating transverse ridge; 6, idem, left mandible (oral view), ar- row indicating “glandular” pores. Ba-mandibular base, La-labrum, Le-mandibular lower edge. Scale bar 100 jm. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fic. 7. Hylephila phyleus phyleus (Drury). First instar larva and characteristic damages in the leaf caused by snipping feeding behavior. Same behavior observed in the other larval instars. Astrid Caldas, Carla Penz, Danuncia Urban, Gerardo Lamas, Robert K. Robbins, and two anonymous reviewers for improving our manuscript. 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CASAGRANDE, Departamento de Zoologia, Univer- sidade Federal do Paranda, CP 19020, Curitiba, Parana 81531-990, Brazil; Email: lycaenidae @ig.com.br Received for publication 8 January 2001; revised and accepted 30 January 2003. NOTES ON THE HISTORIC RANGE AND NATURAL HISTORY OF ANAEA TROGLODYTA FLORIDALIS (NYMPHALIDAE) Additional key words: Croton, Florida, West Indies, seasonal forms, parasitism. Populations of the Florida leafwing, Anaea troglodyta floridalis F. (Comstock & Johnson) (Fig. 1), a butterfly endemic to south Florida and the lower Florida Keys, have become increasingly localized as its pine rockland habitat is lost or altered through an- thropogenic activity (Baggett 1982, Hennessey & Habeck 1991, Schwarz et al. 1995, Salvato 1999, 2001). Croton linearis Jacq., (Euphorbiaceae) a sub- tropical species of Antillean origin, is the sole host plant for A. ¢. floridalis (Opler & Krizek 1984, Schwartz 1987, Minno & Emmel 1993, Smith et al. 1994). Once common throughout the pinelands of the lower Florida Keys (Dickson 1955), C. linearis now oc- curs only on Big Pine Key (Monroe Co.) and in frag- mented populations on the southeast Florida mainland as far north as Jupiter Island (Martin Co.) (Salvato 1999). However, as host plant availability and appro- priate habitat have declined, there is little recent evi- dence that A. t. floridalis ventures further north than southern Miami (Miami-Dade Co.) to make use of these fragmented host populations (Baggett 1982, Smith et al. 1994, Salvato 1999). Salvato (1999) has found few-documented field sighting records or mu- seum collection specimens of A. t. floridalis from areas north of Monroe and Miami-Dade counties suggesting that this species may not have been common further north historically. Delineating the precise historic range of A. t. flori- dalis has been further complicated by its confusion with Florida’s other resident Anaea species, Anaea an- dria Scudder (Opler & Krizek 1984, Hennessey & Habeck 1991). An extremely tolerant species climati- cally, A. andria is widely distributed in the United States and Mexico (Pyle 1981, Opler & Krizek 1984). In Florida, Hernando County appears to represent the southern boundary for A. andria and this may corre- spond with the distribution of its host plants (Salvato 1999). Anaea andria uses several different Croton host species throughout its range, as opposed to A. t. flori- dalis which is stenophagic and will only use Croton lin- earis (Opler & Krizek 1984, Schwartz 1987, Hen- nessey & Habeck 1991, Smith et al. 1994, Worth et al. 1996). In northern Florida, A. andria primarily uses Croton argyranthemus Michx. (Glassberg et al. 2000) as a host, but will also feed on C. capitatus Michx (Opler & Krizek 1984, Salvato 1999). Salvato (1999), in preliminary feeding studies, found that when of- fered a variety of Croton species (C. capitatus, C. lin- earis and C. argyranthemus), A. t. floridalis larvae (n = 5) would only accept C. linearis as a food source. Anaea andria larvae (n = 5), when given the same se- lection, preferred C. argyranthemus as well as C. cap- itatus but refused to feed on C. linearis. The prefer- ence of A. andria for only northern occurring Croton species may explain why the butterfly has not estab- lished itself farther southward in the state. The appar- ently strict diet requirements of A. t. floridalis and pos- sibly an inability to tolerate the colder winter climate of north Florida keep it from expanding northward. Croton grandulosus Michx. is the prevalent Croton species in the central part of Florida where neither butterfly occurs. Both Anaea species refused this plant as a host when offered it in feeding trials. Salvato is currently conducting continued feeding studies with A. andria and A. t. floridalis to establish larger sam- pling sizes. However, it does appear that an allopatric relationship occurs between A. andria and A. t. flori- dalis within Florida, one similar to that observed be- tween other members of the genus within the West In- dies (Smith et al. 1994). Figure 2 indicates the documented distribution of A. t. floridalis and A. an- dria in Florida. Anaea t. floridalis maintains an appearance charac- teristic of the genus and the taxonomy of this sub- JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fic. 1. The Florida leafwing, Anaea troglodyta floridalis 4 January, 2003 in Long Pine Key (Everglades National Park), Florida (photo H. L. Salvato). species has been well described elsewhere (Comstock 1961, Baggett 1982, Opler & Krizek 1984, Smith et al. 1994, Worth et al. 1996). Briefly, its upperwing surface is red to red-brown, the underside gray, with a tapered outline, cryptically looking like a dead leaf when the butterfly is at rest. Anaea t. floridalis exhibits sexual di- morphism, with females being slightly larger and with darker coloring along the wing margins than the males (Fig. 3). The species also appears to demonstrate seasonal polymorphism (Fig. 3). Comstock (1961) employed the terms “summer” and “winter” morph to differenti- ate between seasonal forms within the genus. Riley (1980, 1988a, b) found that the length of photoperiod exposure experienced by fifth-instar larvae (several days prior to pupation) as well as the influence of sea- sonal moisture, were key factors in determining the seasonal forms of A. andria. The summer Anaea form, (wet-season or long-day form) (late May to Septem- ber), of the genus tends to have forewing margins which are blunt and a hindwing with a less pro- nounced tail; their colors also tends to be brighter. The winter Anaea form, (dry-season or short day form), (October to early May) tends to have the opposing characters, these being pronounced tails and crescent- shaped forewings. VOLUME 57, NUMBER 3 @ Anaea andna © Anaea troglodyta floridalis Pokk Osceola Hileborough / | Inckan River ‘ Hardee Marge St Lucie Sarasota eee Marin PREM. Nee Hendry Paim Lee ((_)) Fan ciliata ad - ecole — ee i X Fac ess he ait hier! \ jeratilimbed ss ae Peo t y ae Pom a pee ame Fic. 2. Historic distribution of Anaea species, by county in Florida. Distribution based on verified records of specimens collected or pho- tographed for each county (J. V. Calhoun pers. com.). Although a great deal of research has been con- ducted to explain opposing wing characteristics of sea- sonal forms and how they are cued (Comstock 1961, Riley 1980, 1988a, b), more research is needed to un- derstand what implications this change in wing shape has on Anaea biology. One possibility is that the change in wing shape is an adaptation by Anaea to more cryptically blend into its surroundings during given seasons. Muyshondt (1974a, b) indicated that Anaea (Consul) fabius Cramer and A. (Memphis) eu- rypyle confusa Hall appear very inconspicuous amongst vegetation and that these species alight on tree trucks in a slanted position to minimize the shadow they project. Similar behavior was observed with A. t. floridalis while conducting mark-release-re- capture field studies from 14 July 1997-29 August 1998. When at rest on the sides of slash pines, A. f. floridalis adults would angle their bodies, with wings closed, in such a way that it seemed to mimic the raised and peeling bark of the pine trees. Anaea t. floridalis caught during the winter/spring months (October to early May) of the 1997-95 study (n = 46), always maintained well-developed hindwing tails and anal angle projections, as well as forewings JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fic. 3. Demonstration of seasonal and sexual dimorphism in Anaea t. floridalis. Males (on the left), females (right). Butterflies on the top row are the winter-morph, those on the bottom, the summer-morphs (photo M. H. Salvato). with an acute and falcate apex. Likewise, those marked in the summer/fall months (late May to September) (n = 85), possessed shortened tails on the hindwing, reduced anal angle projections, and forewings that were not apically falcate. Several larvae (n = 15) were reared from field-collected specimens in the winter months (January—March) of 1999. These all produced the winter form. Table 1 indicates the seasonal forms observed in the field-marked A. ¢. floridalis in 1997-98. Further field studies are required to deter- mine the precise periods of change from winter to summer-morph (and from summer to winter-morph). An abrupt change from winter to summer-morph indi- cated in field-captured specimens in April and May 1998 suggests that this is the period of change to the summer-morph for A. ¢. floridalis. There was no evi- dence of intermediate forms between the seasonal morph types. Field-marked adults and museum exam- ined specimens showed characters that were distinctly one of the two-morph patterns. Whether the bi-annual change in wing shape is an adaptive response that pro- duces appropriate seasonal camouflage and/or aerody- namic advantages to flight remains an interesting topic for future study and discussion. Behavior and life cycle observations documented during this study are consistent with what has been re- ported previously for this subspecies (Baggett 1982, Opler & Krizek 1984, Schwartz 1987, Smith et al. 1994, Worth et al.1996). The adults are rapid, wary fliers. The species is extremely territorial, with both sexes flying out to pursue other butterflies (Baggett 1982, Worth et al. 1996). The occurrence of adults consistently perching on the same spot, on a tree or sign post, as well as using the same specific host speci- men for oviposition, suggests these areas are continu- ally suitable and recognized. This behavior was partic- TABLE 1. Monthly overview of seasonal wing pattems observed in marked and released Anaea troglodyta floridalis between 14 July 1997 and 29 August 1998 on Big Pine Key and Long Pine Key, Florida. Winter- Summer- Month n morph morph January 11 11 0) February 0 7 0 March 2 2 0 April 14 14 0 May 12 2 10 June 18 0) 18 July 17 0 17 August 29 0 29 September 2 (0) 2 October 2 0 2 November 7 10 ii December 0 0 0 VOLUME 57, NUMBER 3 ularly well observed (8 occurrences on different survey dates) in the Watson’s Hammock area of Big Pine Key during 1997-98. Anaea t. floridalis is multivoltine, with an entire life cycle of about 60 days (Hennessey & Habeck 1991), and maintains continuous broods in south Florida throughout the year (Salvato 1999). Pre- cise number of broods per year remains unknown, but A. t. floridalis has been recorded in every month (Baggett 1982, Opler & Krizek 1984, Minno & Emmel 1993, Salvato 1999) in south Florida. Males, especially those newly emerged, were frequently flushed from their perches in response to a fluorescent-colored cloth, either by waving it the air or simply placing it in a shirt pocket (Salvato 1999, Salvato 2003). Females lay eggs singly on both the upper and lower surface of the host leaves, normally on developing terminals (Baggett 1982, Hennessey & Habeck 1991, Worth et al. 1996, Salvato 1999). Eggs are spherical and light cream-yellow in color (Worth et al. 1996). Worth et al.(1996) and Salvato (1999) visually estimated that fe- males may fly more than 30 meters in search of a suit- able host and usually requires less than a minute to oviposit each egg. During egg surveys conducted in 1988-89 in both Everglades National Park and Big Pine Key, egg den- sity was approximately 11-66 per ha on sparse patches of host plants scattered throughout the pine rocklands (based on an estimated 80 ha of Croton-bearing habi- tat on Big Pine and 1068 ha in the Everglades) (Hen- nessey & Habeck 1991). Eggs of many Neotropical charaxine species similar to Anaea, such as Memphis Hubner and Consul Hubner are heavily parasitized by chalcid wasps (Muyshondt 1974a, b, 1975a, b, 1976a, b, DeVries 1987). Within the pine rocklands A. ¢. flori- dalis eggs experience a high level of parasitism from trichogrammid wasps (Hymenoptera: Trichogrammi- dae). Once attacked by the wasps, the Anaea eggs turn black (Muyshondt 1975b, Hennessey & Habeck 1991, Salvato 1999). The frequency of these “black eggs” was noted to be as high as 100% in 1988-89 surveys for A. t. floridalis eggs on host terminals both in the Ever- glades National Park and at Watson’s Hammock on Big Pine Key (Hennessey & Habeck 1991). Trichogramma sp. near pretiosum Riley “Naranja species” was identi- fied as the parasitoid and appears to be a key mortality factor for A. t. floridalis (Hennessey & Habeck 1991, Salvato 1999). Hennessey & Habeck (1991) found the larval hatch rate in the field for all survey areas during their 1988-89 studies, including all mortality sources, ranged from 0-33%, depending on location and year. On two occasions (6 July 1988 and 10 October 1989) the mite Balaustium sp. (Acari: Erythraeidae) was ob- served preying upon eggs of A. t. floridalis within the Everglades (Hennessey & Habeck 1991). Crab spiders (Aranea: Thomisidae) were frequently observed in 1988-89 and 1997-98 surveys on C. linearis and may prey upon eggs of A. t. floridalis as well as the Bartram’s hairstreak, Strymon acis bartrami Comstock and Hunt- ington (Lycaenidae). Matteson (1930) recorded ants as predators of A. t. floridalis eggs in Miami. Because the host is dioecious, sex of the plant was noted when eggs were marked (by placing flagging tape on the plant) in order to determine whether there was an oviposition preference by the females (Hen- nessey & Habeck 1991). Of 31 plants recorded with eggs between 10 March and 5 July 1989, 14 (45%) were male plants and 17 (55%) were female plants. Female A. t. floridalis showed little preference for fe- male over male plants as oviposition sites (Hennessey & Habeck 1991). However, further studies are re- quired to determine if there is any preference for host plant sex in A. t. floridalis oviposition behavior. The natural history of the larval stages of A. t. flori- dalis is well described elsewhere (Baggett 1982, Opler & Krizek 1984, Schwartz 1987: Smith et al. 1994, Worth et al. 1996, Salvato in press). Unlike other members of Anaea and similar genus such as Memphis (Muyshondt 1974b, 1975a, b, DeVries 1987) and Con- sul (Muyshondt 1974a, DeVries 1987), larvae of A. t. floridalis do not make frass chains or roll plant leaves into tubes to evade parasites and predators. Caldas (1996) found fifth instar larval parasitism by tachinid flies to be as high as 53% for Anaea (Memphis) ryphea Cramer. Muyshondt (1974b) estimated larval mortality from tachinid flies to be 40% for A. (M.) e. confusa. Ta- chinid flies were noted as a principle mortality factor for A. (C.) fabius (Muyshondt 1974a). DeVries (1987) indicated that larvae of Anaea aidea (Guerin- Meneville) experience parasitism from tachinid flies as well as chalcid wasps. Tachinid flies appear to be a par- asitoid on the larval stages of A. t. floridalis, laying their eggs on the host plant, which are subsequently ingested. Hennessey & Habeck (1991) collected a moribund fifth-instar A. ¢. floridalis larva at Long Pine Key (Everglades) on 14 November 1988. The speci- men was host to four larvae of Chetogena sp. (Diptera: Tachinidae) that emerged from it in the laboratory; these larvae pupated and became adults. Muyshondt (1975b) obtained a large tachinid species (Archytas sp.) from the pupa of Anaea (Memphis) pithyusa R. Felder. Hennessey & Habeck (1991) encountered an A. t. floridalis pupa on Big Pine Key that was in the process of being consumed by ants (species not speci- fied). Muyshondt (1975a) suspected heavy predation on larvae of Anaea (Memphis) morvus boisduvali Comstock from spiders after witnessing spiders in the proximity of leaves where larvae had been feeding. Spiders appear to be a predator on the adult A. t. flori- dalis as indicated from a photograph in Glassberg et al. (2000) of a lynx spider (Aranea: Oxyopidae) with a cap- tured adult. However, Rutkowski (1971) watched a spider (species not specified) quickly release an adult A. t. floridalis from its web after an initial taste. This suggests A. t. floridalis may be chemically protected from certain predatory species. Adults are not frequently attracted to flowers (Baggett 1982, Opler & Krizek 1984, Worth et al.1996) but have been observed feeding on rotting fruit and dung (Baggett 1982, Opler & Krizek 1984, Minno & Emmel 1993). DeVries (1987) reported that both sexes of A. aidea feed on rotting fruits and dung, while males would engage in puddling. Hennessey & Habeck (1991) ob- served an adult feeding at senescent flowers of saw pal- metto, Serenoa repens Bartr. alongside scarab beetles (Coleoptera: Scarabaeidae) in Watson’s Hammock dur- ing 1988. A sliced orange placed at one of the survey transects in the early evening provided the only observa- tion (August 1998) of feeding by adults during 1997-98 field studies (Salvato 1999). Although the species is known to be easily captured in bait traps (Smith et al. 1994), such traps set out at several locations failed to at- tract any A. t. floridalis during the 1997-98 field study. Lenczewski (1980) observed A. t. floridalis (sexes not specified) at the edges of mud puddles in the Ever- glades. Puddling behavior was also observed on 6 occa- sions during 1997-98, by males on Big Pine Key and in the Everglades. Adults reared and kept in captivity also did not feed on provided flowering plants, but frequently fed on artificial sources provided (especially beer). The authors thank T. C. Emmel, D. H. Habeck, J. H Frank, D. M. Griffin III and J. C Daniels for advice and encouragement while conducting these studies. J. V. Calhoun critically reviewed this manuscript and provided range map information for both A. t. floridalis and A. andria. D. Serage (Sanibel-Captiva Conservation Foundation) for rearing and mating facilities, R. M. Baranowski for providing laboratory facilities, J. Y Miller and L.D. Miller for ac- cess and examination of specimens at the Allyn Museum of Ento- mology (Sarasota, Florida) and J.B. Heppner for similar examina- tion of specimens at the Florida State Collection of Arthropods (Gainesville, Florida), J. Bayless for consultations on the collec- tions at ENP, J. Gilmore for field assistance, C. Campbell and A. Williams for plant identification. The authors thank the following individuals for their identification of various arthropod species; N. E. Woodley USDA-SEL (Tachinidae); W. C. Welbourne of the Florida State Collection of Arthropods (Erythraeidae); J. D. Pinto of the University of Southern California-Riverside (Trichogrammi- dae). The authors thank K. A. Schwarz and R. A. Worth for their encouragement, suggestions and insights on this research, and the natural history of these organisms. The authors would also like to thank the staff of National Key Deer Wildlife Refuge, particularly B. Stieglitz, J. Watson, D. Holle, T. Wilmers and B. Frakes, D. Gordon, M. Folk and L. Flynn of The Nature Conservancy, and Everglades National Park for permitting and various technical sup- port. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY LITERATURE CITED BaccETT, H. D. 1982. Order Lepidoptera. In Franz, R. (ed.), In- vertebrates. In Pritchard, P. C. (ed.), Rare and endangered biota of Florida. Vol. 6. Invertebrates, 78-81. Univ. Pr. Florida, Gainesville. 131 pp. Catpas, A. 1996. Fifth instar parasitoids of Anaea ryphea (Nyphal- idae): the missing link. J. Lepid. Soc. 50:89-90. Comstock, W. P. 1961. Butterflies of the American tropics. The genus Anaea, Lepidoptera:Nymphalidae. Amer. Mus. Nat. Hist., New York. 214 pp. DEVRIES, P. J. 1987. The butterflies of Costa Rica and their natural history: Papillionidae, Pieridae, Nymphalidae. Princeton Uni- versity Press, Princeton, New Jersey. 327 pp. Dickson III, J. D. 1955. An ecological study of the Key Deer. Florida Game and Freshwater Fish Commission. Pittmann- Robertson Project Tech. Bull. 3. 104 pp. GLASSBERG, J., M. C. MINNO & J. V. CALHOUN. 2000. Butterflies though binoculars: Florida. Oxford University Press, New York. 242 pp. Henney M. K. & D. H. Hasecx. 1991. Effects of mosquito adulticides on populations of non-target terrestrial arthropods in the Florida Keys. Gainesville: U.S. Fish and Wildlife Service and the Univ. of Florida Cooperative Wildlife Research Unit (unpublished, available from Fish and Wildlife Reference Ser- vice http://fa.r9.fws.gov/r9fwrs/) LENCZEWSKI, B. 1980. Butterflies of Everglades National Park. Homestead: S. Fla. Res. Ctr. Rep. T-588, Natl. Park Serv., Ever- glades Natl. Park. 110 pp. MATTESON, J. H. 1930. Anaea portia—the leaf-wing and a list of the Rhopalocera of Miami, FI. Privately printed. 16 pp. MINNO, M. C. & T. C. EMMEL. 1993. Butterflies of the Florida Keys. Scientific Publ., Gainesville. 168 pp. MuysHonpt, A. 1974a. Notes of the life cycle and natural history of butterflies of El Salvador. II. Anaea (Consul) fabius Nymphalidae). J. Lepid. Soc. 28 (2):81-89. . 1974b. Notes of the life cycle and natural history of but- terflies of El Salvador. IV. Anaea (Memphis) eurypule confusa Nymphalidae). J. Lepid. Soc. 28 (4):306-314. . 1975a. Notes of the life cycle and natural history of but- terflies of El Salvador. V. Anaea (Memphis) morvus boisduvali Nymphalidae). J. Lepid. Soc. 29 (1):32-39. . 1975b. Notes of the life cycle and natural history of but- terflies of El Salvador. VI. Anaea (Memphis) pithyusa Nymphalidae). J. Lepid. Soc. 29 (3):168-176. . 1976a. Notes of the life cycle and natural history of but- terflies of El Salvador. VII. Archaeoprepona demophon cen- tralis (Nymphalidae). J. Lepid. Soc. 30 (1):23-32. . 1976b. Notes of the life cycle and natural history of but- terflies of El Salvador. VIII. Archaeoprepona antimache gulina, Siderone marthesia, Zaretis callidryas and Consul electra (Nymphalidae). J. Lepid. Soc. 30 (3):159-168. Op_eR, P. A. & G. O. KrizEK. 1984. Butterflies east of the Great Plains. J. Hopkins Univ. Pr., Baltimore. 294 pp. PytrE, R. M. 1981. The Audubon Society field guide series to North American butterflies. Alfred A. Knopf, New York. 916 pp. Riwey, T. J. 1980. Effects of long and short day photoperiods on the seasonal dimorphism of Anaea andria (Nymphalidae) from cen- tral Missouri. J. Lepid. Soc. 34 (4):330-337. . 1988. Effect of photoperiod on incidence of adult seasonal forms in Anaea andria (Lepidoptera: Nymphalidae). J. Kansas Entomol. Soc. 61 (2):224—227. . 1988. Effect of larval photoperiod on mating and repro- ductive diapause in seasonal forms of Anaea andria (Nymphal- idae). J. Lepid. Soc. 42 (4):263-268. RurKowskI, F. 1971. Notes on some South Florida Lepidoptera. J. Lepid. Soe. 25 (2):137-139. SatvaTo, M. H. 1999. Factors influencing the declining populations of three threatened butterflies in south Florida and the Florida Keys. M.S. Thesis, Univ. Florida. — — — = tice pi cic ae a wre =e =} = mn VOLUME 57, NUMBER 3 . 2001. Influence mosquito control chemicals on butterflies (Nymphalidae, Lycaenidae, Hesperiidae) of the lower Florida Keys. J. Lepid. Soc. 55 (1):8—14. . Inpress. Butterfly conservation and hostplant fluctuations: the relationship between Strymon acis bartrami and Anaea troglodyta floridalis on Croton linearis in Florida. Hol. Lepid. § (2):53-57. . 2003 Lifestyles of the scaled and beautiful: the Florida leafwing. Amer. Butterflies. 11 (1):36-41. ScHwartTz, A. 1987. The butterflies of the Lower Florida Keys. Milwaukee Pub. Mus. Contrib. in Bio. and Geo., No. 73. 34 pp. SCHWARZ, K. A., R. A. WorRTH & T. C. EMMEL. 1995. Conservation of two threatened south Florida butterflies and their host plant (Lepidoptera: Lycaenidae, Nymphalidae). Hol. Lepid. 3:59-61. SmiTH, D. S., L. D. MILLER & J. Y. MiLLer. 1994. The butterflies of the West Indies and South Florida. Oxford Univ. Pr., Oxford. 264 pp., 32 pl. Journal of the Lepidopterists’ Society 57(3), 2003, 249-250 249 Worth, R. A., K. A. SCHWARZ & T. C. EMMEL. 1996. Notes on the biology of Strymon acis bartrami and Anaea troglodyta flori- dalis in south Florida. Hol. Lepid. 3:52-65. Mark H. SAtvato, University of Florida, Depart- ment of Entomology and Nematology, P.O. Box 110620, Gainesville, Florida 32611, USA; Email: anaea_99@yahoo.com AND MICHAEL K. HENNESSEY, United States Department of Agriculture, APHIS, PPQ, Center for Plant Health Science and Technology, Raleigh, North Carolina 27606-5202, USA; Email: mike.k. hennessey @aphis.usda.gov Received for publication 20 September 2002; revised and accepted 18 April 2003. EDITH’S COPPER, LYCAENA EDITHA (LYCAENIDAE), CONFIRMED FOR CANADA Additional key words: Thomas Baird, Alberta. The status of Edith’s Copper, Lycaena editha (Mead), in Canada has been a matter of conjecture for some time, particularly in Alberta. Bowman (1934, 1951) included this species in his annotated lists of Al- berta Lepidoptera, giving High River as the locality without further comment. This represented the only known Canadian record of Edith’s Copper; its known range is restricted to the western US, from California to Montana eastward to Wyoming and Colorado (Scott 1986). Subsequent works (e.g., Ferris & Brown 1981, Scott 1986) also indicated this species as part of the Al- berta fauna, presumably based on Bowman's list. Bird et al. (1995) were unable to authenticate this record and rejected it. Layberry et al. (1998) also treated this as a dubious record, and did not include L. editha as part of the Canadian fauna. While curating the butterflies in the University of Alberta Strickland Museum collection in 2001, BCS discovered the putative High River specimen in a sep- arate teaching collection, where it had gone unnoticed these many years. It is a male specimen, missing the left antenna but otherwise in excellent condition, with a label reading “High River, Alta / Baird” (Fig. 1). “High River” and “Baird” are handwritten on a printed Donald Mackie label, and “Edmonton” and “D. Mackie” are crossed out (Fig. 1). Comparison of the handwriting to other Donald Mackie labels shows that the specimen was labelled and likely pinned by Mackie after he received the unpinned specimen from Baird. A small amount of glue is visible on the ventral thorax and on the pin, further suggesting that the specimen was not pinned fresh. Donald Mackie made extensive Lepidoptera collections, primarily from the Edmonton region, in the early to mid-1920’s, and the specimen was likely either sent or given to him by Baird. Thomas Baird came to High River from Woodstock, Ontario in about 1896, and worked there for many years as a cob- bler. He was an ardent and versatile collector of all groups of insects, though he appears to have been par- ticularly partial to Diptera. F. H. W. Dod, in his series of “Further notes on Alberta Lepidoptera” (Dod 1914, 1915a, b) made frequent reference to Baird's collec- tions. Among the moths that Baird collected, espe- cially at light, were a number of taxa that were new to science. The precise location where the High River speci- men was collected is impossible to determine, but there is no reason to believe it was not collected in the general vicinity of the town of High River (50°35/N, 113°52’W). Suitable Canadian Zone valley bottom wet meadow habitat that L. editha is reported to frequent (Scott 1986) occurs in the Rocky Mountain foothills west of High River, and it is entirely possible that the specimen originated there. Other butterfly species col- lected by Baird and labeled as “High River” are re- stricted to montane habitats rather than the prairie habitat found at High River, suggesting Baird named his collection localities to the nearest major settle- ment, as did many early collectors. Although it is possible that this specimen is misla- beled, there is no evidence to suggest this. Further- more, there are no accounts of, or insect specimens Teen we. ALTA, (CH RAVER BAIRD |a-Maceir Fic. 1. Specimen representing the only comfirmed Canadian record of Lycaena editha (Mead) (ventral view). collected by, either Baird or Mackie to suggest they collected in the western U.S. (within the main range of editha) or exchanged specimens with other collectors. Lycaena editha is present in Glacier Co., Montana just south of Waterton National Park (Ferris & Brown 1981) and attempts to locate populations of this species in the province should be concentrated in the Waterton to Crowsnest area (Bird et al. 1995) in July and August, during L. editha’s flight period (Scott 1986). Since there is no evidence to suggest that this specimen was not collected in the vicinity of High River, Alberta, Edith’s Copper should be added to JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY both the Alberta and Canadian butterfly faunal treat- ments. Charley Bird kindly provided biographical information on Thomas Baird. We thank Norbert Kondla for confirming the speci- men identification, and Danny Shpeley and Felix Sperling of the University of Alberta Strickland Entomology Museum for providing access to the specimen collections. Page charges for this note were funded by a Natural Sciences and Engineering Research Council of Canada grant to F.A.H. Sperling. LITERATURE CITED Birp, C. D., G. J. Hitcute, N. G. Konpia, E. M. PIKE & F. A. SPERLING. 1995. Alberta butterflies. Provincial Museum Al- berta, Edmonton. 349 pp. Bowman, K. 1934. Annotated checklist of the macrolepidoptera of Alberta, additions and corrections. Canad. Entomol. 66:131-132. Bowman, K. 1951. An annotated list of the Lepidoptera of Alberta. Can. J. Zool. 29:121-165. Dop, F. H. W. 1914. Further notes on Alberta Lepidoptera. Canad. Entomol. 46:93-403. . 1915a. Further notes on Alberta Lepidoptera, with de- scription of a new species. Canad. Entomol. 47:1-8, 33-42. . 1915b. Further notes on Alberta Lepidoptera. Canad. En- tomol. 47:122-134. Ferris, C. D. & F. M. Brown. 1981. Butterflies of the Rocky Mountain states. University of Oklahoma Press, Norman, Okla- homa. 442 pp. LayBERRY, R. A., P. W. HALL & J. D. LAFONTAINE. 1998. The but- terflies of Canada. University of Toronto Press. 280 pp. Scorr, J. A. 1986. The butterflies of North America: a natural his- tory and field guide. Stanford University Press, Stanford. 