"Dy 28 Fs: ra Tati ale Hae it SN Fan Rao “te tintin te hy REAL LEIS bacoraat i . _—— asia netcenion Oe TL Le ee ee SRDS MERA PP REEL SEP ise CE te ae ee ae Dt Le a ate ee ee "ane a i en eT + behy ¢ < sy ‘) > » Sgt oa pe ot * Me Oe hao oe IORI apne At Wid Red splat alpen ame ae DS, Ei Sy eb ie er ad bit Eh aca aoa wie ay A RAE aN re Clarke ex L Volume 32 1978 Number 1 JOURNAL of the LEPIDOPTERISTS’ SOCIETY Published quarterly by THE LEPIDOPTERISTS’ SOCIETY Publié par LA SOCIETE DES LEPIDOPTERISTES Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 7 April 1978 THE LEPIDOPTERISTS’ SOCIETY EXECUTIVE COUNCIL J. W. Trpen, President KENELM W. Putuip, Vice President I. F. B. Common, Ist Vice President JuLian P. DoNAHuE, Secretary Lionet Hiccrns, Vice President RONALD LEUSCHNER, Treasurer Members at large: F. S. CHEW R. A. ARNOLD J. F. EMMEL D. F. Harpwick E. D. CAsHATT R. R. GATRELLE J. B. ZrEcLER R. E. STANFORD A, P.\ Prats The object of the Lepidopterists’ Society, which was formed in May, 1947 and formally constituted in December, 1950, is “to promote the science of lepidopterology in all its branches, . . . . to issue a periodical and other publications on Lepidoptera, to facilitate the exchange of specimens and ideas by both the professional worker and the amateur in the field; to secure cooperation in all measures” directed towards these aims. Membership in the Society is open to all persons interested in the study of Lepidoptera. All members receive the Journal and the News of the Lepidopterists’ Society. Institutions may subscribe to the Journal but may not become members. Prospective members should send to the Treasurer full dues for the current year, together with their full name, address, and special lepidopterological interests. In alternate years a list of members of the Society is issued, with addresses and special interests. There are four numbers in each volume of the Journal, scheduled for February, May, August and November, and six numbers of the News each year. Active members—annual dues $13.00 Student members—annual dues $10.00 Sustaining members—annual dues $20.00 Lite members—single sum $250.00 Institutional subscriptions—annual $18.00 Send remittances, payable to The Lepidopterists’ Society, and address changes to: Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 U.S.A. Memoirs of the Lepidopterists’ Society, No. 1 (Feb. 1964) A SYNONYMIC LIST OF THE NEARCTIC RHOPALOCERA by Cyrit F. pos Passos Price: Society members, $5.00 U.S.; non-members, $7.50 U.S. Paper covers, revisions of the Melitaeinae and Lycaenidae supplied separately. Order: Mail to Charles V. Covell, Jr., Memoirs Editor, Department of Biology, Uni- versity of Louisville, Louisville, KY 40208, U.S.A. The Lepidopterists’ Society is a non-profit, scientific organization. The known office of publication is 1041 New Hampshire St., Lawrence, Kansas 66044. Second class postage paid at Lawrence, Kansas, U.S.A. 66044. Cover illustration: Dasychira dorsipennata larva, dorsal and lateral views. From Fascicle 22.2, “Lymantriidae,” by Douglas C. Ferguson, in Moths of America North of Mexico. The drawing was done by E. R. Hodges, Scientific Illustrator, Department of Entomology, Smithsonian Institution. (Reproduced by permission of the author.) JOURNAL OF Tue LEpPIDOPTERISTS’ SOCIETY Volume 32 1978 Number 1 Journal of the Lepidopterists’ Society 32(1), 1978, 1-2 ieee OF PARNASSIUS CLODIUS GALLATINUS (PAPILIONIDAE ) STEVE KOHLER Montana Department of Natural Resources and Conservation, Division of Forestry, 2705 Spurgin Road, Missoula, Montana 59801 ABSTRACT. Types of Parnassius clodius gallatinus Stichel, 1907, were discov- ered in the remnants of the Elrod collection, at the University of Montana, Missoula. Proper labels have been attached to the specimens, and they have been placed in the collection of the American Museum of Natural History. The description of Parnassius clodius gallatinus Stichel was based on a pair illustrated by Elrod (1906). The actual specimens were never seen by Stichel (1907). While preparing a paper clarifying the nomenclature of Parnassius clodius Ménétriés subspecies found in the Rocky Mountains (Ferris, 1976), Cliff Ferris contacted me in an attempt to locate the types of gallatinus. A search through the remnants of the Elrod Collection, housed at the University of Montana, Missoula, led to the discovery of the two specimens matching El!lrod’s 1906 illustrations. They were figured on page 16 of “The Butterflies of Montana.” At the suggestion of Ferris, the two specimens have been placed in the collection of the American Museum of Natural History. The follow- ing labels are affixed to the specimen pins: Holotype Male: A white label partially machine printed in black ink and partially hand lettered in red ink which reads: Gallatin Co. Mont./ Eley. 6800/ Col. E. Koch/ 6-27 1900, and a red label hand lettered in black ink which reads: Holotype ¢/ Parnassius clodius/ gallatinus Stichel/ ex. Elrod Coll. Univ./ Mont. S. Kohler 1976. Allotype Female: A white label partially machine printed in black ink and partially hand lettered in red ink which reads: Gallatin Co. Mont./ Elev. 6800/ Col. Cooley/ 6-27 1900, and a red label hand lettered in bo JOURNAL OF THE LEPIDOPTERISTS SOCIETY te Fig. 1. Parnassius clodius gallatinus Stichel: a) holotype male, dorsal; b) same, ventral; c) allotype female, dorsal; d) same, ventral. Photos approximately 7 natural size. black ink which reads: Allotype ?/ Parnassius clodius/ gallatinus Stichel/ ex. Elrod Coll. Univ./ Mont. S. Kohler 1976. The specimens are in good condition except for some minor dermestid damage to the abdomen of the female. An additional pair collected by Cooley at the same locality and on the same date was located in the collection of Montana State University, Bozeman, for Ferris by Dr. Nor- man L. Anderson. These are not designated as paratypes. Because of the scarcity of copies of Elrod’s “The Butterflies of Mon- tana, the type specimens have been illustrated in Figure 1. ACKNOWLEDGMENTS I would like to thank Dr. James H. Lowe and the University of Mon- tana for allowing the type specimens to be placed at the American Museum of Natural History, and also Cliff Ferris for comments and assistance, LITERATURE CITED IeLnop, M. J. 1906. The butterflies of Montana. Univ. Montana Bull. 10: 1-174. Fennis, C. D. 1976. A note on the subspecies of Parnassius clodius Ménétriés found in the Rocky Mountains of the United States (Papilionidae). J. Res. Lep. 15: 65-74. SticHEL, H. 1907. Lepidoptera Rhopalocera Fam. Papilionidae Subfam. Parnas- siinae. Wytsman Gen. Ins. Fasc. 58: 1-60. Journal of the Lepidopterists’ Society 32(1), 1978, 3-19 SPECIFICITY, GEOGRAPHIC DISTRIBUTIONS, AND FOODPLANT DIVERSITY IN FOUR CALLOPHRYS (MITOURA) (LYCAENIDAE) Kurt JOHNSON Department of Biology, City University of New York, City College, Convent Avenue and 138th Street, New York, New York 10031 ABSTRACT. The species C. siva, gryneus, hesseli, and turkingtoni are examined. Genitalic evidence of their non-conspecificity is provided along with discussion of particular localities of sympatry. Detailed distributional data are illustrated and a documented table of foodplant diversity included. C. siva and gryneus are oligoph- agous On numerous species of Juniperus (Cupressaceae) which replace each other geographically across the United States. C. hesseli is monophagous on Chamaecyparis thyoides (Cupressaceae); the foodplant of turkingtoni is unknown. Evidence indicates that all local populations are specific to one foodplant species. Callophrys ( Witoura) nelsoni ( Boisduval), C. siva (Edwards), C. loki (Skinner), C. gryneus (Hiibner), and C. hesseli (Rawson & Ziegler), aside from taxonomic descriptions, have been subject to several biological and regional studies, but published works (Anderson, 1974; Johnson, 1972; Pease, 1963; Rawson et al., 1951; Remington & Pease, 1955) are very heterogeneous in content and comprehensiveness. During the last four years I have been compiling data on their dis- tributions and larval foodplants as a base for taxonomic studies of the group. I have also been studying the genitalia of all Nearctic and Neotropical Callophrys (Mitoura) in detail (Johnson, 1976a). The pur- pose of this paper is to present detailed distributional data for three of these species (C. siva, C. gryneus, and C. hesseli), demonstrate that C. siva and C. gryneus are not conspecific, and summarize data on larval foodplants, a number of which are new to the literature. What bio- geographical data are known on the newly named C. turkingtoni Johnson (Johnson, 1976b) will also be presented. The specificity of C. siva and C. nelsoni involves several complex problems in the northwestern United States and will be treated in a separate paper (Johnson, 1977). METHODS AND MATERIALS Using the collection of the American Museum of Natural History as a basis, additional information on localities and possible local foodplants was gathered by correspondence and recorded county by county. Speci- mens or photographs were solicited in cases of peripheral or isolated populations, and available published records were included. The re- search aimed at definitive treatment on the species level only. Genitalic + JOURNAL OF THE LEPIDOPTERISTS SOCIETY studies of males and females were performed in areas where C. siva and C. gryneus were reportedly sympatric. These genitalia were compared with those from many parts of the ranges of C. siva and C. gryneus, as well as with dissections of other congeners. The number of these speci- men dissections included: C. siva, 78; C. gryneus, 46; C. hesseli, 14; C. turkingtoni, 1; C. nelsoni, 83; C. rosneri, 46 (Johnson, 1976a); C. barryji, 19 (Johnson, 1976a); C. byrnei, 9 (Johnson, 1976a), and C. loki, 15. Geographic ranges were studied to discover areas of insect distribution not coinciding with present published foodplant knowledge, and efforts were then made to make the list for each species complete by identifica- tion of exact plants with which the adults were associated by perching behavior (Johnson & Borgo, 1976) or on which oviposition or larvae were observed. Full documentation of each of these methods is given in the foodplant table (Table 1) since a degree of fallibility has been demon- strated in each (Brower, 1958; Downey & Dunn, 1965). An ongoing effort to compile foodplant specimens at one institution was initiated, and plants collected thus far are cited in the table. Since the perching behavior of these insects limits general flight patterns to the vicinity of the foodplant, and since data not only in this study but another (John- sonn, in prep. ) indicate that C. siva and C. gryneus are exclusive Juniperus- feeders, some useful evidence on larval foodplants in areas where only one juniper species was regionally present could be culled from identifi- cation of the plants at the locality indicated on the specimen labels. The list of plants established as the only Juniperus species present in a region (R) or at a locality (L), source of butterfly data (B), source of plant data (P) is: C. siva: Juniperus deppeana Steud., (LL) 10 mi. NW Pine Springs, Culberson Co., Texas, (B) R. O. Kendall, (P) Herbarium, University of Texas, Austin; J. deppeana, (1.) 5 mi. W of McDonald Observatory, Jeff Davis Co., Texas (Bk. 'O: Kendall, (P) Herbarium, University of Texas, Austin. Juniperus occidentalis oc- cidentalis Hook. x J. osteosperma Torr. (Little), (R) Washoe Co., Nevada (Reno and vicinity westward), Ormsby and Douglas cos., (B) P. Herlan, (P) Vasek, 1966. Juniperus monosperma (Engelm.) Sarg., (LL) Sycamore Canyon, NW of Nogales, Santa Cruz Co., Arizona, (B) Share and Clark (American Museum of Natural History (AMNH)), (P) Herbarium, Arizona State University, Tempe. Juniperus pinchotii Sudw., (R) Reeves Co., Texas, (B) D. Stallings and M. R. Turner, (P) Adams, 1972, R. P. Adams, pers. comm. C. gryneus: Juniperus virginiana L., (R) Cass Co., Texas, (B) R. O. Kendall, (P) Adams & Turner, 1970, R. P. Adams, pers. comm. Juniperus ashei Buchholz, (R) McLennan Co., Texas, (B) R. O. Kendall, (P) Adams, 1972; Adams & Turner, 1970; R. P. Adams, pers. comm. Juniperus pinchotii Sudw., (R) Pecos Co., Texas, (B) R. O. Kendall, (P) Adams, 1972, R. P. Adams, pers. comm. Juniperus deppeana Steud., (L) Huejotitlan, Chihuahua, Mexico, (B) AMNH, (P) Little, 1971. J. deppeana, (i) Baboquivari Mountains, S of Baboquivari Peak, Pima iGo Arizona,” (ByenieaD: Gunder (AMNH), (P) Herbarium, Arizona State University, Tempe. Juniperus virginiana L. * J. horizontalis Moe nch., (1) Lynxville, along Mississippi River, VoLUME 32, NUMBER 1 Aste I. Larval foodplants established by the identification of exact plants. Foodplant taxa and specimens Butterfly taxa and specimens Callophrys (Mitoura) siva Juniperus scopulorum Sarg." Plants, PL: Van Haverbeke, N-2; hybrid index at site—63 + 8% J. scopulorum; SL: Johnson, 1972.” Plant specimens, K. Johnson #2 (Smiley Canyon = VanH. N-2; #1 (Chadron), Royal Ontario Museum, Toronto (ROM ).° Juniperus scopulorum var. columnaris Fasset Plants, PL: Van Haverbeke ND-S, I: T. McCabe, (H) North Dakota State Univ., Fargo; SL: Van Haverbeke, 1968. Plant specimens, Van Haverbeke ND-8, Univ. of Nebraska (UN). Juniperus scopulorum Sarg. X J. virgini- ana L. Plants, PL: Van Haverbeke N-7; hybrid index at site—48 + 6% J. scopulorum; Range of use of hybrids (Johnson, 1972 )—70 + 4% to 36 + 4% J. scopulorum. Plant specimens, Van Haverbeke N-7, UN; K. Johnson #3 (Sizer, Keith Co. ) ROM. Juniperus virginiana L. Plants, 17 mi S of PL: Van Haverbeke N-4; hybrid index at site—36 + 4% J. scopulorum; SL: Johnson, 1972. Plant specimens, Van Haverbeke N-4, UN; K. Johnson #4 (locality as above ) ROM. Prostrate morph of J. scopulorum Sarg. x J. virginiana L. x J. horizontalis Moench. s. siva; Dawes Co., Nebraska (Smiley Canyon ), W of Fort Robinson; Catholic Cemetery, Chadron.* Butterthes, PE (ACS@: B)*: K. Johnson (AMNH).° s. siva; Slope Co., North Dakota (Amidon, along burning coal vein). Butterflies, PC: T. McCabe. s. siva; Garden Co., Nebraska (bluffs above N. Platte River, nr. Lewellen). Butterflies, PC (AC): L. Running, AMNH. s. siva; Rock Co., Nebraska (Long Pine Rec. Area). Butterflies, PC (AC, LC, B): K. Johnson, L. Running, AMNH. s. siva; Saskatchewan, Canada (Val Marie, near Rosefield along Frenchman River ). pm 1 Taxon of foodplant used (according to nomenclature of USDA (1953) and Little (1971)). 2Source of plant data: PL — butterflies were specifically collected at a_ particular locality studied by Cupressaceae taxonomists; their designation of the site is noted along with the date of their study. “Hybrid index” refers to these studies’ calculation of the degree of hybridity in plants at these areas. Plant identifications are noted as “7”: I, exact substrate plant identified pen Ee —Rerbarum at ==»), and! location of voucher specimens; I,, plant identified by data sent to ______ by ; I,, foodplant established in original description of butterfly, citation given; I,, plant identified from specimens sent to ——_——_; I,, substrate plant established by matching herbarium specimens with butterfly data and establishing that no other species co- occurs, herbarium cited. An “*” following this category (‘‘Plants’”’) means this foodplant usage is well 3 The label number and place of deposition of plant specimens collected in this study An “* means collection in progress at time of this writing. known; SL = other literature which supports this identification. >? 4Taxon of butterfly concerned (as designated in annotated list), with state and exact locality. 5 Source of butterfly data: PC = “personally collected by Letters in parentheses following mean: AC, adults commonly observed perching; AI, adults perching but not commonly observed; LC, larvae collected; LO, larvae observed; B, behavior studied in detail; O, oviposition observed. 6 Location of specimens if not aforementioned person (AMNH =the American Museum of Natural History, New York). M =museum specimens were used as the source of data; ver. means verified by , and method. TL = type locality of the insect. 6 JOURNAL OF THE LEPIDOPTERISTS SOCIETY TAsrE Continued Foodplant taxa and specimens Plants, Schurtz (1971) indicates this area would be included in his tri- parental swarm. Van Haverbeke (pers. comm.) supports this evaluation; I: (H) Univ. Saskatchewan, Regina; I: D. F. Van Haverbeke (data from R. Hooper, K. Johnson); SL: Little, 1971; Van Haverbeke, 1968; Fassett, 1945. Plant specimens, (H) Univ. Sask. (Sask. Prairie Park); K. Johnson #5* ( Hooper ) ROM. Juniperus osteosperma ( Torr.) Little Plants, I;: D. F. Van Haverbeke; SL: Emmel & Emmel, 1973; Johnson, 1977; Little, 1971. Plant specimens, K. Johnson #15 (Running, locality as above) AMNH. Juniperus californica Carr. Plants +, I: D. F. Van Haverbeke; SL: Comstock, 1927; Emmel & Emmel, 1973. Plant specimens, K. Johnson #16 (Leone, locality as above) AMNH. Juniperus occidentalis occidentalis Hook. Plants, I: D. F. Van Haverbeke; SL: Johnson, 1977, Little, 1971. Plant specimens, K. Johnson #17 (Buckingham, locality as above). Juniperus occidentalis australis Vasek Plants, I: John H. Lane; SL: Vasek, 1966. Butterfly taxa and specimens Butterflies, PC (AC): R. Hooper (ver. K. Johnson, photo ). s. ssp.; White Pine Co., Nevada (nr. McGill Junction). Butterflies, PC (AC): L. Running, AMNH. s. juniperaria; Los Angeles Co., Cali- fornia (Mint Canyon). Butterflies, PC (AC): M. Leone, AMNH. s. ssp.; Jefferson Co., Oregon (nr. Warm Springs, W on road to Twin Buttes ). Butterflies, PC (AC): F. Buckingham, AMNH. s. ssp.; Tulare Co., California (vic. Kennedy Meadows ), San Bernardino Co., California (Big Bear Lake). Butterflies, PC (AC): John H. Lane. Callophrys (Mitoura) gryneus Juniperus virginiana L. Plants, I: K. Johnson; SL: Klots, 1951, Little, 1971. Plant specimens, K. Johnson #21 (locality as above). Juniperus silicicola (Small) Bailey Plants, I: F. D. Fee, (H) Univ. Florida, Gainesville; L: (H) Univ. Gainesville; SL: Klots, 1951; Little, 1971. Plant specimens, K. Johnson #10 (St. Augustine locality, Univ. Florida) ROM. Juniperus scopulorum Sarg. * J. virgini- ana LL. Plants, PL: Van Haverbeke M-1; hybrid index near site—27 + 4% J. scopulorum; Range of use of hybrids (Johnson, g. gryneus; Ulster Co., New York ( West Park, Holy Cross Publications ). Butterflies, PC (AI): K. Johnson, AMNH. g. sweadneri; St. Johns Co., Florida (along Ocean Rt. A1A, St. Augustine ). Butterflies, PC (AC): F. D. Fee. g. gryneus; Jackson Co., Missouri (general ). Butterflies, PC (AC): J. R. Heitzman. VoLUME 32, NuMBER 1 TABLE l. Continued Foodplant taxa and specimens Butterfly taxa and specimens 1972 )—38 + 4% J. scopulorum to 27 + 4% J. scopulorum. Plant specimens, K. Johnson #11 (Heitzman, Independence ) ROM. Juniperus ashei Buchholz Plants, I: J. R. Heitzman; SL: Little, 1971. Plant specimens, K. Johnson #12* (Heitzman, ?) ROM. Juniperus pinchotii Sudw. Plants [two examples], L: R. P. Adams (Scott, Roever); SL: Adams, 1972. I: R. O. Kendall; SL: Little, 1971; Adams & Turner, 1970. Juniperus virginiana L. x J. horizontalis Moench. Plants, I: (by reason of Schurtz, 1971) D. F. Van Haverbeke; SL: Schurtz, 1971; Little, 1971. Plant specimens, K. Johnson #13 (locality as above). g. ssp.; Barry Co., Missouri (Eagle Rock ), also McDonald Co.; Washington and Carroll cos., Arkansas. Butterflies, PC (AC): J. R. Heitzman. g. castalis [two examples]; Armstrong Co., Texas (just below N rim of Palo Duro Canyon, 15-16 mi. S Claude). Butterflies, PC (AC): M. Toliver, H. A. Freeman, J. M. Burns, K. Roever (R. O. Kendall); J. Scott. Bexar Co., Texas (Reo Seco Road, off U.S. Hwy. 281 N of San Antonio). Butterflies, PC (AC, O): R. O. Kendall. g. gryneus; Dane Co., Wisconsin (10 mi. W of Madison). Butterflies, PC (AC): W. Sieker. Callophrys ( Mitoura) hesseli Chaemaecyparis thyoides (L.) B.S.P. Plants*, I: S. Hessel, G. W. Rawson, J. B. Ziegler; I,: Rawson et al., 1952; Rawson & Ziegler, 1950 (therein det. by I. M. Johnston, Harvard Univ. ). hesseli; Ocean Co., New Jersey (Lakehurst, TL). Butteniess 2G. ( ACO. LEG) S. Hessel, G. W. Rawson, J. B. Ziegler. Lacrosse Co., Wisconsin and 5 mi. W of Sauk City, Sauk Co., Wisconsin, (B) F. Amold and W. E. Sieker, (P) Ross & Duncan, 1949; Schurtz, 1971; D. F. Van Haverbeke, pers. comm. C. turkingtoni: Juniperus flaccida Schlecht., (R) 10 mi. E of Namiquipa, Chihauhua, Mexico, (B) W. Gertsch and M. Cazier (AMNH), (P) Little, 1971; Herbarium, University of Mexico, Mexico City. RESULTS Genitalia of C. gryneus and C. siva Genitalia of males and females were studied in three regions where these species were reportedly sympatric (Davis Mountain, Texas; Guada- lupe Mountains, New Mexico and Texas; and Baboquivari Mountains, Arizona) and found to be easily separable. However, some traditionally 8 JoURNAL OF THE LEPIDOPTERISTS SOCIETY Figs. 1-9. Female genitalia of selected Nearctic Callophrys (Mitoura) spp. C. siva siva: 1, topotypical; 2, heavily sclerotized areas of lamellae and eighth sternite; 3, showing tufts of “hair” ailowing diagnosis by naked eye. C. gryneus: 4, topotypical. C. hesseli: 5, topotypical. Genital plates of sympatric species near Alpine, Texas: 6, C. gryneus and 7-9, C. siva. used wing-pattern characters for distinguishing these species (ventral secondaries: post basal spots or pattern of mesial band) were shown to be less reliable (also noted in Johnson, 1976a, 1977). The diagnostic genitalic characters are as follows: Females (Figs. 1-9). C. gryneus (Figs. 4, 6): ductus bursa longer and not “club-ended” as on siva; lamellae tapering caudad from antrum, not shouldered as on siva, lamellae postvaginalis nearly as long as broad; juncture of lamellae and eighth abdominal sternite not heavily sclerotized or connected, C. siva (Figs. 1-3, 7-9). Ductus bursa shorter than gryneus and “club-ended”; lamellae distinctly shouldered, lamella postvaginalis much broader than long. Junc- ture of lamellae and eighth abdominal sternite heavily sclerotized, in area between VOLUME 32, NUMBER 1 9 Figs. 10-12. Male genitalia of selected Nearctic Callophrys (Mitoura) spp., lateral and posterior views with tip of aedeagus (right) and falces (left): 10, C. gryneus castalis, topotypical; 11, C. siva siva, topotypical; and 12, C. hesseli, topo- typical. 1. postvaginalis and 1. antevaginalis forming bulkly ridges and convolutions at their juncture, these binding lamellae tightly with eighth abdominal sternite and con- taining many spines. [C. hesseli (Fig. 5). Easily recognized by unique shape of the lamellae and broad cephalad tapering from the antrum (figured for reference). C. turkingtoni: female unknown. ] Males (Figs. 10-12). C. gryneus (Fig. 10). Valvae, lateral shape: only barely concave between dorsal and ventral articulation with vinculum; valvae, caudad saccus (dorsal or ventral view): rounded and indented, vaguely shouldered caudad. Saccus: long and broad. C. siva (Fig. 11). Valvae, lateral shape: deeply concave and rounded between dorsal and ventral articulation with vinculum; valvae, caudad saccus (dorsal and ventral view): parabolic and unindented, no shouldering caudad. Saccus: short and much less broad than gryneus. [C. hesseli (Fig. 12). Lateral shape of valvae less broad, quite concave between articulations with vinculum, and much longer caudad; valvae caudad saccus broadly round, indented, and extremely shouldered caudad (figured for reference). C. turkingtoni (Johnson, 1976a), easily recognized by extremely long caudad extension of valvae and by heavily sclerotized and spiny area of valvae, caudad saccus.] JouRNAL OF THE LEPIDOPTERISTS SOCIETY 10 ‘CLE ‘Suepy pUue ‘F/6T “OpUT ‘OLET “eum, Y swepy WoT poajdepe suor} -NQLySIp JUL[G “SeUI[ UsyoIq pue soUTeU Aq UAOYS S¥ UOOIPUL [VUOISe1 [e1OUNS OF SMOT[e ATWO SoOUL JO aSs¥sN UOUTWIOD JUOLINS UT UOTSNy -UO0ZD) *S}1eR4} OI[e}USS Aq posouseIp Vas (*P) ‘DO pue snaufus (*‘W) *O Jo suorjeindod o1jzeduiss oyeorpur sorenbs perte}g ‘szpdhovanwpyy pue snvadiwnf ‘syuejdpoof [eArey Iey} pue yassay (‘W) ‘DO pue snaufus (vinopyy) shiydoyjpD Jo suoTNGLNSIp ooIeeN “CT ‘Sly STTeRSeD septohy2 stredhoeerueyD pue PTOOTOTTTsS °c seprohy2 stredAseeuey)D pue euerurbira *f Trzo0youTd sniedrunc By Teyse sniedrunr ia BTOOTOTTIs snzedrunr nT ul eueTUTHhITA snisdrune S| (satoeds Aue) ATuO AINJeIVIT[ WOLF sp10sey 7 poquerdsuer2 Atqtssod ‘uotzetndog T uotjerndod pequetdsuer32 umMouy | setqunod [Tews sfdr3tnw *rressay (°K) °d ¥ Tressey (*W) °“D & satqunod [Tews etdr3rnw *“snauhz5 (*n) -»@ VoLUME 32, NuMBER 1 ai & — J. virginiana = v 6) gs J. flaccida ZN J. californica = J. pinchotii poles occidentalis ee communis horizontalis BY J. scopulorum Se J. monosperma fi J. deppeana EE Eo J. osteosperma FA) preceding 4 ia J. ashei y ee 7 gryneus | I & ¢ Se YW a & C. (M.) siva siva JT Known transplanted (x) ¢- (M.) siva juniperaria @ green morph Bop aeton: Oc. (M.) siva mansfieldi @ brown morph 2? ee OH CEE (Oc. (M.) siva, inner coastal relationship with Danpenbaoeninoenh green-brown Cc. (M.) nelsoni complex. z ae we Cc. (M.) siva, high altitude morph (M.) turkingtoni contact zone Sympatric C. (M.) siva and alc. (M.) gryneus, diagnosed by genitalia. @c. Mexican populations assigned to species as indicated. Fig. 14. Nearctic distributions of Callophrys (Mitoura) siva and its larval food- plant Juniperus spp., and known range of C. (M.) turkingtoni. Plant distributions adapted from Little, 1971. Distribution of Juniperus horizontalis shown only as it exceeds J. scopulorum northward; hybrid swarms of Juniperus spp. illustrated in Pig, 1d. 12 JouRNAL OF THE LEPIDOPTERISTS SOCIETY ae re ew were ee ene pent es tors an North Dakota r South Dakota Wyoming Nebraska Mig. 15. Bi-parental and tri-parental swarms of divergence in Nearctic Juniperus: top, localities indicated by Van Haverbeke (1968) as “hybrid” J. virginiana x J. scopulorum; center, parental areas of J. horizontalis indicating possible centers for VoLUME 32, NuMBER 1 13 Geographic Distributions and Available Larval Foodplants Figs. 13 and 14 show the Nearctic distributions of species of Callophrys (Mitoura) in relation to the ranges of available or established larval foodplants. Fig. 15 shows areas of Juniperus ranges that have been botanically demonstrated as “hybrid swarms” (Van Haverbeke, 1968; Schurtz, 1971). Summary of Data and Current Taxonomic Usages The following is a review of the current common usage of trinomens in each group with a summary that includes distribution, foodplant(s) as established in this paper, general comments on the phenotype, and notes on the particular significance of each population. Where populations are under study by other lepidopterists, and especially where they are plan- ning to assign new names, | have called these subspecies “ssp.” and in- cluded the appropriate investigator's name in brackets. Annotated List Callophrys (Mitoura) siva C. (M.) siva siva. Type locality: Fort Wingate, McKinley Co., New Mexico. Distribution: Workers have named populations distinct from this taxon only on the West Coast, although others are undoubtedly present. Phenotype: There are two general morphs, based on ground color of the ventral secondaries. Great Basin populations (Fig. 14, squares) are brown beneath, whereas others (Fig. 14, plain black circles) are green. Populations of green-browns and mixed greens and browns occur in western Utah (Fig. 14, overlapping square and circles). The brown morph, which Peter Herlan, H. K. Clench, and I have investigated (Johnson, in prep.), is separately treated below. Foodplants: Many western Juniperus species (see Table 1) replace each other geographically. “Hybrids” (see Summary and Con- clusions) of J. virginiana x J. scopulorum, J. virginiana x J. horizontalis, and J. virginiana X J. scopulorum x J. horizontalis (possibly also J. occidentalis x J. osteo- sperma) occur in western Nevada, adjacent California, and eastern Oregon. All populations of Callophrys siva are on erect trees except for one local population (Val Marie, near Rosefield, Saskatchewan, along Frenchman River) on prostrate plants. C. (M.) siva ssp. [Johnson, in prep.]. The research of Herlan, Clench, and Johnson involves naming this Great Basin population. Distribution: southern Nevada northward to Idaho; brown eastward to Salt Lake City; brown westward to southeast Oregon; broad interface with green morph C. (M.) siva siva in western Utah (e.g., Eureka, Dividend, Provo, western Millard Co.). Foodplants: Herlan reported J. osteosperma, but probably J. osteosperma x J. occidentalis near Reno, Nevada (Vasek, 1966). Apparently not J. scopulorum where it is available, al- << “hybrid” status with J. virginiana; bottom, localities indicated by Schurtz (1970) as “hybrid” J. virginiana J. virginiana (see Summary and Conclusions). Particular note: sometimes collected on nectar sources with C. (M.) hesseli, but foodplants are segregated by habitat in nature and not interchangeable. C. (M.) gryneus sweadneri (Chermock). Type locality: St. Augustine, St. John’s Co., Florida. Distribution: Florida, perhaps southern Georgia, and north along the Atlantic Coast where J. silicicola occurs. Phenotype: green morph. Foodplant: J. silicicola. C. (M.) gryneus castalis (Edwards). Type locality: McLennan Co., Texas. Distribution: mainly Texas, but also Chihuahua, Mexico, and areas west of the Mississippi River “gap” in juniper ranges; in addition, used by some workers as a form name within eastern United States populations. Phenotype: green morph. Foodplants: J. virginiana, J. ashei, and J. pinchotii, replacing each other westward. J. deppeana and possibly J. flaccida in Mexico. C. (M.) gryneus ssp. [Johnson, in prep.]. Distribution: the Baboquivari Moun- tains eastward into Cochise Co., Arizona, and possibly southward in disjunct ranges of J. deppeana. Phenotype: green morph. Foodplant: J. deppeana suspected. VoLUME 32, NuMBER 1 15 Callophrys (Mitoura) hesseli C. (M.) hesseli. Distribution: see Fig. 13. Phenotype: green morph. Food- plant: Chamaecyparis thyoides. Callophrys (Mitoura) turkingtoni C. (M.) turkingtoni, a single specimen known from Namiquipa, Chihuahua, Mexico, in habitat of J. flaccida. Phenotype: brown morph. SUMMARY AND CONCLUSIONS Interspecific relations. Studies of C. siva and C. gryneus at several sympatric localities (21 specimens from the Baboquivari Mountains, Pima Co., Arizona; Cochise County (general), Arizona; Guadalupe Mountains, Eddy and Otero cos., New Mexico, Culberson Co., Texas; and Alpine, Brewster Co., Texas) confirmed that they are separable by genitalia of the males and especially the females (Johnson, in prep.). Since town, county, or mountain range is the only data available on some of these specimens, the extent of their microallopatry or microsympatry remains unknown. Biogeographic data suggest that the species may be altitudinally separated at some localities in Texas (C. siva on higher altitude J. deppeana, C. gryneus on lower altitude J. pinchotii), but it is likely that interspecific competition occurs at some locations. Sharing of nectar sources may occur, as reported in C. gryneus and C. hesseli (J. B. Ziegler, pers. comm.). These two species are generally segregated by the habitats of their foodplants. The female genitalia of C. hesseli have not been previously figured in the literature and are included in Fig. 5. Foodplant relations. C. siva and C. gryneus utilize a broad spectrum of related and equally acceptable Juniperus species, which replace or exceed each other in geographic distribution over the Nearctic Realm. There is evidence that every species of Juniperus in the Nearctic is utilized, with two exceptions: J. communis L. and J. horizontalis Moench. Van Haverbeke (1968) and especially Schurtz (1971) have shown that J. horizontalis is actually part of a broadly distributed “swarm of diver- gence’ which involves the parental stock to which the names J. virginiana, J. scopulorum, and J. horizontalis have been applied. Van Haverbeke (pers. comm.) prefers Schurtz’s interpretation that each of these merits species status but that they are tied by their evolutionary histories, J. virginiana being an eastward evolutionary manifestation of J. scopulorum and J. horizontalis being a northward evolutionary manifestation of this biparental parent stock. Thus, there is little chemical or morphological reason (unless it is the number of needles versus fleshy leaves) that would prevent use of J. horizontalis by these Callophrys (Mitoura) especially where it is sympatric with utilized J. virginiana or J. scopu- 16 JouRNAL OF THE LEPIDOPTERISTS SOCIETY lorum. Johnson & Borgo (1976) have shown that the perching behavior of C. siva and C. gryneus is distinctly patterned and preferenced for heights. They postulate that the nature of this patterned perching be- havior selects against prostrate morphs and is at least a partial boundary on their usage as a larval foodplant (Johnson & Borgo, 1977). The im- portance of the number of needled leaves on both J. horizontalis and J. communis needs investigation since first instar larvae burrow into these to feed. Knowledge of the local specificities of the two oligophagous species is quite incomplete, although preliminary evidence from several localities indicates that populations are specific to particular plant species. In Palo Duro Canyon (Randall and Armstrong cos., Texas) J. scopulorum, J. pinchotii, J. monosperma, and hybrids of the latter two occur (Adams, 1972, and pers. comm.). Field data from collectors of C. gryneus indicate that J. pinchotii is the only foodplant. However, verification is needed by someone who can test this hypothesis directly. Peter Herlan (pers. comm.) reports that the Great Basin brown morph of C. siva feeds ex- clusively on J. osteosperma. Perhaps this is true, but Vasek (1966) has suggested that this species introgresses with J. occidentalis westward, and the taxonomic relationships of C. siva in the northwest basin are now indicated as including two, largely disjunct subspecies, one feeding on J. occidentalis in central and eastern Oregon and the other on J. osteosperma in Nevada eastward to Utah. In Missouri and Arkansas, C. gryneus populations are located on J. ashei where it occurs as “islands” within the range of J. virginiana. Other C. gryneus populations are on J. virginiana. This is another location ideal for specificity studies, as are the areas of diversity of juniper species in Arizona and New Mexico. In California, John Lane reports (pers. comm.) C. siva juniperaria perching on both J. osteosperma and J. occidentalis in an area where J. occidentalis has been reported as the foodplant. Thus, foodplant relations in C. siva and C. gryneus mirror situations reported in Burns (1964), Downey (1966), and Downey & Dunn (1965). Local specificities are due to oviposition by the female on the plant species it fed on as a larva. Thus, according to the familiar “Hopkins’ Host Principle,” specificity is main- tained. However, it is obvious that alterations do occur through time and space (as the above authors also indicate), and this is why such species show catholicity when their foodplant usage is viewed as a whole. The mechanism of ovipositional specificity and the nature of chance alterations need further elucidation. Downey & Dunn (1965) suggest that the patterning of Hopkins’ Host Principle is not genetic but physio- logical and undergoes divergence, convergence, and parallelism through VoLUME 32, NUMBER 1 17 time and space. The present study indicates that similar foodplants available as replacers offer opportunity for divergence, since nearly all barriers posed to these insects by replacer plants have been crossed. Similarly, the remarkable coincidence of distinguishable morphs or sub- species generally within the distribution of one or another foodplant or foodplant relative suggests that foodplant adaptations play an important role in subspeciation. Callophrys gryneus sweadneri inhabits the areas of J. silicicola, C. gryneus gryneus those of J. virginiana, and C. gryneus castalis those of the transition of the latter plant to the ranges of J. ashei and J. pinchotii. Relations in the C. siva complex, although trinomial knowledge is less complete, are equally distinctive. If one assumes monophagous C. hesseli evolved through adaptations of some populations of early C. gryneus stock to Chamaecyparis thyoides, a similar mechanism is imaginable, especially since C. thyoides and J. virginiana have under- gone a change in their degree of sympatry through time (M. Rosenzweig, pers. comm.) in which populations of C. thyoides are now somewhat disjunct, and those of C. hesseli apparently extremely so. Laboratory foodplant experiments with these species have not been extensive, and such data is of limited use in drawing inferences about foodplant utilization or preference in nature (Downey & Dunn, 1965; Downey & Fuller, 1962). However, studies to date indicate that quite divergent Cupressaceae species are at least nutritionally adequate and otherwise edible by some of the Callophrys (Mitoura) species. There is a need to further clarify the reported acceptance of J. virginiana by larvae of C. hesseli. Distributional relations. One comment on the distribution of these insects, with regard to the frequency of transplanted populations is ap- propriate at this time. Cupressaceae species are widely used both in agricultural and landscape planting, and a number of transplanted Callophrys (Mitoura) populations have been noted (Figs. 13 & 14). Taxonomists should be especially aware of this when studying the com- parative morphology of these butterflies. The occurrence of C. siva in planted forest well isolated in central Nebraska, where juniper is raised from Rocky Mountain stock, is an extreme example, as is the occurrence of this insect in a shelter belt along the Missouri River. ACKNOWLEDGMENTS Many lepidopterists and botanists contributed data for this study. I owe a special debt to John Lane, Oakley Shields, and Arthur M. Shapiro (Univ. of California, Davis) for discussing Callophrys (Mitoura) species with me, and to Frederick H. Rindge (American Museum of Natural 18 JOURNAL OF THE LEPIDOPTERISTS SOCIETY History, New York) for providing the facilities of the museum and editing this paper. For general support I am grateful to the staff of the Museum of Natural History, University of Wisconsin, Stevens Point, especially Charles A. Long, Director, and Robert Freckmann, Curator of the Herbarium. Debt is owed John C. Downey (University of Northern Iowa, Cedar Falls ) who introduced me to this group and suggested topics for research. The following provided data for which I am very grateful: R. A. Anderson, J. F. Gates Clarke, Ernst J. Dornfield, J. Donald Eff, George Ehle, Scott Ellis, Frank D. Fee, Clifford D. Ferris, Mike Fisher, Richard Funk, Richard Guppy, Lucien Harris, J. Richard Heitzman, Peter Herlan, Sidney Hessel, Ronald Hooper, Roderick R. Irwin, Roy O. Kendall, H. L. King, Alexander B. Klots, Steve Kohler, Henry A. LeBeau, Bryant Mather, James R. Maudsley, Tim McCabe, Lee D. Miller, James Mori, John S. Nordin, Roger Pease, Richard Priestaf, Kilian Roever, Michael Rosenzweig, R. E. Sanford, James Scott, Ernest M. Shull, William Sieker, Michael J. Smith, J. Bolling Sullivan III, Fred Thorne, and J. W. Tilden. I am indebted to the herbaria at Arizona State University (Tempe); Colorado University (Boulder); University of Florida (Gainesville); Uni- versity of Mexico (Mexico City); University of Saskatchewan (Regina); and University of Wisconsin (Stevens Point) and especially to Cupres- saceae specialists R. P. Adams (Colorado State Univ., Fort Collins) and David F. Van Haverbeke (University of Nebraska, Lincoln) for their comments and aid. LITERATURE CITED ApaMs, R. P. 1972. Chaemosystematic and numerical studies of natural popula- tions of Juniperus pinchotii Sudw. Taxon 21: 407-427. Apams, R. P. & B. L. Turner. 1970. Chaemosystematic and numerical studies of natural populations of Juniperus ashei Buch. Taxon 19: 728-751. ANDERSON, R. A. 1974. Southern records of Mitoura hesseli (Lycaenidae). J. Lepid. Soc. 28: 161. Brower, L. P. 1958. Larval foodplant specificity in butterflies of the Papilio glaucus group. Lepid. News 12: 103-114. Brown, F. M., D. Err, & B. Rorcer. 1957. Colorado butterflies. Denver Mus. Natur. Hist., Denver, 368 p. Burns, J. M. 1964. Evolution in skipper butterflies of the genus Erynnis. Univ. Calif. Publ. Ent. 27: 1-216. Comstock, J. A. 1927. Butterflies of California. By author. 344 p. Downey, J. C. 1966. Host-plant relations as data for butterfly classification. System. Zool. 11: 150-159. Downey, J. C. & D. B. Dunn. 1965. Variation in the lycaenid butterfly Plebejus icarioides III. Additional data on foodplant specificity. Ecology 45: 172-178. Downey, J. C. & W. C. Futter. 1962. Variation in Plebejus icarioides (Lycae- nidae ). I. Food-plant specificity. J. Lepid. Soc. 15: 34-42. EMMEL, T. C. & J. F. Emmet. 1973. The butterflies of southern California. Nat. Hist. Mus. Los Angeles Co. Sci. Ser. 26: 1-148 p. VOLUME 32, NUMBER 1 19 Fassett, N. C. 1945. Juniperus virginiana, J. horizontalis, and J. scopulorum— IV. Hybrid swarms of J. virginiana and J. horizontalis. Bull. Tor. Bot. Club 72: 379-384. Jounson, K. 1972. Juniperus (Cupressaceae) speciation and the ranges and evolution of two Callophrys (Lycaenidae). J. Lepid. Soc. 26: 112-116. Jounson, K. 1976a. Three new Nearctic species of Callophrys (Mitoura), with a diagnostis (sic) of all Nearctic consubgeners (Lepidoptera: Lycaenidae). Bull. Allyn. Mus. No. 38, 30 p. 1976b. A new species of Callophrys (Mitoura) from Mexico (Lepidop- tera: Lycaenidae). Pan-Pacific Ent. 52: 60-62. Callophrys (Mitoura) nelsoni (Boisduval) and siva (Edwards) in the northwestern United States, Lycaenidae. (Ms. in prep.). Jounson, K. & P. M. Borco. 1976. Patterned perching behavior in two Callophrys (Mitoura), Lycaenidae. J. Lepid. Soc. 30: 169-183. Perching behavior as a partial boundary of distribution and foodplant utilization in Callophrys (Mitoura) (Lepidoptera: Lycaenidae). (in review). Kxuors, A. B. 1951. A field guide to the butterflies of North America, east of the Great Plains. Houghton Mifflin Co., Boston, xvi + 349 p. LirtLe, J. B. 1971. Atlas of United States trees. U.S.D.A. Misc. Publ. No. 1146. Pease, R. W. 1963. Extension of known range of Mitoura hesseli. J. Lepid. Shen a Romer a Rawson, G. W., J. B. Zrecier, & S. A. Hesset. 1951. The immature states of Mitoura hesseli Rawson & Ziegler. Bull. Brooklyn Ent. Soc. 46: 123-130. Remincton, C. L. & R. W. Pease. 1955. Studies in foodplant specificity. 1. The suitability of Swamp White Cedar for Mitoura gryneus (Lycaenidae). Lepid. News. 9: 4-6. Ross, J. G. & R. E. Duncan. 1949. Cytological evidences of hybridization be- tween Juniperus virginiana and J. horizontalis. Bull. Tor. Bot. Club 76: 419- 429. - ScHurtz, D. L. 1971. A tri-parental swarm of Juniperus L. Unpubl. PhD Thesis, University of Nebraska, Lincoln. 234 p. U.S.D.A. 1953. Checklist of native and naturalized trees of the United States (including Alaska). U.S.D.A. Agric. Handb. No. 41. 472 p. Van HaversBEKE, D. F. 1968. A population analysis of Juniperus in the Missouri River Basin. Univ. Nebr. Studies, No. 38. 82 p. Vasex, F. C. 1966. Distribution and taxonomy of three western Juniperus. Brit- tonia 18: 350-372. Journal of the Lepidopterists’ Society 32(1), 1978, 19 IMPORTANT NOTICE TO CONTRIBUTORS Authors should submit an abstract for all articles to be published in the Journal beginning with Volume 32, issue No. 1. The abstract should summarize the im- portant contents and conclusions of the paper in concise and specific sentences. It should not exceed 1-3% of the paper's total length, and should indicate the ob- jectives, methods, and topics covered by the paper. References to literature, illustra- tions, and tables should be omitted from the abstract, since this should be com- pletely self-explanatory. These abstracts will be printed at the front of each paper, and also will be forwarded to Biological Abstracts for possible inclusion therein. Such abstracts should accompany articles only. They are not required for general notes. Authors who now have papers in press should forward such abstracts to the new editor at their earliest convenience. Journal of the Lepidopterists’ Society 32(1), 1978, 20-36 FOODPLANT, HABITAT, AND RANGE OF CELASTRINA EBENINA (LYCAENIDAE) WarRREN HERB WAGNER, JR. Department of Botany, The University of Michigan, Ann Arbor 48109 T. LAwRENCE MELLICHAMP Department of Biology, University of North Carolina, Charlotte 28233 ABSTRACT. The larval foodplant of the recently described Celastrina ebenina Clench is Aruncus dioicus (Walt.) Fernald (Rosaceae), the Goat’s-beard. Over 150 adults were raised from eggs and young larvae. The range of the butterfly coincides nicely with that of the plant, from Pennsylvania to North Carolina and Missouri. The habitat for plant and butterfly is moist, rich forest. The closely related C. pseudargiolus Boisduval & LeConte is vastly more abundant and ubiq- uitous than W. ebenina, and has a wide variety of larval foodplants. Larvae of C. ebenina differ in several respects from those of C. pseudargiolus, including color pattern and stellate processes. Also described and discussed are the plant and butterfly associates of C. ebenina, flower visitations of the adults, experiments on foodplant specificity, feeding characteristics of the larvae, broods, botany of the foodplant, and geographical distributions, including a number of new locality records. A guide for discovering new colonies of this rare eastern American butter- fly is provided. Except for brief reports (Clench, 1972; Wagner & Showalter, 1976), little has been published on the biology of the poorly known Dusky Blue Butterfly, Celastrina ebenina Clench, of the eastern United States. This lycaenid is notable for several reasons. Interpreted for over a century as an aberration or form, it was not recognized as a distinct species until 1972. The colors of the upper surfaces of the males and females are peculiar for being the reverse of the usual situation among plebejine blues in that the males are dull, dark grayish-brown or blackish, while the females are mainly lustrous blue. The insect is regarded as especially rare and local, having been reported previously, usually as just one or a few individuals, from only 12 localities. Knowledge of its foodplant, behavior, habitat, and geographical distribution has been incomplete or lacking. The present paper records the results of research in 1976. We now understand the ecology of C. ebenina far better than we did in the past, and we believe that we have an explanation for the geographical distri- bution and sporadic occurrence of the species. At the outset of this study, as botanists, we entertained the possibility that the peculiarities of oc- currence of C, ebenina might be due to specialized larval foodplant preference, Because of earlier reports of the species there, the area chosen for our field investigations was in the Daniel Boone National Forest, in and VoLUME 32, NUMBER 1 2] around the Red River Gorge in Powell and Menifee cos., Kentucky, a few miles north of the town of Slade. We found that habitats suitable for C. ebenina occur extensively, though sporadically, throughout this area, and in some places the butterfly is common or abundant, though extremely localized, flying with the much more numerous and ubiquitous Common Blue or “Spring Azure,” C. pseudargiolus Boisduval & LeConte. The topography in the Red River Gorge is made up of steep, abrupt hills and valleys, ranging from 700’ altitude in the river and stream beds to over 1300’ at the tops of the highest hills. Commanding cliffs of light tan or whitish sandstone crop out at the crests of some of the hills, but in the valleys where C. ebenina flies, the most conspicuous rock is a loose, broken, dark-gray shale. The general area is mainly traversed by narrow dirt roads, especially along the larger streams, and there are only occasional two-lane hardtop roads. Our search for suitable habitats to study was initiated on 17 April 1976. We drove from place to place along the country roads, stopping wherever we encountered roadside puddles or wet streamside flats. In such spots accumulations of butterflies were the rule when it was bright and sunny, especially between the hours of 0900 and 1400. Among the guests at these “puddle parties” we encountered rare males of C. ebenina, prac- tically always with at least several and usually many C. pseudargiolus, the latter being much more conspicuous in flight. In spite of our success in encountering specimens of C. ebenina here and there over an area of perhaps a dozen square miles, at no place were there more than a few individuals. Rare observations of females showed them almost always to be in flight, and following them gave no clue to where their eggs were laid. Finally, at around 1030 hrs the following day, we came upon an excel- lent locality—a moist, steep, rocky, north-facing wooded slope, along a narrow dirt road along the Red River. In only 20 min, ca. three dozen C. ebenina of both sexes were observed, the females numbering about twice as many as the males. The latter were all more or less worn, dull in appearance, and flying along the edges of the road, occasionally alighting on wet muddy spots (Fig. 1, lower photo). Most of the females, however, were in fresh condition and were flitting around the vegetation on the slopes above and below the road. It was obvious that many of them were engaged in oviposition. They flew rapidly in an “exploratory” pattern, pausing often at a single species of plant, the Goat’s-beard, Aruncus dioicus (Walt.) Fernald. Occasionally the females landed on the abundant and conspicuous Wild Hydrangea, Hydrangea arborescens L., but usually only momentarily. JOURNAL OF THE LEPIDOPTERISTS SOCIETY bo bo Fig. 1. Celastrina ebenina habitat along Red River, Powell Co., Ky. Upper: View upstream showing extensive understory growth on forest slope. Butterflies visit Geranium flowers here. Lower: View downstream showing damp ruts in road where males congregate. (Photo by J. M. Beitel. ) VOLUME 32, NUMBER 1 23 Figs. 2-6. Celastrina ebenina adults and eggs: 2, female laying eggs on young Aruncus shoots (R. P. Carr); 3, freshly emerged male, showing blue scaling (T. L. Mellichamp):; 4, eggs on leaf and inflorescence primordia of Aruncus (cf. fig. 14) (T. L. Mellichamp): 5, unhatched eggs; 6, eggs (two of them hatched) showing wall pattern detail (R. P. Carr). On Aruncus the butterflies alighted on very young, unfolding leaflets, and then walked around slowly, laying eggs (Fig. 2). After an individ- ual would fly away, we could easily find the eggs, mainly on the lower blade surfaces, on and between the main veins of the leaflets (Fig. 4). At this time of year the main axis of the plant is still embryonic, and the habit is very different from the mature habit with the inflorescence fully developed (cf. Fig. 13, full-grown plant, and Fig. 14, stage at time of oviposition). When freshly laid, the eggs showed a grayish blue color. 