583 pp. Gary G. ANWEILER AND B. CHRISTIAN SCHMIDT, Strickland Entomological Museum, Dept. of Biological Sciences, University of Alberta, Edmonton, Alberta T6G 2E9, Canada. Received for publication 26 November 2002; revised and accepted 29 April 2003. BOOK REVIEWS Journdl of the Lepidopterists’ Society 57(3), 2003, 251-252 PTEROPHOROIDEA & ALUCITOIDEA, by C. Gielis, World Catalogue of Insects 4: 1-198, H. van der Wolf (Editor), Apollo Books; Publication: 2003; hardback; ISBN 87-88757-68-4; Price excluding postage and 10% discount if ordered directly from Apollo Books: DKK 320,00 (about 46 US$). See www.apollobooks.com This is the fourth volume of the World Catalogue of Insects series and the first to treat Lepidoptera. The treatment covers all known taxa of Pterophoroidea (1139 species) and Alucitoidea (205 species), the plume moths and the many-plumed moths respectively. The book is divided in 11 parts. The first one is a short summary where unfortunately the name Agdistopidae is introduced in error for Macropirati- dae. Six new synonyms are recorded in an inconsistent manner as each pair of names should have been men- tioned in their original combination, which is the case for two of the six pairs. In two cases where the new combination of the oldest name is mentioned, the parentheses are missing. And at least one new syn- onym, encountered on page 21, is not listed in the summary. Also, the author writes that a new species is mentioned whereas what is actually mentioned is a new name; consequently, the abbreviation nom. nov. should have been used instead of spec. n. The second part is a four-page introduction that provides background information, an explanation on how the data are presented in the Catalogue, a useful list of the family-group names and genera with their respective numbers of species, and the acknowledge- ments. Here the reader will find that the author has elected to go against Article 31.2 of the International Code of Zoological Nomenclature (ICZN) and use the original spelling of species names in lieu of making them agree in gender with their generic name. There has been and still is debate around that question and a number of people don’t like this Code’s article, but un- til the Code is changed, I believe that it should be fol- lowed. Moreover, this decision is not consistently ap- plied as exemplified by the combination Diacrotricha lanceata (Arenberger) and others. Part of the intro- duction is dedicated to the delimitation of the seven biogeographical regions used throughout the Cata- logue to record distribution data. One of these, the Pa- cific region, is said to include New Zealand, Microne- sia, the Hawaiian and Galapagos archipelagos, etc. As far as is known the Lepidopteran fauna of the Galapa- gos is 100% Neotropical in origin, so the decision to exclude them from this region is not scientifically sound. My last comment regarding this introduction relates to the quality of the language, which is poor, with several spelling mistakes. For example, the word “catalogue” is misspelled twice in the first paragraph. Parts 3 and 4 of the book are the catalogues of Pterophoroidea and Alucitoidea. The Pterophoroidea are divided here into Pterophoridae and Macropirati- dae, the latter containing only the three species of Agdistopis. The Alucitoidea include the Alucitidae (186 species) and the Tineodidae (19 species), two groups that are not divided into subfamilies. The Pterophoridae are divided into four subfamilies fol- lowing the author's revision of the group (Gielis 1993). The subfamilies, the tribes of Pterophorinae and all genera are arranged phylogenetically, and the species are all arranged in alphabetical order. The presentation of each valid name in bold face with proper indentations for the following list of syn- onyms as well as hostplant and distribution data makes for an easy consultation. The list of taxa appears com- plete, but it is unfortunately tinted by the presence of a few of the author's new names that are in press in other publications. Their appearance here will only confuse recorders of nomenclature in the future. I have checked for the accuracy of only a few entries and I have found that some information is missing, such as the new distribution and hostplant data for the species I treated in my second Galapagos Pterophori- dae paper (Landry 1993). In addition, spelling mis- takes are unfortunately rather frequent for country and hostplant names, and the names of the hostplant descriptors are not consistently mentioned. Also, if a line reaches the margin of the page and a word is cut, the required hyphen is consistently missing, and I no- ticed a problem in the use of the diacritic marks for the name of I. Capuse (see pp. 73, 74). One error in the list of Alucita names is that A. montana is attrib- uted to Cockerell, while it was actually made valid by Barnes and Lindsey. Part 5 lists the only known fossil species for the two superfamilies while part 6 is the “Comprehensive Ref- erence List,” which indeed seems comprehensive, but here also the cut words at the end of sentences are not hyphenated. Parts 7 to 11 are the indexes to the Dipterous parasites, the Hymenopterous parasites, the hostplants by generic name, which reduces its useful- ness, the taxa of Alucitoidea, and the taxa of Pterophoroidea inclusive of synonyms. I believe the last two indexes should have been fused, to improve the use of the Alucitoidea index, which is presented before that of the Pterophoroidea. This new resource on the World species of Aluci- toidea and Pterophoroidea will be necessary to a wide range of people interested in these taxa because it ap- pears to fulfill the three most important criteria for usefulness of this type of publication: 1—the systemat- ics is up-to-date; 2—the available scientific names are all recorded; and 3—the orthography of these names is correct except for the agreement in gender of the species names. Moreover, the citations of original pub- lication information of the moths’ names also appears recorded free of errors and the list of references is comprehensive. However, there is some missing infor- mation in the hostplant and distribution data, and poor editing of these data and other parts, which is very un- fortunate given the costs and efforts involved, as this may cause one to become suspicious of the quality of the rest of the information presented. I can only rec- JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY ommend that the future installments of the series be reviewed and edited more carefully. On a final positive note, the quality of the binding and paper are excellent. I thank Ivan Lébl and Jeff Wells for their comments on the man- uscript. LITERATURE CITED GIELIS, C. 1993, Generic revision of the superfamily Pterophoroidea (Lepidoptera). Zoologische Verhandelingen, Leiden 290:1-139. Lanpry, B. 1993. Additions to the knowledge of the Pterophoridae (Lepidoptera) of the Galapagos archipelago, Ecuador, with de- scriptions of two new species. Zoologische Mededelingen, Lei- den 67:473-485. BERNARD LANDRY, Muséwm dhistoire naturelle, C.P. 6434, CH-1211 Geneva 6, Switzerland Date of Issue (Vol. 57, No. 3): 29 September 2003 EDITORIAL STAFF OF THE JOURNAL Caria M. Penz, Editor Department of Invertebrate Zoology Milwaukee Public Museum Milwaukee, Wisconsin 53233 USA flea@mpm.edu Put DeVries, Book Review Editor Center for Biodiversity Studies Milwaukee Public Museum Milwaukee, Wisconsin 53233 USA pjd@mpm.edu Associate Editors: Gerarpo Lamas (Peru), Kenetm W. Puinie (USA), Ropertr K. Rospins (USA), Feuix Spertinc (Canada), Davin L. Wacner (USA), Curister WikLuND (Sweden) NOTICE TO CONTRIBUTORS ontributions to the Journal may deal with any aspect of Lepidoptera study. Categories are Articles, Profiles, General Notes, Techni- omments, Book Reviews, Obituaries, Feature Photographs, and Cover Illustrations. Obituaries must be authorized by the president e society. Requirements for Feature Photographs and Cover Illustrations are stated in Volume 44(2):111 and on the Society's web at http://www.lepsoc.org/. nd Journal submissions to the editor at the above address (or electronically to: flea@mpm.edu). 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Au- thors of Book Reviews and Obituaries are exempt from page charges. _ Correspondence: Address all matters relating to the Journal to the editor. Address book reviews directly to the book review editor. =, PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A. CONTENTS MIbD-WINTER FORAGING OF COLONIES OF THE PINE PROCESSIONARY CATERPILLAR THAUMETOPOEA PITYOCAMPA SCHIFF. (THAUMETOPOEIDAE) TT. D. Fitzgerald and Xavier Panades I Blas ------ 161 DEFENSE MECHANISMS IN PyRALIDAE AND CHOREUTIDAE: FECAL STALACTITES AND ESCAPE HOLES, WITH REMARKS ABOUT COCOONS, CAMOUFLAGE AND APOSEMATISM Annette Aiello and M. Alma Solis --- 168 HYBRIDIZATION OF CHECKERSPOT BUTTERFLIES IN THE GREAT BASIN George T. Austin, Dennis D. Murphy, John F: Baughman, Alan E. Launer and Erica Fleishman ------------------------- 176 THE BIOLOGY OF MELANIS LEUCOPHLEGMA (SticHEL, 1910) (RiopinmDaAez) IN WESTERN PERU Curtis John Callaghan ------------------------------------~---9 2-02 -nn nnn nnn 193 A REVIEW OF THE SCHINIA REGIA (STRECKER) SPECIES COMPLEX WITH A DESCRIPTION OF A NEW SPECIES (Nocrurpae: Heiotuinazr) Michael G. Pogue and Charles E. Harp ----------------------- 197 THE HISTORY AND TRUE IDENTITY OF MeziracA ISMERIA (NYMPHALIDAE): A REMARKABLE TALE OF DUPLI- CATION, MISINTERPRETATION, AND PRESUMPTION John V. Calhoun -------------------------------- 204 LATE-INSTAR SHIFT IN FORAGING STRATEGY AND TRAIL PHEROMONE USE BY CATERPILLARS OF THE NEOTROP- ICAL MOTH ARSENURA ARMIDA (CRAMER) (SATURNIIDAE: ARSENURINAE) James T: Costa, Dietrich A. Gotzek and Daniel H. Janzen --------------------------------------=-=--=-2= 29-2 nn nnn 220 - A NEw SPECIES OF EprBLeMA (TorTRICIDAE) FORMERLY MISIDENTIFIED AS E. WALSINGHAMI (KEARFOTT) AND E. inreLix Heinrich Donald J. Wright and Charles V. Covell Jr. ----------------------- 230 Toucu AFRICAN MODELS AND WEAK MIMICS: NEW HORIZONS IN THE EVOLUTION OF BAD TASTE BP J. DeVries --------------------------------------------- === === 5-3 sn nnn nnn nnn nn nn 235 GENERAL NOTES NOTES ON LARVAL MANDIBLE MORPHOLOGY OF HYLEPHILA PHYLEUS PHYLEUS (Drury) (HESPERIIDAE, Hesrerunaz) Marcelo Duarte, Olaf H. H. Mielke and Mirna M. Casagrande ------ 239 NOTES ON THE HISTORIC RANGE AND NATURAL HISTORY OF ANAEA TROGLODYTA FLORIDALIS (NYMPHAL- wae) Mark H. Salvato and Michael K. Hennessey -------------------------------------- 243 Epity’s Copper, LycagNA EDITHA (LYCAENIDAE), CONFIRMED FOR CANADA Gary G. Anweiler and B. Christian Schmidt. ------------------------------------------------------------------==- 249 Book Reviews PreropHoroiDeEa & Atucitoipea Bernard Landry ------------------------------------==-----=- 251 This paper meets the requirements of ANSI/NISO Z29.48-1992 (Permanance of Paper). () DCL Ow zs el BK Volume 57 Number 4 Beat 9 December 2003 ce NT ISSN 0024-0966 _ Journal of the Lepidopterists’ Society A JAN 0 7 2004 Published quarterly by The Lepidopterists’ Society THE LEPIDOPTERISTS’ SOCIETY EXEcuTIVE CouNCcIL LawrENceE F. Gat, President Scorr E. Mituer, Vice President J. Donatp Larontane, Immediate Past President Konrap Fiepier, Vice President Joun H. Accorn, Vice President Kexty Ricuers, Treasurer Ernest H. Winuiams, Secretary Members at large: J. Bolling Sullivan James K. Adams William E. Conner Philip DeVries Wayne F. Wheling Rebecca Simmons David Ahrenholz Jeffrey R. Slotten Charles V. Covell Jr. EpirortaAL BoaRD Rosert K. Rossins (Chairman) Carta M. Penz (Journal) Lawrence F. Gaur (Memoirs) Puiu J. Scuaprert (News) Joun A. Snyper (Web) Honorary Lire MEMBERS OF THE SOCIETY Cuarves L. Reminctron (1966), E. G. Munroe (1973), Ian F. B. Common (1987), Joun G. Franctemont (1988), Lincotn P. Brower (1990), Douctas C. Fercuson (1990), Hon. Miriam Roruscumtp (1991), Ciaupe 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 pro- mote the science of lepidopterology in all its branches, . . . to issue a periodical and other publications on Lepidoptera, to facilitate the exchange of specimens and ideas by both the professional worker and the amateur in the field; to secure cooperation in all mea- sures” directed towards these aims. Membership in the Society is open to all persons interested in the study of Lepidoptera. All members receive the Journal and the News of The Lepidopterists’ Society. 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For information about the Society, contact: Ernest H. Williams, Department of Biology, Hamilton College, Clinton, NY 13323. To order back issues of the Memoirs, write for availability and prices to Kenneth R. Bliss, 28 DuPont Avenue, Piscataway, NJ 08854. The additional cost for members outside the U.S. is to cover mailing costs. Journal of The Lepidopterists’ Society (ISSN 0024-0966) is published quarterly by The Lepidopterists’ Society, % Los Angeles County Museum of Natural History, 900 Exposition Blvd., Los Angeles, CA 90007-4057. Periodicals postage paid at Los Angeles, CA and at additional mailing offices. POSTMASTER: Send address changes to The Lepidopterists’ Society, % Natural History Museum, 900 Exposition Blvd., Los Angeles, CA 90007-4057. Cover illustration: Final instar caterpillar of Lirimiris meridionalis (Schaus) (Notodontidae) feeding on cacao (Theobroma cacao Lin- naeus) in Belize. Photo by Allen M. Young. ee eo ee en —_— ae A ee ee a ee ee eS ee a ee JOURNAL OF Tue LepiporTeRIstTs’ SOCIETY Volume 57 2003 Number 4 Journdl of the Lepidopterists’ Society 57(4), 2003, 253-269 AN ILLUSTRATED GUIDE TO THE ORTHOCOMOTIS DOGNIN (TORTRICIDAE) OF COSTA RICA, WITH SUMMARIES OF THEIR SPATIAL AND TEMPORAL DISTRIBUTION 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, USA. E-mail: jbrown@sel.barc.usda.gov ABSTRACT. Ten species of Orthocomotis Dognin are reported from Costa Rica: O. ochracea Clarke; O. herbacea Clarke (=O. subolivata Clarke, new synonymy); O. longicilia Brown, new species; O. magicana (Zeller); O. chaldera (Druce); O. herbaria (Busck) (=O. cristata Clarke, new synonymy; = O. wragia Razowski & Becker, new synonymy); O. phenax Razowski & Becker; O. similis Brown, new species; O. nitida Clarke; and O. altivolans Brown, new species. Orthocomotis herbacea has been reared from avocado (Persea americana) and O. herbaria from Nectandra hihua, both in the Lauraceae, suggesting that this plant family may act as the larval host for other species of Orthoco- motis. A portion of a preserved pupal exuvium associated with the holotype of O. herbacea suggests that the pupae of Orthocomotis are typical for Tortricidae, with the abdominal dorsal pits conspicuous in this stage. Adults and genitalia of all species are illustrated, and elevational oc- currence is graphed. Orthocomotis herbaria and O. nitida are species of the lowlands (ca. 0-800 m); O. altivolans is restricted to the highest el- evations (ca. 2000-3000 m); the remainder of the species occupy the middle elevations (ca. 800-1800 m). Five of the 10 species documented from Costa Rica appear to be restricted to this Central American country. Additional key words: Neotropical, systematics, identification, elevation, morphology, biodiversity, avocado, Lauraceae. The genus Orthocomotis Dognin includes 34 de- scribed species restricted to the New World tropics, ranging from central Mexico and the Caribbean (Clarke 1956, Razowski 1999) south to Argentina (Ra- zowski & Becker 1990, Powell et al. 1995); numerous undescribed species are present in the major entomo- logical collections worldwide. The monophyly of the group (i.e., Orthocomotis plus the monotypic Paraco- motis Razowski) is well supported by the presence of paired subdorsal pits on abdominal segments 2 and 3 in both sexes (Brown 1989), a greatly expanded patch of chaetosemata that extends in a narrow band across the entire vertex of the head in both sexes (Brown 1989), a finely and densely spined anellus that is firmly attached to the dorsum of the aedeagus in the male genitalia (Razowski 1982), and dense, long scales on the dorsum of the abdomen. In contrast, the tribal as- signment has remained enigmatic. Clarke (1956) treated Orthocomotis as a member of Tortricinae with- out specific tribal placement. Razowski initially consid- ered the group as Archipini but later (Razowski 1982) transferred it to Polyorthini on the basis of the minutely spined dorsal portion of the anellus and the presence of a dorsal sclerite in the distal, membranous portion of the aedeagus. Powell (1986) assigned Or- thocomotis and Paracomotis to Euliini (Tortricinae). Brown (1989) then transferred them to Schoenotenini on the basis of an unusual modification of the chaeto- semata, and Razowski and Becker (1990) again argued for their placement in Polyorthini. Powell et al. (1995) followed Brown (1989) in the checklist of the Neo- tropical Lepidoptera, placing the group in Schoeno- tenini. Horak (1999) questioned this placement, stating that the absence of the band of chaetosemata in the more primitive Schoenotenini argued against the homol- ogy of the structure between Orthocomotis plus Paraco- motis and the more advanced schoenotenines that pos- sess it. Currently there is no consensus, and Horak (1999) provisionally has returned the group to Euliini. Adults of Orthocomotis are relatively large and col- orful; nearly all species have patches of metallic green or blue-green scales on the upper surface of the forewing. In facies and size they are similar to many large Neotropical Chlidanotini (Chlidanotinae), partic- ularly larger species of Auratonota Razowski; the two genera frequently are mixed in collections of Neotropi- JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TABLE 1. Diagnostic morphological characters and summary of elevational distribution. Unique features in bold. HW pecten Haripencil Cornuti HW color ochracea absent absent large dark brown herbacea absent absent large brown longicilia absent absent medium brown magicana absent absent small brown chaldera absent present minute gray herbaria present present small dark brown phenax absent present small brown similis absent present small dark brown nitida absent present absent dark brown altivolans absent present absent white cal Tortricidae. However, adults of Orthocomotis always can be separated from similar appearing taxa by the conspicuous band of chaetosemata mentioned above. Although the genus is widely distributed throughout the New World tropics, little is known of the biology or the temporal and geographic distributions of the species. One species has been recorded from avocado (Persea americana Mill.; Lauraceae) on at least two oc- casions in Costa Rica and a second from Nectandra hi- hua (Ruiz & Pav.) Rohwer (Lauraceae) once, suggest- ing that other species of Orthocomotis may use Lauraceae as well. No other hosts are known for the genus. The relatively thorough sampling of the genus in Costa Rica provides the opportunity to examine spa- tial and temporal distributions of species in this coun- try. The purposes of this paper are to present a survey of the genus in Costa Rica, provide illustrations of adults and genitalia to facilitate identifications, resolve a number of taxonomic difficulties, identify prelimi- nary temporal and geographic distributions of the species in Costa Rica, and describe three new species that appear to be restricted to Costa Rica. MATERIALS AND METHODS Specimens of Orthocomotis from Costa Rica were borrowed from or examined at the following institu- tions: BMNH, The Natural History Museum, London, England; INBio, Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica; UCB, Essig Museum of Entomology, University of California, Berkeley, California, U.S.A.; USNM, National Mu- seum of Natural History, Washington, D.C., U.S.A.; and VBC, Vitor Becker personal collection, Planaltina, Distrito Federal, Brazil. Specimens were sorted by forewing pattern and sex. The resulting groups then were examined for differ- ences in male and female genitalia, which have been shown to provide the most reliable morphological fea- tures for distinguishing among related species of Tor- tricidae. Types of all species were examined to verify Frons FWlengthd FW length? Cilia Elevation tan 9.6—-11.0 12.3-13.3 short 1000-1750 tan 9.2-12.5 12.0-12.8 short 1000-1750 tan 10.5-11.5 JULY long 1000-2000 white 10.1—-11.0 11.0-13.0 short 500-1500 tan 13.1-15.5 16.5-19.5 short 1000-2750 tan 10.0-11.1 11.6-12.8 short 0-750 tan 10.5-11.2 12.4-12.8 short 500-1750 tan 10,.5-12.5 12.5-16.0 short 1000-1750 tan 9.7-10.1 11.6-12.5 short 0-750 tan 12.0-13.5 13.3-15.0 short 2250-3000 identifications. Preparation of genitalia follows the methodology summarized in Brown and Powell (1991). Because of the large size of Orthocomotis adults, an attempt was made to evert the vesica of the aedeagus by extracting it from the distal end with a pair of fine forceps. Adult specimens were examined using a Wild M3Z™ dissecting microscope; genitalia were examined using the dissecting microscope and a Zeiss™ compound microscope. Illustrations of geni- talia were drawn with the aid of a Ken-A-Vision™ mi- croprojector (model X1000-1). Unless indicated other- wise, genitalia illustrations are of a single preparation. Text descriptions of all taxa are composite, based on all available specimens. Measurements of forewing were made with the aid of an ocular micrometer mounted in a dissecting microscope under low power (x10-16). Forewing length was measured in a straight line from the base of the wing to the apex, including the fringe. Terminology for wing venation and genitalia struc- tures follows Horak (1984). Abbreviations and symbols are as follows: HT = holotype; PT = paratype; ca. = circa (approximately); n = number of individuals ex- amined; X = arithmetic mean; N, E, S$, W = compass points; P.N. = Parque Nacional; Est. Biol. = Estaci6én Biol6gica; Fca. = Finca; ALAS = Arthropods of La Selva (parataxonomists). In the “specimens examined” sections, months of the year are abbreviated using the first three letters. A histogram illustrating elevational occurrence was developed for each species based on the available label data. The number of specimens collected at intervals of 250 m, starting at sea level (i.e., 0-250, 250-500, 500-750, etc.), was tallied. Where ranges in elevation are given on the specimen labels (e.g., 1400-1700 m), 0.5 specimen was used for each of the two elevation categories (i.e., 0.5 specimen for the 1400-1650 m cat- egory, and 0.5 specimen for the 1650-1900 m cate- gory). A comparable method was used for species that were collected at the category “break-point” (i.e., 250 m, 500 m, etc.). VOLUME 57, NUMBER 4 TABLE 2. Species distribution by province. ALA = Alajuela; CAR = Cartago; GUA = Guanacaste; HER = Heredia; LIM = Limon; PUN = Puntarenas; SAN = San José. Species Provinces # of provinces ochracea ALA, CAR, HER, PUN 4 herbacea CAR, GUA, HER, PUN, SAN 5 longicilia ALA, CAR, GUA, HER, PUN, SAN 6 magicana ALA, CAR, GUA, HER, PUN 5 chaldera CAR, GUA, HER, PUN, SAN 5 herbaria ALA, GUA, HER, LIM, PUN, SAN 6 phenax GUA, HER, PUN, SAN 4 similis CAR, GUA, SAN 8 nitida ALA, GUA, HER, LIM, PUN 5 altivolans ALA, CAR, HER, LIM, SAN 5 A brief list of morphological characters useful in dis- tinguishing the species is presented in Table 1; details are presented below. For most species, comparison with the photographs of adults (Figs. 1-12) will pro- vide accurate identifications, which can be confirmed using Table 1. For worn or damaged specimens, geni- talia dissections usually are required, and comparison with the illustrations of genitalia should provide accu- rate determinations. Table 1 includes nine of the most conspicuous fea- tures for distinguishing the species of Orthocomotis treated herein. Hindwing pecten (“HW pecten’) refers to a dense row of somewhat stiff, erect scales at the base of the hindwing along vein CuP. This charac- ter separates O. herbaria from all other species. “Hair- pencil” refers to the presence of a dense fascicle of elongate scales that extends from the metathorax to an unusual lateral pouch bearing scent scales in the sec- ond abdominal segment in males only. This character may define a species group within Orthocomotis; it is present in 6 of the 10 species treated. “Cornuti” refers to the size of cornuti in the vesica of the aedeagus in the male genitalia. Although the categories are qualita- tive (large, medium, small), the cormuti of O. ochracea and O. herbacea are comparatively long, needle-like spines, while those of most other species are short and thorn-like, less than half as long as those of O. ochracea and O. herbacea; the cornuti are absent nearly altogether in O. nitida and O. altivolans. Hind- wing color (“HW color’) refers to the color of the scales on the dorsal surface of the hindwing. While the categories (gray, brown, dark brown, white) are some- what subjective, the hindwing of O. herbaria, O. ni- tida, O. similis, and O. ochracea is darker than that of other species, and the hindwing of O. altivolans is nearly white. “Frons” refers to the color of the scaling on the upper portion of the frons, which is variable: tan, cream, or brownish. However, two species are relatively distinct in this feature: the frons of O. magi- cana is white and that of O. nitida is bicolored (yellow and red-brown). Forewing length (“FW length 3” and “FW length 2”) provides a general range to help dis- tinguish relatively large from relatively small species. “Cilia” refers to the relative length of the cilia from the male antenna. It is moderately uniform and short (ca. 0.40.6 times the diameter of the flagellomere) in all species except O. longicilia, which has conspicuously longer cilia (ca. 1.0-1.2 times the diameter of the fla- gellomere). “Elevation” refers to the general range in elevation (excluding outliers) occupied by each species based on collection records. While most species oc- cupy a relatively broad elevational range, O. herbaria and O. nitida are clearly species of the lowland, and O. altivolans is restricted to the highest elevations. Table 2 presents data on the spatial distribution of Costa Rican Orthocomotis species by Province. No single species has been recorded from all seven provinces, and no province supports more than § of the 10 known species of Orthocomotis. Eight species are known from Guanacaste, Heredia, and Puntarenas provinces, 7 species from Cartago and San José provinces, 6 species from Alajuela Province, and three species from Limén Province. Of the 10 species of Or- thocomotis documented from Costa Rica, five appear to be restricted to Costa Rica. SPECIES ACCOUNTS Orthocomotis ochracea Clarke (Figs. 8, 13, 23, 33) Orthocomotis ochracea Clarke, 1956:144; Razowski & Becker 1990:350. Holotype 2, Costa Rica, Cartago Province, Juan Vifias, Wm. Schaus, USNM. Diagnosis. The absence of the male abdominal hairpencil and the presence of a patch of large cornuti in the vesica of the aedeagus (Fig. 13) suggest that O. ochracea may be most closely related to O. herbacea (Fig. 14). The male genitalia are characterized by ex- tremely broad, pendant socii and a short, wide, hook- shaped distal portion of the gnathos. Orthocomoits ochracea is distinguished superficially from all other congeners by the red-brown forewing reticulation, the darker brown hindwing, and the comparatively small forewing size of males (Table 1). Specimens examined. Alajuela Province: Rio Sarapiqui, 6 air km S San Miguel, 800 m, 7 Jun 1988 (1 4), J. Brown & J. Powell (UCB). Rio Sarapiqui, 2 km SE Casablanca, 700 m, 28 Mar 1992 (1 3), J. McCarty & J. Powell (UCB). Cartago Province: Turrialba, Monumento Nacional Guayabo, 1100 m, Jul 1994 (1 d), Sep 1994 (1 3), G. Fonseca (INBio). Rio Aquiares, nr Santa Cruz, 9 km NW Tur- rialba, 1500 m, 31 May 1985 (1 2), J. Powell (UCB). Paraiso, P.N. Tapanti-Macizo de la Muerte, 300 m N & 100 m W Mirador, 1350 m, Jan 2000 (2 ), R. Delgado (INBio). Parafso, P.N. Tapanti, Sect. La Represa, 300 m SE Puente del Rio Porras, 1660 m, Sep 1999 (1 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 1-12. Adults of Orthocomotis. 1, O. similis (2); 2, O. herbacea (2); 3, O. magicana (3); 4, O. nitida (3); 5, O. herbaria (2); 6, O. herbaria (3); 7, O. altivolans (¢); 8, O. ochracea (2); 9, O. longicilia (3); 10, O. chaldera (6); 11, O. chaldera (2); and 12, O. phenax (8). 3), R. Delgado (INBio). Tapanti, Rio Grande de Orosi, 1300-1400 m, 9°46/N, 83°50’W, 23 Jan 1985 (1 2), D. Janzen & W. Hallwachs (INBio). Juan Vifias, [no date] (HT ¢), Wm. Schaus (USNM). Here- dia Province: El Angel Waterfall, 8.2 kin downhill Vara Blanca, 1350 m, 5 Aug 1981 (1 2), D. Janzen & W. Hallwachs (INBio). 16 km SSE La Virgen, 10°16’N, 84°05W, INBio-OET-ALAS transect, 1050-1150 m, 12 Feb 2001 (2 3), M. Epstein (INBio), 15-21 Mar 2001 (1 3), 18 Mar 2001 (1 d), D. Wagner & J. Rota (INBio), 10 Apr 2001 (1 4), blacklight trap, 11 Apr 2001 (1 d), 12 Apr 2001 (1 4), 13 Apr 2001 (1 4), D. Davis (INBio), 17 Apr 2001 (2 4), J. Brown (IN- Bio). Puntarenas Province: Est. Biol. Las Alturas, 12 air km NE San Vito, 1550 m, 22-24 Jan 1993 (3 4), J. Powell (UCB). Coto Brus, Est. Biol. Las Alturas, 1550, 15-24 Mar 1999 (5 d, 1 9), G. Rodriguez (INBio). Coto Brus, Zona Prot. Las Tablas, Est. Biol. Las Alturas, 1550 m, 16-23 Mar 1999 (3 d), E. Phillips (INBio). A.C.L.A.P. Coto Brus, Zona Prot. Las Tablas, Est. Biol. Las Alturas, 1550 m, 15-24 Mar 1999 (3 d), R. Delgado (INBio). Fea. Cafrosa, Est. Las Mel- lizas, P.N. Amistad, 1300 m, Jan 1991 (12), M. Chavarria & G. Mora (INBio). Las Cruces, nr San Vito, 24 Apr 1965 (1 d),S.S. & W.D. Duckworth (USNM). Unknown Province: Palo Verde, 5200’, “.20” [1920] (1 3), [no collector] (BMNH). Geographic and temporal distribution. Ortho- comotis ochracea is known only from Costa Rica where it ranges from about 700 to 1500 m (Fig. 33) in the central and western portions of the country. It has been collected in January (n = 7), February (n = 2), March (n = 16), April (n = 5), May (n = 1), June (n = 1), July (n = 1), August (n = 1), and September (n = 2). Remarks. This species was described from a single female erroneously identified as a male. Associated with the holotype female is a slide which has the male genitalia of O. chaldera. Based on the incorrectly asso- ciated slides, Clarke (1956) concluded that “The male genitalia of ochracea and chaldera are almost indistin- guishable . . .” Actually, the male genitalia are most similar to those of O. longicilia. Orthocomotis herbacea Clarke (Figs. 2, 15, 24, 34) Orthocomotis herbacea Clarke, 1956:151. Holotype ¢ (herbacea), Costa Rica, San José Province, San Pe- dro de Montes de Oca, ex-larva, 22 Dec 1932, em: 15 Jan 1933, rf. avocado [Persea americana], C. H. Ballou, USNM. Orthocomotis subolivata Clarke, 1956:148; Razowski & Becker 1990:350, new synonymy Holotype ¢ (subolivata), Costa Rica, Tuis, 5800’ [elevation prob- ably in error; Tuis is ca. 2400’], 28 Aug 1908, Wm. Schaus, USNM. VOLUME 57, NUMBER 4 Diagnosis. The forewing pattern (Fig. 2), with a large dark brown or black patch in the distal third of the wing and a distinct, small, dark brown semicircular patch near the middle of the costa, distinguishes O. herbacea from other species, except possibly O. magi- cana, which has considerably more greenish metallic scaling. The triangular process representing the termi- nation of the sacculus in the male genitalia of O. herbacea is extremely variable, ranging from a rounded nub to an elongate spine. The illustration in Fig. 15 represents the extreme in spine development; an additional illustration can be found in Clarke (1956) for O. subolivata. The aedeagus of O. herbacea is char- acterized by an extensive patch of large spinelike cor- nuti, similar to that of O. ochracea. Specimens examined. Cartago Province: R. Grande de Orosi, desde Puente Rio Dos Amigos, hasta la represa, 1400-1800 m, Mar 1996 (1 d), R. Delgado (INBio). Monumento Nacional Guayabo, 1100 m, Oct 1994 (1 4), G. Fonseca (INBio), Jul 1994 (1 3), G. Fonseca (INBio). Paraiso, P.N. Tapanti, Sect. La Represa, 300 m SW Puente del Rio Porras, 1660 m, May 1999 (1 ¢), Nov 1999 (2 3), R. Delgado (INBio). Paraiso, P.N. Tapanti-Macizo de la Muerte, 300 m SE Rio Porras, 1660 m, Jan 2000 (6 3), Feb 2000 (2 2), May 2000 (4 3), Jun 2000 (2 3), Nov 2000 (1 3), Aug 2001 (1 3), R. Del- gado (INBio). Paraiso, P.N. Tapanti, Sect. La Represa, 300 m SW Puente del Rio Porras, 1660 m, Feb 2000 (1 2), May 1999 (1 4), Nov 1999 (1 3), Aug 2001 (1 3), Nov 2001 (3 3), R. Delgado (INBio). Paraiso, P.N. Tapanti-Macizo de la Muerte, 300 m N & 100 m W del Mirador, 1350 m, Jan 2000 (1 2), R. Delgado (INBio). P.N. Tapanti- Macizo de la Muerte, Est. Quebrada Segunda, al costado Ofic., 1200 m, Dec 1999 (1 4), R. Delgado (INBio). P.N. Tapanti, Est. Quebrada Segunda, 1200 m, Oct 2000 (1 2), R. Delgado (INBio). Turrialba, Tayutic, P.N. Barbilla, Sector Cerro Tigre, 1617 m, Jan 2002 (1 ¢), L. Chavarria (INBio). Tapanti, 1200-1700 m, 20 Aug—15 Sep 1999 (4 3, 12), V. Becker (VBC). P.N. Tapanti, 1200-1700 m, 20 Aug—15 Sep 1999 (5 d), V. Becker (USNM). Tapanti, 1500 m, 30-31 Aug 2000 (3 3), V. Becker (VBC). Santa Cruz, Turrialba, 1500 m, Aug 1981 (1 2), V. Becker (VBC). Volcan Turrialba, 1800 m, 13 Aug 1972 (1 2), V. Becker (VBC). Villa Mills, 2840 m, 26-28 Oct 2000 (1 3), V. Becker (VBC). Tuis, 5800’ [elevation probably in error; Tuis is ca. 2400’], 28 Aug 1908 (AT ¢ of subolivata), W. Schaus (USNM). Guanacaste Province: Est. Cacao, S side Volc4n Cacao, P.N. Guanacaste, 1000-1400 m, 8-29 Jul 1991 (1 4), C. Chaves (INBio). Est. Cacao, 1100 m, 17-18 Feb 1995 (1 d), E. Alfaro (INBio). Sector Las peut e 5 km SW Volcdn Rincén de la Vieja, 800 m, 24 Jun—10 Jul 1995 ( ), 23 Jul-6 Aug 1995 (1 d), K. Taylor (INBio). Sector Las ae Le Rincén de la Vieja, 1400 m, 6-26 Jun 1994 (1 2), K. Taylor (IN- Bio). Faldas, SW Volcan Cacao, 1150-1250 m, Jun 1996 (1), I. Vil- legas & C. Moraga (INBio). Derrumbe, Est. Mengo, W side Volcan Cacao, 1400 m, 5 Jun 1988 (1 2), 11 Jul 1988 (1 2), D. Janzen & W. Hallwachs (INBio). Heredia Province: El Angel Waterfall, 8.2 km downhill Vara Blanca, 1350 m, 3 Jan 1981 (2 2), D. Janzen & W. Hallwachs (INBio). Mount Pods [2350 m], [no date] (PT ¢ of herbacea), W. Schaus (USNM). 16 km SSE La Virgen, 10°16’N, 84°05/W, INBio-OET-ALAS transect, 1050-1150 m, 12 Feb 2001 (1 3), M. Epstein (INBio). 6 km ENE Vara Blanca, Braulio Carrillo Nat. Park, 10°11’N, 84°07’W, INBio-OET-ALAS transect, 2000 m, 14 Feb 2002 (1 2), 16 Feb 2002 (2 4), 19 Feb 2002 (1 ¢), 20 Feb 2002 (1 3), J. Brown & J. Powell (INBio). 6 km ENE Vara Blanca, 2000 m, 7 Oct 2002 (1 3), K. Nishida, MV light (USNM). Cerro Gina Res. Biol. Chompipe, R. F. Cord. Vol. Cent., 2100 m, 11 Jul 1991 (1 2), J. F. Corrales (INBio). Puntarenas Province: Est. Pittier, ian m, 22 Sep-9 Oct 1995 (1 3), M. Moraga (INBio), 23 Aug—13 Sep 1995 (2 3), 23-27 Oct 1995 (1 3), 26 Sep-10 Oct 1995 (1 3), E. Navarro (INBio). Sector Altamira, 1 km S Cerro Biolley, bo OL =I A.C. Amistad, 1300 m, 2-20 Apr 1995 (1 d), L. Angulo (INBio). Las Cruces, nr. San Vito, 19-20 Mar 1965 (1 ¢), 24 Apr "1965 ( ilies LO) Se S. & W. D. Duckworth (USNM). Fea. Cafrosa, Embalse, N Tigra, 800 m, 13-21 May 1996 (1 4), E. Navarro (INBio). Est. Biol. Las Cruces, 6 km SE San Vito, Rio Jaba, 1150 m, 20-21 Jan 1993 (1 3), J. Powell (UCB). Buenos Aires, La Amistad, Sector Altamira, Nov 1993 (1 d, 1 9), R. Deleacod INBio). Est. Altamira, Buenos Aires, 15 Sep-14 Oct 1993 (19), R. Delgado (INBio). A.C.L.A.P. Coto Brus, Zona Prot. Las ee Est. Las Alturas, se m, 16-23 Mar 1999 (3 3), M. Moraga (INBio), 16-23 Mar 1999 (1 2), E. Phillips (INBio). Coto Brus, Est. Las Alturas, 1550 m, i 1991 (1 3), M. Ramirez (INBio), Oct 1997 (1 3), B. Gamboa (INBio), 15-24 Mar 1999 (1 2), G. Rodriguez (INBio). Est. Biol. Las Alturas, 1500 m, Aug 1991 (2 3), M. Ramirez (INBio), 1540 m, 28-30 Oct 1997 (1 2), B. Gamboa (INBio). Est. Biol. Las Alturas, 12 air km SE San Vito, 1550 m, 22-24 Jan 1993 (9 Z) J. Powell (UGB). Monteverde, 1500 m, 29-30 Jul WSUS 2), 10-11 Dec 1979 (1 ¢), D. Janzen (INBio), 15-16 May 1980 (1d), D. Janzen & W. ae (INBio), 14 Sep 1999 (3 d) V. sae (VBC), 14 Sep 1999 (1 3), V. Becker (USNM). Las Nubes, 11 km NW Monteverde, 10-11 Dec. 1979 (1 2), D. Janzen (INBio), 31 Jul 1981 (1 2), D. Janzen & W. Hallwachs (INBio). Fila eae 35 km S Palmar Norte, 8°45’N, 83°20’W, 150 m, 7-8 Jan 1983 (1 2), D. Janzen & W. Hallwachs (INBio). San Ie Province: San uae alo de Dota, 7200-7500’, 20 Feb 1996 (1 2), D. & J. Pow- ell (UCB). Est. Zurqui (el Tunel), P.N. Braulio ati 1500 m, 10°04’N, 84°01’W, Nov 1985 (1 ¢), I. & A. Chacén (INBio). Est. Santa Bee Viejo, Santa Elena, Las Nubes, 1210 m, 21-24 Nov 1995 (1 3), E. Alfaro (INBio). San Pedro de Montes de Oca, ex- larva, 22 Dec 1932, em: 15 Jan 1933 (HT ¢ of herbacea), r.f. avocado [Persea americana], C. H. Ballou (USNM). Geographic and temporal distribution. This species ranges from Guatemala (VBC) south through Costa Rica to Ecuador (VBC). In Costa Rica it is known from P.N. Guanacaste to Juan Vifias, from ca. 800 to ca. 2840 m elevation, with the majority of specimens from 1200-1700 m (Fig. 34). Captures range throughout the year: January (n = 22), Febru- ary (n = 5), March (n = 7), April (n = 3), May (n = 8), June (n = 6), July (n = 6), August (n = 5), September (n = 6), October (n = 5), November (n = 7), and De- cember (n = 1). Orthocomotis herbacea has been reared twice in Costa Rica from avocado (Persea americana; Lau- raceae). The anterior portion (head, thorax, and ab- dominal segments 1—3) of a pupal exuvium is pinned beneath the reared holotype. From the exuvium it is clear that the abdominal dorsal pits are conspicuous on the pupa, as in other genera that possess dorsal pits in the adult stage (e.g., Amorbia Clemens, Archips Hiibner), and the rows of dorsal spines on the abdomen are typical for Tortricidae, at least on segment 3. Remarks. Orthocomotis subolivata was described from a single male that is almost certainly conspe- cific with O. herbacea. It is likely that Clarke (1956) did not recognize this because of the paucity of ma- terial and mislabeled slides (see remarks under O. herbaria below). The male genitalia figured Clarke (1956) as O. herbacea belong to a different species. 