24 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Sometimes several eggs are laid in the same spot, but usually they are laid separately (Figs. 4, 5, & 6). Egg-laying occurred over 1% hours of observation, and there was no sign of abatement after 1200 hrs when we left the site. For careful observations, a total of 18 young cuttings like those in Fig. 14 were randomly collected. The two oldest leaves overtop the main shoot at this stage. The main shoot, with its very young leaves and inflorescence primordium, is only about one-fourth the length of the oldest leaf (the large, bipinnate leaf on the right side of the figure) and one-half the length of the next oldest leaf. The softest, most embryonic tissues are those of the primordial main shoot. We found a total of 133 eggs altogether on the collected shoots—35 on the oldest leaves, 26 on the next oldest, and 66 on the young main shoots. Thus, our evidence suggests that the butterflies prefer to lay eggs on the youngest tissues. The number of eggs averaged seven per cutting, but one had 18. Naturally we wondered whether at least some of the eggs we found did not represent the closely related C. pseudargiolus, which flies in large numbers at this locality, a species noted for its polyphagy. There- fore, we decided a couple of weeks later to conduct an experiment de- signed to help us answer this question, as will be described below. The plant community on this slope is a rich mixed-mesophytic forest. We recorded a total of 31 trees and shrubs and 56 herbs (including ferns and graminoids ) in the area where C. ebenina was ovipositing. Most of these plants are typical associates of Aruncus dioicus, and some of the more prominent ones will be enumerated later in our discussion of this plant. The best place to find C. ebenina adults is in association with other mud-loving butterflies (C. pseudargiolus, Callophrys henrici, Erynnis spp., and Papilio spp.) in damp spots along dirt roads and gravelly, sandy, or muddy river flats. Practically all of the “mudding” individuals we observed were males, sometimes as much as a quarter of a mile from the foodplant, although usually much closer. On only two occasions did we find females landing on wet soil. We disturbed one of them several times, but each time it returned. The only flower which seemed to attract C. ebenina at this locality was the Wild Geranium, Geranium maculatum. The showy rose-purple flower has a flat five-petalled corolla 3.0-3.5 cm across. Bearded nectaries occur between and at the bases of the petals. The butterflies walk over the top surfaces of the corolla and probe between the petal bases. Later 'A complete list of the associated plants at this locality will be sent upon request to readers. VOLUME 32, NUMBER 1 25 Figs. 7-12. Celastrina ebenina immature stages: 7 & 8, instar 1 caterpillars (J. G. Bruce III); 9 & 10, instar 2 caterpillars (J. G. Bruce III); 11, mature caterpillars, showing pale, poorly contrasting pattern and characteristic leaf damage; 12, pupa attached to Aruncus leaf (R. P. Carr). we discovered that the bulk of individuals obtain their nectar from underneath the flower! Both sexes flit from flower to flower, landing on the peduncle or on the underside of the perianth, then walking toward the sepal bases where they insert their proboscises. So positioned on the flowers, the butterflies are invisible from above.” We recorded all of the species of butterflies we found in association * Curious to see whether other butterflies behaved in the same manner, we discovered (in Michigan in the middle of May) that Erynnis juvenalis displayed the same routine on Geranium flowers. JOURNAL OF THE LEPIDOPTERISTS SOCIETY pecaaimnnanat e —.rrtrs——C 1O 6M. (‘a —s 13, habit drawing of fully grown Aruncus dioicus showing inflorescence (apex) and form y); 14, young shoot of Aruncus at time of oviposition by C. ebenina (see text) (T. L. ~, Vg , Vial of WEN Nia Fog a ; AS . RSS Cor ) A neil Figs. 13-14. Celastrina ebenina foodplant: of compound leaves in mid-June (Del. J. G. Lac Mellichamp ). VoLUME 32, NuMBER 1 27 ee = *\\ Fig. 15. Geographical distribution of Aruncus dioicus (stippling) and docu- mented localities for Celastrina ebenina (dots) (T. L. Mellichamp). 28 JouRNAL OF THE LEPIDOPTERISTS SOCIETY with C. ebenina on 18 April. The following list is a compilation, since no one locality had all of the species (those marked with an asterisk oc- curred with C. ebenina on wet soil): Papilio glaucus,* P. troilus,* Battus philenor, Pieris virginiensis,* Colias eurytheme, C. philodice, Polygonia comma, P. interrogationis, Boloria bellona, Phyciodes tharos,* Callophrys henrici,* Celastrina pseudargiolus,* Erynnis brizo,* E. juvenalis,* Amblyscirtes samoset, A. aesculapius, and Epargyreus clarus. On several later visits to the study area by Loran D. Gibson, Amos H. Showalter, and ourselves, the following additional species were observed, most of them, however, well beyond the normal flight period of C. ebenina, which ended the first week of May: Limenitis astyanax, Speyeria cybele, Asterocampa celtis, Ewptychia hermes, E. cymela, Lethe ~ creola, L. anthedon, Autochton cellus, Thorybes pylades, and Poanes hobomok. Experiment on Foodplant Specificity The question we addressed ourselves to was whether C. pseudargiolus also shared the foodplant of C. ebenina. The former is abundant in the Red River Gorge area and is in constant association everywhere with C. ebenina. Although we did not observe C. pseudargiolus ovipositing upon Aruncus, this would not preclude the possibility that it occurs. The num- ber of Celastrina eggs we observed was so great that it seemed reasonable to assume, because of their morphological similarity, that perhaps some of them belonged to the abundant species, especially since C. pseu- dargiolus is noted for its varied bill of fare. Literature records show that no less than 10 families of flowering plants contain larval foodplants for this species, and we have discovered early stages of it on such different families as Caprifoliaceae and Cornaceae. It would not have surprised us, therefore, to find C. pseudargiolus utilizing Aruncus, especially since its larvae have been reported on Spiraea, another closely related genus in the same subfamily of Rosaceae. On 2 May, accordingly, we revisited the Red River Gorge area and collected from seven to nine shoots with attached eggs and minute larvae from each of five colonies of the foodplant, all of them a mile or so separate from one another. These shoots were brought back to The Uni- versity of Michigan in Ann Arbor, and the eggs and young larvae were raised to adulthood. Extra foodplant shoots were kept in plastic bags and refrigerated to preserve them until they were needed, and additional foodplants were grown for further supplies. The cut bases of the shoots of Aruncus bearing eggs and larvae were inserted in jars of water in | x 1 x 2 ft glass aquaria covered with a plastic material to keep the VoLUME 32, NUMBER 1 29 larvae from escaping. Some of the caterpillars drowned when they walked down the main stems into the water, others escaped and were lost, and some died as a result of cannibalism. Nevertheless, a total of 153 butterflies were raised to maturity by the end of the second week in June, 76 males and 77 females. Another eight butterflies came from larvae which escaped from their containers. The first caterpillars pupated on 16 May, 14 days after the field collection, and the first adults to emerge appeared on 23 May. This second brood apparently does not occur in nature (see below). The results of this experiment were striking. Not only were none of the “checkery” caterpillars of C. pseudargiolus noticed among the paler, less contrasty larvae of C. ebenina (Figs. 7-9), but not a single one of the adults was C. pseudargiolus; all were C. ebenina. Furthermore, the 14 additional butterflies from our 18 April eggs were all C. ebenina. Also, the four individuals that emerged from dormant chrysalids the following winter were all C. ebenina. With C. pseudargiolus as abundant as it is in the localities where our collections were made, one would expect at least some evidence of its occurrence on Aruncus if it fed on that plant at all. It is interesting to note that ca. one-third of the males display blue scales (Fig. 3), at least at the time of emergence. Although the original description by Clench (1972, p. 37) records the male upperside as “uniform blackish brown when fresh,’ blue scales are conspicuous in these individuals, especially on veins R, to R; and the discal portions of M;, Cu,, and Cu, on the fore wings, and scattered in the discal area of the hind wings. The occurrence of blue scaling on the males may con- stitute a regional difference. The ground color above of the males varies from blackish gray to pale slate gray. The females vary greatly in ground color (whitish blue to fairly intense blue) and the amount of dark mark- ing. There is no confusing any of these specimens, however, with any of the forms of C. pseudargiolus known to us. To make preliminary comparisons of the caterpillars of C. ebenina and C. pseudargiolus, we obtained eggs and first instar larvae of the latter in two localities near Ann Arbor in Washtenaw Co., Michigan. They were readily found on the young inflorescences of Grey Dogwood, Cornus racemosa and Red Osier, C. stolonifera (Cornaceae), and Nanny- berry, Viburnum lentago (Caprifoliaceae); the dates of collection were 8 and 22 May. We maintained the larvae under the same conditions as those of C. ebenina but kept them in a separate room to avoid any op- portunity of possible escapees getting mixed up. We did not make detailed comparisons of the larval morphology, but 30 JOURNAL OF THE LEPIDOPTERISTS SOCIETY certain differences were obvious. The final instar caterpillars of C. ebenina (Fig. 11) are considerably more uniform in color than those of C. pseudargiolus. They are pale whitish blue-green, with three slightly contrasting longitudinal stripes of yellowish white, two lateral and one dorsal. Between the dorsal and each lateral stripe is a line of yellow— white dots, and similar dots and dashes are scattered over the body. The caterpillars of C. pseudargiolus, as is well known, are more variable in eround color, ranging from pinkish or pale bronze to yellowish green; the pattern is much more conspicuously blotched as a rule. Earlier larval stages of the two species (young instars of C. ebenina shown in Figs. 7-10) are very similar in their uniform pale green color. In both species the mature larvae are velvety because of elaborate stellate processes, each with a narrow and pointed filament arising from the center. In C. ebenina the filaments are only slightly curved and stand nearly erect. In C. pseudargiolus the filaments are more strongly curved and tend to be arched over such that they are distally nearly parallel to the body surface. This difference applies mainly to the more abundant, smaller filaments; larger ones in both species are more alike, only slightly curved and nearly erect. Another difference between the larvae of the Dusky and Common Blues involves the use of their respective foodplants. Although larvae of C. ebenina occasionaily feed upon the tiny embryonic floral primordia of Aruncus, the bulk of their feeding is upon the blade tissue between the major lateral veins of the leaflets. Their feeding produces charac- teristic elongated perforations in the blades; these persist when the leaflets have achieved their full size, and give valuable clues during late spring and summer for localities of the butterfly. Fig. 11 shows the characteristic perforations. The younger caterpillars were observed on the tops of the leaves more often than the older ones. Feeding between a pair of lateral veins usually begins near the midrib of the leaflet and progresses outward, but only rarely all the way to the margin and in- cluding it. Very large caterpillars in the last instar may eat major lateral veins. In our experience the larvae of C. pseudargiolus feed primarily upon floral primordia. Their eggs and larvae are found in inflorescenses. We tried moving larvae to leaves, but always, after eating a small amount, they would return to their floral clusters where they continued feeding. Larvae of C. pseudargiolus bite into the sides of the closed flower bud or the inferior ovary and eat the entire contents or leave certain parts (e.g., petals). The body of the caterpillar sits motionless on the floral pedicel or the side of the bud, and the extensible head is projected into VOLUME 32, NuMBER 1 31 the cavity in the young flower to feed. From certain angles the larva appears headless. Broods Previous field experience of our own and others suggested either that C. ebenina is univoltine (in the Red River Gorge, flying from the second week of April to the first week of May), or else, if it has more than one brood, that adults of any later broods are so similar to those of C. pseu- dargiolus that they have not been recognized as distinct. The second lab- oratory brood is like the first brood. As it turned out, the vast majority of our pupae (Fig. 12) emerged within a week or two of pupation. Of those that failed to emerge and were kept until the following winter, only a few produced butterflies. The first adult from our 18 April field collections of C. ebenina eggs emerged on 11 May. The first from our 2 May collections of eggs and young larvae emerged on 23 May. Emergence continued until 29 May, when it was stopped by placing the cultures in a coldroom from this date until 4 June to keep more butterflies from appearing while we were out of town. After we returned them to normal temperatures, emergence resumed on 5 June and continued to 12 June. A grand total of 175 butter- flies in our laboratory cultures seemed to demonstrate that there is a second brood in the wild that follows closely upon the first, the second brood flying from the second week in May to the first or second week in June. We therefore returned to the Red River Gorge to determine whether a second brood occurred in nature. The results of our survey were most unexpected. Loran D. Gibson visited there on 26 May and saw not a single C. ebenina, although many other butterfly species were seen (in litt., 2 June 1976). We then reconnoitered the area on 4 June, when the butterflies should have been at their peak abundance, if there is a second brood in nature. We saw no C. ebenina despite the fact that C. pseu- dargiolus was common as well as 16 other species of butterflies. On 9 June, Amos H. Showalter searched the area and reported that “C. pseu- dargiolus was common, but no ebenina” (in litt., 28 June). After 13 June, the 28 pupae that remained in our cultures apparently went into dormancy, and no more butterflies emerged. At the end of June, therefore, we placed them in a coldroom at a temperature of 2°C and left them there until 22 December. We hope that by keeping them thus, in a set-up that we have used for “winterizing” fruits and seeds, we might avoid the destructive effects other workers have had with C. pseudargiolus, which involve either drying out or molding of the chrys- 32 JouRNAL OF THE LEPIDOPTERISTS SOCIETY alids. On 1 January 1977, two females emerged, one of which failed to expand its wings. Another female emerged on 4 January, and still an- other female failed to escape from the pupal skin. All the remaining pupae appeared to have died, eight of them having moldy surfaces and the rest having an unnatural brown color. } What can we conclude regarding the broods of C. ebenina? In the field we found no evidence for a second brood. If there is one, it must be in extremely low numbers, i.e., a small “partial” brood. Somehow the conditions of our laboratory cultures must have caused an abnormal eclosion without the customary prolonged dormancy period. Some dia- pause stimulus that effects C. ebenina must have been weak or missing under the conditions of our experiment, and thus only a small percentage of the chrysalids went into long-term dormancy. Botany of the Foodplant The colloquial name of the foodplant, Aruncus dioicus, may cause some confusion, since its name “Goat’s-beard” is applied also to the un- related Tragopogon pratensis L. in the Asteraceae, a naturalized weed from Europe. The generic name Aruncus comes from the Greek and means literally “goat’s beard.” It is a member of the Rosaceae and is a native eastern American plant of rich, mature forests. It is famous among United States’ botanists as an illustration of convergent evolution, because superticially A. dioicus resembles closely the “False Goat’s-beard,” Astilbe biternata, of the Saxifragaceae. So closely do these plants resemble each other that they are regularly confused, even in herbaria. Ecologically the two look-alikes occupy almost identical niches, and they are both unusual among members of the mesophytic forest association in being dioecious (male and female flowers being borne upon separate plants). They are pollinated not by wind, which is the usual situation in dioecious plants, but rather by insects, mainly small Hymenoptera. The geograph- ical range of Aruncus dioicus is shown in Fig. 10. The range of Astilbe biternata is much narrower, mainly in the mountains of North Carolina and adjacent parts of Virginia, W. Virginia, Kentucky, Tennessee, South Carolina, and Georgia. Aruncus overlaps it completely, so that students of C. ebenina must be warmed of the danger of confusing the “True” with the “False” Goat’s-beards in the area of their sympatry. Accord- ingly, we have prepared a comparison of the two in Table 1, the most obvious characters marked with asterisks. A line drawing of a mature specimen of Aruncus nearly 1’4 m tall is reproduced in Fig. 1. The stage of growth when the plant serves as larval food for C. ebenina is shown in Fig. 2, corresponding to only the two bottom leaves and the lower VOLUME 32, NuMBER 1] Oo TaBLE 1. Comparison of “True” and “False” Goat’s-beards. Aruncus dioicus Astilbe biternatum *1. Stipules Absent Present 2. Terminal leaflet Unlobed 3-lobed 3. Leaf base Attenuate Cordate (heart-shaped ) *4. Veins per leaflet 8-18 pairs 8 or less pairs *5. Leaf and stem hairs Absent Abundant ( glandular ) 6. Marginal teeth Convex Acuminate 7. Sepals per flower 5 5) 8. Petals per flower 5B 0-5 9. Stamens per flower 15-20 8-10 *10. Carpels per flower 3-4 2 sixth of the drawing. The tissues upon which the caterpillars feed are soft, and the earliest instars feed upon the most embryonic parts. The plants grow rapidly and come into flower 5-7 weeks after the butterflies lays their eggs. Both Aruncus and Astilbe have rather massive underground stems that produce large roots or root masses which hold the plants firmly in place on steep slopes (Fig. 3). Many buds are present at the ground level, and some of these may develop into shoots at the crown, producing clumps of as many as eight flowering shoots. Their spreading compound leaves fill in the space where they grow, presumably allowing little growth of other plants beneath them. The Goat’s-beards are most typical of rich, mesic woods, partly shaded roadsides, and sloping sides of streams and rivers. The ancient habitat was probably on steep eroding slopes and stream banks in dark forested areas, but man has stimulated its spread by creating new habitats where roads have been cut through the mesic forest. In a uniformly shaded forest stand only 10% of the plants may flower, but when released from the effects of low light levels, as on road cuts at the forest edge or steep, eroding stream banks, the populations may display up to 100% flowering. The plants require, however, relatively cool, moist conditions, and they exist almost exclusively upon north-facing slopes. Aruncus is known to occur as high as 5500’ altitude in the mountains (Buncombe Co., N.C.), although the average occurrence throughout its range is considerably less than 2500”. Would-be collectors of C. ebenina should seek the foregoing site con- ditions, with the following array of associated species (based upon studies of a number of Aruncus localities by Mellichamp 1976): woody plants—Acer rubrum, A. saccharum, Aesculus octandra, Betula lenta, Carpinus caroliniana, Cornus florida, Fagus grandifolia, Lindera benzoin, 34 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Liriodendron tulipifera, and Tilia americana; herbs—Adiantum pedatum, Athyrium filix-femina, Botrychium virginianum, Carex plantaginea, Cimicifuga racemosa, Geranium maculatum, Impatiens capensis, La- portea canadensis, Tiarella cordifolia, and Trillium spp. The presence of a majority of these species, together with Wild Hydrangea, Hydrangea arborescens, given the topographical conditions cited, especially a north- facing slope, should lead to colonies of Aruncus and therefore C. ebenina. Ranges of Plant and Butterfly There is a remarkable correlation between the known localities where the butterfly has been found and the geographical distribution of Aruncus dioicus. Clench (1972) has already discussed doubtful records for C. ebenina, including New York City and southern Colorado. We should like to add to the list of doubtful records that of Blatchley in Wabash Co., Indiana (Clench, 1972, p. 41), unless an actual specimen is dis- covered. The “black male” he referred to could have been a melanistic male of C. pseudargiolus, a wind-blown stray of C. ebenina, or a mis- labeled specimen. Clench mentions that Edwards had vaguely attributed the species to Tennessee and Georgia, although Clench himself had seen no specimens in these areas. It now seems very likely, as Clench (1972) suggested, that C. ebenina may turn up in both of those states. Aruncus grows far to the west of the localities cited in earlier studies of C. ebenina (Clench, 1972; Wagner & Showalter, 1976), the western- most documented records for which were all east of Cincinnati, Ohio, and Lexington, Kentucky. We can now report records in Illinois, Missouri, and Arkansas. Charles L. Remington took a fresh male south of Elsah, Jersey Co., Illinois, on 15 April 1942 (in litt., 21 Jan. 1977). He found additional males in St. Louis Co., Missouri, in the late 1930’s. Also, J. Richard Heitzman (in litt., 14 July 1976) has informed us that he ob- tained by exchange “a male and a female taken 4 May 1924 at Creve Coeur Lake near St. Louis, St. Louis Co. There is no collector’s name, but it should have been one of the active collectors of the day, probably Ernst Schwarz, E. P. Meiners, or H. I. O’Burme. AII were active at the time and collected often at Creve Coeur.” The dot shown in Fig. 10 for Arkansas is based upon a single male taken in Hickory Flat Hollow, Washington Co., on 2 April 1973 by Edward Gage. Gage writes (in litt., Dec. 1976): “This male particularly stood out as it was flying about a mud puddle with several C. argiolus and Everes comyntas. .. . I im- mediately assumed that it was a melanic form and quickly collected it. The site was in close proximity to a draw or shallow canyon. About 90% of the immediate surrounding area is deciduous hardwood. ... Can- VoLUME 32, NUMBER 1 By5) yons and woodland extend all around Beaver Lake from the collecting site. ” CONCLUSIONS Celastrina ebenina may have larval foodplants other than Aruncus dioicus. However, it should be noted that none of the unquestioned localities of this butterfly lies outside the known range of Aruncus. Furthermore, we now have good reason to believe that the abundant Common Blue, C. pseudargiolus, does not share the foodplant of the Dusky Blue, C. ebenina. Our experimental raising of eggs and young larvae on Aruncus from areas in which C. pseudargiolus is abundant revealed not a single specimen of that species. All were C. ebenina. One reason for apparent rarity of C. ebenina in comparison with its near relative is that its geographical range is much more limited. Another is that it is probably monophagous rather than polyphagous. Its food- plant is confined to one habitat—north-facing, richly wooded, shaded slopes, so that the butterfly tends to be highly localized and colonial. The multiple foodplants of C. pseudargiolus occupy many habitats, and the butterfly is therefore practically ubiquitous. Celastrina ebenina is probably often overlooked. The dull males in flight may suggest badly worn individuals of C. pseudargiolus. Celastrina ebenina tends to fly closer to the forest floor (a concomitant of its under- story foodplant?), whereas C. pseudargiolus has a slow, up-and-down flight reaching the shrub and lower tree layer. The bright reflecting blue of C. pseudargiolus males plus their tremulous flight pattern through the woods at heights of roughly 2-10’ make them especially visible. The females of C. ebenina resemble dull females of C. pseudargiolus, but they are even more localized than the males, rarely even visiting mud puddles, occurring rather in the herb layer of the forest in more or less checkered sunlight. We wonder how many collectors (including our- selves!) in the spring have overlooked C. ebenina while they focused instead on such critical genera as Erynnis and Callophrys as well as such widely advertised rarities as Pieris virginiensis and Erora laeta, both of which are now known to fly in association with C. ebenina. If the conclusions of this research are correct, we predict that C. ebenina will be found not only in many new localities in the states from which it is already known, but, in addition, southern Indiana, eastern Tennessee, northern Georgia, western South Carolina, and western Mary- lan. To help achieve this, we propose the following formula: 1. Locate areas of rich mesophytic forest in rolling or mountainous country. 36 JoURNAL OF THE LEPIDOPTERISTS SOCIETY 2. Follow roads or streams and find north-facing, cool, shaded forest slopes with some erosion or disturbance. 3. Look for plant associations including such trees and shrubs as Hydrangea, Acer, Aesculus, Betula, Carpinus, Cornus, Fagus, Lindera, Liriodendron, Rhododendron, and Tilia plus the majority of herbs given above. 4, Explore for large colonies of “Goat’s-beard,” Aruncus dioicus, the larval foodplant, especially on north-facing roadsides and streamside slopes. 5. Visit the area in April and early May in search of C. ebenina— males on muddy spots, females around the foodplant, and both sexes on Geranium flowers (careful!—they may be underneath the petals). 6. Or, if the weather is cloudy or rainy, search the young shoots of Aruncus for greenish blue, rough-surfaced eggs, and (or) pale green caterpillars, the latter evidenced by narrow perforations in the soft leaf tissue between the veins of young leaflets. To sum up, our evidence thus far indicates that Celastrina ebenina is a “specialist,” not a “generalist.” When compared with C. pseudargiolus, it has a narrow range (not broad), one foodplant (not many), a single brood (not several), and an essentially uniform morphology (not many varieties and forms ). ACKNOWLEDGMENTS We thank the following persons for their help in this project: J. M. Beitel, J. G. Bruce III, Robert P. Carr, D. J. Harvey, Loran D. Gibson, J. Richard Heitzman, Janice Glimn Lacy, Amos H. Showalter, Florence S. Wagner, and K. S. Walter. We are especially grateful to Harry K. Clench for his encouragement and critical advice. LITERATURE CITED CLeENCH, Harry K. 1972. Celastrina ebenina, a new species of Lycaenidae (Lepidoptera) from the eastern United States. Ann. Carnegie Mus. 44: 33-44. Me.iicuamp, T. L. 1976. A comparative study of Aruncus (Rosaceae) and Astilbe (Saxifragaceae ), and the problem of their relationships. Doctoral Thesis. The University of Michigan, Ann Arbor. Wacner, W. H., Jr. & AMos H. SHowatrer. 1976 Ecological notes on Celastrina ebenina (Lycaenidae). J. Lepid. Soc. 30: 310-312. Journal of the Lepidopterists’ Society 32(1), 1978, 37-48 STUDIES ON RESTINGA BUTTERFLIES. II. NOTES ON THE POPULATION STRUCTURE OF MENANDER FELSINA (RIODINIDAE) Curtis J. CALLAGHAN IBM do Brasil Ltda., P.O. Box 1830, Rio de Janeirv, Brazil ABSTRACT. The objective of the study was to describe various aspects of the adult behaviour and population dynamics of the riodinid butterfly Menander felsina (Hew). The conclusions were based on four years of field observations and a marking-recapture study conducted over a period of 15 weeks. The population was characterized by low intensive and extensive frequencies (rarity in numbers and space respectively), characteristics shared by many other forest riodinid species. M. felsinad maintained constant population levels of about 19 individuals over the 15 week marking-recapture period, due to 1) longevity above that of most holarctic lycaenids, 2) low egg laying frequency both in time and space, and 3) male territoriality, which results in older males doing most of the mating. Limited adult and larval food sources were discounted as an explanation. The population was found to be distributed in small groups or colonies near food plant localities. The reasons for this were the low extensive distribution of foodplants coupled with high female vagility. Depending upon conditions at each foodplant locality, such as predation and exposure to the elements, each colony could become extinct, only to be reestablished by another wandering female. Studies on the ecology of neotropical butterflies to date have been concerned for the most part with the larger species such as heliconiines (Crane, 1955, 1957; Turner, 1971; Ehrlich & Gilbert, 1973), nymphalines (Benson & Emmel, 1973; Young, 1972), and papilionids (Cook e¢ al., 1971). Smaller butterflies such as riodinids have received only super- ficial treatment, mainly consisting of scattered notes on observed habits and population density. Early writers such as Bates (1864) and Seitz (1913) made reference to the rarity of individuals and the great num- bers of species that characterize neotropical riodinids. This has been confirmed statistically by Ebert (1969), who used the terms low ex- tensive and intensive frequency to denote the rarity of populations and individuals, respectively. However, there has been no attempt to explain the mechanism(s) involved in maintaining populations in spite of such low intensive and extensive frequencies. The opportunity to study this phenomenon was provided to me by the discovery of a reasonably large colony of Menander felsina (Hew.) near Rio de Janeiro, Brazil. Whereas the ecology of this butterfly resembles closely that of other forest- dwelling but less common species, it also provides the basis for con- clusions of a general nature applicable to other members of this group. The purpose of this paper is to describe the adult behavior of M. felsina in connection with feeding, male patrolling, and mating. The 38 JOURNAL OF THE LEPIDOPTERISTS SOCIETY results of a marking-recapture program are presented with an analysis and discussion of the population parameters. METHODS The observations that form the basis of this study were gathered over a period of more than 4 years at Pedra de Itauna, an area near the coast west of Rio de Janeiro, Brazil. The vegetation is typical of the restinga (coastal dune community) described in detail elsewhere (Cal- laghan, 1977). The area for the mark-recapture study was a trail some 125 m in length just inside the south (seaward) side of the low woods surrounding the Itauna Rock. Because of the density of the brush, little collecting was possible elsewhere in the forest. The mark-recap- ture project took place over a 14-week period, from 18 July-15 November 1973. Marking was done every weekend except for five near the end, which were lost as a result of inclement weather. An average of ca. 5 h collecting (900-1400 hrs) per day was devoted to the experiment on study days. Individuals, once captured, received a predetermined mark that indicated the day and were immediately released at the same spot. Occasionally, a butterfly would show signs of shock after marking by fluttering to the ground, in which case it was killed and eliminated from the experiment. For analyzing the data, the Fisher and Ford method (Fisher & Ford, 1947) was used because of the relatively small sample sizes. RESULTS General Behavior: Feeding, Mating, and Patrolling Menander felsina inhabit exclusively the low woods and nearby flats on the seaward side of the forested areas. They prefer open habitats such as trails and small clearings in the woods, shunning completely the deep shade found in the higher forests. Yet, they are one of the few butterflies that fly on cloudy days, albeit in smaller numbers than on sunny ones. Both sexes can be found sunning in the early morning hours with wings outspread on the upper surfaces of leaves 2-3 m above the ground. When disturbed, they fly off with a rapid, jerky flight, circling a few times before settling, wings outspread on the undersides of leaves not far from where they departed. After a few minutes they sometimes return to the same spot. When resting, they appear to thermoregulate, raising the wings to sharper angles depending on the intensity of the sun’s rays. Group feeding takes place in the morning hours from 800-1100, and to a lesser degree in the late afternoon. The M. felsina will visit prac- VOLUME 32, NUMBER 1 39 tically any plant that happens to be in bloom in the habitat, often becoming so “engrossed” in feeding that they can be removed with forceps. Males and females feed together, the latter being found com- monly only at this time. Starting ca. 1230 hours, males begin taking up positions on the edges of trails and clearings to await females. They sit motionless for long periods on the upper or undersurfaces of leaves from 0.2—2 m above the ground with their abdomens slightly raised. Males seldom leave their perches to investigate other species of butterflies. The only ones in- vestigated were small white pierids (Eurema sp.), which to some extent resemble M. felsina in flight. Larger butterflies and skippers are com- pletely ignored, which indicates that sight plays a role in the recogni- tion of rivals or mates. This activity continues until ca. 1500. This behavior is in contrast to the habits of lycaenids (Powell, 1968; Scott, 1974) and skippers (MacNeill, 1964). In these cases the males would investigate any object flying past them, including small rocks (MacNeill, 1964). To determine the extent to which spacing occurs among males, a section of trail some 20 m long was observed on three occasions. In the middle of this area is an opening in the woods that faces outward toward the knee-high vegetation of the flats. Here, an older, slightly worn male took up a station at 1255. From time to time he would take off, flying around in an area some 5 X 3 m and perching for a few minutes on various plants within this area. At 1326, two fresh males moved into the study area, taking up positions to the right and left of the older male. When one of these flew near the older male, the latter rose up, flew around in circles with the other for a few seconds, then returned to his original spot, the fresh male alighting a few yards out- side the area of the older male. At 1405, a female alighted on the outward side of the older male’s area. He immediately flew over to the female, who took off, and followed her out onto the flats. A half hour later, he had not returned, nor had other females appeared in the areas of the fresh males. The next day at the same time, the same older male was again observed in the same area. On another occasion only fresh males were observed. They showed less exclusiveness with regard to patrolling areas than the older males. Another male passing nearby would sometimes be engaged in a circular mutual chase, but both individuals would then settle down on leaves sometimes only a meter apart. Once a male left an area, another would often move in. During the observation periods, no females were seen entering the fresh males’ areas. 40 JOURNAL OF THE LEPIDOPTERISTS SOCIETY TasLe 1. Basic capture-recapture data on a population of Menander felsina. The capture or recapture of each individual butterfly is denoted by 1. Aug. Sept. Oct. Nov. 18 1 9 17 24 a 14 20 15 1 iL it il 1 1 1 1 if il if 1 il 1 Il Il 1 il 1 1 1 i 1 Il il i 1 I 1 Ht 1 il iL 1 Jl 1 1 iH 1 1 it it 1 1 Il il il Hi it il 1 1 1 a I i il i 1 1 1 1 Ju 1 1 1 1 1 1 1 IL 1 1 1 1 iL 1 1 1 Tatal Malaise, 122 kL none 17 ile 19 7 bo i bo iN) I conclude that M. felsina males do engage in spacing, the older individuals defending their chosen areas more vigorously than the younger, more inexperienced males. That the older male was successful in meeting a female suggests that they may choose and defend the preferred “rendezvous” areas. VoLUME 32, NUMBER 1 Al TABLE 2. Type A data trellis derived from data in Table 1. Units under “Date of Marking” refer to marks and not animals. Date of Marking Date Captured Released 18 1 9 7) 24 ve 14 200 t5 18 Aug. 22, 22, 1 Sept. 14 14 2 9 Sept. 15 15 2 4 17 Sept. i ley i! 4 8 24 Sept. 15 15 ip I. 7 Oct. 19 19 1 2; 14 Oct. ba ha 1 20 Oct. 24 24 5 3 15 Nov. 22, 22, 2 5 1 Low captures due to inclement weather. On one occasion, a complete courting sequence was observed. About 1416, a female alighted on a leaf, wings outspread ca. 1 m from a perched male, who immediately flew around her several times, then alighted and, with wings moving slowly up and down, walked to a face to face position. There they remained for ca. 30 sec. Then, the female walked around to the underside of the leaf, the male followed, and copulation was initiated. This lasted for ca. 22 min. On another occasion, a pair discovered in copula under a leaf remained so for 8 min before breaking off. Sexual activity continues until late in life. On 15 November 1973, a male that had been marked three weeks previously was found in copula with a freshly emerged female. Finally, mating appears to be done by older territorial males since, of the three cases observed, two involved males that had been previously marked. POPULATION SIZE AND MorTALITY Tables 1 and 2 show the basic marking and recapture data gathered during the study, after Sheppard & Bishop (1973). It is instructive to note that few individuals were captured more than once, which indi- cates that collecting was not very efficient over time. Low captures on 14 October were due to rainy weather. The survival rate (Fisher & Ford, 1947) was 0.45 per week or ca. 0.91 per day. The average life span was 1.82 weeks by the formula E = il , which as- 1 — survival rate sumes that all deaths occur just before sampling. Potential survival is up to 5 weeks in the field, as confirmed by a recaptured female. Ob- served field survival for males is up to 4 weeks. When the Lincoln index modified by Bailey (1952) is applied to the study data, the weekly 42 JOURNAL OF THE LEPIDOPTERISTS SOCIETY TABLE 3. Population composition over time of new and recaptured butterflies, considering all animals captured and recaptured before and after a determined date to constitute part of the population even though they were not captured on that date. Date Total New % Recaptures % 18 Aug. DD, 22, 100 —- - 1 Sept. 17 WP, WL 5 29 9 Sept. 20 9 45 We 55 17 Sept. 19 4 DAML 116) 79 24 Sept. 16 13 81 3 19 7 Oct. 23 19 82 A 18 14 Oct. 11 6 54 5 46 20 Oct. DB) 16 64 9 36 15 Nov. 22, Ike 68 Tf ol Average 19.4 10.4 = WA! = Standard Error +22.6% +49.2% _ +42.4% ~ estimates of total population size show great variation as a result of the large differences in recaptures from one sampling to the next. These fluctuations were felt to be more due to inaccessibility of individuals because of the thick brush on either side of the trail than to changes in the population level. Therefore, to make the capture data more realistic a third table was mounted assuming that those animals captured in week one and recaptured in week three, for example, formed part of the population in week two even though they were not captured at that time. In Table 1 we see that on 18 August, 22 butterflies were marked and released of which five were later recaptured: two on 1 September, two others on the 9th, and one on 19 September. On the Ist, at least three butterflies were not captured. Therefore, rather than a total of 14 for that date, we have 17 which we know formed part of the population at that time: 12 new plus 2 recaptured plus 3 which were recaptured at later dates. The numbers thus derived were entered in Table 3. The procedure was conducted for each capture period during the study. This results in considerably smoother total capture figures, which show that if allowance is made for inefficiency in collecting, the esti- mates of numbers of individuals in the study area was quite stable over the 14-week period (Table 3) with a mean of 19.4 individuals and a standard error of 4.4 or + 22.6%. More consistent results can be ob- tained by eliminating data for 14 October. When efforts were made to capture all the M. felsina in the study area—22 July 1973 and 6 June 1974—21 and 22 individuals were recorded, respectively, which again suggests that the number of M. felsina in the study area remains stable over long periods with a low number of individuals. VoLUME 32, NUMBER 1 43 Sex ratio data gathered during the study were unsatisfactory as a result of the similar appearance and behavior of males and females, except during oviposition and territorial displays. Because of the delicate nature of these butterflies, they had to be kept in the net during marking and afterwards immediately released. This prevented detailed examina- tion. Females, however, were a small minority of all captured. On the two occasions referred to above, 19 males were captured each time with 2 and 3 females, respectively. The reason is that the females entered the study area sporadically only for mating and feeding, thus not being as accessible as the males. The number of females caught under these circumstances is not truly representative of the female population; thus, for the purpose of this study, males and females were considered together. DIscUSSION AND CONCLUSIONS As noted above, tropical butterfly populations, especially those of theclids and riodinids, are characterized by low intensive and extensive frequencies. The data from the marking-recapture program and other observations on the M. felsina population permits a number of sugges- tions as to why this is so. ‘Low Intensive Frequency Low intensive frequency means that an animal is represented by a small number of individuals in a given population. As shown by the recapture data, the number of M. felsina frequenting the study area remained low and fairly constant for long periods. Other students have made similar observations on neotropical butterfly populations. Benson & Emmel (1973) observed a roost of Marpesia berania (Hewitson) in Costa Rica that maintained constant equilibrium of population size for more than 3 months because of constant rates of recruitment and mortality. Ehrlich & Gilbert (1973) recorded a similar structure for Heliconius ethilla Godt in Trinidad over a period of 2 years, as did Young (1972) for Siproeta epaphus (Latreille) in Costa Rica. Why do these populations have an equilibrium level and what is the mechanism that enables them to maintain it? Ehrlich & Gilbert (1973) explain the constant population level in H. ethilla through the constant recruitment of new adults over time which equals mortality. This was thought to be due to unvarying predator pressure on the immatures and not be- cause of a lack of foodplant, which was quite common in the habitat. The other factor was limited adult nectar resources, which controlled egg production, thereby regulating the production of immatures. Young 44 JOURNAL OF THE LEPIDOPTERISTS SOCIETY (1972) in his study of S. epaphus likewise found that egg and adult numbers remained constant throughout the study period. As a mecha- nism, he suggested low fecundity and low adult vagility as well as the sheltered nature of the forest understory community. Both studies dis- count outmigration as a factor. My observations on M. felsina allow the following conclusions to be made. First, neither foodplant nor nectar availability appeared to be a limiting factor. The larval damage was always small compared with the amount of foodplant available, showing little pressure on foodplant resources. The adult’s preference for nearly anything that is in bloom and its willingness to travel good distances eliminates lack of adult nectar sources as a possibility. The factors that appear to be most significant are low egg-laying frequency, increased longevity, and male spacing. Low egg-laying frequency was observed on several occasions. The maximum number of eggs seen laid in any one afternoon by the same female was four, these being laid singly and widely dispersed on the foodplant (Callaghan, 1977). The results of this were shown in the low numbers of widely separated larvae in all instars that could be found on the same foodplant. Pupation and emergence were likewise staggered, meaning a fairly even flow of adults over time. Since the larvae were attended and protected by ants (Callaghan, 1977), preda- tion was kept down to a minimum and thus did not appear to be a significant factor as it was in the case of H. ethilla (Ehrlich & Gilbert, 1973). For M. felsina, then, the low frequency of oviposition assures the low intensive frequency of its populations. Why egg production should be low could not be determined during the course of the present study. Another factor is longevity. The 1.82 weeks observed for M. felsina is high when compared with holarctic butterflies, but low with respect to those tropical nymphalines that have been studied. Scott (1974) reported an average life span of 4.2 days of Lycaena arota Bois with a potential of 8 days. Labine (1968), Turner (1971), and Cook e¢ al. (1971) reported the life expectancy after marking for 6 holarctic species in the field as from 2.8-12.1 days. Powell (1968) gave a maximum of 16 days for Incisalia iroides (Boisduval). On the other hand, Benson & Emmel (1973) demonstrated that the neotropical nymphaline Marpesia berania has an average longevity of 43.9 days with a potential of at least 157. Heliconids are especially long lived. Benson (1972) reported Heliconius erato petiverana Doubleday in Costa Rica with average observed longevities of 52 days and an individual alive 6 months after VOLUME 32, NUMBER 1 45 marking. Turner (1971) and Ehrlich & Gilbert (1973) reported similar results. Greater longevity ensures low intensive frequency and is of considerable selective value in the tropics since it enables the wide- spread dispersal of eggs in both space and time, which may diminish parasitism and ensure a larger number of offspring reaching maturity (Benson & Emmel, 1973). Also, having individuals in all stages of development would permit survival of a natural disaster that might eliminate one stage but not the others. The causes of adult mortality of M. felsina are not precisely known, since no actual predation or other forms of natural mortality were observed during the study. However, potential predation exists in the form of lizards, spiders, ants, and birds. Rapid flight and hiding beneath leaves are two methods used by adult M. felsina and many other riodinids to avoid predation. On the other hand, its fairly sedentary habits might mean greater predator pressure and lower survival rate for M. felsina than for heliconines and nympha- lines, which enjoy mimicry, distastefulness, and/or strong flight. Lower survival would be more likely true for males because of their conspicu- ous behavior. Finally, male spacing seems to select for longevity, since older males have been observed to be more aggressive and fixed to their habitual areas and thus are easily able to drive off younger, more inexperienced newcomers. The result is that females appearing at the rendezvous area will be more likely to mate with strong, long-lived males. This is turn assures longer-lived offspring, which can effectively distribute their eggs widely in space and time. These three factors combined, then, might provide for the perpetuation of the low intensive frequencies observed in the M. felsina population. Although to date other forest riodinids have not been studied in such detail, I suspect that their low intensive frequencies may be explained on much the same basis. Low Extensive Frequencies Low extensive frequency means that populations are found rarely within a given faunistic region. This is the case with M. felsina as well as other forest riodinids. The key factor here in the case of M. felsina is foodplant distribution. Individuals of Norantea brasiliensis Choisy (Marcgraviaceae), the M. felsina foodplant, are widely and sparingly distributed on the restinga, but this does not prevent their being visited by ovipositing females. There is a tendency on the part of the newly emerging butterflies to establish a colony at a suitable locality near the site of their foodplant. This was observed on several occasions, the most notable of which was 46 JOURNAL OF THE LEPIDOPTERISTS SOCIETY when two males were seen engaging in patrolling behavior on a small clump of bushes near a foodplant some 500 m from the main colony. These same two individuals were there one week later, a little bit worn but still recognizable. Before the next visit, they had disappeared and no others had taken their place. On the nearby foodplant, larvae were discovered in various instars. I did not observe any other case of actual colonization occurring on that particular clump of bushes. Undoubtedly, this attempt failed as a result of the rather harsh, unsheltered conditions in that particular section of the restinga. Females, then, are the col- onizers for M. felsina, traveling considerable distances many times in search of foodplants on which to lay their eggs. This phenomenon has been recorded indirectly with other riodinid butterflies. On numerous occasions a single butterfly will be encoun- tered at a particular place and time, whereas subsequent visits under comparable conditions will fail to turn up additional specimens. In the same woods as the M. felsina habitat, a single male Calydna lusca (Geyer) was captured on 18 July 1973. On 7 July 1972, a male and female Nymphidium lisimon attenuatum Stichel were taken and a single female Leucochimona philemon (Cramer) was taken on 7 July 1973. Before and since, 46 collecting days over 4 years have failed to reveal additional examples from this small wood. The best explanation is that the females are very vagile, always on the move searching for new foodplant localities. Similar conclusions have been reached by other students of tropical riodinids. Ebert (1969) mentions that little species such as riodinids “migrate continuously within a great area of favorable biotypes. . ...” Finally, the question arises concerning the barriers that female rio- dinids will cross in their movement. In the case of M. felsina, open flat areas do not appear to provide a serious obstacle, although this might be expected because of their preference for a low forest habitat. Ob- servations on deep forest riodinids are few but significant. On 19 January 1975, a lone female Nymphidium mantus (Cramer) was ob- served passing through dry secondary shrub near Linhares, Espirito Santo, Brazil, an area very different from its normal habitat in the deep forest near the edges of swamps. Water does not appear to be a sig- nificant barrier since many riodinid populations on either side of large rivers such as the Amazon are virtually indistinguishable (Callaghan, in prep. ). SUMMARY The results of the marking-recapture and observations of adult be- havior allow a number of conclusions to be drawn with respect to the VOLUME 32, NuMBER l AT population structure of the neotropical riodinid butterfly M. felsina. It was found that this butterfly exhibits the structure common to most forest butterflies, that of low intensive and extensive frequencies. The reasons for the former are low egg-laying frequency, longevity, and male spacing. The latter was explained by a combination of high female vagility and low intensive and extensive foodplant distribution, which lead to the establishment of new colonies by females at widely dispersed foodplant localities. Depending on conditions at these localities, such as parasitism and exposure to the weather, the colony may become extinct only later to be reestablished by another wandering female. It is sug- gested that similar population structures for other neotropical forest butterflies, particularly riodinids, may be explained on this same basis. ACKNOWLEDGMENTS I wish to thank Drs. Woodruff Benson and Keith Brown for making many helpful comments on the manuscript and the former especially for his helpful discussion and encouragement during the initial phases of the field work. To Dr. Alfredo de Rego Barros my thanks for use of the facilities of the Museu Nacional do Rio de Janeiro. LITERATURE CITED BartEy, N. T. J. 1952. Improvements in the interpretation of recapture data. J. Animal Ecol. 21: 120-127. Bates, H. W. 1864. The naturalist on the River Amazons. University of Cali- fornia Press (1962 reprint of the second edition, John Murray, London). 