258 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 13-16. Male genitalia of Orthocomotis with valvae spread, aedeagus removed and shown in lateral aspect, and vesica everted. 13, O. ochracea; 14, O. longicilia, 15, O. herbacea; 16, O. magicana. Orthocomotis longicilia Brown, new species (Figs. 9, 14, 25, 35) Diagnosis. Superficially, O. longicilia can be distin- guished from other species of Orthocomotis by its forewing pattern and color, with considerably less metallic pale green overscaling. Males are distin- guished from congeners by the conspicuously longer antennal cilia (1.0-1.2 times the width of the flagel- lomere). The stout, strong, thorn-like cornuti of the aedegus are somewhat intermediate between the long spine-like cornuti of O. herbacea and O. ochracea and the smaller thorns of O. chaldera and O. magicana. Description. Male. Head: Upper frons light beige with red brown, lower frons dingy whitish. Labial palpus light beige on inner VOLUME 57, NUMBER 4 surface, pale brown on outer surface. Antenna with elongate cilia, 1.0-1.2 times width of flagellomere. Thorax: Light beige with red brown, with small patch of white scales at posterior end of dorsal tuft. Metathorax without hairpencil. Forewing length 10.5-11.5 (x = 11.2; n = 8) (Fig. 9); ground color whitish, in fresh specimens en- tirely overscaled with irregular patches of gold and steel gray which are lost when worm; pattern elements dark reddish brown, over- scaled with metallic green; a pair of faint, parallel, oblique fascia from costa near base, the outer of which bends 90° at discal cell, ex- tending toward apex; a small semicircular patch near mid-costa; a narrow, sinuate band in apical portion of subtermen; a dash from near mid-dorsum extending toward middle of costa, reaching ca. halfway across wing. Hindwing dark brown. Abdomen: Densely clothed with long, fine, pale brown scales; second segment without lateral pouches; dorsum of segments 2 and 3 with paired subdorsal pits. Genitalia as in Fig. 14 (drawn from JWB slide 1269; n = 5). Un- cus slightly expanded and weakly flattened in distal two-fifths, with dense patch of fine hairs from venter in apical one-fourth. Socius large, pendant, with limited lobe dorsad of attachment. Gnathos simple, narrow, with relatively short pointed process at distal junc- tion of arms. Transtilla a simple slender arch. Valva relatively broad, nearly parallel-sided, gently arched dorsad throughout, densely cov- ered with short scales in distal one-half of inner side; costa differen- tiated; sacculus not developed. Aedeagus short, stout, curved imme- diately distad of ductus ejaculatoris; vesica densely covered with large, thornlike comnuti. Female. Head and thorax: Essentially as described for male. Forewing length 11.7 mm (n = 1); pattern as described for male. Abdomen: Densely clothed with long, fine, pale brown scales; sec- ond segment without lateral pouches; dorsum of segments 2 and 3 with paired subdorsal pits. Genitalia as in Fig. 25 (drawn from JWB prep. 1287; n = 1). Sterigma unsclerotized; ostium large, rounded. Ductus bursae extremely short. Corpus bursae ovoid, with broad wrinkles; slender accessory bursae arising near junction with ductus bursae; spicules absent. Holotype ¢, Costa Rica, Cartago Province, Tapantf, 1200-1700 m, 20 Aug—15 Sep 1999, V. O. Becker (USNM). Paratypes. COSTA RICA: Alajuela Province: Rio Saripiqui, 6 air km S San Miguel, 800 m, 7 Jun 1988 (1 3), J. Brown & J. Powell (UCB). Cartago Province: Rio Grande de Orosi, Puente Rio Dos Amigos, hasta represa, 1400-1800 m, 22 Aug—15 Sep 1995 (1 d), R Delgado (INBio). Paraiso, P.N. Tapanti, Sector La Represa, 300 m S de] Puente del Rio Porras, 1660 m, Jun 2000 (1 3), Jul 2002 (1), R Delgado (INBio), Jul 1999 (1 d), Feb 2000 (2 3), L. Chavarria (IN- Bio). Paraiso, P.N. Tapanti-Macizo de la Muerte, 300 m SE Rio Por- ras, 1660 m, Sep 1999 (3 3), Nov 1999 (1 3), May 2000 (1 3), Jan 2000 (4 3), Oct 2002 (1 d, 1 2), R. Delgado (INBio). Paraiso, P.N. Tapanti-Macizo de la Muerte, Costado de Casa Admin., 1200 m, Nov 1999 (1 2), Jun 2000 (1 ¢), L. Chavarria (INBio). P.N. Tapanti- Macizo de la Muerte, 300 m N & 100 m S Mirador, 1350 m, Oct 1999 (1 3), R. Delgado (INBio). Paraiso, P.N. Tapanti-Macizo de la Muerte, 300 m N Mirador, 1830 m, Jul 2000 (1 ), R. Delgado (IN- Bio). Paraiso, P.N. Tapanti, Est. Quebrada Segunda, Sendero Catarata, 1450 m, May 1999 (1 d), R. Delgado (INBio). La Represa, Tapanti, 1800 m, Apr 1995 (1 6), R. Delgado (INBio). Tapanti, 1200-1700 m, 20 Aug—16 Sep 1999 (3 d), V. Becker (USNM). Gua- nacaste Province: Rio San Lorenzo, Tierras Morenas, 1050 m, Sep 1993 (1 3), G. Rodriguez (INBio). Rio San Lorenzo, R.F. Cord., 1050 m, Jun 1991 (1d), C. Alvarado (INBio). Tapantf, 1200-1700 m, 20 Aug—15 Sep 1999 (8 3), V. Becker (VBC, USNM). Tapanti, 1500 m, 30-31 Aug 2000 (4 3), V. Becker (VBC). Z.P. Tenorio, Secto Also Los Masis, 1100 m, 10-14 Jan 2002 (1 ¢), L. Chavarria (INBio). Heredia Province: E] Angel Waterfall, 8.2 km downhill Vara Blanca, 1350 m, 3 Jan 1981 (2d), 5 Aug 1981 (1 9), D. eee & W. Hallwachs (INBio). 8 km N Vara Blanca, 25 Jul 1990 (1 4), J. Powell (INBio). 16 km SSE La Virgen, 10°16’N, Meare INBio- OET- ALAS transect, 1050-1150 m, 8 Feb 2001 (1 d), 12 Feb 2001 (1 3), 13 Feb 2001 (1 2), M. Epstein (INBio), 10 Apr 2001 (1 3), 15 Apr 2001 (1 d), 20 Apr 2001 (1 ¢), J. Brown (INBio). Puntarenas Province: La Amistad, Sect. Altamira, Buenos Aires, Dec 1993 (1 3), R. Delgado (INBio). Coto Brus, Zona Prot. Las Tablas, Est. Biol. Las Alturas, 1550 m, 16-23 Mar 1999 (1d), E. Phillips (INBio). Est. Biol. Las Alturas, 12 air km NE San Vito, 22-24 Jan 1993 (4 4), J. bo Ol ice) Powell (UCB). Sendero a Cerro Pittier, 600 m N Estac., 1750 m, 15 Jul 1996 (2 3), M. Moraga (INBio). Est. Altamira, 1 km S Cerro Bi- olley, 1300-1450 m, 12-30 Aug 1996 (1 2), R. Villalobos (INBio). Fca. Cafrosa, Embalse, 800 m N Tigra, 1280 m, 8-10 Feb 1997 (1 3), A. Picado (INBio). San José Province: Est. Zurqui, 50 m antes de tunel, 1600 m, 26 Sep-Oct 1990 (1 d), G. Maass (INBio). Geographic and temporal distribution. Ortho- comotis longicilia occupies the middle elevations of the central cordillera from about 800 to about 1800 m (Fig. 35). It has been recorded only from Costa Rica. Captures range throughout the year. Etymology. The specific epithet refers to the elon- gate cilia of the male antenna. Orthocomotis magicana (Zeller) (Figs. 3, 16, 26, 36) Penthina (Sericoris) magicana Zeller, 1866:150. Eulia magicana: Meyrick 1926:249. Orthocomotis magicana: Clarke 1956:151; Razowski & Becker 1990:351. Holotype 2, Colombia, Bogota, [no date], BMNH. Diagnosis. Orthocomotis magicana has a bold black patch in the apical region of the forewing similar to O. herbacea but has considerably more metallic green overscaling in the white regions between the darker patches (Fig. 3); the maculation is somewhat variable. Orthocomotis magicana is recognized most easily by the immaculate white scaling of the frons (in- frequently with a few scattered pale brown scales) in both sexes, contrasting with a patch of dark somewhat metallic scales on the vertex, and the extensive white scaling of the tegulae. The male lacks both the hind- wing pecten and the thoracic hairpencil (see Table 1). Specimens examined. Alajuela Province: Upala, Bijagua, Al- bergue Heliconias, 700 m, Apr 2000 (1d), G. Rodriguez (INBio). Fca. San Gabriel, 16 km ENE de Queb. Grande, 11-15 Jun 1986 (1 2), I. Gauld & J. Thompson (INBio). Cartago Province: Monumento Na- cional Guayabo, Turrialba, 1100 m, Jul 1994 (2), Sep 1994 (4d), Oct 1994 (2 d), Nov 1994 (1 3), G. Fonseca (INBio). Paraiso, P.N. Tapanti- Macizo de la Muerte, al Castado de Casa Admin., 1200 m, Jun 2000 (1 2), R. Delgado (INBio). Turrialba, 600 m, 25 Oct 1971 (1 3), V. Becker (VBC). Juan Vifias, [no date] (1 2), W. Schaus (USNM), [no date] (1 2), W. Schaus (BMNH). Guanacaste Province: Est. Pitilla, 9 km S Sta. Cecilia, P.N. Guanacaste, 700 m, Jun 1991 (1 2), Apr 1991 (1 3), Aug 1991 (1 6), 2-15 May 1992 (1 2), C. Moraga (INBio). Hda. Santa Maria, 750 m, Sep 1996 (1. 3), D. Briceno, A. Solis, E. Araya, F. Quesada & C. Moraga (INBio). 4 km E Casetilla, P.N. Rincén, 750m, ve 1981 (1 3), 25 Jan 1982 (1 d 3), 2: 2 May 1982 (1 2), 27 Dec 1981 (3 , D. Janzen & W. Hallwachs (INBio). Heredia Province: Sara- ae Zona Prot. La Selva, Est. Biol. La Selva, 50-100 m, 6 Feb 1987 (2 3), I. Chacén (INBio). Braulio Carrillo Natl. Park, 6 km E Vara Blanca, 10°11’N, 84°07’W, INBio-OET-ALAS transect, 2000 m, 16 Feb 2002 (1 3), ALAS (INBio). Puntarenas Province: Est. Altamira, Buenos Aires, 15 Sep—14 Oct 1993 (1), R. RN (INBio). Sector Altamira, Buenos Aires, PILA, 1400 m, Jun 1994 (2 2), Jul 1994 (1d), R. Delgado (INBio). Est. Altamira, 1 km S Cerro Biolley, 1300-1450 m, 20-30 Oct 1996 (1 2), R. Villalobos (INBio). Buenos Aires, PILA, Sector Altamira, A.C. Amistad, 1150-1400 m, May 1994 (1 3), R. Del- gado (INBio). Buenos Aires, La Amistad, Sector Altamira, Nov 1993 (1 9), RB. Delgado (INBio). Buenos Aires, Parque Intemacional Amis- tad, Sendero Gigantes, 1450 m, Sep 2001 (1 2), R. Delgado (INBio). Fca. Cafrosa, Est. Las Mellizas, P. N. Amistad, 1300 m, Oct 1989 (1 3), M. Ramirez (INBio). Fea. Cafrosa, Embalse, 800 m N Tigra, 1280 m, 13-21 May 1996 (2 3), E. Navarro (INBio). Buen Amigo, San Luis JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY / Fics. 17-20. Male genitalia of Orthocomotis with valvae spread, aedeagus removed and shown in lateral aspect, and vesica everted. 17, O. chaldera; 18, O. herbaria; 19, O. phenax; 20, O. similis. Monteverde, 1000-1350 m, Sep 1994 (1 4), Z. Fuentes (INBio). Est. Geographic and temporal distribution. The holo- Biol. Las Alturas, Coto Brus, 1500 m, Aug 1991 (1 4, 1 2), M. Ramirez type (BMNH) is from Colombia: however, all subse- (INBio). Humedal San Joaquin, 1000 m, 10-12 Sep 1996 (1 3), A. é Maroto, M. Moraga, L. Angulo & E. Navarro (INBio). Coto Brus, quently reported See tS (Le., Clarke 1965, Razowski Zona Prot. Las Tablas, Est. Biol. Las Alturas, 1550 m, 16-23 Mar 1999 & Becker 1990) are from Costa Rica. In Costa Rica this (1 2). E. Phillips (INBio). species occurs from the Central Cordillera west, from wa Pa SEs. Ss mea INS OR nS ee VOLUME 57, NUMBER 4 about 700 to 1500 m elevation (Fig. 36). It has been col- lected in all months: January (n = 1), se (im = 8), March (n = 1), April (n = 2), May (n = ), June ( n = 6), July (m = 3), August (n = 3), September Ae = 8), October (n = 6), November (n = 2), and December (n = 3). Remarks. Clarke (1956) referred to specimens of O. magicana collected by William Schaus from Juan Vifias, Mount Pods, and Cachi. I examined the speci- men from Juan Vifias (above); the specimen from Mount Pods may be the one I refer to O. herbacea (above); and the location of the specimen from Cachi is unknown to me. Orthocomotis chaldera (Druce) (Figs. 1@, 1b, 17, By, Sy) Grammophora chaldera Druce, 1889:259. Tortrix chaldera: Walsingham 1914:278. Eulia chaldera: Meyrick 1912:38, 1913:38, 1926:249. Orthocomotis chaldera: Clarke 1956:145; Razowski & Becker 1990:351. Holotype ¢, Costa Rica, San José Province, Volcan de Irazii, 6-7000’, [no chic, Godman-Salvin Collection, H. Rogers, BMNH. Diagnosis. Orthocomotis chaldera is the largest and most commonly collected Orthocomotis in Costa Rica. It can be distinguished from all other Costa Ri- can congeners by its large size (i.e., mean FW length = 14.3 mm in the male, 18.0 mm in the female) and dis- tinct forewing pattern, with a large white blotch in the apical region. A few male specimens from Estacién Mengo, Estacion Cacao, and Turrialba are smaller and darker, with less white scaling, approaching O. eu- chaldera Clarke (from Venezuela) in general aspect. However, the genitalia of these specimens are indistin- guishable from those of other O. chaldera. Hence, I am provisionally including them under this name. Male genitalia are characterized by a relatively short, stout uncus, and socii that have a limited dorsal arch beyond their point of attachment to the tegumen. The female genitalia, with an extremely broad ductus bur- sae immediately posterior to the region of bifurcation, also are fairly distinct. Specimens examined. Cartago Province: Rio Grande de Orosi, desde Puerte Rio Dos Amigos, hasta la Represa, 1400-1800 m, 22 Aug—15 Sep 1995 (1 2), R. Delgado (INBio). Tapanti, Rio Grande de Orosi, 1300-1400 m, 23 Jan 1985 (2 ), D. Janzen & W. Hallwachs (INBio). Orosi, 4000’, “.15” [1915] (1 9), [no collector] (BMNH). Paraiso, P.N. Tapanti-Macizo de la Muerte, 300 m SE del Refugio Porras, 1660 m, May 2000 (1 2), R. Delgado (INBio). Paraiso, P.N. Tapanti, Sect. La Represa, 300 m S del Puente del R. Porras, 1660 m, Nov 1999 (1d), R. Delgado (INBio), Feb 2000 (1 ¢), I. Chavarrfa (INBio). Paraiso, P.N. Tapanti-Macizo de la Muerte, al costado de Oficina [or casa] Admin., 1200 m, Nov 1999 (1 2), Jan 2000 (1 d), R. Delgado (INBio). Paraiso, P.N. Telpemiee Macizo de la Muerte, Est. Quebrada Segunda, 1300 m, Jul 2000 (1 2), R. Delgado (INBio). Paraiso, P.N. Tapanti- -Macizo de la Muerte, Est. Ouelimadh Segunda, al costado Ofic., 1200 m, Dec 1999 (1 2), R. Delgado (IN- Bio). P.N. Tapanti-Macizo de la Muerte, 300 m N & 100 m S del Mi- rador, 1350 m, Oct 1999 (2 ¢), Dec 1991 (14), R. Delgado (INBio), Novy 2000 (1d, 1 2), R. Delgado (INBio). P.N. de la Muerte, 300 m N 261 Mirador, 1480 m, Feb 2000 ( 1 2), R. Delgado (INBio). Tapanti, 1500 m, 30-31 Aug 2000 (1 ), V. noes (VBC). Voleén Turrialba, 1800 m, 13 Aug 1972 (2 ), V. ae (VBC). Santa Cruz, Turrialba, 1500 m, Aug 1981 (1 2), V. Becker (VBC). Juan Vifias, [no date] (1 ¢), W. Schaus (USNM). Guanacaste Province: Derrumbe, Est. Mengo, W side Volcan Cacac ao, 1400 m, 5 Jun 1988 (1 ), 11 Jul 1988 (4 2), 26-27 May 1992 (1 4), D. Janzen & W. Hallwi achs (INBio). Est. ne a- cao, S side Volcan Gee 1000-1400 m, Jul-Aug 1991 (1 ¢), A. Ses INBio). Estac. Pitilla, 9 km $ Santa Cecile a, 700 m, Feb 1989 (1 3), GNP Biodiversity puryeyi\ INBio). Est. Mengo, SW side ee Cacao, 1100 m, Feb 1989 (2 2), GNP Biodive -rsity Surv ey (IN- Bio). Sector Las Pailas, 4.5 - SW ae Rincén de la Vieja, 800 m, 23 Jul-6 Aug 1995 (1 d, 1 2), K. Taylor (INBio). Heredia Province: Est. Barva, PN. Braulio Cae 2500. m, Jan 1990 (1 2), G. Rivera (IN Bio), Silex 1990 (1 ), May 1990 (2 d, 1 2), A. Fernandez (INBio), Jun 1990 (1 2), B. a f G. Varela (IN Bio). El Angel Waterfall, 8.2 km dovralnill : ara Blanca, 1350 m, 3 Jan 1981 (1), 5 Aug 1981 (1 ), D. Janzen & W. Hallwachs (INBio). Mount Pods [2350 m], [no date] (1 3, 1 2), W. Schaus (USNM). 16 km SSE La Virgen, 10°16’N, 84°05’W, INBio-OET-ALAS transect, 1050-1150 m, 11-12 Feb 2001 (1 3), M. Epstein (INBio), 15 Apr 2001 (1 2), 16 Apr 2001 (1 5), J. Brown (INBio). Cerro Chompipe, Ae Biol. cane R. F. Cord. Vol. Cent., 2100 m, 11 Jul 1991 (1 ¢), Oct 1991 (1 2), J. Cor- rales (INBio). Puntarenas Province: o Amistad, ae Altamira Cerro Biolley, A.C. Amistad, 1800 m, ee He (1d), Jan 1994 (2 d), R. Delgado (INBio), 13-26 May 1996 ( _R. Villalobos ee Est. Altamira, Buenos ca 1400 m, ie ee 14 Oct 1993 (2 4), R. Delgado (INBio), Jul 1994 (1 ¢) R. Delgado (INBio). Buenos Aires, PILA, Sector Altamira, A.C. pene 1150-1400 m, Jun 1994 (1 2), R. Delgado (INBio). La Amistad, Sect. Altamira, Buenos Aires, Dec 1993 (1 ¢), R. Delgado (INBio). Est. Altamira, ae SE Cerro Biol- ley, PILA-ACLA, 1450 m, 26 Feb-10 Mar 1995 (1 d), M. Segura (IN- Bio). Est. Altamira, | km $ Cerro Biolley, 1300-1450 m, 28 Juz 7 Aug 1995 (1 2), R. Villalobos (INBio). Est. Pittier, PILA-ACLA, ee m, 5-18 Jan 1995 (1d), M. Moraga (INBio), 23 Aug—9 Sep 1995 (1 3), M. Moraga (INBio), 23 Aug— 13 Sep 1995 (1 d), E. Navarro TNE Sep 1995 (1 GC) seks Navarro (INBio), 13-26 May 1996 (1 2), R. Villallobos (INBio). Est. Pittier, 1670 m, 22 Sep—9 Oct 1995 (1 ¢), M. Moraga (INBio). Est. Pittier, Alrededor de la Estacién, 1670 m, 18-20 Jan 1996 (1 2), M. Moraga (INBio). Est. Biol. Las Alturas, Coto Brus, 1500 m, Aug 1991 (3 d, 2 2), M. Ramirez (INBio). ACLAP, Coto Brus, Zona Prot. Las Tablas, Est. Biol. Las Alturas, 1550 m, 16-24 Mar 1999 (1 2), B. Espinoza (INBio). Sende ro a Cerro Pittier, 1 km N Estaci6n, 1800-2000 m, 11—25 May 1997 (1 3), M. Moraga (INBio). Sendero a Cerro Pittier, 600 m NE ae 1750 m, 5-11 Mar 1997 (12), M. Moraga (INBio). Fea. Cafrosa, Est. Las Mellizas, P.N. La Amistad, ee m, Oct 1989 (1 ¢), M. Ramirez & G. Mora (INBio), May 1991 (1 2), M. Ramirez (INBio). Fea. Cafrosa, Embalse, 800 m N vee a, 80 m, 15 Jul 1996 (1 ¢), L. An- gulo (INBio), 10-29 Jul nee 2), E. Navarro (INBio). Monteverde, 1400 m, 22— ee Jul 1990 (1 18 Meredith & J. Powell (UCB), 29-31 Mar 1992 (1 d), J. McCarty By; Powell (UCB), 30 Mar 1992 a Te Powell Gay 2 km E Monteverde, 1500 m, 31 Mar 1992 (2 2), J. McCarty & J. Powell (UCB). Buen Amigo, San Luis Nentcverds 1000-1350 m, Apr 1995 (1 ¢ }), Z. Fuentes (INBio). Monteverde area, 1400-1700 m, 6-14 Jun 1973 (1 ¢), T. eae & G. Hevel (USNM). Monteverde, 1500 m, 14 Sep 1999 s §), V. Becker (USNM). Mon- teverde, 1400 m, 12-15 Jun 1974 (2 oN Watson (BMNH), 25-26 Jun ee 1d), D. ee (INBio), 15- AG May 1980 (3 3), 30-31 Jul 1981 (1 2), 3 jan 1984 (1 ), D. Janzen & Me Hallwachs (INBio). 2 km E edie ou B ie 1988 (1 2), J. Brown & J. Powell (IN- Bio), 31 Mar 1992 (1 2), J. McCarty KJ. Powell ( UCB). Est. La Ca- sona, Res. Biol. ae 1520 m, Mar ne (1 d), Aug 1991 (1 3), N. Obando (INBio), 30 Jan—18 Feb 1995 (1 2), K. Martinez (IN- Bio). Las Nubles, 11 km NW Monteverde, ef Jul 1981 (2 3), D. Janzen & W. Hallwachs (INBio). Monteverde, 1500 m, 1-4 Sep 1999 (6d), V. Becker (VBC). Alturas de Cotén, 1500 m, 15 Sep 1999 (1 <), V. Becker (VBC). San José Province: Est. Zurqui (el tunel), P.N. Braulio Carrillo, 1500 m, Aug 1985 (3 ¢), Oct 1985 (2 ¢), 1. & A. Chacén (INBio). Est. Santa Elena Viejo, Santa Elena, Las Nubes, 1210 m, 29 Sep 1995 (1 3), A.M. Mardo (INBio). Irazu, 6—-7000’, [no 262 ts rh Lal JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 21-22. Male genitalia of Orthocomotis with valvae spread, aedeagus removed, and vesica everted. 21, O. nitida; 22, O. altivolans. date] (HT 3), Godman-Salvin Collection, H. Rogers (BMNH). Un- known Province: Sixola River, [no date] (1 3), W. Schaus (USNM). Cascajal, ex. Janson, Jan 1924 (1 2), [no collector] (BMNH). Geographic and temporal distribution. Orthoco- motis chaldera ranges from Tamaulipas, Mexico (VBC) south to Ecuador (VBC) and Peri (BMNH). In Costa Rica it has been collected throughout the western half of the country, from 800 to 2500 m, but primarily from 1100-1800 m. It has been recorded throughout the year: January (n = 11), February (n = 7), March (n = 9), April (n = 3), May (n = 13), June (n = 8), July (n = 14), August (n = 18), September (n = 16), October (n = 7), November (n = 4), and December (n = 4). Remarks. Druce (1889) described this species from Volcan Irazti, Costa Rica (ca. 2000 m). According to Clarke (1956), the type should be in BMNH, but he was unable to find it. However, I believe that the spec- imen cited above and labeled “Irazu, 6—7000’, H. Rogers, Godman-Salvin Coll.,’ which I discovered in the undetermined collection at the BMNH, is the type. Orthocomotis herbaria (Busck) (Figs. 5, 6, 18, 28, 38) Sociphora herbaria Busck, 1920:85. Orthocomotis herbaria: Clarke 1956:144. Holotype ¢ (herbaria), Guatemala, Cayuga, Wm. Schaus, USNM. Orthocomotis cristata Clarke 1956:155; Razowski & Becker 1990: 346 (map only), new synonymy Holotype ¢ (cristata), Costa Rica [unknown province], Cachi, [no date], W. Schaus, USNM. Orthocomotis uragia Razowski & Becker, 1990:352, new synonymy Holotype 3 (uragia), Costa Rica, Puntarenas Province, Buenos Aires, 200 m, 25 Nov 1975, V. Becker, VBC. Diagnosis. Orthocomotis herbaria is superficially most similar to O. nitida because of their small size, similar forewing pattern, and dark brown hindwing (Figs. 4-6). However, both sexes of O. herbaria can be distinguished from all other species by the presence of the hindwing pecten (see Table 1); male genitalia (Fig. 18) can be distinguished by the rounded-triangular process that represents the termination of the saccu- lus. The male possesses a thoracic hairpencil and has small cornuti in the vesica. Specimens examined. Alajuela Province: Cerro Campaiia, E slope Volcan Cacao, 650 m, 15 Jun 1988 (1 2), J. Brown & J. Powell (UCB). Area de Conservaci6n Guanacaste, Sector San Cristdbal, Rio Blanco Abajo, ex-larva on Nectandra hihua, 23 May 2001, em: 14 Jun 2001 (1 6), “O1-SRNP-1776,” D. Janzen & W. Hallwachs (USNM). Guanacaste Province: Est. Pitilla, 9 km S Sta. Cecilia, RN. Gua- nacaste, 700 m, 19-23 Jun 1993 (1 4), Jun 1991 (1 3), C. Moraga (IN- Bio). Heredia Province: La Selva Biol. Sta., Puerto Viejo de Sara- piqui, 40 m, Sep 1987 (1 °), M. Chavarria (INBio). Est. Biol. La Selva, 50-150 m, 10°26’N, 84°01’W, 17 Mar 1993 (1 3), 3 Jul 1994 (1 9), ALAS (INBio), Jan 1996 (1 3), J. Powell (UCB), 6 Feb 1996 (1 3), 10 Feb 1996 (1 2), 11 Feb 1996 (1 4), 13 Feb 1996 (1 2), 16 Jan 1998 (13), Jan 1998 (1 2), 26 Jan 1998 (1 9), 15 Apr 1998 (1 2), 16 Mar 1999 (1 d), ALAS (INBio). Lim6én Province: Res. Biol. Hitoy Cerere, Est. Hitoy Cerere, Cerro Bobocara, 770 m, Jun 1999 (1 2), R. Barton (INBio). Manzanillo, RNFS, Gandoca y Manzanillo, 0-100 m, 22 Oct-12 Nov 1992 (1 3), F. Quesada (INBio). Sector Cerro Cocori, Fca. de E. Rojas, Jan 1991 (1 4), Apr 1991 (1 2), E. Rojas (INBio). Puntarenas Province: P.N. Manuel Antonio, Quepos, 80 m, May 1991 (1 3), R. Zuniga (INBio), Aug 1991 (1 d), Oct 1992 (1 3), Nov 1992 (3 d), Oct 1993 (2 3), G. Varela (INBio), Feb 1991 (1 3), R. Zuniga (INBio). Est. Sirena, P.N. Corcovado, 0-100 m, Mar 1991 (1 4), G. Fonseca (INBio). Sirena, Corcovado Nat. Park, Osa Peninsula, 1 May 1984 (1 3), D. Janzen & W. Hallwachs (INBio). Est. Rfo Bonito, 2.3 km W Cerro la Gamba, 110 m, 7-10 Nov 1996 (1 3), E. Fletes (INBio). Rancho Que- mado, Peninsula Osa, Nov 1990 (1 4, 1°), F. Quesada (INBio). A.C.O. Golfito, Reserva Ftal. Golfo Dulce, Proyecto Zamia, Playa Cacao, 130 m, 8-12 Oct 1999 (1 2), 6-11 Nov 1999 (1 4), M. Moraga (INBio). Golfito, 25-28 Apr 1965 (13), S. S. & W, D. Duckworth (USNM). Fila VOLUME 57, NUMBER 4 Esquinas, 35 km S Palmar Norte, 150 m, 7-8 Jan 1983 (1 2), D. Janzen & W. Hallwachs (INBio). Buenos Aires, 200 m, 25 Nov 1975 (1 6, HT d of uragia), V. Becker (VBC). San Jose Province: Est. Bijagual, TGR, Biol. Carara, 500 m, Sep 1990 (1 3), G. Varela (INBio). Unknown Province: Cachi, [no date] (HT 4 of cristata), W. Schaus (USNM). Geographic and temporal distribution. This species ranges from Guatemala (HT of herbaria, USNM) to Costa Rica but is restricted to the lowlands (i.e., from sea level to about 700 m elevation) (Fig. 38). It has been recorded nearly throughout the year: Jan- uary (n = 6), Sees (n = 5), March (n = 3), April (n = 3), May (n = 2), June (n = 4), July (n = 1), August (n = 1), September (n = 2), October (n = 5), and Novem- ber (n = 9). Janzen and Hallwachs (2002) report rearing this species from Nectandra hihua (Lauraceae) in Area de Conservaci6n Guanacaste in northern Costa Rica. Ac- cording to the rearing notes, the caterpillar is brilliant green with white setae and a black and brown head. Remarks. Two slides made by August Busck (USNM) were mixed, apparently based on mislabeling by Clarke or Busck. The slide belonging to the holo- type of O. herbaria (A.B. 1 Feb 1920) was associated incorrectly with a paratype of O. herbacea, and the slide belonging to the O. herbacea paratype (A.B. 18 Feb 1920) was associated incorrectly with the holotype of O. herbaria. Although Clarke's holotype of O. cristata matched the holotype of Busck’s O. herbaria, including the presence of the unique hindwing cubital pecten, the genitalia were clearly different, i.e., the genitalia of Clarke’s specimen were correctly associ- ated, while those of the Busck specimen were O. herbacea. On the basis of the genitalic differences, Clarke described O. cristata, stating that the holotype male of O. cristata is “remarkable for the presence of the cubital pecten,” a feature also present in the holo- type of O. herbaria. On this basis I synonymize the two. Orthocomotis uragia was described from a single extremely worn specimen from Buenos Aires, Costa Rica. A second male taken on the same date was iden- tified correctly by Becker as O. herbaria; apparently Razowski did not see the latter. The genitalia of the holotype match those of O. herbaria, and the presence of cubital pecten provides convincing evidence of their conspecificity. Orthocomotis phenax Razowski & Becker (Figs. 12, 19, 29, 39) Orthocomotis phenax Razowski & Becker, 1990:354. Holotype 4, Costa Rica, San José Province, [Parque Nacional] Braulio Carrillo, 1100 m, Jul 1981, V. Becker, VBC. Diagnosis. Males of O. phenax are most similar to those of O. herbacea. They can be distinguished from 263 all other congeners, except O. similis, by the narrow uncus with fine lateral setae throughout the apical third and the elongate tip of the gnathos. Orthocomo- tis phenax can be distinguished from O. similis by its smaller size, its brighter green forewing overscaling, and the presence of a pale subapical forewing fascia bordered basally by a dark fascia. Specimens examined. Guanacaste Province: Est. Pitilla, 9 km S Sta. Cecilia, P. N. Guanacaste, 700 m, May 1991 (1 ¢), 4-13 Dec 1991 (1 3), C. Moraga (INBio), 6-19 Sep 1993 (1 ¢), Feb 1993 1 2), Jan 1994 (1 2), Nov 1994 (1 2), P. Rios (INBio). Heredia Province: 16 km SSE La Virgen, 10°16’N, 84°05’W, INBio-OET- ALAS transect, 1050-1150 m, 16 Mar 2001 (1 3), 18 Mar 2001 (1 2), 19 Mar 2001 (1 d), D. Wagner & J. Rota (INBio) Puntarenas Province: Fca. Cafrosa, Est. Las Mellizas, P N. Amistad, 1300 m, Jan 1991 (1 2), M. Chavarria & G. Mora (INBio). A.C.L.A.P. Coto Brus, Zona Prot. Las Tablas, Est. Biol. Las Alturas, 1550 m, 15-24 Mar 1999 (1 d), R. Delgado (INBio). Coto Brus, Est. Biol. Las Al- turas, 1550 m, 15-24 Mar 1999 (1 2), G. Rodriguez (INBio). La Es- cuadra, P.N. Amistad, 1340 m, 14 Apr 1989 (1 3), M. Ramirez & G. Mora (INBio). San José Province: Braulio Carrillo, 1100 m, Jul 1981 (5 6, HT 3), V. Becker (VBC). Est. Carrillo, RN. Braulio Car- rillo, 700 m, Jul 1984 (1 2), I. Chacén (INBio). Geographic and temporal distribution. Ortho- comotis phenax is known only from Costa Rica, occur- ring from the Central Cordillera westward, from 700-1550 m elevation (Fig. 39). Capture records are scattered throughout the year. Orthocomotis similis Brown, new species (Figs. 1, 20, 30, 40) Diagnosis. Superficially, O. similis resembles O. ex- olivata from Brazil. However, the genitalia of both the male and female of the former (Figs. 20, 30) are virtu- ally indistinguishable from those of O. phenax. Ortho- comotis similis can be separated from O. phenax by its larger size (mean FW length = 11.5 mm and 14.2 mm for males and females of similis, respectively, vs. 10.8 mm and 12.9 mm for phenax), the more subdued green overscaling of the forewing, the presence of an oblong dark spot from the forewing dorsum near the tornus, and the more pale brown scaling of the hindwing. Razowski and Becker (1990) described O. phenax phobetica from Veracruz, Mexico, and commented that although described as a subspecies because of its similarity to O. phenax, it “probably represents a dis- tinct species, very close to the proceeding [O. phenax].” It is possible that O. phenax, O. phenax pho- betica, and O. similis together represent a complex of closely related species. Alternatively, the three may represent variation within O. phenax. For the present, I prefer the former hypothesis. Description. Male. Head: Upper frons cream with a pair of lat- eral red-brown tufts; lower frons pale cream. Labial palpus mostly cream on inner surface, mostly brown on outer surface. Antenna with brown scales on dorsum of basal two-thirds; cilia ca. 0.5—0.6 times width of flagellomere. Thorax: Mostly pale brown and cream with a few red-brown scales; cream tuft at posterior end. Hairpencil of 20-30 pale cream to white elongate scales originating near base of JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY NO O1 aaanndan M00 Raa nn ~ ee o NO OD) Fics. 23-27. Female genitalia of Orthocomotis. 23, O. ochracea; 24, O. herbacea; 25, O. longicilia,; 26, O. magicana; 27, O. chaldera. VOLUME 57, NUMBER 4 hindwing, extending to second abdominal segment. Forewing length 10.5-12.5 mm (x = 11.5; n = 4) (Fig. 1); ground pale brown with ir- regular patches of metallic green and darker brown overscaling; ground color interrupted by variously defined, pale fascia; subapical fascia from tornus, bifurcating near the upper edge of the DC with one bifurcation extending to costa about 0.65 distance from base to apex, and the other bifurcation curving to costa ca. 0.2 distance from base to apex. Hindwing pale brown, lacking cubital pecten. Ab- domen: Densely clothed with long, fine, pale brown scales; second segment with a pair of shallow lateral pouches, each bearing two rows of dense secondary sex scales; dorsum of segments 2 and 3 with paired subdorsal pits. Genitalia as in Fig. 20 (drawn from JWB preps. 1255 and 1258; n = 4). Uncus comparatively long, broad at base, slender in distal two-thirds, straight, with dense, short, some- what evenly spaced, lateral setae, ca. 15 on each side. Socius large, densely scaled. Gnathos slender, with long terminal portion ending in narrow, slightly hooked tip. Transtilla a slender, gently arched band. Valva short, broad, subrectangular, with rounded distal por- tion; neither costa nor sacculus developed. Aedeagus short, stout, curved just beyond ductus ejaculatoris, with a pair of subdorsal, api- cal, sclerotized prongs, one larger, separated from remainder of coe- cum by membranous region; vesica densely covered by small, short cornuti. Female. Head and thorax: Essentially as described for male, except antennal cilia shorter, more sparse, and hairpencil absent. Forewing length 12.5-16.0 mm (x = 14.2; n = 3); pattern essentially as described for male. Abdomen: Gantella as in Fig. 30 (drawn from JWB prep. 1159; n = 3). Sterigma simple with huge, ovoid os- tium. Ductus bursae moderately Iboel moderately long, with ac- cessory sac from left side immediately posterad junction with corpus bursae. Corpus bursae an ovoid sac; signum absent. Holotype 3, Costa Rica, Est. Cacao, S side Volcan Cacao, 1000-1400 m, 8—29 Jul 1991, C. Chaves (INBio). Paratypes. Cartago Province: P.N. Tapanti-Macizo de la Muerte, 300 m N & 100 m S Mirador, 1350 m, Oct 1999 (1 ¢), R. Delgado (INBio). Paraiso, P.N. Tapanti-Macizo de la Muerte, 300 m SE Rio Porras, 1600 m, Nov 2000 (1 2), R. Delgado (INBio). Tapanti, 1200-1700 m, 20 Aug—15 Sep (1 2), V. Becker (VBC). Gua- nacaste Province: Derrumbe, Est. Mengo, W side Volcan Cacao, 1400 m, 5 Jun 1988 (1 2), D. Janzen & W. Hallwachs (INBio). Est. Mengo, SW side Volcan Cacao, 1100 m, Feb 1989 (1 2), GNP Bio- diversity Survey (INBio). Est. Cacao, S side Volcan Cacao, 1000-1400 m, Jun 1990 (1 2), If Curso Paratoxonomia (INBio), 8-29 Jul 1991 (1 2), C. Chaves (INBio). Heredia Province: Braulio Carrillo Nat. Park, 6 km ENE of Vara Blanca, 10°11’N, 84°07’ W, INBio-OET-ALAS transect, 2000 m, 16 Feb 2002 (1 2), J. Brown & J. Powell (INBio). San José Province: Est. Zurqui, 50 m antes de tunel, 1600 m, 26 Sep-Oct 1990 (1 2), G. Maass (INBio). Geographic and temporal distribution. Orthoco- motis similis is known only from the Central Cordillera of Costa Rica; captures range from 1000-1600 m eleva- tion. Adults have been collected in February, June, July, August, September, October, and November. Etymology. The specific epithet refers to the simi- larity between the genitalia of the new species and those of O. phenax. Orthocomotis nitida Clarke (Figs. 4, 21, 31, 41) Orthocomotis nitida Clarke, 1956:143; Razowski & Becker 1990:346 [referred to in legend of map, but no locations given]. Holotype ¢, Guatemala, Cayuga, “4”, Schaus & Barnes, USNM. Diagnosis. Orthocomotis nitida is one of the small- est members of the genus. It can be distinguished from its congeners by the moderately large, dark, rectangular blotch from near the middle of the forewing costa; the distinctly bicolored frons: cream yellow in the lower half and red brown in the upper; and the color of the scaling of the labial palpi: dark cream yellow, except for the outer edge which is fawn brown. The capitate, strongly spined uncus and the shape of the valva in the male genitalia are characteristic of this species (Fig. 21). Specimens examined. Alajuela Province: Cerro Campana, E side Volcan Cacao, 6 km NW Dos Rios, 650 m, 15 Jun 1988 (1 ¢), D. Janzen & W. Hallwachs (INBio). Guanacaste Province: Derrumbe, Est. Mengo, W side Volcan Cacao, 1400 m, 11 Jul 1988 (1 2), D. Janzen & W. Hallwachs (INBio). Heredia Province: Est. Biol. La Selva, Puerto Viejo de Sarapiqui, 50-150 m, 10°26’N, 84°01’W, 11 Jan 1986 (1 2), D. Janzen & W. Hallwachs (INBio), 7 Feb 1996 (1 2), 10 Feb eae 3), 17 Feb 1996 (1 3), 11 Mar 1996 (1 2), 17 Apr 1996 (1 d), 14 Jan 1998 (1 3), 20 Jan 1998 (1 3), 20 Feb 1998 (1 3), 3 Mar 1998 (2 3), 5 Mar 1998 (1 d), 16 Mar 1998 (1 d), 9 Feb 1999 (1 3), ALAS (INBio), Jan 1998 (1 2), J. Powell (UCB). Lina Province: 9.4 km W Bribri, Suretka, 200 m, 9-11 Jun 1983 (1d), D. Janzen & W. Hallwachs (INBio). ea ieee a Sirena, P. N. Corcovado, 0-100 m, Mar 1991 (1 2), Jul 1991 (1 ), Mar 1993 (1 2), G. Fonseca (INBio). Est. aa, N, C.O. ae PN. Corcovado, 0-100 m, 13-22 Mar 1980 (1 ¢), D. Janzen (INBio). Fca. Cafrosa, Est. Las Mellizas, RN. La Amistad, 1300 m, Nov 1990 (1 3), M. Ramirez & G. Mora (INBio). Golfito, PN. Piedras Blancas, Est. El Bonito, 100 m, Jan-Feb 2002 (1 3), M. Moraga (INBio). Unknown Province: V. Neilly, 800 m, 26 Nov 1973 (1 2), V. Becker (VBC). Geographic and temporal distribution. This species ranges from Guatemala (HT, USNM) to Ecuador (BMNH). It appears to be confined to lower elevations with a majority of the captures from 200 m or below, but there are single records from 800, 1300, and 1400 m. Captures range from January through July, with two records from November. The majority of records (14 of 23 specimens examined) are from La Selva Biological Station, from January through March. Orthocomotis altivolans Brown, new species (Figs. 7, 22, 32, 42) Diagnosis. Orthocomotis altivolans can be distin- guished from its congeners by its forewing color and pattern: copper to cinnamon, with irregular yellow gold to pale green gold (rather than metallic green or kee) bands and spots surrounded by white scaling (Fig. 7). Also, the white hindwing of O. altivolans is unusual in the genus. The valvae are extremely simple and more attenuate than in most other Orthocomotis, reminiscent of those in males of Argentulia Brown. The aedeagus lacks cornuti on the vesica. Description. Male. Head: Upper frons copper to cinnamon, lower frons pale orange. Labial palpus pale cinnamon on inner sur- face, slightly darker on outer surface. Antennal cilia 0.8—0.9 times width of flagellomere. Thorax: Dorsum copper to cinnamon, slightly paler a posterior end, with weakly developed tuft; metatho- rax with dense hairpencil of 25-30 elongate pale cream scales ex- tending to pouch in second abdominal segment. Forewing length 12.0-13.5 mm (x = 12.6; n = 8) (Fig. 7); oround color copper to cin- namon, divided by a group of variably Connected narrow, sinuous, 266 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 28-32. Female genitalia of Orthocomotis. 28, O. herbaria; 29, O. phenax; 30, O. similis; 31, O. nitida; 32, O. altivolans. VOLUME 57, NUMBER 4 33 fo} = NR Ww + Leal fon} oO Oo (2) fo) io) Oo 0 250 500 750 1000 1250 15001750 2000 2250 2500 2750 3000 35 Se wm Sb OO @ 6 0 250 500 750 1000 1250 15001750 2000 2250 2500 2750 3000 Sif See ay CO OO GS SG ee Ss 0 250 500 750 1000 125015001750 2000 2250 2500 2750 3000 39 15 12 Sa & MH © © Q 250 500 750 1000 125015001750 2000 2250 2500 2750 3000 41 Oo WO Gf © SF & & 0 250 500 750 1000 1250 15001750 2000 2250 2500 2750 3000 Fics. 33-42. Elevational distribution of Orthocomotis; x-axis i 34 25 20 15 10 5 (0) 0 250 500 750 1000 125015001750 2000 2250 2500 2750 3000 60 36 50 40 30 20 10 0 0 250 500 750 1000 125015001750 2000 2250 2500 2750 3000 ie 38 8 6 4 2 0 0 250 500 750 1000 125015001750 2000 2250 2500 2750 3000 6 40 5 4 3 2 1 (0) 0 250 500 750 1000 125015001750 2000 2250 2500 2750 3000 18 42 15 12 9 6 3 0 OQ 250 500 750 1000 125015001750 2000 2250 2500 2750 3000 = elevation in meters, y-axis = number of individuals examined. 33, O. ochracea; 34, O. herbacea; 35, O. longicilia; 36, O. magicana; 37, O. chaldera; 38, O. herbaria; 39, O. phenax; 40, O. similis; 41, O. nitida; 42, O. altivolans. white fascia with yellow gold to pale green gold overscaling. Pattern usually including a complete fascia ‘extending from TSE ca. 0.65 distance from Bese to apex, to tornus; frequently with less defined incomplete fascia from costa ca. 0.33 distance from base to apex, and a sinuate fascia from costa at base, extending to dorsum ca. 0.25 distance from base to tornus, then angled tow: ard apex of discal cell. Hindwing whitish with scattered pale gray overscaling, cubital pecten absent. Abdomen: Dorsum less densely scaled than in most congeners: second segment with a pair of shallow lateral pouches, each bearing two rows of dense secondary sex scales: dorsum of segments 2 and 3 with paired subdorsal pits: venter of segments 1-2 with strongly sclerotized V-shaped region: dorsum of segment 8 with narrow, sclerotized crescent-shaped ridge. Genitalia as in Fig. 22 (drawn from USNM slide 92702 and JWB prep. 1267: n = 3). *Taers broad in basal 0.33, slightly flattened dorsov: entrally and densely setose in 0.66, strongly curved at ca. 0.33 distance from base to apex. Gnathos narrow, with fine, pointed process at distal junction of arms. Socius large, parallel-sided, with long dense scal- ing; lateral edge conspicuously sclerotized. Valva extremely simple, costa mostly straight, ventral edge evenly curved, apex rounded: sacculus not developed: costa sclerotized. Transtilla a narrow, simple, sclerotized bridge. Anellus with large bristly portion be- tween transtilla and aedeagus, strongly attached to dorsum of aedeagus. Juxta teardrop-shaped. Aedeagus simple, weakly curved just beyond ductus ejaculatoris; vesica with cornuti represented by tiny punctations. Female. Head and thorax: Essentially as described for male, except antennal cilia short, sparser, and thorax without hairpencil. Forewing length 13.3-15.0 mm (X = 14.2: n = 5); pattern as is male. Abdomen: Densely clothed with long, fine pale brown scales: dor- sum of segments 2 and 3 with paired subdorsal pits, without lateral pouches. Genitalia as in Fig. 32 (drawn from JWB prep. 1268; n = 3). Sterigma simple, weakly sclerotized: ostium ovoid-rounded. Ductus bursae relativ ely broad, ‘short, gradually widening into corpus bursae: an oblong accessory bursae originating from junction of corpus bursa and ductus bursa. Corpus bursae pear-shaped, without spicules: a small rounded accessory sac arising near middle of corpus. Holotype 3, Costa Rica, San José Province, Est. Cuerici, por Quebrada Los Leones, 4.5 km E Villa Mills, 2600 m, 7-10 Dec. 1996, A. Picado (INBio). Paratypes. Alajuela Province: Mount Pods. 2350 m, [no date] (1 2), W. Schaus (USNM), 15 Dec 1982 (1 9), D. Janzen & W. Hallwachs (INBio). Paraiso, P.N. Tapanti-Macizo de la Muerte, 300 m SE Rio Porras, 1660 m, Feb 2000 (1 ¢), Nov 2000 (1 2), R. Del- gado (INBio). Cartago Province: Fca. Los Lagos, 2600 m, 8 Jun 1994 (1 ¢), M. Chavarria (INBio). P.N. Tapanti, El] Guarco, A Isidro. Est. ae 2450-2700 m, Age 2000 (1 ¢), May 2000 ( 3), 28 Feb 2001 (1 3), May 2001 (1 3), R. Delgado CNBia = Guarco, Villa Mills- pees 2840 m, ner Oct 2000 (3 ¢), R. Del- gado (INBio). El ee Bae de la Muerte. Sector de k Esper- anza, 2600 m, Jun 2001 (1 3), ful 2001 (1 2), May 2002 (2 3), R. Del- gado (INBio). Villa ae 2840 m, 26-28 Oct 2000 (2 3), Vv. Becker (VBC). Paraiso, PN. Tapanti- ‘Macizo de la Muerte, 300 m SE Rio Porras. 1660 m. Jan 2000 (1 ¢), R. Delgado (INBio). R.F. Rio Ma- cho, El Guarco, 500 m E Est. de la Esperanza, 2600 m, 13-14 May 2002 (2 3), J. Jiménez & E. Phillips (INBio). R.F. Rio Macho, El Guarco, Macizo de la Muerte, Sector de la Esperanza, 2600 m, Aug 2001 (1 2), R. Delgado (INBio). R.F. Rio Macho, El Guarco, Macizo de la Muerte, 2600 m, Oct 2001 (3 2), R. Delgado (INBio). P.N. Tapanti-Macizo de la Muerte, Est. de la Esperanza. 2600 m, Sep 2002 (1 2), R. Delgado (INBio). P.N. Tapanti, 1200-1700 m, 20 Aug—15 Sep 1999 (1 ¢), V. Becker (VBC). Heredia Province: Est. Barva, P.N. Braulio Carrillo, 2500 m, Nov 1989 (1 2), A. Femandez (INBio). Braulio Carrillo Nat. Park. 6 km ENE of Vara Blanca, 10°11’N, 84°07’W, INBio-OET-ALAS transect, 2000 m, 20 Feb 2002 (1 2), malaise trap (UCB). 6 km ENE Vara Blanca, 2000 m, 7 Oct 2002 (1 ¢), K. Nishida, MV light (USNM). Limon Province: Batsi, Valle del Silencio, 2472 m, 11-12 Oct 2000 (1 2), R. Delgado (INBio). San José Province: Est. Cuerici, 4.6 km E Villa Mills, 2600 m, 21-25 Sep 1995 (1 ¢), A. Picado (INBio). Est. Cuerici, JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Sendero al Mirador, 4.6 km E Villa Mills, 2640-2700 m, 19-20 Apr 1996 (1 2), B. Gamboa (INBio), 21 Jun 1996 (1 2), A. Picado (IN- Bio), 20-22 Jan 1996 (1 ¢), B. Gamboa (INBio). Est. Cuerici, por Quebrada Los Leones, 4.5 km E Villa Mills, 2600 m, 7-10 Dec. 1996 (1 2), A. Picado (INBio). Unknown Province: Cascajal, ex. Janson, Jan 1924 (1 ), [no collector] (BMNH). Geographic and temporal distribution. Ortho- comotis altivolans occupies the highest elevations of any of the Costa Rican Orthocomotis, ranging primar- ily from 2500 to 2700 m; there are a few records from 1600 m (Fig. 42). Captures are scattered throughout the year, with no evidence of a defined flight period. During a week of collecting near Vara Blanca (Heredia Province) in February 2002, we saw no specimens of this species at mercury vapor light or in blacklight traps, but found numerous individuals in malaise traps. Etymology. The species name refers to the fact that the species occurs in high elevations. ACKNOWLEDGMENTS I thank the following for allowing me to examine material in their care: Eugenie Phillips (INBio). Jerry Powell (UCB), Kevin Tuck (BMNH), and Vitor Becker (VBC). The following provided com- ments on the manuscript that improved its clarity and quality: Eu- genie Phillips, Instituto Nacional de Biodiversidad, Santo Domingo, Heredia, Costa Rica: David Nickle, Systematic Entomology Labora- tory, USDA, Beltsville. Maryland, U.S.A.; David Smith, Systematic Entomology Laboratory, National Museum of Natural History, Smithsonian Institution, Washington, D.C., U.S.A.; and Jerry Powell, University of California, Berkeley, U.S.A. Illustrations of the genitalia were prepared by David Adamski, USDA, Systematic En- tomology Laboratory, W ashington, DG. and arranged by Linda Lawrence, USDA. Systematic Entomology Laboratory, W: ashington, D.C. The photograph of the adults was prepared by David W. agner, University of Connecticut, Storrs. Harriett Bramble assisted in or- ganizing the data for the histograms of elevational occurrence. Field work in Costa Rica was supported in part by grants from NSF (ALAS IV, the Arthropods of La Selva) and National Geographic So- ciety to Jack Longino, Evergreen State College, Olympia. Washing- ton. Visits to INBio were funded in part by that institution. LITERATURE CITED Brown, J. W. 1989. New tribal assignment for Orthocomotis Dognin and Paracomotis Razowski based on an unusual modifi- cation of the chaetosema (Lepidoptera: Tortricidae). Pan-Pacif. Entomol. 65:454—457. Brown, J. W. & J. A. POWELL. 1991. Systematics of the Chrysoxena group of genera (Lepidoptera: Tortricidae: Euliini). Univ. Calif. Publ. Entomol. 111. 87 pp: + figs. ; Busck, A. 1920. Descriptions of new Central American Microlepi- doptera. Insec. Inscit. Mentr. 8:53—95. CarkE, J. F. G. 1956. Neotropical moths of the genus Orthoco- motis Dognin (Lepidoptera: Tortricidae). Tone. Royal Ento- mol. Soc. fiandat 107 (1955):139-168. Deuce, H. H. 1889. Lepidoptera, Heterocera, volume 1. In God- man & Salvin (eds.), Biologia Centrali-Americana, Insecta. Horak, M. 1984. Assessment of taxonomically significant struc- tures in Tortricinae (Lep., Tortricidae). Mitt. Schweiz. Entomol. Gesell. 57:3-64. 1999. Tortricoidea, pp. 199-215. In Kristensen, N. (ed.), Lepidoptera, moths and butterflies. Vol. 1. Evolution, systemat- ics, and biogeography. Handbook of Zoology 4 (35), Arthro- poda: Insecta. Walter de Gruyter, Berlin & New York. VOLUME 57, NUMBER 4 JANZEN, D. H. & W. Hatiwacus. 2002. Caterpillars and their adults (Lepidoptera) of the Area de Conservacién Guanacaste (ACG), Costa Rica. Web site http://janzen.sas.upenn.edu/index. html. Meyrick, E. 1912. Family Tortricidae. In Wagner, H. (ed.), Lepi- dopterorum Catalogus, Lepidoptera, Heterocera 10. 86 pp. . 1913. Family Tortricidae. In Wytsman, P. (ed.), Genera In- sectorum, Lepidoptera, Heterocera 149. 81 pp. . 1926. Exotic Microlepidoptera 3 (8):225-256. 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, Chlidanotinae, pp. 138-151. In Heppner, J. B. (ed.), Atlas of Neotropical Lepidoptera, checklist part II: Hy- blaeoidea—Pyraloidea—Tortricoidea. Association for Tropical Lepidoptera, Scientific Publishers, Gainesville, Florida. 269 RAzOwsKI, J. 1982. Notes on Orthocomotis Dognin (Lepidoptera: Tortricidae) with descriptions of new taxa. Bull. Acad. Polon. Sci., Ser. Sci. Biol. 30:29-35. . 1999. Tortricidae (Lepidoptera) from the Dominican Re- public. Acta Zool. Cracoy. 42:307-319. Razowskl, J. & V. BECKER. 1990. Descriptions and notes on Or- thocomotis Dognin (Lepidoptera: Tortricidae). Acta Zool. Cra- cov. 33:345-365. WALSINGHAM, LORD T. DE Grey. 1914. Lepidoptera, Heterocera. Vol. 4. Tineina, Pterophorina, Orneodina, and Pyralidina and Hepialidina (part). In Godman & Salvin (eds.), Biologia Centrali- Americana, Insecta. 482 pp. + 10 color plates. ZELLER, P. C. 1866. Beschreibung einger amerikanischen Wickler und Crambien. Stett. Entomol. Zeit. 27:137—157. Received for publication 5 December 2002; revised and accepted 22 April 2003. Journal of the Lepidopterists’ Society 57(4), 2003, 270-273 SCLEROCONA ACUTELLA (EVERSMANN) (CRAMBIDAE: PYRAUSTINAE), NATURALIZED ALONG THE EASTERN SEABOARD Davip L. WAGNER Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, Connecticut 06269, USA. E-mail: david. wagner@uconn.edu Douc C. FERGUSON! Systematics Entomology Laboratory, USDA, Agricultural Research Service, c/o National Museum of Natural History, Washington, DC 20560-0168, USA JOHN D. GLASER 6660 Loch Hill Road, Baltimore, Maryland 21239, USA ABSTRACT. _ Sclerocona acutella (Eversmann), a Eurasian species previously undocumented from North America, is reported from 14 lo- calities in Connecticut, Massachusetts, Maryland, and Rhode Island. The first North American specimen was captured in 1984 at a coastal lo- cation in Bristol, Massachusetts. Capture and rearing records suggest the moth feeds on common reed, Phragmites australis (Cav.) Trin. ex Steud. (Poaceae), in freshwater and brackish wetlands. Given the increasing abundance of Phragmites along water courses and some upland habitats, Sclerocona stands to become one of the region’s most common wetland moths. Additional key words: alien species, Phragmites, biocontrol, wetlands. Sclerocona acutella is a large, easily recognizable, wetland pyraustine. It is native to Europe and Asia, oc- curring from Spain and Sicily northward to Great Britain (as a stray) and Denmark (rare) east to Siberia, Japan, and China (Inoue et al. 1982, Palm 1986, Karsholt & Razowski 1996, Parenti 2000, Siberian Zo- ological Museum 2002). Ongoing surveys of Lepi- doptera in southern New England and Maryland re- vealed the presence of Sclerocona in a variety of wetlands along the eastern sea board of the United States, seelnd hag estuaries, marshes, fens, swamps, and lake and pond margins. We assume that the species was introduced accidentally, but because larvae feed on Phragmites australis (Cav.) Trin. ex Steud. (Poaceae), an aggressively invasive plant in many northeastern wetlands, there remains the possibility that it was purposefully introduced, although we were unable to locate any literature indicating such. Voucher specimens are deposited in the following in- stitutions and personal collections: John D. Glaser (JGC), Lloyd Center for Environmental Science (LCES), University of Connecticut (UCONN), and Uni- versity of Rhode Island Biological Control Lab (URI). Selected references and a diagnostic description follow. Sclerocona acutella (Eversmann) Crambus acutellus Eversmann, 1842:563. Sclerocona acutella Meyrick, 1890:445. Meyrick proposed Sclerocona with Crambus acutellus Eversmann as the only included species. ‘Deceased. Sclericona acutellus, Marion, 1957:82. Sclerocona acutellus, Palm, 1986:231. Sclerocona acutella, Karsholt & Razowski (eds.), 1996:194. Diagnostic description. Length of forewing (males): 11.5-12.5 mm. The light brown adults of this species superficially resemble their namesake, Nascia acutella (Walker) of eastern North America (see Munroe 1976: pl. 1, figs. 67-71), but Sclerocona adults have much longer porrect palpi, glossy white wing fringe scales, a pale costa, and lack the distinctive pale intervenular streaks of the forewing of Nascia. The unique forewing venation (Fig. 5) of S. acutella males is probably diagnostic within our fauna. The cubital stem bends deeply into the posterior side of the discal cell and back again, reducing the proximal half of the cell to only half the width of the distal half. This makes space, as it were, for a curiously enlarged and modified, comu- copia-shaped retinaculum (Figs. 3, 4), which opens dis- tally and provides a shelter on the wing surface just in- side its opening for a small mat of darker, specialized scales of uncertain function (perhaps the structure and modified scales serve to disperse a sex pheromone). The male genitalia (Fig. 6) are unlike those of the nearctic Nascia acutella but show surprising similarity to those of Oenobotys Munroe (Crambidae: Pyrausti- nae) (Munroe 1976: pl. A, figs. 6, 7; Ferguson et al. 1991: figs. 203a, c), having a similar fanlike cluster of processes on the inner face of the valve that resemble spatulate scales, each of which is expanded and bi- or trifurcate at its outer end. However, Sclerocona acutella also has a separate, prominent, spinose scle- VOLUME 57, NUMBER 4 271 Fics. 1-4. Sclerocona acutella. 1, Adult male, dorsal. 2, Adult male, ventral. 3, Retinaculum (arrow). 4, Cleared forewing. rite (4-5 spines) embedded in the valve beneath (or be- hind) the fanlike structure as viewed on the slide. The uncus is elongate, smooth, and regularly tapered from base to tip. The aedeagus has almost no sclerotized in- clusions (cornuti). The everted vesica shows only one blind sidepocket (diverticulum) at about its midpoint, and a small, low, weakly sclerotized hump farther out. Distribution. Connecticut: Fairfield Co., Danbury, Tarrywile Park, 8.vi.2001, D. L. Wagner (1) (UCONN); Litchfield Co., Canaan, Robbins Swamp, The Nature Conservancy Hollenbeck Preserve, 1/2.vii.1997, M. Volovski (1)(UCONN); Salisbury, Moore Brook, 8/9.vii.1994, D. L. Wagner, V. Giles & M. C. Thomas, MV & blklt (1)(UCONN); same locality, 23-30.vi.1995, D. Wagner, J Trouern- Trend, D. Primozich & M.C. Thomas, MV & blklt (7)( UCONN); Sal- isbury, Twin Lakes, 8/9.vii.1994, D. L. Wagner, V. Giles & M. C. Thomas, MV & blklt (1)(UCONN); same locality, 14.vii.1995, V Giles 3) & A. Valley, MV & blklt (1)(UCONN). Massachusetts: Barnstable Co., Bourne, 23.vi-21.vii.1995, M. Mello, blklt trap (3)(LCES); Bris- tol Co., South Dartmouth, Lloyd Center, 2.vii.1984, M Mello, blklt (1) (LCES); same locality, 26.vi.1989, M Mello, blkt (1) (UCONN); Hampden Co., Brimfield, 14.vi.1996, M Mello, blklt trap (1) (LCES); Hampshire Co., Amherst, 7 May, 1999 (larva), adult issued circa 15 June, 1999, Lisa Tewksbury and Geoffrey Balme (1)(URI); Suffolk Co., Boston, Thompson Island, 24.vi.2002, M Mello, blklt trap (3) (LCES). Maryland: Dorchester Co., Taylor’s Island Wildlife Man- agement Area, 14.vi.1998, J.D. Glaser, blklt (4)(JGC, USNM); Har- ford Co., Bush Wildlife Management Area, 5.vi.1999, J.D. Glaser, blklt (9)(JGC, USNM): Worcester Co., Isle of Wight, 1.vi.2000, J.D. Glaser, blacklight traps (26) (JGC, USNM); Hickory Point Cypress Swamp, 29.v.2002, J.Glaser, blklt (1)(JGC). Rhode Island: Newport Co., Tiverton Pocassett Swamp, 25.vi.2001, M Mello, blklt trap (1); Little Compton, 25.vi.2001, M Mello, blkt trap (1)(LCES). Early stages. Parenti (2000) and Robinson et al. (2002) list Phragmites australis as the host for Sclero- Fics. 5-6. Sclerocona acutella. 5, Wing venation. 6, Male genitalia. bo ~l bo cona acutella. Lisa Tewksbury and Geoffrey Balme (pers. com.) reared a single adult from a larva collected in the lower stem of Phragmites, on May 8-9, 1999, near Amherst, Massachusetts. The Phragmites stem was dead and broken; presumably it had been alive through the fall of the previous season. The single larva is believed to have pupated without further feeding. Common reed, Phragmites australis, grows in both freshwater and brackish wetlands, which is consistent with the range of localities represented among our col- lections. Label data suggest the species is univoltine. Records from southern New England, where the species is best known, range from14 June to 21 July (n = 23), with most collections falling within the last week of June and the first half of July. The Maryland speci- mens were collected between | and 14 June; the four from the latter date being slightly worn, indicating that Sclerocona flies earlier in Maryland. Distribution in eastern North America. North American records are from both inland and coastal wetlands from Massachusetts, Connecticut, Maryland, and Rhode Island. The first North American specimen is believed to be a male taken by Mark Mello in South Dartmouth, Bristol County, Massachusetts, at the Lloyd Science Center in 1984. Although the moth cer- tainly is distributed more widely, correspondence with other microlepidopterists suggests that it may not have spread much beyond the range we circumscribe here. We have written to (microlepidoptera) collectors in Maine (Tony Roberts), Michigan (George Balogh and Brian Scholtens), Ohio (Steve Passoa), Ontario (Jean- Francois Landry), and Quebec (Louis Hanfield)— none have yet taken this species. Etymology. The generic name is a compound of the Greek adjective sclero, meaning tough, hard, and the Latin masculine noun, conus (or the Greek konos), obviously referring to the cone-shaped retinaculum. Meyrick (a classics scholar) intended Sclerocona to be feminine, spelling it with a feminine ending and changing the species name from acutellus to the femi- nine form, acutella. We mention this because some au- thors have continued to use acutellus in combination with the feminine generic name. Remarks. Given the broad distribution of the moth along the eastern seaboard and the scant collections of pyraloids, the moth surely has been established for some time on the East Coast. Short of drawing infer- ences from detailed population genetic studies, there would seem to be no way to establish when, where, or how this species was introduced. Mikkola and Lafontaine (1994) made the observa- tion that several recently introduced moths in the Northeast, e.g., Apamea unanimis (Hiibner), A. ophio- JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 1999 1996 2002 1995 1984, 1989 2001 2001 1994, 1995, 1997 2001 1999 2000, 2002 1998 Fic. 7. Distribution of Sclerocona acutella along eastern seaboard, Locality data appears in text. gamma (Esper), and Rhizedra lutosa (Hiibner) are Phragmites, Phalarus, or other grass-feeders that are common in coastal habitats in Europe. The recently established Oligia strigilis (L.) reported by Handfield (1999) also fits this pattern (J. Don Lafontaine pers. com.). Mikkola and Lafontaine (1994) suggest that the turf and soil being picked up in shipyards, e.g., on the bottoms of large shipping containers, could be respon- sible for the recent spate of European coastal moth in- troductions along our eastern seaboard. Sclerocona acutella, being a Phragmites feeder, is yet another can- didate for their list. Phragmites australis is the focus of conservation ef- forts on both sides of the Atlantic Ocean, but for dif- ferent reasons. In parts of Europe it is a local, pro- tected species that has even been the focus of restoration efforts (e.g., Skuhravy 1978, Ostendorp 1989, Tscharntke 1990, 1992). On this continent, the plant is considered an invasive species that is overrun- ning wetlands, establishing almost pure monocultures of reed in coastal and inland wetlands that previously were floristically diverse (Garcia 1998, Chambers et al. 1999: Orson 1999, Saltonstall 2002). Given the new- found successes of Phragmites, we expect that adult Sclerocona will be become one of the most abundant moths in the vicinity of wetlands in many northeastern states. If the moth proves to be a specialist of Phrag- mites, it might have a future as a biocontrol agent in programs seeking to curb the spread of this grass. VOLUME 57, NUMBER 4 ACKNOWLEDGMENTS Our friend and colleague, Doug Ferguson, passed away on 4 No- vember 2002. He had assembled a partial draft of this paper in 2001. It is with deep sense of loss and admiration for Doug that we com- plete this small paper. René Twarkins prepared the line drawing of the wing venation and helped assemble the figures. Mark Mello sent us data from his numerous collections of Sclerocona. Lisa Tewksbury and Geoffrey Balme supplied observations on their collection and rearing of Sclerocona. Several students, friends, and colleagues helped with the acquisition of the Connecticut specimens: Valerie Giles, Jon Trouern-Trend, Dave Primozich, Michael Thomas, and Monty Volovski. David Grimaldi and Tam Nguyen took the image of the male genitalic capsule using a Nikon D1X digital camera with an Infinity (c) K2 lens, illuminated with fiber optic flashes from MicrOptics, Inc. We thank Peter Touhey and Alma Solis for recovering host rearing infor- mation from a Systematic Entomology Laboratory database and re- laying it to us. Steven Passoa helped us track down European and Asian literature. Funding for surveys that resulted in captures of Sclerocona to DLW was provided by the Connecticut Chapter of The Nature Conservancy, Connecticut Department of Environmental Protection, and Connecticut State Museum of Natural History. LITERATURE CITED CHAMBERS, R. M., L. A. MEYERSON & K. SALTONSTALL. 1999. Ex- pansion of Phragmites australis into tidal wetlands of North America. Aquat. Bot. 64:261-273. EVERSMANN, E.F von. 1842. Quaedam Lepidopterorum, species novae, in Rossia Orientali observatae, nunc descripti et depic- tae. Bull. Soc. Impér. Nat. Moscou 15:543-565. FERGUSON, D. C., D. J. HILBURN & B. Wricut. 1991. Lepidoptera of Bermuda: their food plants, biogeography, and means of dis- persal. Mem. Entomol. Soc. Canada. No. 158. 105 pp. Garcia, L. 1998. Phragmites australis: impact on invertebrate bio- diversity and its invertebrate fauna. MS. Thesis, University of Connecticut. 103 pp. HANDFIELD, L. 1999. Le guide des papillons du Québec. Broquet Inc., Boucherville, Quebec, Canada, 982 pp. INOUE, H., S. Suci, H. Kuroxo, S. Morruti & A. KAWABE. 1982. Moths of Japan. 2 vols. Kodansha, Tokyo, Japan. KarRSHOLT, O. & J. RAZOWSKI (EDS.). 1996. The Lepidoptera of Eu- rope, a distributional checklist. Apollo Books, Stenstrup, Den- mark, 380 pp. Marion, H. 1957. Classification et nomenclature des Pyraustidae d'Europe. Entomologiste 13:82 bo >I ee) Meyrick, E. 1890. On the classification of the Pyralidina of the European fauna. Trans. Entomol. Soc. London 1890:439— 492. MIKKOLA, K. & J. D. LAFONTAINE. 1994. Recent introductions of riparian noctuid moths from the Palearctic region to North America, with the first report of Apamea unanimis (Hiibner) (Noctuidae: Amphipyrinae). J. Lepid. Soc. 48:121—-127 Munrog, E. 1976. Pyraloidea. (in part). Fascicle 13.2A. The moths of America north of Mexico. E. W. Classey Limited and Wedge Entomological Research Foundation, London, England. 78 pp., 4 col pls., 8 bw pls. Orson, R. A. 1999. A paleoecological assessment of Phragmites australis in New England tidal marshes: changes in plant com- munity structure during the last few millennia. Biol. Invas. 1:149-158. OstENDoRP, W. 1989. ‘Die-back’ of reeds in Europe—a critical view of the literature. Aquat. Bot. 35:5-26. PaLM, E. 1986. Nordeuropas Pyralider. Danmarks Dyreliv Bind 3. Fauna Bgger, Copenhagen, Denmark. 287 pp., 8 col. pls. PARENTI, U. 2000. A guide to the Microlepidoptera of Europe. Museo Regionale di Scienze Naturali Torino, Stenstrup, Den- mark, 432 pp., 132 col. pls., 24 bw pls. ROBINSON, G., P. R. ACKERY, I. J . KITCHING, G. W. BECCALONI & L. M. HERNANDEZ. 2002. HOSTS—a database of the hostplants of the world’s Lepidoptera (http:/Avww.nhm.ac.uk/entomology/ hostplants/) (accessed 24 Nov., 2002). SALTONSTALL, K. 2002. Cryptic invasion by a non-native genotype of Phragmites australis into North America. Proc. Nat. Acad. Sci. USA 99:2445-2449. SIBERIAN ZOOLOGICAL MusEuUM. 2002. Website of the Siberian Zoo- logical Museum of the Institute of Animal Systematics and Ecol- ogy, Siberian Branch of the Russian Academy of Sciences (http:// www.bionet.nsc.ru/szmn/index.html) (accessed 24 Nov., 2002). SkuHRavy, V. 1978. Invertebrates (insects): destroyers of common reed (Phragmites communis). Ecol. Stud. Anal. & Synth. 