465 p. Benson, W. W. 1972. Natural selection for Miillerian mimicry in the Heliconius erato in Costa Rica. Science 176: 936-939. Benson, W. W. & T. C. Emmet. 1973. Demography of gregariously roosting populations of the nymphaline butterfly Marpesia berania in Costa Rica. Ecology 54: 326-335. CaLLAGHAN, C. J. 1977. Studies on restinga butterflies. I. The life cycle and immature biology of Meander felsina (Riodinidae). J. Lepid. Soc. 31: 173-182. Cook, L. M., K. Frank, & L. P. Brower. 1971. Experiments on the demography of tropical butterflies. I. Survival rate and density in two species of Parides. Biotropica 3: 17-20. Crane, J. 1955. Imaginal behavior of a Trinidad butterfly Heliconius erato hydara Hewitson, with special reference to the social use of color. Zoologica 40: 167-196. 1957. Imaginal behavior in butterflies of the family Heliconiidae: Chang- ing social patterns and irrelevant actions. Zoologica 42: 135-146. Esert, H. 1969. On the frequency of butterflies in eastern Brazil, with a list of the butterfly fauna of Pocos de Caldas, Minas Gerais. J. Lepid. Soc. 23: Suppl. 3, 48 p. EnruicH, P. R. & L. E. Gimperr. 1973. Population structure and dynamics of the tropical butterfly Heliconius ethilla. Biotropica 5: 69-82. 48 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Fiser, R. A. & E. B. Forp. 1947. The spread of a gene in natural conditions in a colony of the moth Panaxia dominula (L.). Heredity 1: 143-174. LasineE, P. A. 1968. The population biology of Euphydryas editha. VIII. Ovi- position and its relation to patterns of oviposition in other butterflies. Evolu- tion 22: 799-805. MacNeri, C. D. 1964. The skippers of the genus Hesperia in western North America with special reference to California (Lepidoptera: Hesperiidae). Univ. California Publ. Ent. 35: 230 p. PowEL., J. A. 1968. A study of area occupation and mating behavior in Incisalia iroides (Lepidoptera: Lycaenidae). J. N. Y. Ent. Soc. 76: 47-57. Scott, J. A. 1974. Population biology and adult behavior of Lycaena arota (Lycaenidae). J. Lepid. Soc. 28: 64-72. Seitz, A. 1913. Macrolepidoptera of the World. Vol. 5. The American Rhopaloc- era. Alfred Keren Verlag, Stuttgart. 1143 p. SHEPPARD, P. M. & J. A. BisHop. 1973 (1974). The study of populations of Lepidoptera by capture-recapture methods. J. Res. Lepid. 12: 135-144. Turner, J. R. G. 1971. Experiments on the demography of tropical butterflies. II. Longevity and home-range behaviour in Heliconius erato. Biotropica 3: 21-31. Younc, A. M. 1972. The ecology and ethology of the tropical nymphaline but- terfly, Victorina ephaphus. I. Life cycle and natural history. J. Lepid. Soc. 26: 155-170. Journal of the Lepidopterists’ Society 32(1), 1978, 48 THE INTERNATIONAL CODE OF ZOOLOGICAL NOMENCLATURE The draft third edition of the International Code of Zoological Nomenclature is now available for comment by zoologists. Copies may be obtained (price £2.50 surface mail, £5.00 air mail) from the Publications Officer, International Trust for Zoological Nomenclature, c/o British Museum (Natural History ), Cromwell Road, London SW7 5BD, U.K. Comments should be sent as soon as possible, and in any case before 30 November 1978, to the Secretary, International Commission on Zoological Nomenclature, at the above address. A paper explaining the major changes proposed by the Commission’s Editorial Committee to the existing Code has been published in the Bulletin of Zoological Nomenclature, vol. 34, part 3. Copies may be obtained (price 50p) from the same address as copies of the draft Code. Journal of the Lepidopterists’ Society 32(1), 1978, 49-54 AN ANALYSIS OF THE HELIOTHIDINE TYPES (NOCTUIDAE) OF HERMAN STRECKER WITH LECTOTYPE DESIGNATIONS D. F. HAarpwIick Biosystematics Research Institute, Ottawa, Canada ABSTRACT. The authenticity of the nominal type specimens of species of Heliothidinae described by Herman Strecker is evaluated. A number of the nom- inal type specimens are judged to be spurious. Lectotypes for the following Strecker species are selected: Schinia approximata, Schinia dolosa, Heliothis fastidiosa, Schinia labe, Schinia lora, and Schinia pyraloides. In anticipation of future revisionary work on the Heliothidinae I took the opportunity in October of 1976 to examine Herman Strecker’s type material belonging to this subfamily at the Field Museum of Natural History in Chicago. Strecker’s species names have always presented a problem to Lepidopterists, firstly because his original descriptions were often very brief, and secondly because he evidently had the habit of augmenting or replacing his original type series. Thus, although Heliothis regia was described from a single specimen, there are now six specimens each labelled in his hand as “type” of regia. In the earlier years of his career, Strecker evidently had no type con- cept, or at least a very nebulous one. As a result, many of the specimens on which he based his early original descriptions must have been either destroyed or misplaced. In later years, however, with increasing aware- ness of the value of type specimens, Strecker presumably tried to rectify his earlier laxity by labelling specimens other than the “originals” as his types. If such substitutions can be demonstrated, then obviously the spurious types have no status under the “Rules.” Nevertheless such pseudotypes do have value in indicating Strecker’s concept of his species in, the maturity of his later years, and should be considered in any subsequent neotype selection procedures if these are found to be necessary. At the present time the Strecker Collection is housed as a separate entity within the collections of the Field Museum of Natural History, and Strecker’s arrangement of species and his hand-printed labels have been retained. In my discussion of type specimens which follows, the species names are arranged alphabetically. Rhododipsa aden Strecker Strecker, 1898, p. 11. The original description of aden was based on a single specimen. The male labelled as “original type” in the Strecker collection matches the original descrip- 50 JOURNAL OF THE LEPIDOPTERISTS SOCIETY tion well and is evidently the one on which Strecker based his description. It is labelled as follows: “Col.”; “384”; “S. Aden, Orig. Type’. There is a major piece of the left hind wing broken off and the anal angle of the right front wing is missing. A genitalic slide (no. FM Hel 1) has been prepared from the holotype. Schinia approximata Strecker Strecker, 1898, p. 10. This species was described on the basis of three specimens collected by Boll near Dallas, Texas. The three females labelled as “original types” match the orig- inal description well and are assumed to be authentic. Because of its superior condition the specimen numbered 76 is hereby selected as lectotype; it is labelled as follows: “76”, “S. approximata, 374, Orig. Types”. A genitalic slide (no. FM Hel 2) has been prepared from the lectotype. Schinia ar Strecker Strecker, 1898, p. 10. The single male in the Strecker Collection labelled as “original type” matches the original description well, and is evidently the one on which the name was based. The specimen is labelled as follows: “371”; “S. ar, 371, Orig. Type”. A genitalic slide (no. FM Hel 3) has been prepared from the holotype which is in excellent condition. Schinia dolosa Strecker Strecker, 1898, p. 9. The original description was based on two specimens taken near San Antonio by Boll. The two males in the Strecker Collection labelled as “original types” are evidently authentic. I hereby select the slightly larger specimen as lectotype; it is labelled as follows: “tex”; “S. Dolosa, Orig. Type”. A genitalic slide (no. FM Hel 3) has been prepared from the specimen. Heliothis fastidiosa Strecker Strecker, 1876, p. 121. The original description of fastidiosa was based upon two specimens collected by Boll in Texas. The two males in the Strecker Collection match the original description well and undoubtedly represent the specimens on which it was based. I hereby select the smaller specimen bearing the individual “31” label as lectotype. The specimen is labelled as follows: “31”; “S. Fastidiosa, 31, Orig. Types”. A genitalic slide (no. FM Hel 5) has been prepared from the lectotype. Heliothis gloriosa Strecker Strecker, 1877, p. 132. A single specimen in the Strecker Collection is labelled as “original type” and this is evidently the one on which the original description was based. The specimen, a female, is in excellent condition except for lacking a portion of the right antenna. It expands 1%,” and is labelled as follows: “18”; “S. gloriosa Orig. Type”. A genitalic slide (no. FM Hel 6) has been prepared from the holotype. Schinia hanga Strecker Strecker, 1898, p. 9. The species was described on the basis of one specimen collected by Boll at Dallas, Texas. The male in the Strecker Collection labelled as “original type” is VOLUME 32, NUMBER l 5li evidently this specimen. It expands 144” and is labelled as follows: “70”; “393”; “S. Hanga, Orig. Type”. A genitalic slide (no. FM Hel 7) has been prepared from the holotype, which is in excellent condition. Heliothis imperspicua Strecker Strecker, 1876, p. 122. The original description of imperspicua was based upon a single specimen, bear- ing the number 53, which was collected in Texas by Boll. There are two specimens in the Strecker Collection each labelled as “original type” but these are evidently both spurious. One specimen is labelled as having been collected in Colorado; the other is without a locality label and bears the number “49”. Neither specimen differs from the rather generalized original description in any striking detail. The true type of imperspicua must be presumed lost. Heliothis inclara Strecker Strecker, 1876, p. 122. The original description was evidently based upon a single specimen collected by Boll in Texas, which was numbered 46. There are now two specimens in the Strecker Collection labelled as “original types”. One of these bears the number 78 and is considerably smaller than the specimen cited in the original description. The other specimen is without collection number but corresponds well with the original description and may be the true type. Schinia labe Strecker Strecker, 1898, p. 10. The original description of labe was based upon two specimens collected at Dallas, Texas by Boll. The two specimens in the Strecker Collection labelled as “original types” are apparently these. I hereby select the smallest of the two, which bears a separate “372” label, as lectotype. The lectotype is a male ex- panding slightly less than 34” and is labelled as follows: “372”; “S. Labe, 372, Orig. Types’. A genitalic slide (FM Hel 10) has been prepared from the specimen. Heliothis lanul Strecker Strecker, 1877, p. 132. There is a single male in the Strecker Collection labelled as “original type” and this is evidently the specimen on which the original description was based. There is no locality data indicated in the original description nor on the specimen. The holotype is labeled as follows: “85”; “S. Lanul, 85., Orig. Type”. A genitalic slide (no. FM Hel 11) has been prepared from the type. Schinia lora Strecker Strecker, 1898, p. 10. The original description of lora was based on three specimens, two from Boll collected near Dallas, Texas, and one from Heiligbrodt at Bastrop, Texas. Only two specimens in the Strecker Collection are labelled as “original types” and these are apparently authentic. There is another, unlabelled specimen in the collection which may represent the third specimen of the type series. Of the two specimens labelled as “original types” I hereby select the specimen with the separate “73” label as lectotype. The lectotype is a male in generally good condition which bears the following labels “73”; “373”; “S. Lora, 373, Orig. Types”. A genitalic slide (no. FM Hel 12) has been prepared from the lectotype. 52 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Schinia neglecta Strecker Strecker, 1898, p. 10. The single specimen labelled as “original type” in the Strecker Collection matches the original description well and is evidently the one on which the name neglecta was based. According to the original description the holotype was collected at Loveland, Colorado. The specimen is a female, expands 1”, and bears the following labels: “Col.”; “377”; “S. Neglecta 377., Orig. Type”. A genitalic slide (no. FM Hel 14) has been prepared from the holotype. Heliothis nubila Strecker Strecker, 1876, p. 122. Strecker’s original description of nubila was evidently based on a single specimen taken in Texas by Boll (number 48). There are two specimens in the Strecker Collection labelled as “original types”. Neither of these matches the original de- scription very well, there being no red shading on the underside of the wings, and both are numbered “72” rather than “48”. I consider these specimens to be spurious; the three types must be presumed lost. Schinia obscurata Strecker Strecker, 1898, p. 10. The single specimen labelled as original type of obscurata in the Strecker Collec- tion is obviously the one on which the original description was based. The holotype is in good condition except for having a notch in the left forewing. The female specimen is labelled as follows: “St. Vincent, Pa.”; “378”; “S. Obscurata, 378, Orig. Type’. A genitalic slide (no. FM Hel 15) has been prepared from the specimen. Schinia pyraloides Strecker Strecker, 1898, p. 9. The four specimens on which the original description of pyraloides was based are in the Strecker Collection and labelled as “original types”. The type series was taken at Glenwood Springs, Colorado by Bruce. I hereby select the male specimen with the individual “Col”. label as lectotype. The specimen is in generally good condition except for lacking most of the left antenna and having a slit in the right hind wing. The lectotype is labelled as follows: “Col.”; “S. Pyraloides, Orig. Type, Colorado”. A genitalic slide (no. FM Hel 16) has been prepared from the specimen. Heliothis regia Strecker Strecker, 1876, p. 121. So far as can be determined from the original description, the name regia was based upon a single specimen. There are, however, six specimens in the Strecker Collection labelled as “type”. According to the original description the holotype was taken in Texas by Boll but none of the six nominal “types” bears a locality label. I have compared each of the specimens with the original description and one female matches it very well, and I construe this to be the true type. It is labelled as follows: “S. Regia, Type”. A genitalic slide (no. FM Hel 17) has been prepared from the holotype. Heliothis rubiginosa Strecker Strecker, 1876, p. 122. Strecker’s description of rubiginosa was evidently based upon a single specimen taken in Texas by Boll; there are now six specimens, each labelled as “Type” in the VOLUME 32, NUMBER 1 53 Strecker Collection. None of these, however, bear the “50” label mentioned in the original description. One of them, a male, matches the original description well and may represent the true type but the evidence is insufficient to make a definitive judgement. Heliothis siren Strecker Strecker, 1876, p. 122. Strecker’s original description of siren was evidently based on a single specimen collected by Boll in Texas. There are now two specimens in the Strecker Collection labelled as “original type” but neither of these bears the “45” label mentioned in the original description. One of the two is without number label and the other bears an “80” label. The unnumbered specimen matches the original description well and may represent the true type but there is no way of establishing this with certainty. Heliothis spectanda Strecker Strecker, 1876, p. 122. The original description of spectanda was based upon a single specimen taken in Texas by Boll. When the Strecker Collection was examined, no specimen labelled as type of spectanda was found. There was, however, a specimen in the series of Heliothis virescens bearing the number “52” which was cited in the original de- scription as belonging to the type. The specimen matches the original description very well and I construe it to be the holotype. A genitalic slide (no. FM Hel 20) has been prepared from the specimen which is a female. Heliothis sulmala Strecker Strecker, 1878, p. 1862. The original description of sulmala was based upon a single male taken at Pagosa Springs (Colorado). Strecker evidently mislaid the specimen because it was found in 1939 in a drawer of miscellaneous moths; it lacks Strecker’s charac- teristic type label. The specimen matches the original description very well, how- ever, and I construe it to be the holotype. The specimen is labelled as follows: “Heliothis Sulmala Streck., Pagosa Springs Col., (Orig. Type), McCauley, Found (1939) in a drawer with misc. moths.” A genitalic slide (no. FM Hel 21) has been prepared from the holotype. Schinia tanena Strecker Strecker, 1898, p. 10. Strecker described tanena on the basis of a single specimen taken at Bastrop, Texas by Heiligbrodt. There is a single male in the Strecker Collection labelled as “type” and there is no reason to doubt its authenticity. The specimen is labelled as follows: “tex”; “380”; “S Tanena, 380, Orig. Type”. A genitalic slide (no. FM Hel 22) has been prepared from the holotype. Schinia ultima Strecker Strecker, 1876, p. 122. The original description of ultima was evidently based upon a single specimen taken in Texas by Boll. There are two specimens labelled as “original type” in the Strecker Collection, but both specimens bear the number “71” rather than the “49” indicated in the original description and both specimens differ from the original description in several details. I do not consider either specimen to represent the holotype, and the latter must be presumed lost. 54 JoURNAL OF THE LEPIDOPTERISTS SOCIETY ACKNOWLEDGMENTS I appreciate the cordial co-operation of Dr. Rupert Wenzel, Chairman of the Department of Zoology, and Dr. Eric Smith of the Division of Insects, during my visit to the Field Museum of Natural History. LITERATURE CITED SrrRECKER, H. 1876. Lepidoptera, Rhopaloceres and Heteroceres indigenous and exotic. Number 13. Reading, Pa. P. 109-124. 1877. Id. Number 14. P. 125-134. . 1878. Lepidoptera in Annual report of the engineers for 1878. Appendix SS. Washington, D.C. P. 1750-1865. . 1898. Lepidoptera, Rhopaloceres and Heteroceres, indigenous and exotic. Supplement 1. Reading, Pa. P. 1-12, pl. 1, 2. Journal of the Lepidopterists’ Society 32(1), 1978, 54 Letter to the editor: A Comment on Monarchs and a “Tragedy of the Commons” in Science When the paper by Urquhart & Urquhart appeared in this journal (1976, Vol. 30: 153-158), I sat down and wrote a letter criticizing the editorial policy of allowing an observation to be published without providing sufficient information to allow verification by other biologists working with Lepidoptera. While I shared the fear that publication of the exact locale of the Mexican roost would possibly endanger it, I felt that the authors should have at the very least volunteered to disclose the site to responsible qualified scientists researching monarch biology. Subsequent events have made me regret not sending in my original comment. Incredibly, a scientist of international reputation, Lincoln Brower, was denied the locality information by Professor Urquhart. I do not consider such secrecy to be in the spirit of modern science, nor necessary in this particular instance. Anyone familiar with Brower’s body of work on the monarch would not question his scientific stature. Anyone who has seen his environmental film on the Connecti- cut River System cannot doubt his sensitivity to ecological problems. We all respect the effort that Professor Urquhart has put into studying monarch migration. That does not, however, give him territorial rights over monarch roosting areas or free him from the scientific pesponsthality of allowing other scientists to verify his results. Much of the controversy and ill will which apparently has followed L. Brower’s independent visit to the Mexican monarch roosting area might have been avoided had the study of the monarch proceeded as unselfish science rather than a race for glory in glossy magazines. In the future I would hope that this journal will insist that authors be willing to disclose their study sites to responsible colleagues. LAWRENCE E. GILBERT Department of Zoology University of Texas Austin, Texas 78712 Journal of the Lepidopterists’ Society 32(1), 1978, 55-56 ATOPOTHOURES A. BLANCHARD: A SYNONYM OF GOYA RAGONOT (PYRALIDAE) A. BLANCHARD P.O. Box 20304, Houston, Texas 77025 ABSTRACT. Atopothoures ovaliger A. Blanchard becomes Gaya ovaliger (A. Blanchard ), close to, but different from Goya stictella Hampson. Karan and Jay Shaffer, my wife and I went collecting, 17-24 May 1977, at the Welder Wildlife Foundation Refuge, near Sinton, Texas. Dr. Shaffer made a special effort to collect Peoriines and was well satisfied with the results of this trip. On their way back home the Shaffers spent an afternoon with us at Houston, so that he could examine my collection of Peoriines. This is when he discovered that what I had un- fortunately described as Atopothoures ovaliger (Blanchard, 1975) should have gone under the genus Goya Ragonot. The male genitalia of G. ovaliger are extremely close to those of G. stictella Hampson which is not too uncommon in Texas, but the two Figs. 1-5. Goya: 1-4, stictella: 1, male, Welder Wildlife Refuge, Sinton, San Patricio Co., Texas, 30 June 1975 (U.S.N.M.); 2, male genitalia of same (slide A.B. 3828); 3, male genitalia of another male, same location, same date, (slide A.B. 3827; 4, same enlarged to show gnathos. 5, ovaliger, El] Rancho Cima, Hays & Comal cos., Texas, 29 August 1975, slide A.B. 3826 enlarged to show gnathos. 56 JoURNAL OF THE LEPIDOPTERISTS SOCIETY species are definitely distinct. The habitus of G. ovaliger (Blanchard, 1975, Figs. 1-4) is quite different from that of G. stictella (this paper, Fig. 1 and Shaffer, 1968, Fig. 23). The differences between their male genitalia are not so obvious. Fig. 2 shows the genitalia of G. stictella prepared in the conventional manner. In Fig. 3 they are prepared in the manner favored by Shaffer (1968, page 3). Figs. 4 (stictella) and 5 (ovaliger) show the enlarged gnathos and the webs or ribs which sup- port its apical process from beneath; this is where the most obvious difference between the two species is to be found. Dr. Shaffer, who had the opportunity to look at many more specimens than I had, gave me the following information: “These ribs are provided in ovaliger with a double row of teeth (two or three to six in each row). In stictella the number of teeth per row varies from zero to two. Counting is com- plicated by the fact that in both species the size of the teeth varies from large and well developed ones to tiny, barely discernible nubbins.” LITERATURE CITED Biancuarp, A. 1975. A new phycitine genus and species (Pyraloidea). J. Lepid. Soc. 29: 95-97. SHAFFER, J. C. 1968. A revision of the Peoriinae and Anerastiinae (Auctorum) of America north of Mexico. Bull. U.S. Nat. Mus. 280, 124 p. Journal of the Lepidopterists’ Society 32(1), 1978, 56 PROTECTIVE BEHAVIOR IN AMPLYPTERUS GANNASCUS (SPHINGIDAE) During August 1974, I spent about two weeks collecting Lepidoptera on the grounds of the Inter-American Institute of Agricultural Sciences, approximately 45 km SE of San Jose, Costa Rica (near the town of Turrialba). On two separate occasions I witnessed an interesting behavioral response in the sphingid Amplypterus gannascus (Stoll), which was common in the area. A few gannascus would some- times remain resting high up on the whitewashed walls of the Institute buildings until about 1000, having been attracted to these sites by the lights on the buildings the night before. In two cases it was possible to touch individuals by means of tossing a multi-segmented net about twelve feet long at them. The individuals responded to being touched by releasing their grip and sailing slowly to the ground in a slow spiralling descent, with their wings held rigidly in a swept-back V position. Once on the ground the moths remained passive in spite of being nudged, and only attempted to escape after being seized by hand. The appearance of this behavior was strikingly similar to the appearance of a dead leaf wafting to the ground from a tree, and would seem to be a behavioral adaptation to escape predators by imitating an unappetizing plant fragment. Jerr Ross, Department of Museum Science, Texas Tech University, Lubbock, Texas 79403. Journal of the Lepidopterists’ Society 32(1), 1978, 57-58 OENEIS ALBERTA (SATYRIDAE) IN MONTANA Oeneis alberta Elwes has been taken to date in widely scattered colonies from Alberta, Manitoba and Saskatchewan (alberta), Colorado (oslari Skinner), Arizona (daura (Strecker) and New Mexico (capulinensis Brown). On 19 and 20 May 1976, a series of 21 males and 6 females of alberta was taken by the author from the high grasslands in the Little Snowy Mountains of central Montana, Fergus and Golden Valley counties. This is a new state record for the species in Montana. Habitat of the Golden Valley County colony is pictured in Fig. 1. Elevation is approximately 6500 feet. Specimens from the colony are shown in Fig. 2. The Montana colonies represent the nominate subspecies. Additional colonies of alberta will probably be discovered in Montana as suitable habitat in areas east of the Continental Divide is explored at the proper time of year. Colonies should also be expected to occur in Wyoming and Utah, though none have been found thus far. STEVE KoHLER, Montana Department of Natural Resources and Conservation, Division of Forestry, 2705 Spurgin Road, Missoula, Montana 59801. Fig. 1. Habitat of Oeneis alberta in the Little Snowy Mts., Golden Valley Co., Mont. 58 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Mig. 2. Oeneis alberta from the Little Snowy Mts., Golden Valley Co., Mont. (a & c) males, dorsal; (b & d) same, ventral; (e & g) females, dorsal; (f & h) same, ventral. All photos natural size. Journal of the Lepidopterists’ Society 32(1), 1978, 59-60 NEW OR INTERESTING LEPIDOPTERA RECORDS FROM WESTERN TEXAS The Panhandle and South Plains areas of Texas have probably received less close attention from lepidopterists than other areas of the state because of the dearth of resident collectors and the greater number of interesting species in other parts of Texas. In view of the relative lack of information on Panhandle-Plains species, it seems worthwhile to publish certain significant records from my collection and the Texas Tech University collection at this time. I report herein one new Texas record (Pieridae) and additional records of species not usually associated with these areas of Texas. PIERIDAE Kricogonia lyside (Godart) has previously been reported from the Panhandle- Plains area only in October (Kendall and Freeman 1963, The Butterflies and Skippers of Texas: A Tentative List, Sinton, Texas, 6 p.). The Texas Tech University col- lection contains two males of this species from Lubbock, Texas (Lubbock Co.), both of which are in very good condition. One was collected on 12 July 1970 by D. W. Kiser, and the other on 24 September 1967 by U. Barber. In addition to these specimens, I collected one male and two females in fair condition at Palo Duro Canyon State Park, Randall Co., Texas, on 6 May 1977. Phoebis agarithe maxima (Neumoegen). Although this species was not cited in Kendall and Freeman’s checklist as having been recorded from the Panhandle-Plains region of Texas, the Texas Tech collection contains two males, in fair condition, from Lubbock, Texas (Lubbock Co.). One was collected on 7 July 1967 by “E J W,” and the other on 16 September 1970 by P. M. Allen. I observed numer- ous males and females of agarithe maxima in Lubbock, Texas throughout September of 1976, and captured a single worn male on 19 September 1976. Pieris napi (Linnaeus). The Texas Tech collection contains a single perfect male specimen of an undetermined subspecies collected on 17 August 1970, at Canyon, Texas (Randall Co.) by Walt Fournier, a former Tech graduate student. As far as can be determined, this record is a new one for the state of Texas. LYCAENIDAE Lycaeides melissa melissa (Edwards). I collected a single perfect male of this species on 31 August 1975 at the Buffalo Springs Lake Recreation Area (4 mi. E Lubbock, Lubbock Co., Texas). This record tends to support the contention by Rickard and Vernon (1975, J. Lepid. Soc.: 150) ihat this heretofore rarely reported species has probably just been overlooked in the past. NYMPHALIDAE Chlosyne janais (Drury). A single female of this common neotropical species was collected by me at my residence in Lubbock, Texas, on 11 June 1977. The specimen has badly torn hindwings but is in fair condition otherwise. This species has not previously been reported from the Panhandle-Plains region of Texas. SATURNIIDAE Hemileuca hera hera (Warris). Although Douglas C. Ferguson states that hera is “widespread in the West but not known from Texas” (1972, Bombycoidea- Saturniidae in part, p. 106. in R. B. Dominick, et al., The Moths of America north of Mexico, Fascicle 20.2B), the Texas Tech collection contains a single male in very good condition, collected on 16 September 1969 in Dickens, Texas (Dickens Co.) by M. Hughes. Callosamia promethea (Drury). The Texas Tech collection contains one female, 60 JOURNAL OF THE LEPIDOPTERISTS SOCIETY in fair condition, collected on 2 September 1973 at Junction, Texas (Kimble Co.) by Tech graduate student Sandy M. Benbow. According to Ferguson (p. 235), promethea has not been cited previously as occurring west of Tyler, San Jacinto, and Montgomery counties in eastern Texas. Thus this record suggests a possible range extension of several hundred miles into the Edwards Plateau area. Texas Tech University students and faculty collect annually at a field campus in Junction, so it should be possible to determine if promethea is more than just a stray in the area. ACKNOWLEDGMENTS I wish to thank Roy O. Kendall of San Antonio, Texas, for reviewing this paper and confirming the identification of these specimens, and Dr. David E. Foster of the Texas Tech entomology faculty for allowing me to examine material in the Tech entomology collection. Jerr Ross, Department of Entomology, Texas Tech Uniwersity, Lubbock, Texas 79403. Journal of the Lepidopterists’ Society 32(1), 1978, 61-62 ERYNNIS BRIZO LACUSTRA AND HESPERIA COLUMBIA IN THE SIERRA NEVADA Burns (1964, U. C. Publ. Entomol. 37: 1-214) reports no records for Erynnis brizo lacustra Wright for the Sierra Nevada, and MacNeill (1964, U. C. Publ. Entomol. 35: 1-221) lists no records for Hesperia columbia Scudder from there except one female in the AMNH from “Sier. Nev.” Both are indicator species of the coast range serpentine belts north of San Francisco. Until recently, serpentine outcrops have been little collected in the western foothills of the Sierra Nevada of east-central California. Table 1 (next page) lists the new distribution records there. Sometimes the adults may fly a few miles from their serpentine areas to hilltop: e.g., both hilltop on Rocky Ridge, 1700-1900’, N. of Monticello Dam, Yolo Co., a non-serpentine area composed of Upper Cretaceous marine rocks of the Venado Formation. The nearest serpentine occurs in the extensive Mesozoic ultrabasic in- trusive rocks and the Franciscan Formation some 6 miles to the west. Similarly, Footman Ridge, Mariposa Co., is Paleozoic marine (also the area to the N & E), and to the south is Mesozoic granitic rocks, with no serpentine nearby. The nearest serpentine is found 5 mi. W. as Jurassic-Triassic metavolcanic rocks and 8 mi. SW near Mariposa as Mesozoic ultrabasic intrusive rocks. In the meadows, forests, and canyon immediately adjacent to Footman Ridge on the W & N, neither species has ever been collected. On 15 May 1970, E. slope Walker Ridge along Brim Grade, c. 1800’, SW of Leesville, Colusa-Lake Co. line, I noticed a female lacustra ovipositing on the terminal growth of a Quercus durata Jepson bush growing on serpentine soil along a roadbank, at 1125. Bums (1964) says “the skipper invariably occurs in direct association with QO. durata, a serpentine obligate” (see Whittaker et al., 1954, Ecology 35: 258-288). However, in some areas, it may also use Quercus dumosa Nutt. which hybridizes with Q. durata and grows in strictly non-serpentine soils (see Forde & Faris, 1962, Evolution 16: 338-347 ). Heretofore, these skippers were considered more coastal in their California distribution. OaxkLEy SHIELDS, Department of Entomology, University of California, Davis, California 95616. ‘Pu OJUST[eT)-YSlFpog uo JoURNAL OF THE LEPIDOPTERISTS SOCIETY 62 =— STTPMA vant GLOT URN TE = O00E Ve eOdeS) tle Ss sp Sry EO Er speus‘O 8 PT OLE APIN ST sjepsesief 70 AS =: SPleMisn© Vaan PLOT AFINES 009F ‘HUIUNS OSphy UeUl}OO 4 esodiieyy = DIquunjoo “HH SOMES PT OLET ABW ST SplPtys ‘O PT PLOT APIN £3 efepAesie[ FO HS — spyelys ‘O OF ONG PLOI ACW T 009F ‘}IUIUINS OSply UeU}00F esOdiivjy Dijsnovd) “q °F SIEM ©) 2 LE PLOT [dy FI oye sseq f° AN euluedies spperys ‘O OT ZL6I [May €Z 0S0Z “WIUIUUNS [[IE{ OULg opelogd [q viysnov) *q “FT wnyNY “YA FO APULOIA ‘doyry uo 1aATy =< SEES © PT OLGT APN T 00060 SOUICE SILOS) Sis ON) OMAMOLGE IGE RC I uInqny fo “WY “IOATY, UBOLIOUTY “YA e[PPrHN ‘ON JO VoUSNTFUOS JOY TU G4] ‘eg Ss lolepmnyyy UI¥.119} JUTZUOdIS aAoqe do.10yNo ur surddoy[iy sppetys ‘O Ori LLOI YEW 6I 98D JUO}JSOUTT] FO LUTEUS 1IdV[q Dvijzsngv) “gq “7 esp sunuedios pasodxe uo sutddo suits “Y ‘Ss = OL6T [Edy FZ [EYASe10 7 -[[Ey A[wourut09 «sug “YS — GJ61 ARIN 9G OOIP'O "IN ‘TU “pi ¢'g aoR[q Diysnov) *q °Z S}USULULOT) 10}D9T[OD x9S ‘ON a}yeq UOoTITIOD uoTVADT HA AP[VI0T yuno5 i sarads “BIULOFITCC) jo BPPAQN BIIIIS 94} uLOlt DIquin}od piladsa pue DLYSNID] OZ1LG swuuhly jo SplO09Y HOTNNQLOSICT “[ WTidgvy, Journal of the Lepidopterists’ Society 32(1), 1978, 63-64 BOOK REVIEWS THE BUTTERFLIES AND Morus oF HAMPSHIRE AND THE ISLE OF WicHT (being an account of the whole of the Lepidoptera) by B. Goater. 1974. E. W. Classey, Ltd., Faringdon, Oxon, England. xiv + 439 pp. Price: £6.50, postpaid. Perhaps nothing so authoritative as this book has been written on so small an areas fauna. The book is annotated with published records, manuscript notes and personal observations on all of the butterflies and moths of Hampshire and the Isle of Wight since the beginning of collecting there. Make no mistake, this book is an historical document, and as a record of what is (and was) in the County, and on the island, it is invaluable. Goater has drawn records from many contemporary sources, and there are some prominent English entomological contributors to the list: such as D. W. Ffennell, John Heath, E. C. Pelham-Clinton, and the Baron C. G. M. de Worms, names well-known in English lepidopterology. This information provides a regional list unlike any we have seen and the coverage is complete through 1972. Despite the lovely picture of Argynnis paphia on the cover, this is no “coffee table” book. It sticks strictly to business, and those looking for pretty pictures by which to identify British Lepidoptera should be forewarned to stay away from it. As a book of information (isn’t that what we really need? ), it is superb, and from it one can discover when, where and at what time any species of butterfly or moth has been captured in that area, and, if it has been reared, on what foodplant. I suspect that in this alone the book has fulfilled its purpose, and, additionally, it should stimulate the collector in the area to “fill in the blanks”. The nomenclature used is standard perhaps only to the British, since it is derived from the Kloet and Hincks Checklist of British Insects, part 2, Lepidoptera, 1972. In this treatment, the Hesperioidea and Papilionoidea directly follow the Ptero- phoroidea and precede the Bombycoidea. To a North American rhopalocerist this arrangement will seem strange, even incomprehensible, since there are few other classifications that follow this one. Most schemes place the butterflies and skippers above the Noctuidae, the “top” family in Goater’s system. If you are interested in the butterflies, by the way, look on pp. 214-245. The sphingids may be found on pp. 307-312, and the saturiid (there is only one) on p. 248, while Catocala are on pp. 404-406. This gives a bit of a “road map” to the reader just trying the book for the first time (I confess to a great deal of initial confusion ). As stated before, don’t buy this book on the basis of the pretty picture on its cover. Neither is this an identification manual. But if you are interested in a superb compendium of what is known about a limited fauna, by all means get the volume. It sets a fine standard, despite a few typographical errors not alluded to here (they happen to everyone! ), for future lists on small faunas. Lee D. Miter, Allyn Museum of Entomology, 3701 Bay Shore Road, Sarasota, Florida 33580. BUTTERFLIES OF West MALAYSIA AND SINGAPORE by W. A. Fleming. 1975. E. W. Classey Ltd., Farington, Oxon, England. Vol. 1: vii-x+64 pp., pls. 1-54; vol. 2: vii-x + 92 pp., pls. 55-90. Price: £19.50, postpaid. This book, effective as it is, is something of an enigma. I find it impossible to rationalize making it in two volumes if the series is only to be sold as a whole, and not broken into separate volumes, if the buyer so desires. The text is identical in both volumes to page 15, so it is only in the plates and the parts following them that the two volumes differ. A little elementary arithmetic shows that if the volumes 64 JOURNAL OF THE LEPIDOPTERISTS SOCIETY were combined into a single one, there would be a very manageable and useful single book of four prefatory pages, 144 text pages and 90 plates. Surely this would have been a better plan. Maybe others will have the problem I did—I would refer to “the” book to identify a Malaysian butterfly, and I almost inevitably selected the wrong volume. The only excuse I can think of for publishing this work in two volumes is economic, and perhaps £19.50 for a single book would put off some buyers, but with the price of books what it is today, I doubt it. But enough of the complaining. There is a multitude of information in this book, even though the style is such that it takes some acclimation. The nomenclature is up-to-date and applied to the right insects. Species in which the illustrations are not enough for identification are characterized in the text, and in those instances where genitalic dissection is necessary for final determination, the fact is noted, even though the genitalia are not figured. If Fleming had included some biblio- graphic citations to these problem areas, and to the many included foodplants records, the book would have been more authoritative, and the space these refer- ences would have added could not have been that much. The illustrations, however, are where the books truly excel. All of the photographs of specimens illustrate the salient points well and facilitate the identification of the insects in question. Al] of the specimens used are not perfect; some are downright tatty, such as the illustrated female of S12, Lethe europa malaya Corbet on Plate 24, but these were the best specimens available in collecions, and the photographs mercifully have not been “prettied up”. The color fidelity is very high, and at least most of the specimens are fresh, rather than century-old museum relics. Iden- tification of even the difficult Malaysian lycaenids is facilitated by them, though of course it is not made simple—no book could achieve that! I particularly appreciated the accurate citation of the authors of various taxa, even though these names were not bracketed where appropriate. At long last, both of the Felders are cited as the authors of names proposed in the “Reise Novara’, not just a blanket “Felder”. This latter practice seems to have dated from “Seitz” where only Cajetan Felder was given credit for the descriptions in the work, even though the authors themselves cited “nobis” on every new name, rather than the singular “mihi”. On balance, this is an exceilent book, the foregoing criticisms notwithstanding, and one that is remarkably free of typographical errors. The text portions are per- haps a bit too abbreviated, and authority is not given for many statements. I personally would have preferred a single volume about the size of Corbett and Pendlebury’s The Butterflies of the Malay Peninsula, but the present book accom- plishes some things that the earlier authors could not: Fleming has made the identification of Malaysian butterflies considerably easier than before. No more can be asked of any author! If your interests lie in the butterflies of southeastern Asia, by all means buy this book. Lee D. Mitier, Allyn Museum of Entomology, 3701 Bay Shore Road, Sarasota, Florida 33580. EDITORIAL STAFF OF THE JOURNAL Austin P. Puatr, Editor Department of Biological Sciences University of Maryland Baltimore County, 5401 Wilkens Avenue Catonsville, Maryland 21228 U.S.A. Dovuctas C. Frercuson, Associate Editor THEODORE D. SARGENT, Associate Editor NOTICE TO CONTRIBUTORS Contributions to the Journal may deal with any aspect of the collection and study of Lepidoptera. Contributors should prepare manuscripts according to the following instructions. Abstract: A brief abstract should precede the text of all articles. Text: Manuscripts should be submitted in duplicate, and must be typewritten, entirely double-spaced, employing wide margins, on one side only of white, 8% x 11 inch paper. Titles should be explicit and descriptive of the article’s content, including the family name of the subject, but must be kept as short as possible. The first men- tion of a plant or animal in the text should include the full scientific name, with authors of zoological names. Insect measurements should be given in metric units; times should be given in terms of the 24-hour clock (e.g. 0930, not 9:30 AM). Underline only where italics are intended. References to footnotes should be num- bered consecutively, and the footnotes typed on a separate sheet. Literature Cited: References in the text of articles should be given as, Sheppard (1959) or (Sheppard, 1959, 196la, 1961b) and all must be listed alphabetically under the heading LrreRATuRE Crrep, in the following format: SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London. 209 p. 196la. Some contributions to population genetics resulting from the study of the Lepidoptera. Adv. Genet. 10: 165-216. In the case of general notes, references should be given in the text as, Sheppard (1961, Adv. Genet. 10: 165-216) or (Sheppard 1961, Sym. Roy. Entomol. Soc. London 1: 23-30). Illustrations: All photographs and drawings should be mounted on stiff, white backing, arranged in the desired format, allowing (with particular regard to lettering ) for reduction to their final width (usually 4% inches). Illustrations larger than 8% x 11 inches are not acceptable and should be reduced photographically to that size or smaller. The author’s name, figure numbers as cited in the text, and an indication of the article’s title should be printed on the back of each mounted plate, Figures, both line drawings and halftones (photographs), should be numbered consecutively in Arabic numerals. The term “plate” should not be employed. Figure legends must be typewritten, double-spaced, on a separate sheet (not attached to the illustrations), headed ExpLANATION OF FicuRES, with a separate paragraph devoted to each page of illustrations. Tables: Tables should be numbered consecutively in Arabic numerals. Headings for tables should not be capitalized. Tabular material should be kept to a minimum and must be typed on separate sheets, and placed following the main text, with the approximate desired position indicated in the text. Vertical rules should be avoided. Proofs: The edited manuscript and galley proofs will be mailed to the author for correction of printer’s errors. Excessive author’s changes at this time will be charged to authors at the rate of 75¢ per line. A purchase order for reprints will accompany the proofs. Correspondence: Address all matters relating to the Journal to the editor. Short manuscripts such as new state records, current events, and notices should be sent to the editor of the News: Jo Brewer, 257 Common Street, Dedham, Massachusetts 02026 U.S.A. ALLEN PRESS, INC. REUSED LAWRENCE, KANSAS Us he CONTENTS Types OF PARNASSIUS CLODIUS GALLATINUS (PAPILIONIDAE). Steve Bove oO Ses PS CON La aaa i 1 SPECIFICITY, GEOGRAPHIC DISTRIBUTIONS, AND FOODPLANT DIVERSITY IN Four CALLOPHRYS (MITOURA) (LYCAENIDAE). Kurt Johnson 3 FooppLANT, Hapirat, AND RANGE OF CELASTRINA EBENINA (LyY- CAENIDAE). Warren H. Wagner, Jr. and T. Lawrence Mellichamp 20 Strupies ON Restinca Butrerruises. IJ. NoTes ON THE POPULATION STRUCTURE OF MENANDER FELSINA (RiopINDAE). Curtis J. Callaghan, 2 37 AN ANALYSIS OF THE HELIOTHIDINE TyPEs (NocrumAE) OF HERMAN STRECKER WITH LEcToTyPE DesicNaATIoNs. D.F. Hardwick... 49 ATOPOTHOURES A. BLANCHARD: A SYNONYM OF GOYA RAGONOT (Pyratmar). A. Blanchard 0. ee 50 GENERAL NOTES Protective behavior in Amplypterus gannascus (Sphingidae). Jeff Robb 56 Oeneis alberta (Satyridae) in Montana. Steve Kohler __.. 