28:376-388. TSCHARNTKE, T. 1990. Fluctuations in abundance of a stem boring moth damaging shoots of Phragmites australis: causes and ef- fects of overexploitation of food in a late-successional grass monoculture. J. Ecol. 27:679-692. TsSCHARNTKE, K. 1992. Fragmentation of Phragmites habitats, min- imum viable population size, habitat suitability and local extinc- tion of moths, midges, flies, aphids, and birds. Conserv. Biol. 6:530-536. Received for publication 20 December 2002; revised and accepted 26 April 2003. Journal of the Lepidopterists’ Society 57(4), 2003, 274-278 A NEW DESERT SUBSPECIES OF COLIAS OCCIDENTALIS (PIERIDAE) FROM SOUTHEASTERN OREGON PAuL C. HAMMOND Dept. of Zoology, Oregon State University, Corvallis, Oregon 97331, USA AND Davip V. MCCORKLE Dept. of Biology, Western Oregon University, Monmouth, Oregon 97361, USA ABSTRACT. Colias occidentalis Scudder is a complex polytypic species that is widely sympatric with the closely related C. alexandra W.H. Edwards throughout much of western North America. Populations of C. occidentalis have yellow males along the West Coast, orange males in the northern Rocky Mountains and across Canada, and mixed yellow/orange males across the Intermountain region of the Pacific Northwest. A southern isolate of these mixed populations has evolved in the northern Great Basin of southeastern Oregon, and is here described as Colias occidentalis sullivani, new subspecies. This subspecies uses an unique larval food plant, the desert bush pea (Lathyrus rigidus White). It has a relatively limited distribution in Harney and Malheur Counties in Oregon, but may extend into adjacent parts of Idaho and Nevada. In addi- tion, C. o. sullivani is superficially very similar to the sympatric C. alexandra, and has been confused with that species in the past. However, the two species have different larval food plants, and specific distinctions in wing color patterns between them are outlined in the present paper. Additional key words: Lathyrus rigidus, Colias alexandra, foodplants, adaptations, biogeography, variation. The genus Colias belongs to the subfamily Coliadi- nae in the family Pieridae. The most recent mono- graph of the genus (Verhulst 2000) recognizes up to 85 species distributed throughout much of the world, al- though many of these taxa may be regarded as geo- graphic subspecies of more widespread polytypic species. This problem of geographic variation combined with incipient or incomplete speciation is a major issue for many of the North American species, particularly for the species complex discussed in this paper. Colias occidentalis Scudder and C. alexandra W.H. Edwards are closely related legume-feeders that are widely sympatric in western North America, and have been the focus of much taxonomic confusion in the past. Ferris (1993) has provided the most recent re- view of this group. Typical forms of C. occidentalis such as C. o. occidentalis and C. 0. chrysomelas W.H. Edwards, have males colored yellow dorsally with no ultraviolet reflectance, and are distributed along the West Coast from northern California to British Co- lumbia. The allopatric christina group has orange dor- sal color with UV-reflectance, and is distributed across central Canada and through the northern Rocky Mountain region to South Dakota, Wyoming, Utah, and eastern Oregon. Colias alexandra has yellow dor- sal color with mostly no UV-reflectance except for a deeper yellow/orange UV-reflecting patch on the dor- sal hindwing. It is widely sympatric with both the or- ange christina and yellow occidentalis groups, and functions as a fully distinct biological species. The primary confusion has centered on the christina group, which was originally classified as a separate species. Klots (1961) treated this group as geographic subspecies of C. alexandra. However, the widespread sympatry of the christina group with typi- cal C. alexandra made this classification untenable (Ferris 1993). Ferris (1993) divided the christina group into three species based upon slight differences in male UV-reflectance patterns and female color pat- terns, including (1) C. christina W.H. Edwards across Canada and in the northern Rocky Mountains south to Wyoming , (2) C. pseudochristina Ferris in Utah and the eastern Pacific Northwest, and (3) C. krauthii Klots, disjunct between the Black Hills of South Dakota and southwest Yukon and adjacent Alaska. Intergrading populations between C. pseudochristina and typical C. occidentalis in Grant County of central Oregon were already known by Ferris (1993), but he regarded this intergradation as a purely local phenomenon in argu- ing for separate species status for C. occidentalis and the various members of the christina group. Recent field studies strongly challenge Ferris’ clas- sification. Layberry, Hall and Lafontaine (1998) found that the kluanensis Ferris subspecies of C. krauthii forms a complete intergrading cline with C. christina across the southern Yukon. Likewise, extensive field work in the Pacific Northwest and northern Rocky Mountains over the past ten years has shown that the intergradation among C. occidentalis, C. christina, and C. pseudochristina is not restricted to a local phenom- enon, but forms very long, gradual clines that extend from the east slope of the Oregon Cascades eastward through central and eastern Oregon to southeastern Washington and central Idaho, and then north from Wyoming to Alberta and northeast British Columbia (Pyle 2002). Therefore, it is our opinion that dorsal VOLUME 57, NUMBER 4 Fic. 1. Variation in Colias occidentalis sullivani and comparison with spring brood forms of C. alexandra edwardsii. Top row, left to right dorsal views: C. 0. sullivani, Holotype male, yellow form; C. o. sullivani, Allotype female, white form; C. a. edwardsii, male; C. a. edwardsii, fe- male. Second row, left to right ventral views: C. o. sullivani male, olive-green form; C. o. sullivani, female, blue-green form; C. a. edwardsii, male; C. a. edwardsii, female. Third row, left to right ventral views: C. o. sullivani, male, yellow-green form; C. o. sullivani, male, gray-green form with large discal spot; C. o. sullivani, male, yellow form; C. o. sullivani, male, orange form. Bottom row, left to right dorsal views: C. o. sullivani, male, yellow form with slight orange flush; C. o. sullivani, male, light orange form; C. o. sullivani, male, medium orange form; C. o. sullivani, male, darker orange form. UV-reflectance patterns in males of this particular Co- lias group are not useful for the classification of species. We believe that the orange christina group should be treated as a geographic subspecies of Colias occidentalis. A peripheral part of this broader pattern of geo- graphic variation has been the recent discovery of a distinctive new subspecies of C. occidentalis in the deserts of southeastern Oregon where it co-exists in sympatry with C. alexandra. This new discovery is the topic of the present paper. Colias occidentalis sullivani Hammond and McCorkle, new subspecies Male. Forewing length 23-30 mm (x = 26 mm, n = 194). Forewing apex slightly elongate and pointed. Dorsal ground color usually yellow (95%), rarely orange (5%) of the 194 specimens ex- amined. Black border of forewing broad with smooth inner margin and yellow veins. Small black discal spot on forewing usually pres- ent, sometimes absent. Discal spot of dorsal hindwing only faintly evident and yellow. Heavy black basal suffusion present on fore and hindwings. Ventral ground color of hindwing light to dark olive- green, varying to yellow-green or gray-green. Black scaling in medial area of ventral forewing variable, heavy to absent. Discal spot on ventral hindwing usually small and white (55%), but sometimes medium size (31%) and rarely large (14%) of the 194 specimens ex- amined. A red or purple ring around the discal spot is variably pres- ent or absent. Male UV-reflectance. Males are highly variable with nearly 50% showing little or no UV-reflectance on dorsal wings as in typical C. o. occidentalis. Others show a weak, diffuse reflectance on fore and hindwings as in C. 0. pseudochristina, or a bright luminous patch on the hindwing as in C. alexandra, or bright luminous patches on both fore and hindwings as in C. o. christina. Female. Forewing length 25-30 mm (x = 28 mm, n = 116). Dor- sal ground color usually pure white (67%), yellowish white (29%), or rarely yellow (4%) of the 116 specimens examined. Black border of bo =I jo>) dorsal forewing usually absent (71%), vaguely present (23%) or rarely well developed (6%) of the 116 specimens examined. Black discal spot of dorsal forewing large, round to oblong. Discal spot of dorsal hindwing pale orange to white. Ground color of ventral hindwing gray-green to blue-green. Other characters as in male. Etymology. We name this taxon in honor of Barry Sullivan of Salem, Oregon who originally discovered this butterfly, and who has contributed greatly to our knowledge of the butterfly fauna of the Pacific North- west by his extensive exploratory collecting. Types. Holotype: male, Oregon, Harney County, Alvord Desert Road at north end of Steens Mountains, T29S, R36E, sec.25,26; 9 May 2001, Barry Sullivan leg. The holotype is deposited in the American Museum of Natural History, New York, New York, USA. Allotype: female, same data and deposition as holotype. Paratypes: 192 males, 116 females, and 1 gynandromorph, same locality as holotype, 22 April 1990, 3 May 2000, 9May 2001, 25 May 2001, 11 May 2002, 13 May 2002, 15 May 2002, 16 May 2002, P.C. Hammond, J. Harry, D.V. McCorkle, H. Rice, E. Runquist, B. Sulli- van, and A, Warren; 1 male, 1 female, Harney Co., east slope of Stinkingwater Mts. at U.S. Hwy. 20, 10 May 2001, 24 May 2001, D.V. McCorkle; 2 females, Harney Co., U.S. Hwy. 20 nr. Drewsey, 8 June 2001, 15 May 2002, P.C. Hammond, D.V. McCorkle; 7 males, Harney Co., south end of Stinkingwater Mts. east of Crane, 16 May 2002, P.C. Hammond and D.V. McCorkle; 1 male, 1 female, Harney Co., Alvord Desert Road at Ten Cent Lake, 25 May 1950, S.G. Jew- ett, Jr.; 2 males, 1 female, Malheur Co. north end of Sheepshead Mts. on Hwy. 78, 16 May 2002, P.C. Hammond and D.V. McCorkle. Disposition of paratypes as follows: one pair each to the U.S. Na- tional Museum of Natural History, the California Academy of Sci- ences, the Natural History Museum of Los Angeles County, and the Allyn Museum of Entomology (Sarasota) of the Florida Museum of Natural History; five pairs to the Oregon State Arthropod Collec- tion, Oregon State University, additional paratypes are in the private collections of Paul C. Hammond (36 males, 16 females), Jack Harry (2 males, 2 females), David V. McCorkle (40 males, 34 females), Harold Rice (10 males, 6 females), Erik Runquist (6 males, 6 fe- males), Don Severs (7 males, 2 females), Barry Sullivan (60 males, 18 females, 1 gynandromorph), Andrew D. Warren (22 males, 23 fe- males). DISCUSSION Throughout the northern Great Basin and Inter- mountain regions of the Pacific Northwest, sympatric populations of C. occidentalis and C. alexandra exhibit sharp ecological segregation in larval foodplants. Col- ias occidentalis primarily feeds on peas (Lathyrus spp.) and false lupines (Thermopsis spp.), while C. alexandra feeds mostly on milk-vetches (Astragalus spp.) and locoweeds (Oxytropis spp.) (pers. obs.). All of the above genera are herbaceous legumes of the family Fabaceae. Because most species of peas and false lupines are found in moist coniferous forest and montane meadows, populations of C. occidentalis are usually limited to the higher mountain ranges of the West. Colias alexandra often flys with C. occidentalis in these areas, but its larvae feed on Astragalus grow- ing on nearby dry, open hillsides. Previously, it was thought that C. alexandra alone lived on the dry, desert plains and lower mountains of the northern JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Great Basin and Intermountain regions where only As- tragalus and Oxytropis species usually grow. In moist forests of central and eastern Oregon and southeast Washington, the C. 0. occidentalis/pseudo- christina intergrade populations have been observed to oviposit on Lathyrus lanszwertii Kell., L. pauciflorus Fern., and L. nevadensis Wats., all forest peas with a vine-type growth habit, and also on false lupine (Ther- mopsis montana Nutt.). However, there is a desert bush pea (Lathyrus rigidus White) that is found in the low- land sagebrush/bunchgrass steppes of central and east- em Oregon, extending into Adams County, Idaho, Washoe County, Nevada, and Modoc County, California. On 22 April 1990, Barry Sullivan found a population of Colias at the north end of the Steens Mountains in Harney County, Oregon that was associated with Lathyrus rigidus rather than Astragalus. Males are mostly yellow dorsally, and have a dark olive-green ventral hindwing with a very small white discal spot that often lacks a red ring. This population was initially thought to be a peculiar form of the spring brood of C. alexandra edwardsii W.H. Edwards because of these characters. Two specimens of this Colias were first col- lected in this area at Ten Cent Lake by Stanley G. Jew- ett, Jr. on 25 May 1950, but remained identified as C. alexandra in the Oregon State University collection for 50 years. In May of 2000 and 2001, Barry Sullivan and DVM visited this site and noticed aspects of this Col- ias that are more similar to C. occidentalis than to C. alexandra. These include the Lathyrus hostplant asso- ciation and the fact that nearly all females in the popu- lation are albinistic. Indeed, most females are pure white to creamy white in dorsal color, and the black wing borders are usually greatly reduced or completely absent. Female albinism is relatively rare in the sym- patric C. alexandra edwardsii. Additional study revealed a number of subtle wing pattern differences that distinguish this new Colias from C. alexandra, even though both species fly to- gether at most localities. In males of C. o. sullivani: (1) the dorsal black wing border is usually broad with a smooth inner margin, (2) there is heavy black basal suffusion, (3) a deeper yellow/orange patch is usually absent from the dorsal hindwing, (4) the ventral hind- wing ground color is a dark olive-green, and (5) some specimens have a large red-ringed ventral discal spot. The ventral ground color of females varys from gray- green to blue-green. By contrast, males of C. alexan- dra usually have: (1) a narrow black wing border, often with a dentate inner margin, (2) very little black basal suffusion, (3) a deeper yellow/orange patch on the dor- sal hindwing, (4) a ventral hindwing ground color vary- ing from gray to gray-green, and (5) a ventral discal VOLUME 57, NUMBER 4 spot that is usually small to very small. Female ventral ground color is also gray to gray-green. Of particular interest was the discovery that a small proportion (5%) of C. o. sullivani males from the type locality have orange dorsal coloration on both fore and hindwings as in the christina/pseudochristina forms. Except for the strange olive-green to blue-green ven- tral ground color, all of the differences that distinguish this new Colias from C. alexandra are also shared by other races of C. occidentalis, including a high fre- quency of female albinism. Figure 1 illustrates these variations in C. o. sullivani and the differences from C. alexandra. During May of 2001 and 2002, major efforts were made to locate additional populations of this new Col- ias using OSU herbarium records of Lathyrus rigidus as a guide. Extensive populations of the bush pea were indeed located in the John Day valley of Grant County and the Powder River valley in Union and Baker Counties on dry, open prairie. However, these areas are within the general range of forest C. 0. occiden- talis/pseudochristina populations, and these appar- ently never associate with the lowland L. rigidus. At the southwest edge of the Wallowa Mountains in Union County, dry open hills covered with L. rigidus are present at a forest/prairie ecotone where L. nevadensis and L. pauciflorus occur in forest riparian areas along a stream. One apparently stray male of C. o. pseudochristina was collected on the open hillside among the L. rigidus, but we have no further evidence that this subspecies uses L. rigidus at this site. However, additional populations of C. 0. sullivani were located in Harney and Malheur Counties. This part of southeastern Oregon covers a diverse land- scape of desert mountain ranges, lowland plains, and rugged canyonlands. The bush pea was found to be quite local and narrowly restricted in habitat, but was often extremely abundant, especially in areas that had evidence of past fires. The habitat consists of low hill- sides just above the valley floor. Peas were never found on the lowland plains proper or higher in the moun- tains. Vegetation in the habitat is usually a sage- brush/bunchgrass prairie, although at the type locality, the ground is somewhat barren of vegetation except for the pea plants and a rich variety of native herbs. Several thousand butterflies were present at the type locality during 2002, with the adults flying low among the peas. At all other sites observed, the but- terflies were only moderately abundant to very rare. Nectaring usually takes place from the pea flowers or from composites. Many ova and young larvae were col- lected from pea plants during May of 2001 and 2002. The adult flight season lasts from late April to early June, and there is only a single generation per year. Pea plants enter senescence by June, so reproductive efforts must be complete by that time. At the time of this writing, Malheur County is mostly unexplored for C. 0. sullivani, but herbarium records indicate that extensive populations of L. rigidus are present in central parts of the county west of the Owyhee River. It is possible that the butterfly could occur eastward into Owyhee County, Idaho. Also, the Bowden Hills and Trout Creek Mountains in southern Harney and Malheur Counties remain unex- plored. However, a small population of C. o. sullivani was located at the north end of the Sheepshead Moun- tains about 16 km east of the type locality. In the northeast corner of Harney County, addi- tional populations of C. o. sullivani were located along the eastern edge of the Stinkingwater Mountains, both east of Stinkingwater Pass on U.S. Highway 20 and at the south end of the mountains east of Crane. Indeed, populations are probably located along the entire east- er edge of these mountains. More butterflies were found to the east near Drewsey on Highway 20, and it is probable that populations are scattered throughout central Malheur County south of Juntura, but access to many areas is limited. However, a particularly important population was located north of Juntura and east of Beulah Reservoir in northwestern Malheur County near the prairie/for- est ecotone not far from the southeast edge of the Blue Mountains. This population is still basically of the C. o. sullivani type, and is associated with a very large population of L. rigidus on a bunchgrass/sagebrush prairie in a mid-elevation valley. Here the frequency of dorsally orange males of the christina/pseudochristina type is much higher, around 22% compared to 5% at the type locality about 112 km to the south. The fre- quency of medium to large discal spots on the ventral hindwing is also much higher, about 78% compared to 45% at the type locality. Also, about 9% have a yellow or orange ground color on the ventral hindwing in- stead of the greenish color characteristic of C. o.sulli- vani. These character frequencies suggest extensive gene exchange with the C. 0. pseudochristina forest populations to the north in the Blue Mountains. In conclusion, it is hypothesized that C. o. sullivani probably evolved from the C. o. occidentalis/pseudo- christina populations in the Blue Mountains, and that ancestral populations spread southward from the Blue Mountains into the Steens Mountains during a glacial maxima of the Pleistocene, following the forest habitat of Thermopsis and Lathyrus nevadensis southward. As conditions warmed and dried during an interglacial period, the ancestral butterfly probably dwindled and bo =I Oo nearly disappeared along with its forest foodplants in the Steens Mountains, except for a small founder population that was able to switch and adapt to Lathy- rus rigidus. Colias o. sullivani appears to be highly adapted for feeding on this particular foodplant, while the northern populations of C. occidentalis in the Blue Mountains seem largely unable to switch to L. rigidus even when it is locally available at the prairie/forest ecotone. Moreover, the olive-green to blue-green ground color of the ventral hindwing of C. o. sullivani blends perfectly in camouflage with the blue-green fo- liage of the hostplant. One other line of evidence in support of the above evolutionary speculations comes from a large popula- tion of the C. o. occidentalis/pseudochristina inter- grade type in the Aldrich Mountains of Grant County. A sample of 276 males showed a frequency of 70% dorsal yellow color and 30% orange color. However, on the ventral hindwing, the ground color was 66% or- ange, 29% yellow, and 5% green, while discal spot size was 27% large, 44% medium, and 29% small. Like- wise, a sample of 43 females from the same population showed a dorsal ground color of 19% orange, 37% yel- low, 30% yellowish white, and 14% pure white, while the black wing border was heavily developed in 16%, slightly present in 42%, and completely absent in 42%. Thus, a few male and female individuals from this northern population are nearly a perfect match in color phenotype to C. o. sullivani. In addition, it should be noted that male UV- reflectance patterns in C. o. sullivani show the same range of polymorphic variation as in the C. 0. occiden- talis/pseudochristina intergrade populations from Grant County, Oregon illustrated by Ferris (1993). Ap- proximately 50% of males show no UV-reflectance as in typical C. o. occidentalis, while others show a weak, JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY diffuse reflectance as in C. 0. pseudochristina, or a bright luminous patch on the hindwing as in C. alexan- dra, or bright luminous patches on both the fore and hindwings as in C. o. christina. Thus it appears that most genetic traits used in the evolution of this desert subspecies were originally present at low frequencies in the ancestral forest populations. ACKNOWLEDGMENTS Many people assisted with this study. We particularly acknowl- edge Barry Sullivan, Andy Warren, Terry Stoddard, Dana Ross, Harold Rice, and Erik Runquist for their field sampling efforts which helped form the basis of the study. We also thank William Neill, Jon Pelham, Jon Shepard, Don Severns, Jack Harry, Nelson Curtis, and Steve Kohler for allowing us to examine specimens in their collec- tions. The systematic research collections at Oregon State University were also critical to this study, and we thank Aaron Liston and Dick Halse for use of the Oregon State Herbarium, and Darlene Judd and Andy Brower for use of the Oregon State Arthropod Collection. We also thank the Burns District Office of the U.S. Bureau of Land Man- agement for assistance in locating populations of Lathyrus rigidus. Support for this publication was provided by the Harold and Leona Rice Endowment for Systematic Entomology at Oregon State Uni- versity. LITERATURE CITED Ferris, C. D. 1993. Reassessment of the Colias alexandra group, the legume-feeding species, and preliminary cladistic analysis of the North American Colias (Pieridae: Coliadinae). Bull. Allyn Mus. (138):1—-91. Kiots, A. B. 1961. Genus Colias Fabricius, pp. 53-62. In Ehrlich, PR. & A. H. Ehrlich, How to know the butterflies. Wm. C. Brown Co., Dubuque, Iowa. LayBERRY, R. A., P. W. HALL & J. D. LAFONTAINE. 1998. The but- terflies of Canada. University of Toronto Press, Toronto, Canada. 280 pp. Pye, R. M. 2002. The butterflies of Cascadia. Seattle Audubon Society, Seattle, Washington. 420 pp. VERHULST, J. T. 2000. Les Colias du Globe: monograph of the genus Colias. Goecke & Evers, Keltern, Germany. 571 pp. Received for publication 23 September 2002; revised and accepted 6 May 2003. Journal of the Lepidopterists’ Society 57(4), 2003, 279-283 MALE-SPECIFIC STRUCTURES ON THE WINGS OF THE GULF FRITILLARY BUTTERELY, AGRAULIS VANILLAE (NYMPHALIDAE) CASANDRA L. RAUSER! AND RONALD L. RuTOWSK?? Department of Biology, Arizona State University, Tempe, Arizona 85287-1501, USA ABSTRACT. Males of the Gulf Fritillary butterfly, Agraulis vanillae Linnaeus (Nymphalidae), have distinctive structures on certain veins of the dorsal wing surface that appear to be involved in pheromone production. Here we confirm and extend an earlier description of these structures. Observations using light and scanning electron microscopy indicate that in these structures, patches of several scale types alternate with scaleless areas along the veins. Some of these open areas have pores that we suggest might be the route by which the pheromone moves from cells in the wing integument onto brush-bearing scales from which it is disseminated during courtship. We have also found that although the dimensions of the basic units of these structures on the veins are not correlated with body size, larger males do have greater total vein length devoted to these structures. These findings are discussed in light of the courtship of this species and the potential for these structures to be in- volved in mate choice and to be a product of sexual selection. Additional key words: pheromones, androconia, morphology, mating behavior, SEM. In many butterflies and moths, males have struc- tures on their wings or body that produce chemical signals, or pheromones, used by males when courting females (Myers & Brower 1969, Pliske & Eisner 1969, Vane-Wright & Boppré 1993, Kan & Hidaka 1997, Iyengar et al. 2001). In some cases, behavioral experi- ments have confirmed the function of these structures, but more often a phermonal function is inferred from the high surface area of all parts of them, and the ob- servation that they are often brought close to the an- tennae of the female during courtship (Lundgren & Bergstrém 1975, Rutowski 1977, Boppré 1984). Al- though much is known of the morphology and opera- tion of such structures in danaine butterflies, the full diversity of their structure and function in butterflies is generally not well documented. Miiller (1877) first discovered and described pre- sumptive scent-producing structures on the wings of males of the Gulf Fritillary, Agraulis vanillae Lin- naeus (Nymphalidae), a common heliconiine butterfly in the American tropics and subtropics. In particular, he reported that along six of the forewing (FW) veins there were patches of long thin scales each bearing a brush at the distal end, which alternated with rows of normal scales crossing the vein. Our preliminary ob- servations of these structures using light microscopy suggested that the arrangement of scales along these veins while serially repeated, as Miiller (1877) re- ported, was more complex than indicated by his de- scription and so we undertook a more detailed study of their morphology using light and scanning electron microscopy. Our first objective was to repeat and extend Miiller’s observations because we believed more detailed ob- "Corresponding author's current address: Department of Ecology and Evolutionary Biology, 321 Steinhaus Hall, University of Califor- nia, Irvine, California 92697-2525, USA, email: crauser@ uci.edu 2 Email: rrutowski@asu.edu servations could suggest how the scent was produced and disseminated in A. vanillae. The second objective was to determine how much, if any, inter-individual vari- ation there is in the overall size of these structures. This could suggest the extent to which males might vary in their ability to produce critical chemical signals during courtship. Lastly, we wanted to relate Miiller’s and our observations to published descriptions of the courtship behavior of A. vanillae (Rutowski & Schaefer 1984). METHODS We examined the FWs of male and female A. vanil- lae using both light and scanning electron microscopy. Specimens were either laboratory-reared from field- collected larvae and eggs on cuttings of Passiflora spp. or field-caught as adults in Tempe, Arizona. For exam- ination of scale morphology, we chose butterflies with little or no wing wear. In some cases, we removed scales with a small paintbrush to expose the cuticle covering the veins underlying the scales. For light mi- croscopy we mounted FWs onto slides and pho- tographed specialized structures and scales. For scan- ning electron microscopy (SEM), we attached wings to stubs with conducting paint and no coating, and ob- served them with a Philips/FEI XL 20 scanning elec- tron microscope. To determine whether larger males also have a larger androconial area on their FWs, and thus greater pheromone-disseminating ability, we measured the length of the portion of each of the veins containing specialized structures on the male FW. For each spec- imen (n = 20), we then totaled these lengths and plot- ted this total against FW length and subjected the data to correlation analysis. We also determined the density of specialized structures along the veins to investigate whether it also varies with FW size. To do this we esti- mated the number of structures per mm on each of the six veins on a male FW. We then calculated the 280 ca. 10mm Fic. 1. A sketch of the dorsal side of the FW of the male Gulf Fritillary butterfly with the veins containing the presumptive pheromone-disseminating structures labeled according to Scoble’s (1992) notation. The segments of the veins containing the special- ized structures are highlighted. mean number of structures per mm for all 6 veins and compared the mean number of structures per mm to the FW length, also using correlation analysis (n = 20). RESULTS As reported by Miiller (1877), male-specific scales and other structures are found on the dorsal FW of the male on veins M,, M,, M,, CuA,, CuA,, and 1A, + 2A (Fig. 1; wing vein notation, Scoble 1992). On veins that had not been disturbed we observed that, along at least part of each of the six veins, rows of brush-like scales alternated with rows of scales with broad, straight edges (Fig. 2). The average length of each vein segment that contains the male-specific structures is 11.7 + 1.43 mm on M,, 13.4 + 1.51 mm on M,, 13 + 1.57 mm on M,, 13.5 + 1.69 mm on CuA,, 14.7 + 1.74 mm on CuA, and 12.96 + 1.73 mm on 1A, + 2A. The FW length of the 20 specimens we examined averaged 33.7 + 2.6 mm. With the scales removed from the veins, we found that a set of five distinct areas make up a unit 0.24 mm in length that is serially repeated along each of the six veins (Fig. 3). In order from the wing base to the wing edge, the five distinct areas are the inter-scale vein- section 1 (IVS-1), the scale-section 1 (SS-1), the inter- scale section 2 (IVS-2), the scale-section 2 (SS-2) and the porous section (PS) (Figs. 3, 4). As shown in Fig. 4, the IVS-1 (mean length = 0.12 mm) occurs between the repetitive structures and does not contain any scale attachment sites. Every third row of scales occuring on the inter-vein part of the wing runs uninterrupted across the veins. SS-1 (mean length = 0.02 mm) is the scale attachment row after IVS-1 and contains two different types of scales. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY . fi Fic. 2. An SEM of a segment of the vein containing the spe- cialized structures with the scales intact (250x). The brush-like scent scales are attached to the vein at SS-2 and the broad flat scales are at- tached at SS-1. One type is broad at the top, narrow at the base and shiny black, whereas the other type is equally as long, but broad at the base and tip and translucent, shiny gold. Both types of scales are flat and the black scales fit snugly over the gold, scales as shown in Fig. 5. IVS- 2 (mean length = 0.01 mm) is the area containing no scale attachments between SS-1 and SS-2. SS-2 (mean length = 0.04 mm) is a region with the scale attach- ments between IVS-2 and PS. The scales arising in SS-2 are densely packed, long (mean length = 0.4 mm), nar- row, and have brush-like ends with high surface area. In addition, the bases of these scales are black, fol- lowed by a thin transparent area, then by a black, nar- row section which contracts into another thin trans- parent area, followed by a black brush-like apex (Fig. 6). These brush-bearing scales are positioned so that the transparent basal area (mean length = 0.04 mm) overlies PS as shown in Fig. 6. PS (mean length = 0.06 Fic. 3. The repetitive nature of the specialized structures along the vein of a male’s FW is shown with the scales removed (20x). The five distinct sections of the specialized structures are labeled accord- ing to their function: Scale Section 1 and 2: SS-1 and SS-2; Inter- scale Vein Section 1 and 2: IVS-1 and IVS-2: and Porous Section: PS. VOLUME 57, NUMBER 4 Fic. 4. An SEM of the male Gulf Fritillary pheromone- disseminating structure with the five distinct sections labeled (750x). PS and SS-2 are further magnified in order to clearly see the differ- ence between the two areas (1500x). SS-2 is the scale attachment point for the scent scales and PS is the hypothesized area of pheromone secretion. mm) is the porous area containing no scale attachment sites adjacent to SS-2. The total length of the vein segments that contain the presumptive pheromone-disseminating structures is positively correlated with FW length (r? = 0.858, p < 0.0001) (Fig. 7). This means that the longer the FW of the male, the longer the length of the veins containing the pheromone-disseminating structures. However, there is no relationship between the mean number of structures per mm and the length of the FW (p > 0.40) (Fig. 8). Therefore, the number of structures per mm along the veins remains constant regardless of the size of the FW. Collectively, these results indicate that males with longer FWs have more total scent-disseminating structures than males with shorter FWs and thus greater scent disseminating ability. DISCUSSION Our observations confirm and extend Miiller’s (1877) description of male-specific structures found along certain veins on the dorsal FW surface of A. ae Fic.5. An SEM of the two types of scales and their attachment points at SS-1 (1500x). Both scale types are flat and one fits snuggly over the other. The scale type that lies on top is broad at the top, nar- row at the base and shiny black. The other scale type is equally as long, broad at the base and tip and translucent, shiny gold in color. vanillae. In addition, we found that these male-specific structures consist of serially repeated units of scale arrangements along the veins. However, each unit consists of five rather than two distinct elements, as described by Miiller. One of these elements is a patch of brush-bearing scales and rows of scales typical of those found crossing the vein elsewhere on the wing (Magnus 1950). We also found other scale types and three regions without scales that included an area dot- ted with large pores opening onto the wing surface. The specific elements of the structures found along these veins may interact in the following ways to dis- seminate a pheromone. The brush-bearing scales lo- cated in SS-2 lie directly over the porous area (PS) (Fig. 6). These pores could be the openings through which the product of pheromone-producing cells in Fic. 6. A photograph of a pheromone-disseminating structure on the male FW with the scent scales intact at SS-2 (20x). The trans- parent portion of the scent scales closest to the point of attachment lies directly over the porous area (PS). PS is hypothesized to secrete pheromones that disseminate onto the high surface area brush-like scales. bo ee) bo Total vein length with male-specific scales (mm) Forewing length (mm) Fic. 7. The positive relationship between the FW length and the total length of the vein segments containing pheromone- disseminating structures (r? = 0.858, p < 0.001). the wing integument is secreted. This arrangement would allow the compounds to be directly transferred onto the densely packed, high surface area brushes on the overlying scales. The black and gold scales de- scribed in SS-1 lie over the narrow stalk of delicate scent-scales (Fig. 2). These SS-1 scales, which were difficult to remove, appear to form a protective barrier over the relatively fragile and easily removable SS-2 brush-bearing scales. This protective barrier may help prevent the loss of brush-bearing scales and excessive evaporation of pheromone secretions. Rutowski and Schaefer (1984) described the courtship behavior of A. vanillae and reported a male display, which they called the wing clap. During courtship the male positions himself head-to-head alongside the female with his body at about a 45° angle relative to the body of the female. Initially his wings are slightly spread and the female’s ipsilateral antenna comes to lay back between the male’s wings. He then quickly closes and reopens his wings repeatedly catch- ing the female’s antenna between them and bringing the female antenna into brief contact with the distinc- tive structures on the dorsal FW veins. Hence, as in other species, we find a male courtship behavior that strongly suggests that these male-specific, high surface area structures are producing a pheromone that may be important in mate choice by females (Tinbergen et al. 1942, Magnus 1950, Brower et al. 1965, Conner et al. 1980, Pivnick et al. 1992). Our data show that males with longer FWs have a greater total length of veins containing pheromone- disseminating structures (Fig. 7). Because the density of androconia per individual is not affected by FW length (Fig. 8), and FW length is a good measure of male size, larger males should have a greater total number of pheromone-disseminating structures than smaller males. Thus, the number of pheromone- JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Mean number of structures per mm “I oO QO e ee 5.5 + r a 28 30 32 34 36 38 40 Forewing length (mm) Fic. 8. No relationship exists between the number of structures per mm per individual and the FW length. In other words, the den- sity of androconia along the veins remains constant regardless of FW size (p > 0.40). disseminating structures and the quantity of phero- mone secreted are positively correlated with male size. A similar correlation was found in the moth Utetheisa ornatrix and was determined to be a heritable trait (Iyengar & Eisner 1999). If the amount of pheromone disseminated by the male during courtship is critical to female mate choice in A. vanillae, larger males should be more successful at mating than smaller males. This is the case in U. ornatrix where the amount of phero- mone disseminated by the male is the only criterion used by the female when choosing a mate (Iyengar et al. 2001). Similarly, males in Drosophila grimshawi deposit a pheromone to attract females to a mating site by rub- bing their abdomen on the substrate. Males who de- posit the greatest amount of pheromone at their site are the most successful at attracting females and mat- ing (Droney & Hock 1998). The evidence discussed above indicates that the morphology and location of the male androconia in A. vanillae may be a product of sexual selection (Fisher 1958, Baker & Cardé 1979, Eisner & Meinwald 1995). The quantity of the pheromone secreted may be an in- dicator of male quality that females evaluate when choosing a mate (Dussourd et al. 1991, LaMunyon 1997, Iyengar & Eisner 1999). Quality here could mean species identity or ability to provide either mate- rial (i.e., defensive secretions ) or genetic benefits (i.e., genes for large size). Pheromone quantity may also in- dicate a male’s age and/or mating status because scent- scales are likely to be lost with age and with each mat- ing or courtship. In this way a female may be able to assess the physiological state of a male when choosing a mate. However, sometimes the Gulf Fritillary female will mate with a male even when he does not perform the wing-clap display (Rutowski & Schaefer 1984). Per- VOLUME 57, NUMBER 4 haps pheromones are not the only indicator of a male’s quality, female choice is not based on this character, some females are less discriminating than others, or some males have a strong enough odor to elicit a re- ceptive response from the female without using the display. To better understand the functions of these pheromone-disseminating structures, further investiga- tion needs to be undertaken to identify the pheromones, to determine which of these pheromones are behav- iorally active during courtship and mating, and whether pheromone amount affects female mate choice. ACKNOWLEDGMENTS We thank Randi Papke and Jenny Drnevich for their invaluable support, encouragement, and critiques. We would also like to thank Jamal Ruhe for field assistance, Bob Mendoza for helping us with the figures, Michael Demlong of the Phoenix Zoo for allowing us to use the butterfly garden as a field site, and Irene Piscopo for preparing the SEM samples. We are also thankful to John Alcock, Damon Orsetti, Carla Penz, and an anonymous reviewer for help in revisions of this manuscript. Lastly, we would like to acknowledge the Howard Hughes Medical Institute and Arizona State University Biology Research Experience for Undergraduates program for funding. LITERATURE CITED BAKER, T. C. & R. T. CARDE. 