57 New or interesting Lepidoptera records from Western Texas. Jeff Robb 59 Erynnis brizo lacustra and Hesperia columbia in the Sierra Nevada. Oakley Shields i000 ee 61 Noes) AND News 2220230 ies a 19, 48 Book (REVIEWS. i eT aa a 63 Volume 32 1978 Number 2 JOURNAL of the LEPIDOPTERISTS’ SOCIETY Published quarterly by THE LEPIDOPTERISTS’ SOCIETY Publié par LA SOCIETE DES LEPIDOPTERISTES Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 19 July 1978 THE LEPIDOPTERISTS’ SOCIETY EXECUTIVE COUNCIL J. W. TuwpeNn, President KENELM W. Puut.ip, Vice President I. F. B. Common, Ist Vice President JuLtian P. DoNnAHUE, Secretary Lionet Hiccrs, Vice President RONALD LEUSCHNER, Treasurer Members at large: F. S. CHEw R. A. ARNOLD J. F. EMMEL D. F. Harpwick E. D. CasHAaTT R. R. GATRELLE J. B. ZrEcLER R. E. STANFORD A, Po Pian The object of the Lepidopterists’ Society, which was formed in May, 1947 and formally constituted in December, 1950, is “to promote the science of lepidopterology in all its branches, . .. . to issue a periodical and other publications on Lepidoptera, to facilitate the exchange of specimens and ideas by both the professional worker and the amateur in the field; to secure cooperation in all measures” directed towards — these aims. Membership in the Society is open to all persons interested in the study of Lepidoptera. All members receive the Journal and the News of the Lepidopterists’ — Society. Institutions may subscribe to the Journal but may not become members. Prospective members should send to the Treasurer full dues for the current year, together with their full name, address, and special lepidopterological interests. In alternate years a list of members of the Society is issued, with addresses and special interests. There are four numbers in each volume of the Journal, scheduled for February, May, August and November, and six numbers of the News each year. Active members—annual dues $13.00 Student members—annual dues $10.00 Sustaining members—annual dues $20.00 Life members—single sum $250.00 Institutional subscriptions—annual $18.00 Send remittances, payable to The Lepidopterists’ Society, and address changes to: Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 U.S.A. Back issues of the Journal of the Lepidopterists’ Society, the Commemorative Volume, and recent issues of the NEWS are available from the Assistant Treasurer. The Journal is $13 per volume, the Commemorative Volume, $6; and the NEWS, $.25 per issue. Order: Mail to Charles V. Covell, Jr.. Memoirs Editor, Department of Biology, Uni- versity of Louisville, Louisville, KY 40208, U.S.A. The Lepidopterists’ Society is a non-profit, scientific organization. The known office of publication is 1041 New Hampshire St., Lawrence, Kansas 66044. Second class postage paid at Lawrence, Kansas, U.S.A. 66044. Cover illustration: Dasychira dorsipennata larva, dorsal and lateral views. From Fascicle 22.2, “Lymantriidae,” by Douglas C. Ferguson, in Moths of America North of Mexico. The drawing was done by E. R. Hodges, Scientific Illustrator, Department of Entomology, Smithsonian Institution. (Reproduced by permission of the author. ) J OURNAL OF Tue LeprpopreRists’ SOCIETY Volume 32 1978 Number 2 Journal of the Lepidopterists’ Society 32(2), 1978, 65-74 STUDIES ON THE INTERACTIONS OF MORPHO PELEIDES (MORPHIDAE) WITH LEGUMINOSAE ALLEN M. YouNG Invertebrate Division, Milwaukee Public Museum Milwaukee, Wisconsin 53233 ABSTRACT. The butterfly Morpho peleides Kollar is a widespread species throughout tropical America, exploiting several wild genera and species of Leguminosae as larval foodplants. Field studies show that this species feeds on a broad spectrum of wild legumes on a regional basis. This interaction was explored in the laboratory by rearing caterpillars on peanut plants and alfalfa, cultivated legumes. The life cycle is completed successfully on these artificial foodplants, but feeding on alfalfa taken from an expressway led to mass mortality of caterpillars. Apparently the alfalfa was contaminated from some environmental source. In the native habitats of this butterfly, the Leguminosae are both diverse and numerous locally. This suggests that the mo- nophagous feeding habit provides sufficient ecological flexibility for exploiting different genera and species of the family. This is sufficient to maintain breeding populations of M. peleides in secondary habitats. Forest-dwelling species of Morpho are predicted to be experiencing different types of selection pressures favoring polyphagous feeding. In the premontane tropical wet forest life zone (Tosi 1969) of north- eastern Costa Rica, a larval foodplant of the butterfly Morpho peleides Kollar (Lepidoptera: Morphidae) is the vine Machaerium aff. flori- bundum Benth. (Leguminosae). The vine and butterfly occur in stands of mixed primary and secondary tropical wet forest (Fig. 1). It is known that M. peleides utilizes several leguminous woody vines and trees as larval foodplants in Costa Rica (Young and Muyshondt 1973) and the species can be reared on commercially available peanut plants both in Costa Rica and Wisconsin (Young 1974). This present paper examines further the feeding habits of M. peleides larvae, using eggs obtained from a population in premontane tropical wet forest (rather than from montane forest, as in a previous study), and involves plants not studied previously (Young 1974). The results further support the assumption 66 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Mig. 1. Above: mixed primary-secondary tropical wet forest habitat of the butterfly Morpho peleides at Finca El Tigre, near La Virgen, Heredia Province (Sarapiqui region), Costa Rica. At the spot shown here, M. peleides and M. granadensis are microsympatric. Below: Machaerium aff. floribundum, a forest leguminous vine which is a larval food plant of M. peleides (and probably M. granadensis) in the wild. VOLUME 32, NUMBER 2 67 that larvae of M. peleides are monophagous feeders on many temperate and tropical genera of Leguminosae. METHODS A female of M. peleides was captured on bait of rotting bananas placed near the edge of a forest habitat (Fig. 1) on February 14, 1977 at “Finca El] Tigre,’ a farm adjacent to “Finca La Tirimbina,” a few km from La Virgen, Heredia Province (Sarapiqui region), Costa Rica (220 m elev.). She was placed in a clear plastic bag containing a fresh cutting of M. aff. floribundum, and within eight days had produced a total of 40 viable eggs. The female was then preserved, and the eggs were brought to Milwaukee, Wisconsin for rearing. The eggs began to hatch on February 25 in Costa Rica and by the time the morphos arrived in Wisconsin, they were all Ist instar. In Costa Rica these larvae were fed leaves of Dioclea wilsoni (Leguminosae) but they were switched to peanuts (Arachis hypogea L.—Leguminosae) upon arrival at the Milwaukee Public Museum. The larvae were kept on potted peanut plants placed in a covered glass tank in a laboratory. A growth light was kept over this rearing chamber. The rearing program in Wisconsin extended from March 3 through May 25, 1977 (the date of the late eclosion). Records were kept on body lengths and head capsule widths of all caterpillars. The sources of peanut plants used were (1) Olds Seeds from Madison, Wisconsin and (2) Crop Science Department of North Carolina State University (Raleigh). Near the end of the experiment (April 22), the foodplant was switched to alfalfa (Medicago sativa L—Leguminosae); at this time most of the larvae were in the late 4th instar. The alfalfa plants used were obtained from a farm in Waukesha County, Wisconsin. Later (May 2) the remaining 5th instar larvae (several had pupated) were fed alfalfa collected from the side of an expressway in downtown Milwaukee. Like the peanuts, the alfalfa plants were potted, but this time soil brought in with the plants from the field was used. One 4th instar caterpillar was offered a seedling of Erythrina crista-galli L. (Leguminosae) from Brazil. Records were kept on larval survival throughout the study. The adults obtained were kept for further examination. RESULTS Both young and older larvae of M. peleides ted successfully on peanut and alfalfa leaves in the laboratory, followed by normal eclosion (Fig. 2). In addition, at least the 4th and 5th instar larvae will feed on Erythrina. Although some caterpillars feed intermittently throughout the day, the 68 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Fig. 2. Right column: second and fourth instar catepillars of M. peleides on peanut leaves at the Milwaukee Public Museum; note eaten areas of leaf in first illustration. Leaf column: fifth instar caterpillar on peanut plant and freshly-enclosed adult clinging to empty pupa case (at Milwaukee Public Museum ). VOLUME 32, NUMBER 2 69 MEAN AND RANGE OF BODY LENGTH (MM) OF CATERPILLARS 20 3 . . 16 14 tt ae MAR. 8 MAY 2 (DAY 1) SUCCESSIVE DAYS OF OBSERVATION (DAY 38) Fig. 3. Growth and size (body length) patterns for Morpho caterpillars reared in the laboratory. The vertical lines give the range in body lengths. greatest amount of feeding occurred in the late afternoon and early morning (e.g., 16:00-19:30 hrs/C.S.T.). Fourth and 5th instar larvae rested on the rims of the pots containing the peanut plants, and they would crawl up the plants to feed. Younger ones rested on leaves and shoots. Survival both on peanuts and on “farm alfalfa” was 100%. However, larvae fed “expressway alfalfa” showed considerable mortality: between May 4 and May 12, the number of healthy caterpillars dropped from 32 to 13. Very shortly after being fed the expressway alfalfa, many died. Death was preceded by a drastic contraction of body length, and spasmic 70 JOURNAL OF THE LEPIDOPTERISTS SOCIETY TasLe 1. Head capsule size statistics for caterpillars of the tropical butterfly Morpho peleides. No. Mean head capsule Range of head Instar measured width (x = S.D.) capsule widths (mm ) 1 17 1.50 + 0.08 1.4-1.5 2, 34 2.19 + 0.09 2.0—2.3 3 315} BA ae Vpllc 3.3-3.8 4 oll 4,91 +0.11 A4.7-5.1 5 Ia 6.09 + 0.45 5.2-6.7 Note—data obtained from molted head capsules at the end of each instar [distorted (crushed) head capsules are not included in the calculations]. All head capsules were collected within 24 hours after each molt. waves of movements lengthwise. Afflicted caterpillars fell from the plants and wriggled on the bottom of the cage before dying. When attempts were made to replace them on the foodplant, they again lost hold and fell off. Their feces had a reddish-orange component that was quickly absorbed on paper toweling. A few died as prepupae. Initial signs of the affliction included a larva remaining stationary on the plant, with the anterior half of the body hanging off to one side. Such a state lasted a few days before the larva fell off the plant and died. As a result of this sudden and epidemic-like mortality, only those indi- viduals that had already pupated by May 2 survived to emerge as adults. A total of 12 adults, all males, were obtained. Thus, the mortality was 100% among the 19 individuals that were still larvae on May 4. Based on a sample of ten randomly chosen larvae, molting is not syn- chronous. The 5th instar is the longest, and despite changes in the — foodplant type, development proceeds without major accelerations or decelerations in the daily pattern of growth (Fig. 3). With the exception of the Ist instar, there is considerable variation in the body length and the magnitude of this variation is about the same for the four instars (Table 1). Excluding a small bias introduced by unequal sample sizes, the range of variation in head capsule width is very low for all instars (Table 1). Body length and head capsule width are used here as estimates of larval size. The total development time of 91 days is broken down as follows: (1) egg = 11 days; (2) caterpillar = 65 days; (3) pupa = 15 days. The right forewing length for the adult male butterflies ranged from 55 to 65 mm (N = 12), but the exclusion of two individuals reduced this range to 61-65 mm. DIscUSSION The data are useful for discussing (1) feeding behavior of the cater- pillars of M. peleides, and (2) apparent effects of environmental con- VoLUME 32, NUMBER 2 fol tamination of a foodplant of an exotic butterfly. The latter was an unexpected outcome of the study. Morpho accepts peanuts and other legumes, not used as foodplants in the wild, as discussed previously (Young 1974). But to these records I add alfalfa and Erythrina as acceptable foodplants of M. peleides cater- pillars in the laboratory. Erythrina is native to the New World tropics (Bailey 1969), although it is not known if it is a natural foodplant of Morpho. Greenhouse cultures of this plant are usually infested with herbivorous insects, suggesting few effective defenses operative against such attacks. Alfalfa, a near relative of peanut, is commonly cultivated, and occurs as a weed species along roadsides; it is native to the Old World. As a weed species, alfalfa may possess few defenses against herbivores as energy allocation is likely to be directed toward high repro- ductive potential (Lewontin 1965). Thus, cultivars such as peanut and alfalfa, weeds such as alfalfa, and ornamentals, such as Erythrina, are examples of leguminous plant species with few defenses against her- bivores, perhaps making them ideal to serve as food for Morpho larvae. It is not known if Morpho will oviposit on these plants in the laboratory. I observed previously that caterpillars of M. peleides are primarily “dawn-dusk” feeders in the wild (Young 1972a), and this is also true for laboratory cultures experiencing the Wisconsin dawn-dusk cycle (Young 1974; pers. obs.). Apparently the larvae are programmed with a rhythmicity for peak periods of feeding activity in both tropical and temperate situations. In the wild, 4th and 5th instar caterpillars rest on the trunks of the foodplant where they blend in with the background (Young 1972a), and this behavior explains why they rested on the rims of the pots containing the peanut plants. Using a larger sample size, Young (1974) estimated the total develop- mental time for M. peleides on peanuts to be about 105 days, or about 14 days longer than the estimate obtained in the present study. Body length of 5th instars in the previous study, was about 73 mm as com- pared to 70-71 mm in the present study. Several factors may be relevant here: (1) the eggs in the two studies came from different regions. Thus selection pressures could have been different in terms of effects on development time; (2) differences in the foodplant as related to time of the year, and other factors. The discrepancy in the development time is in the length of the larval stage; the egg and pupal stages are the same (Young 1974; pers. obs.). Thus, differences in the foodplant may be involved. Switching to alfalfa (not done in the previous study ) might have accelerated development. To test this, eggs will have to be reared entirely on this plant in a future study. The transfer to alfalfa was done —~l bo JOURNAL OF THE LEPIDOPTERISTS SOCIETY in the fifth instar, the time of greatest food intake. In the previous study, older peanut plants were used, and it may well be that older plants have resistant properties more expressed than younger ones. There is some evidence that the defense systems of peanut plants change with age: in the present study I observed that Morpho larvae refuse to eat the first set of leaves of a peanut seedling, eating only shoot and leaf tissue above this point (S. Borkin and A. Young, pers. obs.). The observed high level of mortality among 5th instar larvae on alfalfa plants taken from the downtown expressway may have been due to a contamination of these particular plants by an industrial or automobile residue. An exotic insect such as Morpho, when exposed to a con- taminated foodplant, may be expected to encounter such mortality. Since only male adults were reared I presume that all the females died as Sth instar larvae, since M. peleides has a sex ratio of unity (Young 1972b, 1973; Young and Muyshondt 1973). Female larval development generally takes longer than that of males. The observed inability of afflicted larvae to grasp the foodplant and feed was very likely due to the contaminant affecting the nervous system. It is not likely that a strain difference affecting feeding ability by morphos exists between the expressway alfalfa and farm alfalfa, since larvae did eat the former and until the time of obvious signs of illness, their feeding behavior appeared normal. The data indicate that feeding flexibility of M. peleides caterpillars is considerable in the sense that it allows this monophagous tropical her- bivore to exploit a broad range of genera and species locally. Foodplant records for M. peleides from Costa Rica and El Salvador indicate that many different wild Leguminosae are used, and allied South American species exhibit similar behavioral flexibility (Otero 1971; pers. comm. ). Secondary forest habitats in the wetter regions of Central America locally support a wealth of leguminous vine, shrub, and tree species, many of which are used by M. peleides. It is therefore not surprising that cater- pillars will feed successfully on allied legumes not used as natural food- plants, including cultivated forms such as peanuts and alfalfa. On a per unit area basis, secondary habitats in the tropics support large patches of Mucuna, Dioclea, Machaerium, Inga, etc. and many patches may occur locally. Thus, in terms of larval foodplants, the environment is very certain or predictable creating selection pressures favoring eco- logical specialization such as monophagy. As foodplant patches increase in number and size in an area, monophagy is considered an optimal feeding strategy for a herbivore (Levins and MacArthur 1969). Some species of Morpho that live in the canopy of primary tropical forest deposit eggs on different families of trees and woody vines (Otero, VoLUME 32, NUMBER 2 73 pers. comm.) and these species may be polyphagous. Miller (1968) lists several Morpho foodplant families, although the degree to which each species oviposits on more than one family has not been determined. Polyphagy in Morpho is adaptive in habitats where individuals of each foodplant species are greatly dispersed over large areas, making it ener- getically difficult to exploit a single family of foodplants. In forest hab- itats where each foodplant species is greatly dispersed over large areas, the environment is less certain in terms of a female butterfly locating suc- cessfully an individual of that particular plant. As more plants are added to the local foodplant niche, the environment becomes more certain; the incorporation of additional local foodplants implies the evolution of polyphagy, since member genera and species of individual plant families in tropical forests are greatly dispersed over large areas. Thus, although M. peleides and its near allies such as M. achilles may exhibit considerable feeding flexibility within the Leguminosae, they are monophagous species; polyphagous species of Morpho are expected to occur in primary forests. These include such likely candidates as M. amathonte, theseus, grana- densis, and cypris in the Central American rain forests. ACKNOWLEDGMENTS This research would not have been possible without the financial support of the Friends of the Museum (of the Milwaukee Public Museum) and James R. Neidhoefer. Susan Borkin and Joan Jass ( Mil- waukee Public Museum) conducted the rearing studies and assisted with taking measurements. Dr. J. Robert Hunter allowed me to work at Finca El Tigre and Dr. Ridgway Satterthwaite of the Associated Colleges of the Midwest provided logistical assistance in various ways. Luis D. Gomez of the Museo Nacional de Costa Rica provided a field vehicle. Janice Mahlberg of the Milwaukee Public Museum provided photo- graphic assistance. The assistance of Dr. Martyn Dibben and Neil Luebke with growing peanut plants at the museum is greatly appreciated. I also thank Joe Sugg, head of the North Carolina Peanut Council and Dr. Harry Cobel (Crop Science, North Carolina State University) for arranging peanut plants to be sent to me when the museum supply was defoliated. I thank Cheryl Castelli for typing the manuscript. To all of these people I am very grateful. LITERATURE CITED Baitey, L. H. 1969. Manual of cultivated plants. MacMillan Co., Toronto. Levins, R. & R. MacArruur. 1969. An hypothesis to explain the incidence of monophagy. Ecology 50: 910-911. 74 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Lewontin, R. C. 1965. Selection for colonizing ability. In The Genetics of Colo- nizing Species (H. G. Baker and G. Stebbins, eds.). Academic Press, New York, 2 11-94. Rien. L. D. 1968. The higher classification, phylogeny and zoogeography of the Satyridae (Lepidoptera). Mem. Amer. Entomol. Soc., No. 24, 174 pp. Orero, L. S. 1971. Instrucoes para criacao da borboleta “Capitao-do-mato” (Morpho achillaena) e outras especies do genero Morpho (“Azul-seda’, “Boia’, “Azulao-branco”, “Praia-grande”). Inst. Brasileiro Desenvolv. Florestal, Rio de Janeiro, 27 p. Tost, J. A., Jr. 1969. Mapa ecologico. Republica de Costa Rica. Centro Cientifico Tropical, San Jose, Costa Rica. Younc, A. M. 1972a. Adaptive strategies of feeding and predator-avoidance in the larvae of the neotropical butterfly Morpho peleides limpida (Lepidoptera: Morphidae). J. New York Entomol. Soc. 80: 66-82. . 1972b. Community ecology of some tropical rain forest butterflies. Amer. Midl. Nat. 87: 146—157. . 1973. The comparative ethology and ecology of several species of Morpho butterflies in Costa Rica. Studies Neotrop. Fauna 8: 17-50. 1974. The rearing of Morpho peleides (Morphinae) on peanuts. J. Lepid. Soc. 28: 90-99. Younc, A. M. & A. MuysHonpt. 1973. The biology of Morpho peleides in Central America. Carib. J. Sci. 13: 1-49. Journal of the Lepidopterists’ Society 32(2), 1978, 75-85 NOTES AND DESCRIPTIONS OF EUPTYCHIINI (LEPIDOPTERA: SATYRIDAE) FROM THE MEXICAN REGION Ler D. MILLER Allyn Museum of Entomology, 3701 Bay Shore Road, Sarasota, Florida 33580 ABSTRACT. Several species of Euptychiini (Lepidoptera: Satyridae) are dis- cussed and/or described. Described as new: Taygetis mermeria griseomarginata (Guerrero, Mexico), Splendeuptychia kendalli (Tamaulipas, Mexico), and Cyllopsis wellingi (Cayo dist., British Honduras). The previously unknown female of Cyllopsis dospassosi L. Miller is described and figured. Some years ago Mr. Roy O. Kendall of San Antonio, Texas sent me a strange euptychiine satyrid from northern Mexico for identification. It was apparent that the specimen was a representative of a new species in the genus Splendeuptychia Forster (1964), a group hitherto known from no further north than Panama. Since the Panamanian species, S. salvini (Butler), was unrepresented in the Allyn Museum collection, I compared the Mexican insect with the colored figures of salvini given by Butler (1866) and by Godman and Salvin (1880 [1879-1901]). Many dis- crepancies between the two insects became obvious, and a hurried call to Mr. Gordon B. Small, Jr. of Balboa, Canal Zone resulted in his sending two males of S. salvini that confirmed the superficial differences between it and the Mexican butterfly as well as genitalic ones. Once Mr. Kendall and his wife had managed to rear the Mexican Splendeuptychia they needed a name on which to base the paper that follows. Accordingly, I am taking this opportunity to describe the new Splendeuptychia and some other euptychiines from the Mexican region. Additional data are given on species that have come to my attention since the publication of portions of my revision of the tribe. Some of the new species described herein are members of genera not covered in the revision yet, but it is felt that publication of these parts may be so far in the future that other workers could benefit by having the names proposed at this time. Taygetis mermeria griseomarginata L. Miller, new subspecies Figs. 1-5 Male: Head, thorax and abdomen clothed with dark brown dorsal and tan to reddish-tan ventral hairs. Palpi reddish-tan, somewhat darker laterad. Antennae dark brown dorsad, reddish-brown checkered with tan ventrad. Legs clothed with brown hairs laterad, reddish-tan ones on inner portions of segments. Wings with acutely falcate forewing apices as in T. mermeria excavata Butler (1868). Upper surfaces of wings dark, rich brown, unmarked except for broad (4-8 mm) grayish overscaling along margins of all wings and more narrowly and 76 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 1.0 mm Figs. 1-5. Taygetis mermeria griseomarginata, n. ssp. 1-2, Holotype ¢ upper (1) and under (2) surfaces; MEXICO: GUERRERO: Acahuizotla (Allyn Mus. photos 101476-7/8); LFW (length of forewing) 47.0 mm. 3-4, Paratype 9 upper (3) and under (4) surfaces; MEXICO: COLIMA: Comala (Allyn Mus. photos 101476-9/10); LFW 55.2 mm. 5, @ genitalia of Holotype; slide M-2732 (Lee D. Miller). VoLUME 32, NuMBER 2 Wale less prominently along forewing costa. Under surfaces of all wings mottled in vari- ous shades of brown, reddish-brown grayish-tan or ochreous (highly variable indi- vidually) with forewing mesial bands poorly developed and only the extradiscal bands of the hindwings well developed (usually delimited by some gray-green scaling distad of the dark brown bands themselves); ocelli of both wings varying from very well developed to obsolescent. Fringes of all wings gray above, tan to reddish-brown below. 