1979. Courtship behaviour of the ori- ental fruit moth (Grapholitha molesta): experimental analysis and consideration of the role of sexual selection in the evolution of courtship pheromones in the lepidoptera. Ann. Entomol. Soc. Am. 72:173-188. BoppreE, M. 1984. Chemically mediated interactions between but- terflies. In Vane-Wright, R. I. & P. R. Ackery (eds.), The biology of butterflies. 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The scent-scales of the male of Dione vanillae. Kosmos 2:38-41. Translation In. LONGSTAFF, G. B. 1912. Butter- fly hunting in many lands. Longmans, Green, and Co., London. Myers, J. & L. P. BRoweR. 1969. A behavioural analysis of the courtship pheromone receptors of the queen butterfly Danaus gilippus berenice. J. Insect Physiol. 15:2117-2130. PIVNICK, K. A., J. LAvolE-DoRNIK & J. N. MCNEIL. 1992. The role of the androconia in the mating behaviour of the European skipper, Thymelicus lineola, and evidence for a male sex pheromone. Physiol. Entomol. 17:260-268. PLISKE, T. E. & T. EISNER. 1969. Sex pheromone of the queen but- terfly: biology. Science 164:1170-1172. Rutowski, R. L. 1977. Chemical communication in the courtship of the small sulphur butterfly Ewrema lisa (Lepidoptera, Pieri- dae). J. Comp. Physiol. 115:75-85. Rutowski, R. L. & J. SCHAEFER. 1984. Courtship behavior of the gulf fritillary, Agraulis vanillae (Nymphalidae). J. Lepid. Soc. 38:23-31. SCOBLE, M. J. 1992. The lepidoptera: form, function and diversity. Oxford University Press, New York. TINBERGEN, N., B. J. D. MEEUSE, L. K. BOEREMA & W. W. VAROSSIEAU. 1942. Die Balz des Samtfalters, Eumenis (=Satyrus) semele (L.). Z. Tierpsychol. 5:182-226. VANE-WRIGHT, R. I. & M. Boppreé. 1993. Visual and chemical sig- naling in butterflies—functional and phylogenetic perspectives. Phil. Trans. R. Soc. Lond., B. 340:197—205. Received for publication 25 November 2002; revised and accepted 7 May 2003. Journal of the Lepidopterists’ Society 57(4), 2003, 284-290 INTERINDIVIDUAL VARIATION IN MITOCHONDRIAL ENZYME ACTIVITY IN MALE MONARCH BUTTERFLIES, DANAUS PLEXIPPUS L. (NYMPHALIDAE) ELENA LEVINE!, OLIVIA BYRON-COOPER, MEGAN DESCH-O’DONNELL AND CARRIE METZINGER Biological Sciences Department, California Polytechnic State University, San Luis Obispo, California 93407, USA ABSTRACT. We determined activity levels for the mitochondrial enzyme succinate dehydrogenase in male monarch butterflies overwin- tering on the central coast of California from October 2001 through March 2002. Mitochondrial activity is important for generating the energy required for metabolically demanding activities such as flight, which is essential for monarch reproduction and survival. There is a genetic com- ponent to variation in flight performance in various species, and individual variation in mitochondrial activity may contribute to these differ- ences, since mitochondrial enzyme levels often correlate with performance abilities. To understand possible functional consequences of mito- chondrial activity, it is first necessary to determine the degree of individual variation within the population. We found a high degree of interindividual variation in enyzme activity in male monarchs, at least a twelve-fold difference between the lowest and highest activities mea- sured, with a coefficient of variation of forty-seven percent. In addition, we investigated possible correlations with season, body weight, body size, and wing damage. Although there were some month-to-month differences, individual variation in mitochondrial enzyme activity is not ex- plained by seasonality or body size, and is not related to the degree of wing damage. The results suggest that interindividual differences in mi- tochondrial enzyme activities are considerable, and worth investigating as a factor in individual performance and success. Additional key words: succinate dehydrogenase, insect flight muscle, overwintering, energetics. Monarch butterflies are known for their dramatic migration across the U.S.A. and overwintering in mas- sive aggregations along the California coast and in cen- tral Mexico (Tuskes & Brower 1978, Brower 1985). Flight is energetically expensive; indeed, insect flight muscle is notable for the highest metabolic rates and power output of any animal tissue (Sacktor 1976, Suarez 2000). In addition to long flights of migration during fall and spring which may cover thousands of miles (Brower 1985), mating involves a short energet- ically intense nuptial flight. Nuptial flight or mate transport to the nearby tree canopy occurs after a male successfully couples with a female (Shields & Emmel 1973). Regulation of metabolism is also important for survival, since monarchs rely primarily on stored lipids during the overwintering period (Chaplin & Wells 1982, Masters et al. 1988, Alonso-Mejia et al. 1997). Understanding metabolic variation in monarchs could therefore provide valuable insight into differences in flight performance, reproductive success and survival. Relatively little is known about monarch flight mus- cle metabolism or about intraspecific metabolic varia- tion in general. Individual variation in flight perfor- mance has been shown to be partly genetic in moths (Parker & Gatehouse 1985). Correlations between flight activity or capacity and metabolic differences have been seen in Agrotis ipsilon moths (Sappington et al. 1995), Colias spp. butterflies (e.g., Watt et al. 1983), Epiphyas postvittana moths (Gu 1991) and Drosophila melanogaster flies (Barnes & Laurie- Ahlberg 1986, Marden 2000). In Colias butterflies, ge- netic differences in metabolic enzymes also correlated with differing mating success by males (Watt et al. ‘Email: elevine@calpoly.edu 1985). There is evidence that monarch butterflies may have individual differences in flight ability at cool tem- peratures (Hughes et al. 1992). Further investigation of metabolic parameters may reveal some of the mech- anisms underlying this individual variation. Mitochondrial activity is a critical aspect of metabo- lism; insect flight muscle metabolism is strongly aero- bic, so most of the energy is produced by the mito- chondria (Sacktor 1976). Mitochondrial enzyme levels appear to be a good indicator of mitochondrial density and overall aerobic metabolism measured by oxygen consumption or oxidative capacity in a variety or or- ganisms (e.g., Holloszy & Booth 1976, Spina et al. 1996, Putnam & Bennett 1983). The enzymes most commonly used as markers of mitochondrial activity are succinate dehydrogenase (SDH) and citrate syn- thase (CS), mitochondrial-specific enzymes with criti- cal roles in the tricarboxylic acid cycle and therefore directly involved in the generation of ATP. Individual mitochondrial enzyme activities can vary considerably and the variation often correlates with performance. The largest interindividual variation re- ported is an approximately twenty-fold difference seen in CS activity in leg muscles of 20 toads (Bufo mari- nus) (Longphre & Gatten 1994). More commonly, dif- ferences of three- to five-fold are seen in CS activity among individuals in a variety of non-insect species (e.g., Walsberg et al. 1986, Garland & Else 1987). Less is known about individual variation in insect mitochon- drial activity, but in one study including 19 insect species, citrate synthase in flight muscle varied as much as three-fold even with only three individuals from each species (Alp 1976). Functional correlates of variation in mitochondrial enzyme activities have been demonstrated in a variety of organisms, most com- VOLUME 57, NUMBER 4 monly in terms of response to exercise. In a variety of mammals, amphibians and reptiles, correlations have been seen between mitochondrial activity and maxi- mum aerobic speed, endurance, and degree of im- provement after exercise training (Longphre & Gatten 1994, Garland & Else 1987, Cummings 1979). Variables which might be related to mitochondrial activity include seasonality and physical characteristics. Seasonal variation in mitochondrial enzyme activities has been reported in several species, including agamid lizards (Garland & Else 1987), iguanid lizards (John- Alder 1984), turtles (Olson 1987) and snapper (Majed et al. 2002); in all cases the changes are thought to re- flect seasonal patterns of activity and energetic re- quirements. In several organisms including lizards and fish, mitochondrial enzyme activity is related to body size (Garland & Else 1987, Somero & Childress 1980). Wing damage may be related to activity levels, as more damage is likely to occur during activities such as mat- ing. Higher wing damage has been seen in male mon- archs attempting to mate or drinking dew, as com- pared to those remaining in clusters (Frey et al. 1998, Oberhauser & Frey 1999, Frey et al. 2002). It is possi- ble that these males represent a more active subset of the population, which could be related to metabolic differences such as mitochondrial enzyme activity. Characterization of mitochondrial activity in overwin- tering monarchs in central California will provide basic information about metabolic variation in this popula- tion, which may be important for understanding critical variables for successful overwintering and reproduction. The purposes of this study were to determine the extent of variation in succinate dehydrogenase (SDH) activity in the male overwintering monarch population and in- vestigate possible correlations with season or physical characteristics. MATERIALS AND METHODS This study was carried out at an overwintering site at Pismo Beach State Park, Pismo Beach, California. Male monarch butterflies were collected monthly at approximately 0800 h on the following dates: 20 Octo- ber, 26 October, 21 November, and 16 December 2001; 17 January, 19 February and 14 March 2002. In all cases this was before the temperature reached the flight threshold, so all monarchs were still in clusters (Frey et al. 2002). Butterflies were collected from clus- ters using a standard butterfly net on a long pole, and 20 males were randomly selected from the net. The butterflies were placed in small ziplock bags and stored in a styrofoam cooler with icepacks until they reached the lab at California Polytechnic State Uni- versity, approximately fifteen miles away (Frey et al. 2002). At the lab we measured weight to the nearest milligram and forewing length to the nearest millime- ter from the thorax to the longest extension on the forewing. We recorded the number of damaged wings containing rips, punctures, or parts missing, and as- signed each butterfly a wing condition value of 1-4 based on lack of scales, brightness of color, and dam- age, with 1 being the worst condition and 4 the best. Each butterfly was assigned an identification number and placed in a ziplock bag to be stored at —60°C for an average of 29 days. There was no relationship be- tween storage time and enzyme activity (regression analysis, R? = 4.8%, n = 118). Succinate dehydrogenase (SDH) activity was mea- sured in a sample of thorax muscle tissue from each butterfly using the rate of reduction of the artificial elec- tron acceptor 2,6-dichlorophenolindophenol (DCIP) by procedures modified from Singer and Kearney (1957). Muscle was obtained by removing all ap- pendages from the thorax, freezing it in liquid nitro- gen, and then removing 9-11 mg thorax muscle. The tissue sample was weighed and then homogenized with a ground-glass homogenizer for 5 minutes on ice in 250 ul homogenizing buffer (0.3 M mannitol, 0.02 M phosphate, pH 7.2). The homogenate was cen- trifuged at 600 g for 10 minutes at 4°C to remove par- ticulates and large organelles. Each butterfly tissue was assayed in triplicate, using ninety-six well mi- croplates and a SPECTRAmax Microplate Spec- trophotometer (Molecular Devices Corp., Sunnyvale CA). Reactions were carried out in a total volume of 200 ul with 30 ul tissue homogenate, 1.5 x 10-4 M DCIP, 0.002 M sodium azide and 0.01 M sodium suc- cinate in assay medium (0.3 M mannitol, 0.02 M phos- phate, 0.01 M potassium chloride, 0.005 M magne- sium chloride, pH 7.2). Reactions were started by adding diluted homogenate to the other reagents, and a kinetic assay was immediately run at 600 nm every 15 seconds for 10 minutes. Only the initial linear data was used for calculations, typically the first 3 minutes. SDH activity in umoles-min™-g tissue! was calculated from the slope of the initial reaction, the extinction co- efficient of DCIP (19,100), the measured path length, and the volumes and weights used (activity = slope x l/e x 1/b X 60sec/min x assay volume/sample homog- enate vol x total homogenate volume/tissue weight x 1000 mg/1g). All statistical analyses were performed using Minitab (Minitab Inc.). RESULTS Succinate dehydrogenase (SDH) activity was deter- mined for a total of 120 butterflies. The overall dis- tribution of SDH activity for the overwintering season Frequency 3 a 00 O58 10 415 20 25 30 3.6 SDH activity SDH activity Oct. Nov. Dec. Jan. Feb. Mar. Month Fic. 1. SDH activity in male overwintering monarchs. SDH ac- tivity is given as Lmoles-min™-g tissue. A, Frequency distribution of SDH activity for the entire sampled population. B, Boxplot of monthly SDH activity: the horizontal line represents the median and the ends of the box represent the upper and lower quartiles. is shown in Fig. 1A, and ranged from 0.23-—3.40 umoles-min!-g tissue’, approximately a fifteen-fold difference. Ninety percent of the values fall between 0.5 and 2.7 umoles-min™!-g tissue, a five-fold range. Since SDH activity was measured in triplicate samples from each butterfly, the standard error associated with the mean SDH value for each gives some indication of the analytical error; the mean standard error is 0.16, approximately eleven percent on average. Given this degree of analytical uncertainty, a conservative esti- mate stills yields at a minimum a four-fold range for ninety percent of the enzyme activities and a twelve- fold range for all values. To illustrate the degree of variation in SDH activity, we calculated the coefficient of variation (CV = SD-x!-100). The overall CV for the population over the entire overwintering period was forty-seven percent while monthly CVs ranged from thirty-six percent in February to fifty-two percent in November (Table 1). Monthly distributions of SDH activity are shown in Fig. 1B and Table 1; there was clearly considerable variation within each month. Season does have a sig- nificant association with SDH activity, as seen in a one- JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TABLE 1. Basic statistics for SDH activity, body weight, and wing length. SDH activity is given as umoles-min“-g tissue. Coefficient of variation (CV) is the standard deviation divided by the mean, mul- tiplied by 100. Subscripts (a, b) for SDH activity means are based on Tukey’s pairwise comparisons; the only significant differences are be- tween October and November, and November and February. Sam- ple sizes are n = 20 for each month except for February n = 19. Mini- Maxi- Mean SD mum mum CV(%) SDH activity October 1.73, 0.77 0.57 3.40 44 November 1.07, 0.55 0.23 2.23 52 December 141, 0.58 0.60 DD 4] January 1.43, 0.73 0.43 2.98 51 February 180\ = 0166) 0573) mo CONES March 142, 071 0.49 2.86 50 All butterflies IS O70 O28 840 47 Body weight (mg) October OM 2 2 iB November 536 68 490 705 13 December 529 73 395 745 14 January 539 68 434 670 13 February 557 65 430 660 12 March 478 87 330 650 18 All butterflies 539 81 330 745 15 Wing length (mm) October 51.1 3.0 45 56 6 November 50.7 2.5 45 55 5 December 51.9 2.3 46 56 4 January 50.9 2.1 47 5D 4 February 48.2 2.8 42 53 6 March 50.1 FS 45 53 5 All butterflies 50.5 LY 42 56 5 way ANOVA for SDH activity vs. month, F = 3.33, p = 0.008 (Table 2). Post-hoc pairwise comparisons re- vealed that the only months with significant differ- ences are October vs. November and November vs. February (Table 1). In terms of seasonal pattern in the means, we regressed SDH activity on centered month data up to a fifth order polynomial, and only the third order polynomial was significant (p = 0.001); however this pattern explains very little of the variation (R° = 9.8%). Differences in the population at the beginning and end of the season may affect the results: in October only about 10% of the butterflies have arrived com- pared to the peak population in December and Janu- ary, and by March less than 10% of the butterflies are left at the overwintering site (D. Frey pers. com.). Therefore, we also analyzed the data for the core over- wintering season when the population is more com- plete. Consideration of the centered data only from November through February reveals a linear pattern (p = 0.001), but this still only explains a fraction of the individual variation (R?2 = 13.5%). The estimated in- crease in SDH activity per month is 0.22 + 0.06 umoles-min '-g tissue”. VOLUME 57, NUMBER 4 TABLE 2. ANOVA for a general linear model of SDH vs. month and all physical variables (body weight, wing length, wing damage, and wing condition). Calculations were performed for the full model and for month alone, with all physical variables removed. Source df Ss MS F p Month 5 7.4036 1.4807 3.33 0.008 Physical variables 4 1.8438 0.4610 1.04 0.391 Error 109 48.4266 0.4443 — = Total 118 57.6740 We examined two measures of size (body weight and wing length) and two measures of wing damage (number of damaged wings and overall wing condi- tion) for possible correlation with SDH activity; size data is summarized in Table 1. Mean wet body weight was found to decline from October to March, with considerable variation apart from the linear decrease (simple linear regression analysis, R* =10.4%, t = 3.69, p = 0.000). Mean wing length fluctuated through the overwintering period and declined slightly in February and March (simple linear regression analysis, R* = A £ 2 3) oO at a) 7) 300 350 400 450 500 6550 600 650 700 750 Body weight (mg) Cc e e e = ! A 2 ° i © i ° oO i 8 Ba i 2 fa) 5 ¢ 8 09) t 4 © e 8 e 8 : ) 4 2 3 4 Number of wings damaged 6.0%, t = 2.74, p = 0.007). The coefficient of variation was fifteen percent for body weight and five percent for wing length, and the CVs within each month are comparable to the overall population. None of the physical variables contributed significantly to SDH ac- tivity (comparison of ANOVA general linear model with all variables to model with only month, F = 1.04, p = 0.391; Table 2). DISCUSSION Interindividual variation in succinate dehydro- genase activity. We found a large degree of interindi- vidual variation in SDH activity in the male monarch population overwintering at Pismo Beach, at least a twelve-fold range of enzyme activities. This degree of variation is greater than that seen in other physical characteristics measured; the coefficient of variation for SDH activity is forty-seven percent compared to a CV of fifteen percent for body weight and five percent for winglength. Some of the calculated variation in SDH activity B e 2 i 2 8 © a AC e ja) ° D 8 8 e 40 45 50 55 Wing length (mm) D ® e = r e } 3 cRingeltceabtie® ie : 8 g a) 8 : j 7) 8 C e Sun iiclaabién| e e : 1 2 3 4 Wing condition Fic. 2. SDH activity and morphological variables. SDH activity is given as Umoles-min“!-g tissue’. A, SDH activity and butterfly body weight. B, SDH activity and butterfly wing length. C, SDH activity and the number of wings damaged. D, SDH activity and wing condition on a scale from 1 (worst) to 4 (best). (umoles-min '-g tissue) may be an artifact of normal- izing by tissue weight. Enzyme activities are most commonly normalized by wet tissue weight, but dry tissue weight, protein content, DNA content, and en- tire organ weight have also been used (Pelletier et al. 1995, Cascarano et al. 1978, Spina et al. 1996, Long- phre & Gatten 1994). Monarchs are known to vary in their state of hydration; in Mexico and southern Cali- fornia more dehydrated males are seen at the end of the overwintering season (Chaplin & Wells 1982, Calvert & Lawton 1993), while this pattern has not been seen in central California (D. Frey pers. com.). Individual differences in hydration status may have in- troduced some variation into individual SDH activity values, but there is no overall correlation between lower body weight and higher SDH activity; such a correlation might be expected if dehydration led to a decrease in body weight but an increase in the number of cells per tissue weight. Normalization by protein content would address this issue but could introduce other problems, since lipid reserves may be depleted at the end of the season leading to a breakdown in pro- tein (Chaplin & Wells 1982, Masters et al. 1988, Alonso-Mejia et al. 1997). Future studies will investi- gate the effects of different normalization approaches and possible changes in protein content at the end of the overwintering season. The degree of interindividual variation in SDH ac- tivity seen here in male monarchs is within the ranges found in other organisms; more importantly, it is large enough to have functional consequences since as little as a two-fold difference has been correlated with per- formance differences (Holloszy & Booth 1976). Relationship between succinate dehydroge- nase activity and other variables. Seasonal effects are small in comparison to the large variation in indi- vidual SDH activity levels in butterflies collected on the same date. Mean enzyme activity in November is significantly lower than in either October or February, and there are no significant differences between any other months. It is possible that the higher mean SDH activity in October could reflect earlier arrival by a subset of butterflies with higher mitochondrial activity. An increase in mitochondrial activity from November to February could be adaptive since the strenuous ex- ertion of mating occurs late in the season (Tuskes & Brower 1978); an increase could also be a conse- quence of prior flight activity, analogous to the exercise training effects seen in other studies. The March mean is not significantly different from February, and the downward trend seen could reflect preferential emi- gration of butterflies with higher mitochondrial ac- tivity, since only a small fraction of the overwintering JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY population remains in March. It is also possible that mitochondrial activities in November were lower than in other months because of unknown environmental factors; preliminary analysis of air temperature data did not reveal any correlations with SDH activity or un- usual occurrences in November (data not shown), but careful analysis of potential weather factors has not been done. Further investigation will be necessary to confirm that the seasonal patterns are seen consistently in different years and to determine possible causes. None of the physical characteristics examined ap- pear to be important in determining mitochondrial en- zyme activity in male overwintering monarchs. Other studies have found more linear declines in body weight during the overwintering season, but the de- gree of weight loss depends on the specific overwin- tering colony examined (Tuskes & Brower 1978, Chaplin & Wells 1982, Calvert & Lawton 1993), and it is possible that the Pismo Beach site allows better maintenance of body weight. The pattern of wing lengths seen is similar to that reported by Calvert and Lawton (1993) at Mexican overwintering sites except that they observed a more dramatic drop in late Feb- ruary and March after stability for the rest of the sea- son. They suggest that the decrease at the end of the season could be due to larger butterflies leaving the colony first. Our observation that mean body weight is lower in March but wing length is not, suggests that the decreased weight may be due to depleted lipid re- serves, dehydration, and/or breakdown of protein in tissues. Since each of these factors could affect mito- chondrial enzyme activities, the SDH activity data for March are less easily interpreted than for the rest of the season. The lack of a correlation with wing damage suggests that if there are subsets of the male monarch population with different behaviors and corresponding degrees of wing damage, mitochondrial activity is not a critical determinant nor is it significantly affected by behavioral differences which lead to differing wing damage. Since we have not examined activities di- rectly, it is still possible that differing mitochondrial enzyme levels do affect activity level, performance or mating success. Mitochondrial activity could be affected by other environmental or physical variables. Age can affect mi- tochondrial activity and flight performance; changes in mitochondrial structure, an increase in mitochondrial damage and changes in levels of some metabolic en- zymes have been reported with aging in the flight mus- cle of a variety of insect species (Sohal 1976, Ross 2000). We do not have data on the age of the monarchs in this study, but they likely vary by at least a month based on emigration data from late summer popula- VOLUME 57, NUMBER 4 tions (K. Oberhauser pers. com.). However, we found no evidence for an effect of aging on SDH activity since the entire population is aging considerably dur- ing the overwintering period and there was not a uni- directional seasonal trend. Conclusions. The primary conclusion of this study is the high degree of interindividual variation in ac- tivity of a mitochondrial enzyme in the male monarch butterflies overwintering at Pismo Beach on the cen- tral California coast. This variation is not explained by seasonality or body size and is not related to the de- gree of wing damage. Individual mitochondrial en- zyme activity variation may be partly due to genetic, nutritional, and behavioral (especially in regards to previous activity levels) differences. Mitochondrial ac- tivity has potential functional consequences for flight performance, and in the case of monarch butterflies, potential consequences for survival and reproduction. The existence of such substantial individual variation suggests that this could be an important factor in indi- vidual performance and success. Future experiments should investigate possible correlations with flight per- formance and examine mitochondrial activity in fe- male monarchs, especially with regard to energetically demanding reproductive development. In addition, further analysis of metabolic parameters at the end of the overwintering season would provide valuable infor- mation about the requirements for successful survival and reproduction during the overwintering period. ACKNOWLEDGMENTS We would like to thank Dennis Frey for guidance and help with monarch collections and morphometrics; Andrew Schaffner for help with statistical analysis; Chris Kitts for use of the mi- croplate spectrophotometer; and Sara Epperson, Mandy Lawless and Kristen Lamb for previous work developing laboratory tech- niques. Monarchs were collected under a permit from the Cali- fornia Parks and Recreation Department. We are also grateful to David Keeling, Dennis Frey, Andrew Schaffner, Maria Florez- Duquet, Carla Penz and Robert B. Srygley for review of the man- uscript. LITERATURE CITED ALONSO-MEJIA, A., E. RENDON-SALINAS, E. NONTESINOS-PATINO & L. Brower. 1997. Use of lipid reserves by monarch butterflies overwintering in Mexico: implications for conservation. Ecol. Appl. 7:934-947. ALP, P. R., E. A. NEWSHOLME & V. A. ZAMMIT. 1976. Activities of citrate synthase and NAD*-linked and NADP*-linked isocitrate dehydrogenase in muscle from vertebrates and invertebrates. Biochem. 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Adaptation at specific loci. IV. Differential mating success among glycolytic allozyme genotypes of Colias butterflies. Genetics 109:157— LB, Wart, W. B., R. C. Cassin & M. S. SwAN. 1983. Adaptation at spe- cific loci. II. Field behavior and survivorship differences among Colias PGI genotypes are predictable from in vitro bio- chemistry. Genetics 103:725-739. Received for publication 31 October 2002; revised and accepted 13 May 2003. Journal of the Lepidopterists’ Society 57(4), 2003, 291-294 ABUNDANCE OF CHLAMYDASTIS PLATYSPORA (ELACHISTIDAE) ON ITS HOST PLANT ROUPALA MONTANA (PROTEACEAE) IN RELATION TO LEAF PHENOLOGY AURORA BENDICHO-LOPEZ,! IVONE REZENDE DINIZ? AND JOHN Du VALL Hay? Instituto de Ciéncias Biolégicas, Universidade de Brasilia, ICC Sul, Térreo, Sala ATI 49, Campus Universitario Darcy Ribeiro, Asa Norte, 70910-900, Brasilia, Distrito Federal, Brazil ABSTRACT. Chlamydastis platyspora (Elachistidae) is a bivoltine species whose larva is a specialist on Roupala montana Aubl. (Pro- teaceae), a common tree in the cerrado. We studied the presence of larvae in relation to leaf phenology of its host plant in the cerr: ado sensu stricto (savannah-like vegetation) of the Field Station Agua Limpa, belonging to the University of Brasilia, Federal District, Brazil. We exam- ined 3600 plants in an area of 16.2 ha between Nov onalbten 1999 and October 2000. The host plant purtsuces leaves asynchronously during the year, and individuals present one of three leaf phenological phases at a given time: (1) new leaves only, (2) mature leaves only and (3) both ma- ture and old leaves. Larvae were found on 273 of the examined plants. Larvae were encountered between January and Mz arch (first generation) and were found only on plants of the third phenological group. Larvae were also encountered between June and October (se cond generation) and occurred predominantly on the third group. Although the host plant has a high abundance in the cerrado area the presence of ibsar ae of C. platyspora is apparently limited by the abundance of plants that simultaneously have mature and old leaves. Additional key words: Brazil, caterpillar, cerrado, feeding specialist. The center of distribution of the family Proteaceae is South Africa and Australia. This family contains 72 genera and about 1400 species but only three genera occur in Brazil: Grevillea, Euplassa and Roupala (Joly 1993, Mendonga et al. 1998). The species of Roupala occur mainly in the cerrado, but also are found in other biomes, such as the Atlantic forest. Roupala montana Aubl. is common in the cerrado sensu stricto and is found from the APA of Curiat (Amapa) (00°20’N 51°03’W) to Jaguariaiva (Parana) (24°09’S 50°18’W) (Ratter et al. 2000). The highest production of leaves of R. montana in the cerrado of central Brazil occurs during September and October, a transitional period from dry to wet season (Franco 1998). However, leaf production may occur in some individuals during the whole year, corresponding to the pattern found in several woody species of the cer- rado (Morais et al. 1995). In a study conducted in a cerrado near Brasilia in central Brazil, Diniz & Morais (1995) showed that Chlamydastis platyspora (Meyrick, 1932) (Elachisti- dae) was locally restricted to R. montana. Chlamy- dastis platyspora is bivoltine and its first generation occurs between November and April and its second between May and October (Bendicho-Lopez 2000). In spite of R. montana being common in the cerrado near Brasilia (Ratter 1980), the larvae of this moth are not found in great numbers, or with high frequency (Diniz et al. 2001). Since resources may be concentrated both in time and space plant phenology may affect the dis- persion of herbivores (Solomon 1981). The objective of the present study was: to quantify the abundance of lar- vae of C. playtspora in relation to plant leaf phenology. 'Pés-graduacao em Biologia Animal. > Departamento de Zoologia. 3 Departamento de Ecologia. MATERIALS AND METHODS Study area and its host plants. The study area, a cerrado sensu stricto, was located on the Field Station Fazenda Agua Limpa (15°55’S, 47°55’W) of the Uni- versity of Brasilia, Federal District, Brazil, at 1100 m in elevation. This region has two well-defined seasons, a dry one from May to September and a wet one from October to April (Fig, 1A). Sampling took place three times per month over 12 months from November 1999 to October 2000. Over the study period we inspected 3600 individuals (300 per month) of R. montana in an area of approximately 16 ha. All individuals inspected were between 0.5 and 1.5 m in height and there was no repetition of individ- uals over the sampling period. As C. platyspora is bi- voltine this period comprised both generations. The first generation occurs in the wet season and the sec- ond occurs during the transition period from the end of the dry season to the beginning of the wet season (Bendicho-Lopez 2000). On each collection date, 100 individuals of R. montana were surveyed for the pres- ence of larva of C. playtspora. When encountered, the developmental stage of each larva was recorded and an evaluation of the phenological stage of the leaves on the individual was also made. Detailed in- formation on the identification of the developmental stages of the larvae is given in Bendicho-Lopez & Diniz (in press). The phenological stage of the leaves was based on the density of trichomes on their abaxial surface. This characteristic was used as indicator of the relative age of the leaves, classifying them into three categories: New leaves—expanding leaves or recently expanded leaves still totally covered by trichomes on both sur- faces. This stage lasts for less than seven weeks (Fig. QA): 292 —® Precip . oll ORE 100 =| Ba cia = 250 m < 200 60 s s = 100 es (>) o & 50 20 0 0 Ark DPE Bw as ZAses eS S27 ZA6 Month —*—- NL B + ML —@— MOL 4 250 Ce} = 200 i @ 150 = 2 100 g = 50 =} Zz 0 ae Oo Ff 8 he YS fe SS s Z2AaSES TESS 246 Month Fic. 1. Mean monthly precipitation and relative humidity dur- ing the study period and leaf phenology of the host plant. A, Mean monthly precipitation (Precip) and relative humidity (R.H.) (Nov/1999-Oct/2000), data from the [BGE Meteorological Station, Brasilia; B, Variation in foliar phenophases of the individuals exam- ined of Roupala montana. NL = new leaves, ML = mature leaves, MOL = mature and old leaves. Mature leaves—expanded leaves, which had already begun to lose their trichomes. This stage lasts be- tween eight and nine months (Fig. 2B); Old leaves—leaves lacking trichomes. This stage lasts for up to two months (Fig. 2C). Individuals of R. montana were classified in three phenological groups: (1) plants with all new leaves (NL), (2) plants with only mature leaves (ML) and (3) plants with mature and old leaves at the same time (MOL). To compare the residence period and the develop- ment time of a larva on its host we followed the devel- opment of larvae on marked host plants. The resi- dence period and development time of larvae in the first generation was studied using eggs or first instar larvae found on 15 individuals of R. montana in Janu- ary. These larvae were observed twice a week through the pupal state until the emergence of the adults. This JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fic. 2. Relative age of Rouwpala montana leaves showing the abaxial face: A, New leaves: B, Mature leaves and C, Old leaves. procedure was repeated in June using larvae found on another 15 individuals of R. montana to accompany the residence period and development of larvae from the second generation. Statistical analyses were done using Statistix 7 (Ana- lytical Software 2000). RESULTS Leaf phenology of Roupala montana. January, middle of the wet season, had the largest proportion of plants with new leaves; while the maximum level for mature leaves was May (beginning of the dry season). Also at the beginning of May, there was an increase in the number of plants of the group mature and old leaves with the maximum number recorded in August (Fig. 1B). > ps at Oe © ae VIE , | as = VOLUME 57, NUMBER 4 Number of plants NL ML MOL Leaf phenological stage Number of plants NL ML MOL Leaf phenological stage Wi with larvae Dwithout larvae Fic. 3. Abundance of larvae of Chlamydastis platyspora found on leaves of Roupala montana in different phenological categories in a cerrado in central Brazil. NL = new leaves, ML = mature leaves, MOL = mature and old leaves. A, First generation (January to March); B, Second generation (June to October). Over the whole year the phenological phase with highest frequency was ML (37%), NL was next (36%) and the least abundant was MOL (27%). There was a significant difference among these values (x* = 18.79, p = 0.0001). Larval phenology and relationship with R. mon- tana. Overall we found 475 larvae on 273 of the 3600 ex- amined plants (7.58%). Among the plants used by the larvae, 4 (2%) belonged to the group with only NL, 43 (19%) on plants with OL and 226 (79%) occurred on plants MOL group. Larvae of C. platyspora were found in only 8 of 12 months and there was no overlap between the two generations. First instar larvae of the first gener- ation were found in January and developed until March, when they passed to the pupal stage. The second larval generation began in June and extended into October. Thus, the larval phase was longer in the dry season than in the wet season. In the first generation, 70 larvae were found on 46 of the 900 inspected plants (5.1%) and all individuals with larvae were members of the third phe- nological group (Fig. 3A). In the period of the second generation, 405 larvae were found on 227 of 1500 exam- ined plants (15.1%) and were present on all three phe- 293 nological groups (Fig. 3B). A test of proportions showed a significant difference in the proportion of plants with larvae between generations (z = —7.42, p = 0.000). Also the abundance of larvae differed among the three leaf phenophases in both the first (y* = 45.75; p = 0.000) and second (x° = 115.72: p = 0.000) generations. All monitored larvae used in the study of residence period and development time, from the first to the last instars, remained on their monitored plants. In both trials, all of the monitored plants belonged to the MOL phenological group. DISCUSSION Larvae of C. platyspora used the MOL phenological group of plants with the highest frequency. Therefore, for this species the results do not corroborate what is common for insect herbivores of moist tropical forests, namely that the majority use new leaves (Coley & Barone 1996). Herbivores that can use old leaves, with low nutri- tional quality, may be able to take advantage of a pe- riod of low predator density (Moran & Hamilton 1980). They also avoid physical defenses such as leaf trichomes that are present on new leaves (Pullin & Gilbert 1989, Paleari & Santos 1998). As reported by Morais et al. (1999) previous studies in the cerrado have shown a lower density of predators and para- sitoids during the dry season. The nutritional quality of leaves varies among species and over their life cycle, and young leaves gen- erally have a higher content of nitrogen ail water than mature ones, which are more fibrous. Marquis et al. (2001) showed these trends for 25 plant species in the cerrado. Herbivores are affected by nutritional quality, by the content of water and fiber, and by leaf tough- ness (Coley & Barone 1996). Mature and old leaves used by larvae of the first generation are physiologi- cally “younger” than those used by the larvae of the second generation since these leaves have been pro- duced more recently. The leaves used by the second generation could have a lower nutritional content and this could have an influence on the duration of the lar- val development. Foliar analyses of R. montana (Medeiros & Haridasan 1985) showed higher concen- trations of K and P and lower concentrations of Al, Mg and Ca in younger leaves compared to older leaves Additionally, the larvae that develop in the dry season face more extreme climatic conditions, such as the ab- sence of rain, and low relative humidity, as well as the lowest temperatures of the year. The slower larval dey lopment observed in the dry season versus the wet season is not exclusive to C. platy- spora. In the same study area, similar growth rates were recorded for larvae of Cerconota achatina (Zeller) (Elachistidae), which feeds on Byrsonima coccolobifolia, B. pachyphylla (=B. crassa) and B. verbascifolia (Mal- phigiaceae). Generally, larvae of this species collected in the dry season and raised in the laboratory took twice the time to develop as those collected during the wet season (Morais et al. 1999). Here abiotic variability was reduced so differences were due to differences in leaf quality. The number of larvae in the second generation was seven times higher than that of the first generation. This result is similar to that in another study of C. platyspora larva on R. montana done by another col- lector (B. Cabral, unpublished). These results obtained for C. platyspora coincided with the seasonal pattern of the lepidopteran larvae established for the cerrado by Morais et al. (1999), who showed a largest proportion of plants with larvae in May to July (dry season). Our results showed the close relationship of C. platyspora larvae with leaf phenology of R. montana. This association appears to affect the size of the popula- tions of both generations and can explain the low occur- rence of larvae on its host plant, in spite of the high lo- cal density of R. montana. The limiting resource are plants belonging to the third phenological group (ma- ture and old leaves) at the time of oviposition (May and December). Thus, the proportion of individuals of R. montana bearing different leaf phenophases can explain the low occurrence of this specialist larva in the cerrado. ACKNOWLEDGMENTS We would like to thank Dr. Vitor O. Becker, who identified the moth species, offered information and bibliography for the comple- tion of this work. Dr. Helena C. Morais (Universidade de Brasilia) and Dr. Aldicir Scariot (EMBRAPA) made valuable suggestions on an early draft of the manuscript and many valuable suggestions were made by an anonymous reviewer. Mr. Kiniti Kitayama and Mr. Fausto Goncalves took the photographs CAPES provided a M.S. scholarship to the first author. LITERATURE CITED ANALYTICAL SOFTWARE. 2000. Statistix 7.0. Analytical Software, Inc. Tallahasse, 333 pp. BENDICHO-LopEZ, A. C. 2000. 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SOLOMON, B. P. 1981. Response of a host-specific herbivore to re- source density, relative abundance, and phenology. Ecology 62:1205-1214. Received for publication 14 October 2002; revised and accepted 10 June 2003. SS ee ee ee ae eee ee Se a GENERAL NOTES Journdl of the Lepidopterists’ Society 57(4), 2003, 295-298 NEW RECORDS ON THE DISTRIBUTION AND ECOLOGY OF COMMON GEM BUTTERELY, PORITIA HEWITSONI HEWITSONI MOORE FROM THE LOWER WESTERN HIMALAYAS: A LESSER KNOWN TAXA Additional key words: geographical distribution, seasonality, abundance, habitats, habits, larval food plant. The common gem butterfly, Poritia hewitsoni hew- itsoni Moore (1866) (Poritiinae: Lycaenidae), is en- demic to the Oriental (Indo-Australian) region (Fig. 1). Its distribution extends from Kumaon in northern India in the west, up to north Thailand in the east, through the lower Himalayan tracts in Nepal, Sikkim, W. Bengal (Dargeeling), Bhutan up to parts of north- east India (Assam and Meghalaya [Khasi hills]), ex- treme south-east Bangladesh (Chittagong hill tracts) and north Myanmar (Chin, Arakan and Karen hills, Chindwin, Pegu) (De Niceville 1890, Bingham 1907, Swinhoe 1910-11, Evans 1932, Wynter-Blyth 1957, d Abrera 1986, Mani 1986, Haribal 1992). W. Doherty collected one male and one female of this species from Kali river valley at Garjighat near Kumaon-Nepal bor- der (approx. 80°07’E and 29°12/N) and this record is considered to be the western most limit in the distri- bution of this species (Hannyngton 1910). In east to central Nepal, P. h. hewitsoni occurs in lower midlands from 160 m to 1050 m (Lamjung, Rupandehi, Chitwan districts) as a locally abundant, fairly common species found during winter. It has also been recorded in March, April, August, September, November and De- cember months, on trees in jungle clearings, riverine and sal, Shorea robusta flowers from Nepal (Smith 1989, 1997). However, in Sikkim it is not easily recorded presumably as it flies high among the trees and goes unnoticed as it flies around rapidly to settle on leaves in jungle country at low elevations (Mangan and Rangpo areas) during October and November (Wynter-Blyth 1957 & Haribal 1992). In Darjeeling (north Bengal) one male was collected in March (Maude 1949). Its life history and food plants have so far not been recorded and only its egg has been de- scribed by W. Doherty as ‘truncate pyramid in shape, half again as long as wide with two vertical and sloping and two horizontal faces, reticulate above as is usual in the family Lycaenidae’ (De Niceville 1890). The tuft of hairs present on the hind wings of this butterfly are known to produce a pleasant perceptible odor (Haribal 1992). Recently, this butterfly was collected from the New Forest campus (8 individuals in August 1988 on a guava tree) and adjoining forested slopes of Tons valley (10+ recorded on November 1989 on a mango tree in an open and mixed sal forest) (Singh 1999). Both the places lie in the Dehra Dun valley (77°40’E to 78°15’E and 30°00’N to 30°35/N), in Uttaranchal state of northern India, which lies further west to Ku- maon, the known western most limit for the distribu- tion of this species. Later, this species was also col- lected from Paonta valley (4 individuals from a sal forest edge at Rajban in July 1996) and Nahan (77°20’E to 30°33’N) (one specimen [male] observed in a mixed sal forest with Terminalia tomentosa trees besides the road near Shambuwala in November 1999). These places lie in the Sirmaur district of Hi- machal Pradesh state, which is further west to Dehra Dun district. Even Mackinnon and De Niceville (1899) who had studied butterflies of Mussoorie and neigh- boring regions during all the seasons for 11 successive years (1887-98), had not record this species in Dehra Dun district. One reason could have been non assess to Paonta valley and Nahan due to poorly developed road communication at that time. As there were no previous records of this butterfly from the western Himalayas, I decided to carry out ex- tensive surveys in Dehra Dun valley to know more about the distribution and ecology (seasonality, food plants, breeding time, habits, habitat, life history, etc.) of this lesser known butterfly species in the lower west Himalayan tracts of Uttaranchal state. Study area. The Dehra Dun valley lies between the west Himalayan mountain ranges in the north and the Shiwalik range running parallel to it in the south at a mean altitude of 485 m and covers an area of ca. 1920 kim’. In the west it is bordered by the river Yamuna and in the east by the river Ganga. The valley is also well watered by perennial streams. The mountain slopes on the north and south sides of the valley are covered with pure and mixed forests dominated by sal, Shorea ro- busta (tropical moist deciduous sal forests or TMDSF; Champion & Seth 1968). These forests cover 51-58% of Dehra Dun valley (FSI 1995). Mixed stands have Terminalia tomentosa, T.belerica, Adina cordifolia, Lagerstromia parviflora, Mallotus philippensis, Lannea cormondalica, Syzygium cumini trees, as other domi- nant species besides sal. The valley receives ca. 200 cm rainfall annually, mostly during the monsoons (June-September). The temperature fluctuates be- 296 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 60°E 70°E ore 80°E 90°E 100°E aa ee | 2 PEE valley (HP) P Paonta valley (H.P.) . Mangan & hra Du Rangpo j sigh (Sikkim) CHINA {Uttranchal) ee Ne Ree district BHUTAN 30°N ° a / SS a oo Me, S J *e °° sam aN 5 “SeNEP. un the we ww 4 J oF ‘eo @! 7 i @ hasi lp ) ons & > mre (Meghalaya) Gharjighat, Kali valley (Kumaon -Nepal er gee 2 JD galoh wh Pender Rupendehi & ‘ & - . _ Q lievamdia 2 Ie : vain @ Previous distribution records ---— Previously known westward distributional limit A New records (present survey) Westward distributional limit of larval food plants inTMDSF , * -Continuous 4 , ‘ Vv Shan states hittag6ng Hills ie 20°N = Chisthills er @vor THAILAND Arakan Hills é $ ‘ : Wore Ve patches a sth SRA, oir cio a Fic. 1. Map depicting the geographical distribution of common gem butterfly, Poritia hewitsoni hewitsoni Moore in the Oriental region and the location of collection sites in Tropical moist deciduous sal forests (TMDSF) from where it was recently recorded. tween —1°C to 43.9°C from winter to summer. The dis- tribution of the seasons in the area is as follows: Spring (March-April); Pre-Monsoon (May-June); Summer/ Monsoon (July-August); Post-Monsoon (September— October); Autumn (November—December); Winter (January-February ). Study sites and sampling. A total of 5 sites (Baar- wala, Jhajra, Thano, Timli & Karvapani forest areas) each covering a continuous area of 4 km? and repre- senting the TMDSF, spread over the valley were se- lected for sampling. Sampling of each site for butter- flies was done visually by walking and counting the number of individuals of butterfly species on a line transect for 30 minutes during sunshine. In all 8 line transects were covered in each site totaling to 4 h of sampling period in 2 successive days (2 h/day in a stretch). All the three strata (canopy, middle story and ground level) were sampled for butterflies with the help of binoculars and butterfly nets. Only a few voucher specimens were collected for identification of difficult species. Destructive sampling was kept to the minimum. Each site was thus sampled once in two months for two successive years (July 2000—August 2002), based on the methodology adopted by Blair and Launer (1997). Seasonality and abundance. P. h. hewtisoni spec- imens (both male and female) were recorded from all the 5 sites. This species was found to be relatively lo- cally abundant as compared to other butterflies, being collected in almost half (46%) of the total samplings. The data on the number of individuals collected from different sal forest sites in Dehra Dun valley is given in the Table 1. The flight period of P. h. hewitsoni in the lower western Himalayas, as recorded in this study, is from spring to autumn seasons with higher abundance in July-August (monsoons) when it also breeds. Habits and Habitat. Most of the collections were made in edges/ openings of sal forest. Large assem- blages of this butterfly were recorded (a) while nectar feeding on flowering Syzygium operculata trees grow- ing besides a stream (riverine) in the company of Large Oak Blue, Aropala amantes Hewitson and Com- mon Silverline, Spindasis vulcanus Fabricius, butter- flies (Baarwala); (b) in the edge of a sal forest growing VOLUME 57, NUMBER 4 TABLE 1. busta forest sites in Dehra Dun valley, the lower western Himalayas Year Season Month Baarwala 2000 Monsoon July August 31 Post-Monsoon Sept October Autumn November 1 December 2001 Winter January February Spring March April Pre-Monsoon May June Monsoon July August 1 Post-Monsoon Sept October Autumn November 2 December 2002 Winter January February Spring March April Pre-Monsoon May June Monsoon July August 297 Common gem butterfly, Poritia hewitsoni hewitsoni Moore individuals recorded* from tropical moist deciduous sal, Shorea ro- Sites Jhajra Thano Timli Karvapani 1 ] 2 18 22 4 3 23 16 i) it 1 1 1 1 5 * Recorded in 4 h of sampling time period in 2 successive days and covering 8 transects in an area of 4 km? for each site. ? to} in mixed association with tall Teminalia tomentosa trees (in flowering) and Mallotus phillipiensis trees oc- cupying lower story below it (Jhajra); (c) degraded, ex- tensively lopped open, pure sal forest (Thano); (d) in small openings in a dense, mixed sal forest having closed canopy, on bushes and dry leaves present on the forest floor in late spring (Karvapani and Timli, both sites lo- cated on the Shiwaliks). During November it was ob- served basking on tree tops (canopy) of medium to tall trees (Timli). It was not recorded outside sal forest ar- eas. Adults were recorded being predated by spiders. Larval food plants and breeding. Fourth and fifth instar larvae were recorded feeding on tender and mature leaves of sal, Shorea robusta and Sain, Termi- nalia tomentosa trees during the rainy season (in Au- gust at Karvapani , Jhajra and Baarwala). Brief life history. Larvae: As many as 45 fourth and fifth instar larvae recorded feeding together in a group (like a pack of cigarettes), half in line above the leaf surface while rest of the half below the leaf surface in such a way that all the mouths feed together in a line, on a leaves of S. robusta and T. tomerntosa trees, dur- ing day time (Karvapani, August) (Fig. 2). This may be an adaptation for protection against natural enemies by giving them a confusing effect as collectively they appear to be one single mass covering the leaf surface from both the sides, making it very difficult to judge its actual size and shape. Pupae: Pale in color with a line of black spots on the 2 margins, 10 mm long which were found attached to the upper surface of fresh leaves of young sal trees in an open forest (Bar- wala, August) and also on the leaves of a climber Mil- letia auriculata in sal forest (Timli, September). Pupal RIG, 2. Common gem butterfly, Poritia hewitsoni hewitsoni Moore larvae feeding on sal (Shorea robusta) leaf. 298 period was recorded to 2-3 days in August and 3-4 days in September. Adults: Wing span: 28-39 mm. In April and August males appear to be fresh with bril- liant metallic blue colors. Sex ratio of adult butterflies on emergence from one group of 45 larvae brought from the field (August-Karvapani), was found to be 1:1. Adult longevity in laboratory ranged from 6-16 days (August) when kept in breeding cages and fed honey-sugar solution. The present study is part of a project (FRI-145/FED-9) on ‘but- terflies and land use in the lower western Himalayas’ being carried out by Entomology Division, Forest Research Institute (FRI), Dehra Dun, India being funded by the Indian Council of Forestry Research and Education. The author is thankful to the Director (FRI) and Head, Entomology Division (FRI) for extending their support and facilities. Thanks are also due to B.C. Pandey and R. Kumar (Technical Assistants) for extending their help in collection of data and insect material form the field. LITERATURE CITED BINGHAM, C. T. 1907. Fauna of British India, Ceylon and Burmah. Vol. II Taylor and Francis, London. Buair, R. B. & A. E. LAUNER. 1997. Butterfly diversity and human land use: species assemblage along an urban gradient. Biologi- cal Conservation 80:113-125. CuamPion, H. G. & S. K. SETH. 1968. Forest types of India. Gov- ernment of India Publication, Delhi. D’ABRERA, B. 1986. Butterflies of the Oriental region. Part-II (Ly- caenidae and Riodinidae). Hill House, Australia. Evans, W. H. 1932. The identification of Indian butterflies. Bom- bay Natural History Society, Bombay. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY FSI. 1995. The state of forest report. Forest Survey of India, Kaulagarh Road, Dehra Dun. HANNYNGTON, F. 1910. The butterflies of Kumaon. Parts I & II. Journal of Bombay Natural History Society 20:130-142, 361-372. Haripat, M. 1992. Butterflies of Sikkim and their natural history. Sikkim Nature Conservation Foundation. Thompson Press, New Delhi. MACKINON, P. W. & L. DE NICEVILLE. 1899. List of butterflies of Mussoorie in the Western Himalayas and neighboring region. Journal of Bombay Natural History Society 11:205-221, 368-389, 585-605. Mani, M. S. 1986. Butterflies of the Himalaya. Oxford and IBH Publishing Co. Ltd., London. MAuDE, E. W. 1949. List of butterflies caught in the Darjeeling district between 1700ft and 4400ft. Part III. Journal of Bengal Natural History Society, Darjeeling 24 (1):31-34. SINGH, A. P. 1999. New Forest, Dehra Dun, India: a unique man- made habitat for butterflies in the Lower Western Himalayas. Indian Forester 125 (9):913-922. SMITH, C, 1989. Butterflies of Nepal (Central Himalayas). Crafts- men Press, Bangkok. . 1997. Butterflies of Royal Chitwan National Park, Nepal. Tecpress Books, Bangkok, Thailand. SWINHOE, C. 1905-1913. Lepidoptera indica. Parts VIII-X. Lovell Reeve Co. Ltd., London. WynTER-BLYTH, M. A. 1957. Butterflies of the Indian Region. Bombay Natural History Society, Bombay. ARUN P. SINGH, Entomology Division, Forest Re- search Institute, ICFRE, P.O. New Forest, Dehra Dun, Uttranchal, India—248 006; email: singhap @icfre.org Received for publication 25 September 2002; revised and accepted 2 June 2003. Journal of the Lepidopterists’ Society 57(4), 2003, 299-303 ALEXANDER DOUGLAS CAMPBELL FERGUSON (1926-2002) Douglas C. Ferguson, 1926-2002 1979, National Museum of Natural History Staff Directory Dr. Douglas C. Ferguson (Doug to everyone who knew him), a charter member, past president, and hon- orary member of The Lepidopterists’ Society, died on 4 November 2002 following surgery on 16 October. Doug was born in Halifax, Nova Scotia on 17 February 1926, attended local schools, and received a B.S. from Dalhousie University in 1950. His M.S. (1956) and Ph.D. (1967) were awarded by Cornell University. His doctoral thesis was a revision of the green Geometridae. He was a field assistant to J. H. McDunnough in 1946; Curatorial Assistant, Curator of Entomology, and Chief Curator (Science Division) at the Nova Sco- tia Museum (1949-63): Research Associate in Ento- mology (Peabody Museum of Natural History) then Research Staff Biologist and Lecturer (Department of Biology) and Curatorial Associate in Entomology (Peabody Museum of Natural History), Yale Univer- sity (1963-69); and Research Entomologist, System- atic Entomology Laboratory U.S.D.A. at the National Museum of Natural History (1969-96). Upon retire- ment he continued as a Collaborator of the U.S. De- partment of Agriculture and Research Associate of the Smithsonian Institution. Doug's interest in natural history began in childhood when he seriously watched birds and discovered the nests of most local species. After reading W. J. Hol- land’s account of sugaring for moths in The Moth Book in 1941, he tried it on the trees around his home and was thrilled to catch five species of Catocala the first night. Halifax was a small city with many collecting sites within walking or cycling distance, and it had a mu- seum with a collection of local Lepidoptera, a library, and a helpful director. Doug's initial involvement with the Lepidoptera increased exponentially and resulted in The Lepidoptera of Nova Scotia, part 1, Macrolepi- doptera in 1954. He was deeply influenced by Mc- Dunnough, W. T. M. Forbes, Charles Remington, and John Franclemont during his formative years. Throughout his career Doug was an avid, know- ledgeable collector. Field trips were directed to learn and document the fauna of particular areas. He used black and incandescent light and bait as attractants, and In later years he used traps to augment the array of species sampled in an area. Despite being behind on spreading, sometimes he would collect butterflies during the day. Doug collected in the southern parts of the Provinces and all States but Hawaii. He spread and labelled an estimated 200,000+ specimens during his career. These specimens have augmented significantly the holdings of the U.S. National Museum of Natural 300 te Doug Ferguson and Paul Opler collecting at Pefia Blanca Lake, Arizona (August 1999). Photo courtesy Evi Buckner-Opler. History, the Peabody Museum of Natural History, and the Nova Scotia Museum. Doug and I had several joint field trips in South Carolina, Texas, Utah, Colorado, and Nebraska. We would stay in a “permanent” base and collect in several sites within reasonable driving distance. I was respon- sible for the evening meal while he handled the clean- up. During the day we would sit and spread moths, of- ten in silence, until some chance thought, often about the tentative identity of a specimen, elicited conversa- tion. Optimally, a public radio station was available that enabled us to enjoy classical music. Because Doug recognized so many moths, his collecting resulted in series of uncommon or unknown entities and three or four pairs of common species. He was extremely inter- ested in learning the life history of species and reared to the adult stage more than 600 species, documenting many of them with 35 mm slides of the larvae and adults. Often, he would bring fertile females, which were collected late in a trip, home and effect the rear- ing there. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Doug Ferguson preparing plate for a MONA fascicle at Wedge Plantation (1978). Photo courtesy Charles V. Covell, Jr. A chance meeting in 1967 with Richard B. Do- minick, a Yale alumnus and Lepidopteran enthusiast, at the Peabody Museum led to several collecting trips at The Wedge, Dick’s estate near McClellanville, South Carolina. Here began the Moths of North America project and subsequently the establishment of the Wedge Entomological Research Foundation, which funds and publishes the series. Doug enlisted the participation of John Franclemont, Eugene Munroe, and me for the project, originally projected to be a synoptic update of Holland’s moth book. Stud- ied consideration led to the project’s present scope of an anticipated 130+ fascicles to treat the estimated 16,000+ species in the area. Doug contributed fasci- cles on the Saturniidae, Lymantriidae and Geometri- nae and had the text and line drawings completed for a major revision of the geometrid tribes Cassymini and Macariini before his death. Doug was an excellent field biologist who interacted and collaborated with many Lepidopterists. As well, he aided many collectors by identifying specimens and occasionally describing species whose identity was needed for economic or biologic purposes. Doug had VOLUME 57, NUMBER 4 two students: Roger Heitzman (Ennominae) and Alma Solis (Pyraloidea). He was very generous with his knowledge and would drop what he was doing to an- swer their questions. Doug was a quiet, thoughtful, well-read person who had many interests, history, gar- dening, and music among them. He often brought a different and valued view to discussions. Doug is survived by Charlotte, his wife of 49 years, daughters Stephanie and Caroline, and six grandchil- dren. Additionally, he is held in high esteem and is sorely missed by many friends and colleagues. RON HODGES, 85253 Ridgetop Drive, Eugene, Ore- gon 97405-9535, USA Received and accepted for publication 3 June 2003. PUBLICATIONS — D. C. FERGUSON Fercuson, D. C. 1950. Collecting a little-known Papilio (Lepi- doptera, Papilionidae). Lepidopterists’ News 4:11—12. FERGUSON, D. C. & L. R. RUPERT. 1951. The results of a collecting trip to the Gaspé Peninsula. Lepidopterists’ News 5:53-54. Fercuson, D. C. 1953. On the identity and status of Eubaphe lamae Freeman (Lepidoptera: Arctiidae). Canadian Entomol. 85:371-373. . 1953. Revision of the occiduaria-argillacearia complex of the genus Itame, with descriptions of new races (Lepidoptera: Geometridae). Canadian Entomol. 85:453-461. . 1954. The Lepidoptera of Nova Scotia, part 1, Macrolepi- doptera. Proc. Nova Scotian Inst. Sci. 23:161—375, pls. 1-16. . 1954. A revision of the genus Hypenodes Doubleday, with descriptions of new species (Lepidoptera: Phalaenidae). Cana- dian Entomol. 86:289-298. . 1955. A nearctic race of Hiibner, with remarks on the sta- tus of montana Packard (Lepidoptera: Phalaenidae). Bull. Brooklyn Entomol. Soc. 50:23-27. . 1955. The status of Perizoma grandis Hulst (Lepidoptera: Geometridae). Bull. Brooklyn Entomol. Soc. 50:54-56. . The North American species of Calocalpe Hiibner (Lepi- doptera: Geometridae). Canadian Entomol. 87:325-330. . 1957. Investigations of the Lepidoptera of Newfoundland, I, Macrolepidoptera (H. KRoGERUS, 1954, Acta Zoologica Fen- nica 82:1-80). Lepidopterists’ News 10:175-176. (Book Re- view). . 1958. Notes on Larentiinae—new records and corrections (Lepidoptera: Geometridae). Canadian Entomol. 90:42-43. . 1958. Entomology at the Nova Scotia Museum of Science. Canada Dept. Agr. Science Serv., Entomol. Div. Newsletter 36 (4):1-2. (Popular Publication). . 1963. Leucania comma in North America (Lepidoptera: Noctuidae). Canadian Entomol. 95:105—107. . 1963. Taxonomic and biological studies of the noctuoid and geometroid moths of Florida. American Phil. Soc. Year- book, 1962:285-288. (Report). . 1963. James Halliday McDunnough. Nova Scotia Museum Newsletter 3 (4):44-46. (Obituary). . 1963. Field work in the South. Nova Scotia Museum Newsletter 3 (4):46-55. (Popular Publication). . 1963. James Halliday McDunnough—a biographical obit- uary and bibliography. J. Lepid. Soc. 16:209-228. . 1963. Immature stages of four nearctic Notodontidae (Lepidoptera). Canadian Entomol. 95:946-953. . 1965. A new North American noctuid of the genus Anomogyna (Insecta: Lepidoptera). Postilla (Yale) 89:1-11. 301 . 1967. Insect studies in the Crazy Mountains, Montana. Discovery (Yale Peabody Museum magazine) 2:11—20. (Popular Publication). 1969. A revision of the moths of the subfamily Geometri- nae of America north of Mexico. Yale Peabody Museum Bull. 29:1-251, pls. 1-49. (PhD. thesis). . 1971. Saturniidae (part 1), In Dominick, R. B. et al., The moths of America north of Mexico, fasc. 20.2A, Bombycoidea. E.W. Classey Ltd. and RBD Publications, Inc., London. Pp. 1-153, pls. 1-11. . 1972 Saturniidae (part 2), In Dominick, R. B. et al., The moths of America north of Mexico, fasc. 20.2B, Bombycoidea. E.W. Classey Ltd. and RBD Publications, Inc., London. Pp. 154-275, xv—xxi, pls. 12-22. . 1972. Two new conifer-feeding species of the genus Semiothisa (Lepidoptera: Geometridae). Canadian Entomol. 104:563-565. . 1972. The occurrence of Chloroclystis rectangulata (L.) in North America (Geometridae). J. Lepid. Soc. 26:220-221. . 1972. New records of Lepidoptera from the United States (Arctiidae, Geometridae, Epiplemidae). J. Lepid. Soc. 26:222-995. . 1973. The identity of Macaria inaptata Walker and Itame varadaria (Walker) (Geometridae). J. Lepid. Soc. 27: 288-290. . 1973. An index to the described life histories, early stages and hosts of the Macrolepidoptera of the continental United States and Canada (H. M. TierTz, 1972, 1041 pp. in 2 vols. Allyn Museum of Entomology, Sarasota, Florida). J. Lepid. Soc. 27:309-310. (Book review). . 1973. The species of the genus Tacparia Walker (Lepi- doptera, Geometridae). Proc. Entomol. Soc. Washington 75:467-478. . 1974. The relationship of Holomelina costata (Stretch) and H. intermedia (Graef), with revised synonymy (Arctiidae). J. Lepid. Soc. 28:14. . 1974. Moths of the Semiothisa signaria complex (Lepi- doptera: Geometridae). Canadian Entomol. 106:569-621, figs. 1-235. . 1974. A new species of the genus Semiothisa from the southeastern United States (Geometridae). J. Lepid. Soc. 28:297-301. BLANCHARD, A. & D. C. FERGUSON. 1975. Rostrolaetilia—a new North American genus of the subfamily Phycitinae, with de- scriptions of seven new species (Pyralidae). J. Lepid. Soc. 29:131-150. SaBROsky, C. W. & D. C. FERGUSON. 1975. A challenge to the family name Attacidae (Insecta: Lepidoptera). Bull. Zool. Nomenclature 32:149-153. FERGUSON, D. C. 1975. Host records for Lepidoptera reared in eastern North America. U.S. Dept. Agr. Tech. Bull. 1521:1—49. . 1976. The correct name for the gypsy moth. U.S.D.A. Coop. Plant Pest Report 1 (9):83-84. . 1976. The butterflies of the Far East USSR (A.I. KURENT- ZOV, 1970, Acad. Sci. USSR, Siberian Div.). Proc. Entomol. Soc. Washington 78:490-491. (Book Review). . 1977. A new North American species of Apamea formerly confused with A. verbascoides (Guenée) (Noctuidae). J. Lepid. Soc. 31:57-62. . 1978. Winter Moth, Operophtera brumata (L.) (Lepi- doptera: Geometridae)—a contribution to the series entitled Pests not known to occur in the United States or of limited Dis- tribution. U.S.D.A. Coop. Plant Pest Report 3:687-694. . 1978. Lymantriidae. In Dominick, R. B. et al., The moths of America north of Mexico, fasc. 22.2, Noctuoidea. E. W. Classey and The Wedge Entomological Research Foundation, London. 110 + x pp., 23 text figs., 9 pls. . 1979. A new ghost moth from the southern Appalachian Mountains (Hepialidae). J. Lepid. Soc. 33:192-196. FLETCHER, D. S., I. W. B. Nye (British Museum) & D.C. FERGU- SON. 1980. Lymantriidae Hampson, 1893 (Insecta, Lepi- 302 doptera). Proposed precedence over Orgyiidae Wallengren, 1861, and Dasychiridae Packard, 1864. Z.N. (S) 2216. Bull. Zool. Nomenclature 37:40-48. FERGUSON, D. C. 1980. Response to J.C.E. Riotte’s review of the lymantriid fascicle of The Moths of America North of Mexico. J. Res. Lep. 17:265-267. FERGUSON, D. C. & V. A. BRou. 1981. A new species of Automeris Hiibner (Saturniidae) from the Mississippi River Delta. J. Lepid. Soc. 35:101-105. FEeRGuSON, D.C. 1982. The moths and butterflies of Great Britain and Ireland, Vol. 9, Sphingidae-Noctuidae (Part 1) (John Heath, A. Maitland Emmet et al., 1979. Curwen Books, North Street, Plaistow, London, E13 9HJ, England). J. Lepid. Soc. 35:331. (Book Review). . 1982. First occurrence of Perizoma alchemillata (Lepi- doptera: Geometridae) on the mainland of North America. Canadian Entomol. 114:543. . 1982. Butterflies and moths of Newfoundland and Labrador. the Macrolepidoptera. (R. F. Morris, 1980. Agricul- ture Canada, Research Branch, Publ. 1691. 407 pp.). Bull. En- tomol. Soc. Canada 14:56. (Book Notice). LAFONTAINE, J. D., J. G. FRANCLEMONT & D. C. FERGUSON. 1982. Classification and life history of Acsala anomala Benjamin (Arc- tiidae: Lithosiinae). J. Lepid. Soc. 36:218-226. FERGUSON, D. C. 1982. A revision of the genus Meropleon Dyar, with descriptions of a new species and subspecies (Lepidoptera: Noctuidae). Entomography 1:223-235. . 1982. A revision of the genus Macrochilo Hiibner (Lepi- doptera: Noctuidae). Entomography 1:303-332. . 1983. Families Thyatiridae, Drepanidae, Geometridae (except Sterrhinae), Epiplemidae, Sematuridae, Uraniidae, Sat- urniidae, and Lymantriidae. Pp. 88-109, 119, 120 In Hodges, R. W., et al., A check list. of the Lepidoptera of America north of Mexico. E. W. Classey and The Wedge Entomological Research Foundation, London. 284 pp. . 1983. A new genus and species of geometrid moth from Texas (Lepidoptera: Geometridae). J. Lepid. Soc. 37:24-28. . 1983. The Cutworm Moths of Ontario and Quebec (E.W. Rockbume & J. D. Lafontaine, 1976. Agriculture Canada, Re- search Branch, Publ. 1593). J. Lepid. Soc. 37:96. (Book Review). . 1983. The identity of two monotypic geometrid genera wrongly attributed to the nearctic fauna (Lepidoptera: Geometridae). J. Lepid. Soc. 37:146-147. . 1983. On the status of Pseudothyatira expultrix (Grt.) and Euthyatira pennsylvanica J.B. Smith (Thyatiridae). J. Lepid. Soc. 37:179-180. . 1983. Butterflies and moths of Newfoundland and Labrador. the Macrolepidoptera. (R. F. Morris, 1980. Agricul- ture Canada, Research Branch, Publ. 1691. 407 pp.). J. Lepid. Soc. 37:189-192. (Book Review). FERGUSON, D. C., A. BLANCHARD & E. C. KNUDSON. 1983. A new genus and species of Geometridae (Lepidoptera) from Big Bend National Park, Texas. Proc. Entomol. Soc. Washington 85:552-556. FEerGuSON, D. C. 1983. Pests not known to occur in the United States or of limited distribution, No. 40: Bean Pod Borer, Maruca testulalis (Geyer). APHIS-PPQ, U.S. Dept. Agriculture. 6 pp. . 1984. Two new generic names for groups of holarctic and palearctic Arctiini (Lepidoptera: Arctiidae). Proc. Entomol. Soc. Washington 86:452—-459. FERGUSON, D. C., A. BLANCHARD & E.. C. KNUDSON. 1984. A new species of Neodavisia Barnes and McDunnough (Lepidoptera: Pyralidae) from southem Texas. Proc. Entomol. Soc. Washing- ton 86:769-772. FERGUSON, D. C. 1983. The preparation and use of fermented peach moth bait. Maryland Entomol. 2:52-53. (Technique). . 1985. Geometridae, subfamily Geometrinae. In Domi- nick, R. B., et al., The moths of America north of Mexico, fasc. 18.1. The Wedge Entomological Research Foundation, Wash- ington. 131 + 5 unnumbered pp., 4 col. pls. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY . 1985. Contributions toward reclassification of the world genera of the tribe Arctiini, part 1—introduction and a revision of the Neoarctia-Grammia group ( Lepidoptera: Arctiidae: Arc- tiinae). Entomography 3:181—275, 155 figs. COVELL, C. V., D. C. FERGUSON & G. B. STRALEY. 1986. Ennomos alniaria (Lepidoptera: Geometridae), a European moth re- cently discovered in British Columbia. Canadian Entomol. 118:499-501. FERGUSON, D. C. 1986. Biogeography and ecology of the Island of Newfoundland (G. R. SouTH et al., 1983. W. Junk Publishers, The Hague, Netherlands). Bull. Entomol. Soc. America, Fall 1986, pp. 176-177. (Book Review). . 1986. Mounting specimens, and parts of other chapters. In G. C. Stayskal, W. L. Murphy & E. M. Hoover (eds.), Insects and mites: techniques for collection and preservation. USDA Misc. Publ. 1443. (Technique). FERGUSON, D. C. & E. C. KNUDSON. 1987. Four new United States records of moths from Texas. J. Lepid. Soc. 40:353-354. FERGUSON, D. C. 1987. Xanthorhoe clarkeata (Geometridae), a new species and possible endemic of the Queen Charlotte Is- lands, British Columbia. J. Lepid. Soc. 41:98-103. Davis, H. G., L. M. MCDONNOUGH & D. C. FERGUSON. 1987. Sex attractant for Scoparia biplagialis (Lepidoptera: Pyralidae). Proc. Entomol. Soc. Washington 89:500-501. WHITTLE, K. & D. C. FERGUSON. 1987. Melon Moth, Diaphania indica (Saunders). Pests not known to occur in the United States or of limited distribution, No. 84. APHIS-PPQ. 8 pp. WHITTLE, K. & D. C. FERGUSON. 1987. Eggplant Fruit Borer, Leu- cinodes orbonalis Guenée. Pests not known to occur in the United States or of limited distribution, No. 85. 9 pp. FercUSON, D. C. 1988. Systematics of Stamnodes animata (Pearsall) (Lepidoptera: Geometridae). Pp. 7-10, In M. L. Fur- niss, D. C. Ferguson, K. W. Voget, J. W. Burkhardt, A. R. Tiede- mann & J. R. Oldemeyer, Life history and ecology of the geometrid moth, Stamnodes animata (Pearsall), on curlleaf mountain mahogany in Nevada. Fish and Wildlife Research 3:i-iv, 1-26. . 1988. New species and new nomenclature in the Ameri- can Acronictinae (Lepidoptera: Noctuidae). J. Res. Lep. 26: 201-218. . 1989. “Feature Photograph.” J. Lepid. Soc. 43:79. [Repre- sents first natural interfamilial mating recorded in the Lepi- doptera]. TEPEDINO, V. J., A. K. KNApp, G. C. EICHwort & D, C. FERGUSON. 1989. Death Camas (Zigadenus nuttalli) in Kansas: pollen col- lectors and a florivore. J. Kansas Entomol. Soc. 62:411—-412. WuiTtLe, K. & D. C. Fercuson. “1988” [1991]. Spotted Stalk Borer, Chilo partellus (Swinhoe). Pests not known to occur in the United States or of limited distribution, No. 96. APHIS- PPQ. 10 pp. WHITTLE, K. & D. C. FEerGusON. “1988” [1991]. Asiatic Rice Borer, Chilo suppressalis (Walker). Pests not known to occur in the United States or of limited distribution, No. 97. APHIS- PPQ. 10 pp. FERGUSON, D.C. 1991. Review of the genus Epimorius Zeller and first report of the occurrence of E. testaceellus Ragonot in the United States (Pyralidae: Galleriinae). J. Lepid. Soc. 45:117-123. . 1991. The identity of Arctia obliterata Stretch (Lepi- doptera: Arctiidae). Proc. Entomol. Soc. Washington 93:828— 833. . 1991. First record of the genus Acrapex from the New World, with description of a new species from the Carolinas and Virginia (Noctuidae: Amphipyrinae). J. Lepid. Soc. 45:209-214. FEeRGusON, D. C., D. J. HitpurN & B. WRIGHT. 1991. The Lepi- doptera of Bermuda, their food plants, biogeography, and means of dispersal. Mem. Entomol. Soc. Canada 158; 1-106, 1 col. pl., map, 204 figs. FERGUSON, D. C. 1991. An essay on the long-range dispersal and biogeography of Lepidoptera, with special reference to the VOLUME 57, NUMBER 4 Lepidoptera of Bermuda. Pp. 67-79, In FERGUSON, HILBURN & Wricut, The Lepidoptera of Bermuda .. . etc. Mem. Entomol. Soc. Canada 158. ———. 1991. Adult moths. Pp. 231-244 In Gorham, J.R. (ed.), In- “sect and mite pests in food, an illustrated key. Agriculture Handbook 655, U.S. Dept. Agr., Washington, D.C. 767 pp. in 2 vols. . 1992. Franclemontia interrogans (Walker), a new genus for an old species (Lepidoptera: Noctuidae). J. New York Ento- mol. Soc. 100:257-266. . 1993. A revision of the species of Nematocampa 303 (Geometridae: Ennominae) occurring in the United States and Canada. J. Lepid. Soc. 47 (1):60-77. POWELL, J. A. & D. C. Fercuson. 1994. A new genus of winter moths (Geometridae) from eastern California and western Nevada. J. Lepid. Soc. 48:8-23. FERGUSON, D. C. 1997. Review of the New World Bagisarinae, with description of two new species from the southemm United States (Lepidoptera: Noctuidae). J. Lepid. Soc. 51:344-357. PocuE, M. G. & D. C. FERGUSON. 1998. A revision of the cypress- feeding moths of the genus Cutina Walker (Lepidoptera: Noc- tuidae). Proc. Entomol. Soc. Washington 100:331—-352. BOOK REVIEW Journal of the Lepidopterists’ Society 57(3), 2003, 304-305 AN OBSESSION WITH BUTTERFLIES: OUR LONG LOVE AFFAIR WITH A SINGULAR INSECT, by Sharman Apt Russell. ISBN 0-7382-0699-7. Publication: 2003, Perseus Publishing, Cambridge, Massachusetts US$24. Sharman Apt Russell's An Obsession with Butterflies is an impressionistic guide to some of the treasures available to those who pursue butterflies. The book traces a sort of natural history of butterfly obsessions in 15 satisfyingly brief, informative, and eager—if some- what breathless—essays. The essays, on such topics as metamorphosis, intel- ligence, color, parenting, ecology and the environ- ment, are, on the whole, nicely detailed and enjoyable. I thought the essay on the natural history museum quite successful, though I found confusing the attempt to trace conservation efforts for the El Segundo Blue in Los Angeles. It is possible to read these essays all at once in a few hours, and thus to emerge with a reason- ably satisfying, many-angled view—a Cubist por- traitP—of the butterfly. The essays are also fine as quick reads; in a few minutes with one essay you can learn something about butterfly migration, say, or the composition of a butterfly’ wing. The bibliographical essay at the end is an enjoyable and useful approach to the resources made available to the writer. Russell's approach to the subject has us both implic- itly and explicitly comparing what we know about things in our lives to the sort of things butterflies do. The writing is gently anthropomorphic, as some of the chapter headings attest: You Need a Friend, Love Sto- ries, The Single Mom. These butterflies think, remind themselves of things, wait for “love” or “destiny.” There is enough assuring distance and savvy, however, to keep the author from ascribing more than figurative associa- tions between our species. After all, for some of us but- terflies are not just bugs but metaphors on the wing. This guide to some of the many ways in which our lives intersect with those of butterflies could have ben- efited from a clearer focus. Russell flits about from one subject to another, to the point of giving each indi- vidual essay a somewhat different tone from the oth- ers. Much of the book introduces individual scientists and collectors, past and present, who have themselves been obsessed with butterflies in one way or another. Other parts portray various different butterflies and caterpillars, their behavior throughout the life span in different environments, and the cultural surround by which butterflies are assimilated by those of us who are not obsessed. Obsessions are, like butterflies, fascinating things. They are resolutely individualistic, and should prove excellent guideposts both to our human nature and to whatever it is—butterflies in this case—we might be- come obsessive about. Whatever the obsessions of the collectors and scientists briefly sketched in this book, however, they must be assumed, because they have not been demonstrated or evoked. The characters in- troduced in these essays, even the author, never seem to emerge from the pages. This means that their indi- vidual curiosities and interests merge with everyone else’s, and are rarely satisfyingly portrayed. The relationship between the natural world and our human obsessions seems, at times, obscure. She writes, for example, I like the numbers, the big numbers. More is better. More but- terflies are better than fewer butterflies. A river of butterflies is a wonderful thing. Millions of butterflies are the jackpot. I like the largesse, the almost casual gesture, as if a generous earth were whispering into my ear, “See how I replenish myself, see how I birth and birth and birth and darken the skies and fill the waters and cover the ground and still I have more to give.” I find a little of this sort of thing goes a long way. It is true, I suppose, that most who read this book will do so for what it has to say about butterflies, not about our selves in nature, or as obsessive beings, or because the Greek word for this creature is our word for “soul.” But I had wished for more. I confess that the book taught me a great deal about butterflies I had not known; I further confess, how- ever, that I knew very little about them before I read it. I teach in the humanities; my most compelling in- terest in butterflies has been my interest in the work of Vladimir Nabokoy, a writer known to all lepidopterists and a subject fit for any consideration of obsessions with butterflies. As an artist and as a scientist, Nabokov was keen on design; he enjoyed exploring and toying with the intricate, devious, tricky plays of meanings which patterns reveal and conceal. It is little wonder he followed butterflies, or that Russell quotes him in her book. Nabokov is worth quoting because he con- veyed his personal obsessions to the written page with enormous finesse and skill as a writer, in both scientific and fictional contexts. Russell is wise to borrow his ob- servations—and perhaps to avoid his more mystical musings on mimicry. The essays are composed in a dramatic style, which, to my taste, relies too much on the sort of flourish cre- ated by paragraphs of a single sentence. The impres- sionistic details in many of the essays can charm up close, but often fail to contribute to a satisfying sense of the whole. The opening character sketch, of 17th cen- tury collector Eleanor Glanville, is tantalizingly indis- tinct and ill-focused. Here is someone with a genuine VOLUME 57, NUMBER 4 obsession for butterflies, and a tragic story to go with it, yet her portrait here is never achieved, and her role in the book, as heroine, guide, or symbol, is never made clear. I had concerns with the illustrations. I suppose no matter how well illustrated any book on butterflies is likely to be, we will always wish for more. The black and white illustrations (uncredited) seem to be very well drawn, but are small, sparse, and appear poorly printed. 305 Although it can be enjoyed by any reader, I would feel most comfortable recommending this book to curious youngsters who have already found something in na- ture—it wouldn't have to be butterflies—and who would like to know, and find, more. The book itself is not likely to instill that curiosity—that’s up to the butterflies. KENNETH Hope, Columbia College of Chicago, 600 S. Michigan Ave., Chicago, Illinois 60605, USA ERRATA Journal of the Lepidopterists’ Society 57(4), 2003, 305 THE HISTORY AND TRUE IDENTITY OF MELITAEA ISMERIA (NYMPHALIDAE): A REMARKABLE TALE OF DUPLICATION, MISINTERPRETATION, AND PRESUMPTION In the above article by John V. Calhoun (Journal of the Lepidopterists’ Society 57(3):204-219), page 208, first paragraph, line 17 should read: “Volume 16 contains 130 paintings and is dated 1804 (V. Veness pers. com.)” Journal of the Lepidopterists’ Society 57(4), 2003, 306-308 Abaera, 168 Abbot, John, 204 abundance, 295 Accintapubes, 121 acoustics, 1 activity patterns, 161 adaptations, 274 Africa, 1, 235 Agraulis vanillae, 279 Aiello, A., 168 Alberta, 249 alien species, 270 Amarillo S., A., 54 Amauris albimaculata, 235 Anaea troglodyta floridalis, 243 Andes, 62 androconia, 279 angular appendices, 147 Anoplolepis, 1 ant-association, | Anweiler, G. C., 249 aphytophagy, 1 Apiaceae 36, 153 aposematism, 168 Arceuthobium, 47 Arsenura armida, 220 Arsenurinae, 220 Aster, 144 Asteraceae, 153, 197 Asterope markii, 68 Austin, G. T., 75, 76, 176 avocado, 121, 253 avocado-feeding, 121 Bacillus thuringiensis, 92 Baird, Thomas, 249 Bassus, 113 Baughman, J. F., 176 behavior, 43, 161, 279 Bendicho-Lopez, A., 291 Berenbaum, M. R., 36 biocontrol, 270 biodiversity, 253 biogeography, 86, 274 biological control, 113 biology, 197 birdwing butterflies, 17 bivoltine, 291 black swallowtail, 149 Blas, X.P.1., 161 Bolivia, 54 Brazil, 100, 291 Brenthia, 168 British Columbia, 150 Brown, J.W., 113, 253 Bt, 92 buckmoth, 137 burn, 137 butterfly mimicry, 235 Byron-Cooper, O., 284 INDEX FOR VOLUME 57 (new names in boldface) Calhoun, J. V., 204, 308 Callaghan, C. J., 193 camouflage, 168 Canada, 249 Caribbean, 121 Casagrande, M. M., 239 Catalunia, 161 caterpillars, 239, 291 Celastrina, 150 cerrado, 291 Chacin, M. E., 86 checkerspot butterflies, 176 chemical communication, 220 Chionodes, 47 Chlamydastis platyspora, 291 Chlosyne gorgone, 204 Chlosyne nycteis, 204 Choreutidae, 168 Chrysothamnus, 144 Citrus, 43 Claassens, A.J. M., 1 Clavijo, J. A., 86 cocoons, 168 Colias alexandra, 274 Colias occidentalis sullivani, 275 Collins, M. M., 159 Colombia, 54, 62 common gem butterfly, 295 common palm butterfly, 147 conifers, 47 Conlan, C. A., 54 Copaxa lunula, 54, 59 Copaxa orientalis, 54 Copaxa semioculata, 54 corn, 92 Costa Rica, 113, 121, 253 Costa, J. T., 220 Covell Jr., C.V., 230 Crambidae, 270 Crematogaster, | Croton, 243 Cucullia asteroides, 144 Cucullia montanae, 144 Cucullia similaris, 144 Cuculliinae, 144 Cyaniris ladon var. quesnellii, 150 Cymothoe herminia, 235 Danaus plexippus, 92, 284 De Marmels, J., 86 defense mechanisms, 168 Depressaria, 36 Desch-O’ Donnell, M., 284 detritus, 1 DeVries, P. J., 77, 157, 235 diapause, 25 Dichrorampha acuminatana, 152: Dichrorampha petiverella, 152 Dichrorampha vancouverana, 152 Diniz, I. R., 291 distribution, 176, 253, 295 DNA, 17 Donovan, J., 25 Duarte, M., 239 dwarf mistletoe, 47 Eantis thraso, 43 early stages, 54, 68 eastern seaboard, 270 ecology, 295 Ecuador, 43, 54, 62, 68, 193 Edith’s copper 249 Elachistidae, 36, 291 elevation, 253 Elymnias hypermnestra undularis, 147 energetics, 284 entomophagy, 1 enzyme activity, 284 Epiblema gibsoni, 230, 233 Epiblema infelix, 230 Epiblema walsinghami, 230 Epipaschiinae, 121 escape holes, 168 Euchlaena, 107 Eucosmini, 230 Euliini, 113 Euphydryas, 176 Euphydryas anicia, 176 Euphydryas colon, 176 Euptychia peculiaris, 100 Euthalia, 47 evolution, 1, 235 fecal pellets, 168 fecal stalactites, 168 feeding specialist, 291 Ferguson, D. C., 270, 299 fiery skipper, 239 Filatima natalis, 47 fire, 137 fire-adaptation, 137 Fitzgerald, T. D., 161 Fleishman, E., 176 Florida, 243 food plant record, 149 food plant, 47, 274 foodplants, 274 foraging, 161, 220 Formicidae, | Freitas, A.V.L., 100 Galechiidae, 47 genetically modified plants, 92 genital plate, 147 genitalia, 147, 176 geographical distribution, 295 Geometridae, 107 Georgia, 204 Glaser, J. D., 270 Goldstein, P. Z., 153 VOLUME 57, NUMBER 4 Goztek, D. A., 220 grasslands, 137 grass-specializing feeders, 239 Great Basin, 176 Greeney, H. F., 43 gregarious caterpillars, 161, 220 Grindelia, 144 group foraging, 220 gulf fritillary, 279 Guppy, C. S., 150 habitat, 295 habits, 295 Hammmond, P. C., 274 Harp, C. E., 197 Hay, J. D. V,, 291 Heath, A., 1 Heliconiinae, 279 Heliothinae, 197 Hemileuca eglanterina, 137 Hennessey, M. K., 243 Hereau, H., 71 Hesperiidae, 43, 239 Hesperiinae, 239 heterospecific mating, 71 Hill, R. 1., 68 Himalayas, 295 Hodges, R. W., 299 holarctic, 152 Hope, K. 252 Hossler, E. W., 107 Hossler, F. E., 107 host plant associations, 153 host plants, 25, 36, 43, 47, 68, 153, 168, 176, 197, 291 Hutchinson, W. D., 92 hybridization, 25, 71, 176 Hylephila phyleus, 239 identification, 253 immature stages, 54, 100 immigrant, 152 India, 147 Inga, 193 insect flight muscle, 284 interspecific copulation, 71 Janzen, D. H., 220 Kawahara, A. Y., 81 Koch, R. L., 92 Kondla, N. G., 150 Kondo, K., 17 Landry, B., 251 larvae, 43, 54, 68, 92, 107, 121, 144, 153, 193, 204, 239, 291 larval behavior, 43, 161, 168, 220 larval endophagy, 113 larval food plants, 43, 68, 149, 295 larval mortality, 92 larval shelters, 43 Lathyrus rigidus, 274 Launer, A. E., 176 Lauraceae, 121, 253 leaf phenology, 291 lectotype, 150 Lemaire, C., 54 Levine, E., 284 Libythea collenettei, 81 Libytheinae, 81 life cycles, 43 life history, 1, 43, 68, 100, 107, 113, 144, 193 Lomatium, 36 lucia, 150 lunula, 54 Lycaena editha, 249 Lycaenidae, 1, 47, 150, 249, 295 Lytrosis permagnaria, 107 Lytrosis sinuosa, 107 Lytrosis unitaria, 107 mandible, 239 Marquesas Islands, 81 maternal investment, 137 mating behavior, 279 Matsuka, H., 17 McCorkle, D. V., 274 Mckenna, D. D., 36 Melanis leucophlegma, 193 Melitaea ismeria, 204 Metzinger, C., 284 Michigan, 25 Mielke, O. H. H., 239 milkweed, 92 Miller, J. Y., 62 Miller, L. D., 62 Miller, W, E., 152 mimicry, 107, 235 mimics, 235 mistletoe, 47 mitochondria, 284 mitochondrial enzyme activity, 284 mitochondrial gene, 17 Mitoura, 47 models, 235 molecular systematics, 17 monarch butterfly, 92, 284 Monoloxis flavicinctalis, 168 Mooney, K. A., 47 morphological analysis, 25 morphology, 81, 239, 253, 279 Murphy, D. D., 176 myrmecophily, 1 natural history, 81, 243 natural hybrids, 25 ND5 gene, 17 Neotropical, 54, 100, 113, 193, 200, 253 Nevada, 176 new genus, 100 new species, 62, 63, 113, 197, 201, 230, 253 new subspecies, 86, 90, 275 nigrescens, 150 Nishida, K., 113 307 Noctuidae, 144, 153, 197 North America, 152 Nymphalidae, 47, 62, 68, 81, 92, 100, 147, 176, 204, 235, 243, 279 obituary, 299 Ocotea veraguensis, 121 Olethrentinae, 152 Olethreutinae, 230 Oregon, 137 Orthocomotis, 253 Orthocomotis altivolans, 265 Orthocomotis longicilia, 258 Orthocomotis similis, 263 overwintering, 161, 284 oviposition behavior, 193 Oxypolis rigidior, 149 Palaearctic, 152 palatability spectrum, 235 Panama, 168 Pantepui, 86 Papaipema, 153 Papilio canadensis, 25 Papilio glaucus, 25 Papilio glaucus, 71 Papilio polyxenes, 71 Papilio polyxenes, 149 Papilionidae, 17, 71, 149 parasitism, 243 parasitoid, 113 Paullinia, 68 Pedaliodes chrysotaenia form fassli, 62 Pedaliodes fassli, 62 Pedaliodes gustavi, 62, 63 Pedaliodes negreti, 62 Pedaliodes pheretias form griseola, 62 Peigler, R. S., 157 Pereute lindemanae pemona, 86 Pereute lindemanae piaroa, 86 Persea americana, 121 Persea, 54 Peru, 54, 193 pests species, 113 Petit, J. C., 144 Petit, M. C., 144 phenology, 176 pheromones, 279 Phragmites, 270 phylogenetic analysis, 121 phylogenetic tree, 17 Pieridae, 86 pine processionary caterpillar, 161 Pinus, 161 Poaceae, 100 Pogue, M. G., 197 pollen, 92 polyphagy, 153 Poritia hewitsoni, 295 prairie, 137, 230 prescribed burn, 137 pre-zygotic reproductive isolating mech- anisms, 71 308 processionary behavior, 161 Promylea lunigerella glendella, 47 Pronophilini, 62 Proteaceae, 291 Pseudacraea lucretia, 235 Pseudodebis, 100 Pseudopieris viridula mimaripa, 86, 90 pupae, 43, 54, 68, 107, 204 Pyralidae, 47, 121, 168 Pyraustinae, 270 Pyrginae, 43 Quinter, E. L., 153 range, 243 Rauser, C. L., 279 Riodinidae, 193 risk assessment, 92 Roberts, M. A., 152 Roupala montana, 291 Rubbus spp., 113 Rutaceae, 43 Rutowski, R. L., 279 Salvato, M. H., 243 Sapindaceae, 68 Saturniidae, 54, 137, 220 Satyrinae, 62, 100, 147 scanning electron microscopy, 239, 279 Schinia regia species complex, 197 Schinia regina, 197, 201 Schmidt, B. C., 249 Scriber, J. M., 25, 71 seaboard, 270 seasonal forms, 243 seasonality, 295 SEM, 279 Seticosta rubicola, 113, 114 Severns, P.M., 137 sex-ratio, 176 sexual selction, 71 Sharma, N. 147 shelters, 43 Shinkawa, T., 17 signa, 147 Singh, A. P,, 295 snout butterfly, 81 social caterpillars, 161, 220 Solis, M. A., 121, 168 sound-producing organ, 86 South Africa, 1 South America, 121 South Carolina, 204 Spain, 161 spatial distribution, 253 Styer, L., 121 subspecies, 86 succinate dehydrogenase, 284 swallowtail butterflies, 71 systematics, 17, 54, 81, 100, 113, 253 taxonomy, 81, 86, 113, 197, 204 Taydebis, 100 Taygetis, 100 Date of Issue (Vol. 57, No. 4): 9 December 2003 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Thaumetopoea pityocampa, 161 Thaumetopoeidae, 161 thermal regulation, 161 tiger swallowtail, 25 toothed brachia, 147 Tortricidae, 113, 152, 253 trail following, 161, 220 trail pheromone, 220 transgenic corn, 92 trophallaxis, 1 tropical dry forest, 220 twig mimicry, 107 type locality, 150 variation, 274 Venette, R. C., 92 Venezuela, 54, 86 Viloria, A. L., 62 Wagner, D. L., 107, 270 Wallacea, 17 Warren, A. D., 43 Weibull model, 92 West Indies, 243 Western United States, 36 wetlands, 270 Williams, A.H., 149 wing toughness spectrum, 235 wings, 235, 279 Wisconsin, 149 Wolfe, K. L., 54 Wright, D. J., 230 EDITORIAL STAFF OF THE JOURNAL Carta M. Penz, Editor Department of Invertebrate Zoology Milwaukee Public Museum Milwaukee, Wisconsin 53233 USA flea@mpm.edu Put DeVries, Book Review Editor Center for Biodiversity Studies Milwaukee Public Museum Milwaukee, Wisconsin 53233 USA pjd@mpm.edu Associate Editors: Gerarpo Lamas (Peru), Kenetm W. Puitie (USA), Ropert K. Rossins (USA), Feuix Srertinc (Canada), Davin L. Wacner (USA), Curister WikLuNb (Sweden) NOTICE TO CONTRIBUTORS i: eaieivutions to the Journal may deal with any aspect of Lepidoptera study. Categories are Articles, Profiles, General Notes, Techni- ~ eal Comments, Book Reviews, Obituaries, Feature Photographs, and Cover Illustrations. Obituaries must be authorized by the president _ of the society. Requirements for Feature Photographs and Cover Illustrations are stated in Volume 44(2):111 and on the Society’s web site at http://www.lepsoc.org/. 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Au- thors of Book Reviews and Obituaries are exempt from page charges. Correspondence: Address all matters relating to the Journal to the editor. Address book reviews directly to the book review editor. PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A. CONTENTS AN ILLUSTRATED GUIDE TO THE OrrHocomoris Docnin (TorrriciDAE) OF Costa Rica, WITH SUM- MARIES OF THEIR SPATIAL AND TEMPORAL DISTRIBUTION John Wise = D5 (3 SCLEROCONA ACUTELLA (E\VERSMANN) (CRAMBIDAE: PyrRAUSTINAE), NATURALIZED ALONG THE EASTERN SEABOARD David L. Wagner, Doug C. Ferguson and John D. Glaser ----------------------- 270 A NEW DESERT SUBSPECIES OF COLIAS OCCIDENTALIS (PIERIDAE) FROM SOUTHEASTERN OrEGoN Paul C Hammondiand David\V. McCorkle === ee 274. MALE-SPECIFIC STRUCTURES ON THE WINGS OF THE GULF F'RITILLARY BUTTERFLY, AGRAULIS VANILLAE (NYMPHALIDAE) Casandra L. Rauser and Ronald L. Rutowski --------------------==== 279 INTERINDIVIDUAL VARIATION IN MITOCHONDRIAL ENZYME ACTIVITY IN MALE MONARCH BUTTERFLIES, Danaus pLexiepus L. (NympHaLipaE) Elena Levine, Olivia Byron-Cooper, Megan Desch- O’Donnell and Carrie Metzinger--------------------------------------------=--=-----==-------=--- 284 ABUNDANCE OF CHLAMYDASTIS PLATYSPORA (E,LACHISTIDAE) ON ITS HOST PLANT ROUPALA MONTANA (PRO- TEACEAE) IN RELATION TO LEAF PHENOLOGY Aurora Bendicho-Lopez, Ivone Rezende Diniz and John Du Vall Hay--------------------------------------------------------------------=--=--=--==- 291 GENERAL NOTES NEW RECORDS ON THE DISTRIBUTION AND ECOLOGY OF COMMON GEM BUTTERELY, PoRITIA HEWITSONI HEWITSONI MOORE FROM THE LOWER WESTERN HIMALayas: A LESSER KNOWN TAXA Arun P. Singh-----------------------------------------------=--------9 2-2-9222 2a 2a nanan nnn 295 OBITUARY ALEXANDER Doucias CAMPBELL FERcuson (1926-2002) Ron Hodges ------------------------ 299 Book REvIEw AN OBSESSION WITH BUTTERFLIES: OUR LONG LOVE AFFAIR WITH A SINGULAR INSECT Kenneth Hope 304 FQRRATA -===-==== === === a 305 INDEX FOR VOLUME 57 --------------------------9-- 2-2 2222222 nnn nnn nnn nnn nn nnn nna 306 @) This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanance of Paper). +; Hh ane'y ha ee an CF, 4 Il | ARIES | 1 BR, | l il | {| 3 9088 01073 8 SMITHSONIAN INSTITUTION LI ! et. teens Fae MI Let R38 Asad dy Pi pornrser aR gases wiete SPL Bay Mates thy resewy