6 genitalia similar to those of other Mexican specimens (excavata) with some- what stubbier valvae than those of South American representatives. Length of forewing of Holotype ¢ 47.0 mm, those of the 21 ¢ Paratypes rang- ing from 44.6 to 53.3 mm, averaging 50.35 mm. Female: Similar in appearance to the ¢, differing chiefly in the paler coloration both dorsally and ventrally and by the presence of a poorly defined transcellular band of the hindwings beneath that is not shown by the ¢. Lengths of the forewings of the seven 2 Paratypes range from 54.1 to 58.0 mm, averaging 55.95 mm. Described from 29 specimens, 22 males and seven females, from the western slope of the Sierra Madre Occidental, Mexico. Holotype ¢@: Mexico: Guerrero: Acahuizotla, ix.1957 (T. Escalante); @ genitalia slide M-2732 (Lee D. Miller). Paratypes: all Mexico. GuERRERO: same locality as Holotype, 14 viii.1957, 5 4 ix.1957, 16 x.1957, 12 viii.1958, 12 iii.1958 (all T. Escalante); Tierra Colorado, 126 42 viii-ix.1971 (all A. Diaz Frances). Nayarir: vic. Compostela, 14 Posen bo Kiots). Comma: Colima, 14 11.1.1968; Comala, 14 31.x.1967, 12 14.41.1968 (all R. Wind). Disposition of type-series: Holotype ¢, 17é¢ and seven @ Paratypes in Allyn Museum of Entomology; single ¢ Paratypes will be placed in the American Museum of Natural History, the National Museum of Natural History, Carnegie Museum and the British Museum (Natural History ). The name of this subspecies refers to the broadly gray dusted margins of both wings on the upper surfaces. This situation is only hinted at in specimens of T. m. excavata in which the maximum development of this marginal gray scaling is about 1-1.5 mm on the forewing and virtually absent on the hindwing. In the present subspecies this gray marginal scaling is most prominent on the hindwing, but the forewing scaling is more extensive than on any excavata specimen. This gray-margined subspecies is apparently restricted to the western slopes of the Sierra Madre Occidental from at least Nayarit to Guerrero. While I have not seen material from all of the states in this area, I feel confident that griseomarginata will be found in Jalisco, Michoacan and possibly southernmost Sinaloa. A single specimen in the Allyn Museum collection from Chiapas (Tuxtla Gutierrez, 13.viii.1961, leg. “M. S.”) that was part of the Jae collection is referable to griseomarginata, but since all material from Oaxaca and Chiapas that I have seen has been referable to only excavata, I have excluded this Chiapas specimen from the type-series. It may have been mislabelled, or it may represent a genetic “throwback”, but it certainly is not typical of Chiapas-Oaxaca before me. All of the specimens I have seen from the Nayarit to Guerrero 78 JOURNAL OF THE LEPIDOPTERISTS SOCIETY range have been referable to griseomarginata, and the presence of a single specimen from outside this range should not be taken as “proof” that the subspecies does not exist. Splendeuptychia salvini (Butler), 1866 Figs. 6-8 Euptychia salvini Butler, 1866: 498 (type locality: Lion Hill, [Canal Zone], “Panama” ). Forster (1964: 128 ff.) erected the genus Splendeuptychia for 23 Neo- tropical species, most of which are restricted to South America. The species included in the present genus are among the loveliest of the Euptychiini, and their pattern is unmistakable. Only S. salvini has thus far been reported from Central America, and it is restricted to the Canal Zone and adjacent Panama, possibly as far south as the Darien. S. salvini seems to be a rare butterfly, at least in collections. I suspect that this appearance of rarity is real, since the insect is one of the more spectacular Satyridae and should not be overlooked by even a casual collector. Two males were obtained from Gordon B. Small, Jr. for examination and comparison with the Mexican species described below. The differ- ences are cited under the new species, but it suffices to say here that the two are not conspecific. Splendeuptychia kendalli L. Miller, new species Figs. 9-13 Male: Head, thorax and abdomen clothed with gray-brown dorsal and ochreous- tan ventral hairs. Palpi pale gray laterad and dark gray ventrad and dorsad. An- tennae brown dorsad, reddish-brown ventrad; tip of club slightly darker. Legs clothed with gray hairs, but those of tarsi tan. Wings above dull brown with three dark brown marginal lines separated by tan; otherwise unmarked, but the markings of the under surface showing through vaguely on this surface. Forewings below gray-brown in proximal half, tan in distal half; two thickened rust-brown lines, one across cell, the other just outside cell dividing the gray-brown from the tan ground color; three almost straight dark brown marginal lines; between the marginal lines and the distal band is a row of black-edged silver spots from M,—M:. to Cu:—2A, the whole spothand surrounded by a thin brown ring. Hindwings below with gray-brown proximal and tan distal ground color; thickened rust-brown bands of forewing continued on hindwing; three thin, dark brown marginal lines following the slightly crenulate wing outline; a mesial to submarginal yellow patch from Rs—M; to Cu.-2A encompassing silvered spots in the interspaces, those in Rs—M:, M.-M2 and Cu,-Cu: with well defined black irides; a subsidiary black line between the marginal lines and the yellow patch from Cu, to the tornus; along M, and Cu; are two black submarginal patches. Fringes pale gray above, tan below. genitalia as figured, differing from those of S. salvini (Fig. 8) in many re- spects, especially the shorter gnathos arms and the simpler valvae. Length of forewing of Holotype ¢ 17.8 mm, those of the 374 Paratypes rang- ing from 16.7 to 18.8 mm, averaging 17.67 mm. VoLUME 32, NUMBER 2 79 0.5 mm Figs. 6-8. Splendeuptychia salvini (Butler). 6-7, ¢ upper (6) and under (7) surfaces; PANAMA: PANAMA: Bayano, nr. Pina (Allyn Mus. photos 010677— 11/12); LFW 15.9 mm; G. B. Small, Jr. collection. 8, 4 genitalia of same speci- men; slide M-3418 (Lee D. Miller). Female: Very similar to the ¢, differing chiefly in size, slightly paler coloration and the more extensive yellow patches of the hindwing under surface. Lengths of forewings of the 36 2 Paratypes range from 17.3 to 21.0 mm, averag- ing 18.84 mm. Described from 74 specimens, 38 males and 36 females, from the Mexican states of Tamaulipas and San Luis Potosi. Holotype ¢: Mexico: Tamauuipas: Gonzalez Ranch, nr. Los Kikos, ex ovum on Bambusa aculeata, emerged 9.i.1975 (R. O. and C. A. Kendall); chromosome specimen no. 3A-32-M; ¢ genitalia slide M-3651 (Lee D. Miller). Paratypes: all Mexico. TamMau.ipas: same locality as Holotype, 2¢ 19 Momeni ovat LOV3., 194 M39 x 1973, 26 29 11974, Lea wl974, 36 49 xi.1974, 26 12 xii.1974, 16 792 i.1975, reared from Bambusa aculeata (all col- lected or reared by R. O. and C. A. Kendall or W. W. McGuire). San Luis Porost: Ciudad Valles, 29 vii.1970, 14 vii.1972, 36 39 vii.1973 (all collected by H. A. Freeman); El Naranjo, 3¢ 1@ ii.1976 (all collected by R. O. Kendall); Tama- zunchale, 1@ vii.1951 (T. Escalante). 10) JoURNAL OF THE LEPIDOPTERISTS SOCIETY 0.5 mm ho Figs. 9-13. Splendeuptychia kendalli, n. sp. 9-10, Holotype ¢ upper (9) and under (10) surfaces; MEXICO: TAMAULIPAS: Gonzalez Ranch, nr. Los Kikos (Allyn Mus. photos 010677-16/17); LFW 17.8 mm. 11-12, Paratype 2 upper | (11) and under (12) surfaces; same locality as Holotype (Allyn Mus. photos 010677-14/15); LFW 19.6 mm. 13, ¢ genitalia of Holotype; slide M-3651 (Lee D. Miller). Disposition of type-series: Holotype 2, 10¢ and 109 Paratypes in the collec- tion of the Allyn Museum of Entomology; nine ¢@ and 10 2 Paratypes returned to R. O. Kendall; 194 and 1592 Paratypes returned to W. W. McGuire. These series will be divided later among other museum collections. I take great pleasure in naming this distinctive Mexican satyrid for VoLUME 32, NUMBER 2 81 Mr. Roy O. Kendall who reared the Holotype and several other examples in the type-series. His work on the life histories of various Mexican, as well as Texan, butterflies has been of the greatest value to lepidop- terology and promises even more future benefits to the science. The Tamazunchale specimen came from the Escalante collection and bore a cryptic determination label in an unknown hand identifying the specimen as S. salvini. I had previously discounted salvini as the name for the Mexican butterfly, and the receipt of true salvini confirmed my previous analysis. The genitalia are somewhat aberrant for members of Splendeuptychia (Forster, 1964: figs. 161-164), especially in regard to the aborted enathos. In addition to the genitalic dissimilarities between kendalli and salvini, the former may be distinguished by the following superficial characters: 1) the ground color of kendalli is browner, both dorsad and ventrad; 2) the marginal lines on the upper surface are better developed in kendalli; 3) the ventral forewing of salvini bears four dark bands proximad of the silver spotband, whereas in kendalli the basal of these is missing al- together and the distal band is merely a thin line forming part of the ring around the silvered spots; 4) the ventral hindwing of kendalli also lacks the basalmost band that is prominent in salvini; 5) the yellow patch of the ventral hindwing is more extensive in kendalli, whereas in salvini this patch is poorly developed to absent posteriad of vein Cu,; and 6) the silver spotband of the ventral forewing which extends posteriad as far as 2A in kendalli reaches no further posteriad than Cu, in salvini. This species is apparently restricted to the mesic environments found in a few places in the eastern foothills of the Sierra Madre Oriental. Thus far the butterfly has been found in a very few localities from Tamazunchale north to Tamaulipas where colonies of B. aculeata grow. What we know about the bionomics of S. kendalli is given in a following paper (Kendall, 1978). Obviously, the insect is multivoltine. Members of Splendeuptychia are almost uniformly rare. I suspect this is a real occurrence, since they are much more attractive than are most Euptychiini. Perhaps the relative abundance of S. kendalli and its as- sociation with Bambusa will make possible the discovery of greater num- bers of other species of Splendeuptychia. The association of this genus with bamboo is further confirmed by S. S. Nicolay who brought me several specimens of an as yet undetermined Splendeuptychia that he took in a bamboo thicket in eastern Ecuador. 82 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Figs. 14-15. Cyllopsis dospassosi L. Miller, @ upper (14) and under (15) surfaces; MEXICO: SAN LUIS POTOSI: El Salto Falls (Allyn Mus. photos 010677-2/3); LFW 17.7 mm; R. O. Kendall collection. Cyllopsis dospassosi L. Miller, 1969 Figs. 14-15 Cyllopsis dospassosi L. Miller, 1969 (“1968”): 53; 1974: 84-86 (type locality: 52 mi. E Ciudad Victoria, Tamaulipas, Mexico ). The type of this species remained unique until Mr. and Mrs. Kendall collected one at El] Salto Falls, San Luis Potosi on 16.i.1975. This speci- men is the second known example of dospassosi and, fortunately, is the first female. It is quite comparable to the male, but the ground color of the upper side is slightly darker, and that of the under surface is some- what less olivaceous. Nevertheless, the maculation of the under surface is comparable to that of the male with the addition of an ochreous outer element to the extradiscal band of the hindwing. The “gray patch” en- closing the ocelli of the ventral hindwing is obscure, as in the male. The length of the forewing is 17.7 mm. I have not done a genitalic dissection of this specimen since the female genitalia are not diagnostic in Cyllopsis (L. Miller, 1974: 4). The Kendalls’ specimen of this species (which is in their collection) extends the known range of C. dospassosi from the Sierra de Tamaulipas to the dry eastern flanks of the Sierra Madre Oriental, presumably of Tamaulipas, as well as San Luis Potosi. The range of C. dospassosi may be much wider than previously thought, and its rarity in collections may be attributable to the usual lack of collecting of the smaller Euptychiini. Cyllopsis wellingi L. Miller, new species Figs. 16-20 Male: Superficially like C. nayarit (R. Chermock), but differing in the follow- ing particulars: somewhat larger, approaching size of C. pephredo (Godman); VOLUME 32, NUMBER 2 83 0.5 mm. ne | Figs. 16-20. Cyllopsis wellingi, n. sp. 16-17, Holotype ¢ upper (16) and under (17) surfaces; BRITISH HONDURAS (BELIZE): Cayo District: Pine Ridge, Thousand Foot Falls (Allyn Mus. photos 021777-1/2); LFW 17.3 mm. 18-19, Paratype 2 upper (18) and under (19) surfaces; same locality as Holotype; (Allyn Mus. photos 021677-1/2); LFW 18.5 mm. 20, @ genitalia of Holotype; slide M-3667 (Lee D. Miller). 84 JOURNAL OF THE LEPIDOPTERISTS SOCIETY under surface bands on both wings redder than in either species; transcellular bands of both wings below less well developed than in nayarit; ochreous markings of hind- wing below much more extensive than in nayarit (these are only hinted at in peph- redo); and ocelli not edged inwardly with ochreous in gray patch area, as in nayarit. 4 genitalia as figured, not at all resembling those of pephredo. The genitalia do bear some resemblance to those of C. pseudopephredo (R. Chermock) (L. Miller, 1974: fig. 141), a species that is otherwise quite distinct from wellingi and other members of the pephredo subgroup in its lack of an androconial patch. The valvae of the present species are somewhat broader than those of pseudopephredo, but the characteristic inwardly directed teeth (which usually appear as dorsally diverted ones) are very similar. Length of forewing of Holotype ¢ 17.3 mm, those of the 11 ¢ Paratypes rang- ing from 17.0 to 18.2 mm, averaging 17.48 mm. Female: Differs from the @ of C. nayarit in much the same manner as does the ¢, with the additional characteristic of a reddish flush on the upper surface of some specimens that is not shown in other members of the pephredo subgroup. Lengths of forewings of the eight 2 Paratypes range from 17.9 to 19.1 mm, averaging 18.45 mm. Described from 20 specimens, 12 males and eight females, from British Honduras ( Belize ). Holotype ¢: British Honpvuras: Cayo District: Pine Ridge, Thousand Foot Falls, 650 m, 2.ix.1976 (E. C. Welling M); @ genitalia slide M-3667 (Lee D. Miller ). Paratypes: all same locality as Holotype, 2—3.ix.1976 (E. C. Welling M.), 11é 82 (males all determined genitalically ). Disposition of type-series: Entire type-series placed in Allyn Museum of Ento- mology, but the series may be subdivided later. It is with great pleasure that I name this little satyrid for Sr. Eduardo C. Welling M. of Mérida, Yucatan, Mexico. He has consistently been available to collect specimens of Euptychiini for me, and has often placed undescribed and unexpected species at our disposal for systematic work. This species is something of a puzzle. One male of thirteen in front of me was a specimen of C. pephredo, but all of the others were the new insect, as demonstrated by genitalic dissections. I have no idea why the single pephredo was intermingled with wellingi at the type locality of the latter, and this single specimen represents the first record of pephredo from British Honduras. These two species now bring the total number of Cyllopsis from that country to three (C. gemma freemani {Stallings and Turner] also occurs there). ACKNOWLEDGMENTS I am most grateful to Messrs. Roy O. Kendall, E. C. Welling M., Gordon B. Small, Jr. and Alberto Diaz F. and Drs. W. W. McGuire and Tarsicio Escalante for providing the material on which this is based. Mr. A. C. Allyn took the photographs used to illustrate the paper. Mr. Allyn and my wife and colleague, Jacqueline, read and suggested upon the paper. To all of these individuals I owe a great debt of gratitude. VoLUME 32, NUMBER 2 85 LITERATURE CITED Butter, A. G. 1866. A monograph of the genus Euptychia, a numerous race of butterflies belonging to the family Satyridae; with descriptions of sixty species new to science, and notes on their affinities, &c. Proc. Zool. Soc. London, [1866]: 458-504; ill. 1868. Catalogue of the diumal Lepidoptera of the family Satyridae in the collection of the British Museum. London, Trustees British Mus.: vi + 211 pp.; ill. Forster, W. 1964. Beitrage zur Kenntnis der Insecktenfauna Boliviens. XIX. Lepidoptera III. Satyridae. Ver6dff. Zool. Staatssamml. Miinchen, 8: 51-188; ill. Gopman, F. D. & O. Satvin. 1879-1901. Biolcgia Centrali-Americana. Insecta. Lepidoptera-Rhopalocera. London: 2 vols. + 1 vol. of ill. KENDALL, R. O. 1978. Larval foodplant, life history notes and temporal distribu- tion for Splendeuptychia kendalli (Satyridae) from Mexico. J. Lepid. Soc., 32: 86-87. Minter, L. D. 1969. On Mexican Satyridae, with description of a new species. J. Res. Lepid., 7 (“1968”): 51-55; ill. . 1974. Revision of the Euptychiini (Satyridae). 2. Cyllopsis R. Felder. Bull. Allyn Mus., (20): 98 pp.; ill. Journal of the Lepidopterists’ Society 32(2), 1978, 86—87 LARVAL FOODPLANT, LIFE HISTORY NOTES AND TEMPORAL DISTRIBUTION FOR SPLENDEUPTYCHIA KENDALLI (SATYRIDAE) FROM MEXICO! Roy O. KENDALL? Route 4, Box 104-EB, San Antonio, Texas 78228 ABSTRACT. Larval foodplant, Bambusa aculeata, Gramineae, rearing notes, ecologic and temporal distribution at its northern distributional limit, are recorded for Splendeuptychia kendalli Miller. Field-collected adults of the satyrid, Splendeuptychia kendalli Miller, were found at 2 locations: 1) along the Rio Sabinas at Rancho Pico de Oro near Ciudad Mante, Tamaulipas, and 2) along the Rio Salto at El Naranjo, San Luis Potosi. This species is closely associated with its larval foodplant, Bambusa aculeata (Ruprecht) Hitchcock, Gramineae. It is doubtless that this insect will be found at other locations along water courses where its larval foodplant grows. Adults were taken in January, February, July, October, November, and December over a 4- year period. The areas were not visited during the other months. It is therefore unknown whether this multivoltine species is continuous brooded,; it may have a reproductive diapause. Rearing. At Rancho Pico de Oro, 21 December 1972, I observed a 2 deposit a single egg on a juvenile leaf of B. aculeata. The ? was not captured, but the egg was recovered and preserved. Again at this location on 22 January 1974, two females were collected and kept for egg production. Between 23 January and 2 February, 51 eggs were deposited in confinement on B. aculeata. Most of the eggs were deposited by 1 female which died 2 February. The other female was killed at this time, and both adults were preserved in alcohol to- gether with 6 eggs. The remaining eggs hatched between 28 January and 7 February. Larval losses were rather high resulting from an in- adequate supply of fresh food. Attempts to keep the plants fresh in a refrigerator were only moderately successful. Earlier, several specimens of the foodplant were transplanted to the Los Arcos Courts gardens at Ciudad Mante, our field headquarters, but they did not survive. In an attempt to circumvent a 60-mile trip every few days for larval food, the larvae were offered bermuda grass, Cynodon dactylon (L.) Pers. The ' Contribution No, 380, Bureau of Entomology, Division of Plant Industry, Florida Department of Agriculture and Consumer Services, Gainesville 32602. * Research Associate, Florida State Collection of Arthropods, Division of Plant Industry, Florida Department of Agriculture and Consumer Services. VoLUME 32, NUMBER 2 87 larvae ate the bermuda grass, and it was thought a laboratory solution had been found for rearing this species. However, the larvae soon began to die. The bermuda grass may have been toxic to the larvae, but the lack of proper nourishment in the grass was suspect. On 6 April 1974 all remaining larvae (24) were preserved. Once again a 2 collected 23 November 1974 from this location de- posited 27 eggs between 24 and 29 November and died 3 December 1974. These eggs hatched between 28 November-l11 December, and a maximum effort was made to rear them. Numerous trips were made to the collection site for fresh bamboo. Even so, there were several larval casualties attributed to rapid desiccation of the cut bamboo. Eleven larvae pupated between 27 December 1974 and 12 January 1975. Adults emerged (2 6, 72) between 7 and 23 January 1975. Two larvae and 2 pupae (one deformed ) were preserved. Field-Collected Adults. In addition to the above, other field collections include: Rancho Pico de Oro, 21 December 1972 (24,12), 9 January 1974 (32 ), 22 Janu- ary 1974 (146), 22 February 1974 (14), 23 November 1974 (44, 19), 4 December 1974 (14), 6 December 1974 (39 ), and 8 January 1975 (1¢), all leg. Roy O. and C. A. Kendall. At the same location, 27 December 1972 (124), 18 July Poaee ee 20nfuly 1973 (2° ), 22 October 1973 (176, 82), 25 October 1973 (36, 22) all leg. W. W. McGuire. At E] Naranjo, 13 February 1976 (1@), 14 February 1976 (224), and 29 February 1976 (14) all leg. Roy O. and C. A. Kendall. ACKNOWLEDGMENTS Mrs. Kendall and I wish to thank Sr. and Sra. Carlos Gonzales for permission to conduct field research at their rancho, and for their warm hospitality. To Sr. and Sra. Fernando Reyes Bugarin and their family we are most grateful for the comfortable field headquarters provided and for the use of their botanical gardens in our research. Journal of the Lepidopterists’ Society 32(2), 1978, 88-96 NOTES ON THE LIFE CYCLE AND NATURAL HISTORY OF VANESSA ANNABELLA (NYMPHALIDAE) THomMas E. Dimock? 111 Stevens Circle, Ventura, California 93003 ABSTRACT. Observations on the life history of Vanessa annabella (Field) show the early stages to be quite variable: the eggs in rib structure, and the later larval stages in color pattern and behavior. Immature and adult behavioral charac- teristics are similar to those of other Vanessa. V. annabella is usually present throughout the year in coastal southern California. Vanessa annabella (Field), the West Coast Lady, is a common and familiar butterfly in western North America. Because it can usually be found throughout the year in coastal southern California, opportunities to study its life history are almost always present. However, there are few published records available and none has included photographs of the complete life cycle. Of published reports, Dyar (1889) gave one of the more complete written accounts; Huguenin (1921) made some general observations on the life cycle and natural history; and Coolidge (1925) described the egg in detail and listed the larval foodplants. More recently Emmel & Emmel (1973) illustrated paintings of a light form of the last instar larva and the pupa and gave brief descriptive notes. Specimens used for the present descriptions of the life cycle stages were collected as freshly laid ova by following an ovipositing female at the type locality in Ventura, California (Dimock, 1972). The leaves on which these eggs were laid were placed in plastic containers 11 cm square by 4 cm deep. Humidity was maintained by dampened tissue paper placed on the container bottom. The containers were kept indoors in a room temperature which varied from 17 to 25°C. Photographs and measurements were made of each stage. Other specimens were reared upon cut stalks of nettle placed in water so that leaf shelter construction and other activities could be observed. Afternoon sunshine provided direct and ambient light. Full descriptions of the adults are given by Field (1971); thus, the following adult descriptions are limited to those characteristics which help distinguish V. annabella from related North American species. Life Cycle Stages _ Ege (lig. 1). Barrel-shaped, light green, with 10 to 14 transparent vertical ribs. Measurements (Coolidge, 1925): 0.72 mm tall, 0.52 mm wide, tapering to 0.30 mm at base and 0.26 mm at top. Duration 4 days. ‘Museum Associate in Entomology, Natural History Museum of Los Angeles County, 900 Ex- position Blyd., Los Angeles, California 90007. ; VoLUME 32, NUMBER 2 89 Figs. 1-12. Vanessa annabella (Field): (1) egg, ca. 0.6 mm wide; (2) first instar larva, 3 mm; (3) second instar larva, ca. 4 mm: (4) third instar larva, ca. 9 mm; (5) fourth instar larva, ca. 13 mm; fifth instar larvae, all ca. 30 mm: (6) dark morph, black and yellow, (7) intermediate morph, orange and gray, (8) rusty orange morph, (9) light morph, gray; (10) head, fifth instar, light tan morph; (11) head, fifth instar, dark morph; (12) prepupa. First instar larva (Fig. 2). Head shiny black, setae and thoracic legs black. Ground color grayish brown after feeding for 2 days. When mature, body with vague brownish mottling. Segments A-2, A-4, and A-6 with a pair of light yellow spots between subdorsal and supralateral setae. Grows to 3 mm in 5 days. Second instar larva (Fig. 3). Head shiny black. Ground color mottled dark brown. Short branched spines black except for middorsal spines on A-4 and A-6 and subdorsal spines on A-2, A-4, and A-6, which are yellow. A narrow pair of vague yellow lines divided by a narrow middorsal line of dark ground color running from about T-1 to A-8. Grows to 4.5 mm in ca. 3 days. Third instar larva (Fig. 4). Head shiny black with black setae arising from black chalazae. Ground color usually black, but may begin to lighten as in lighter morphs. Spines black except for subdorsal spines on A-2, A-4, A-6, and usually 90 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Wj ee. Z iY Wy, » ty Fig. 13. Vanessa annabella (Field): fifth instar larval nest on Urtica holosericea Nutt. middorsal spines on A-4 and A-6, which are yellow with black tips and yellow bases confluent with dorsal double yellow lines. These paired lines, variable in expression, separated by a middorsal line of ground color, are interrupted by ground color at bases of non-yellow spines. Grows to ca. 9 mm in ca. 3 days. Fourth instar larva (Fig. 5). Head black with bronze hightlights. Head cap- sule width 1.5 mm. Ground color and markings nearly as variable as in fifth instar (see following description). Grows to ca. 14 mm in ca. 3 days. Fifth instar larva (Figs. 6-9). Head blackish or brownish black (Fig. 11) with bronze highlights, or less often with vertical whitish tan stripes in light morphs (Fig. 10). Capsule width 2 mm. Extremely variable in ground color and mark- ings. Ground color varies from black to greenish white or grayish white, including various browns and tans. Light markings present in fourth instar here vary from dark rusty reds and oranges to yellow or various browns and tans. Extent of mark- ings and lateral line varies independently of color; these tend to disappear alto- gether in morphs of whitish ground color. Lateral line may be absent in any morph. tusty orange spots may appear between subdorsal and supralateral spine bases, varying in extent from absence to confluence with other pattern elements. Spines branched, variable from black in dark morphs to whitish in light morphs, or dark anteriorly and light posteriorly in intermediate morphs. Arrangement of spines: middorsal row on segments A-1 to A-8, subdorsal rows on T-2 to A-8, supra- lateral rows on T-2 to A-10, and lateral rows on A-1 to A-8. Body shape thickest at midabdominal segments. Grows to ca. 25-30 mm in ca. 5 days. Prepupa (Fig. 12). Light markings darken somewhat. Larva becomes slightly shorter and thicker. Duration 1 day. Pupa (Figs. 14-16). Head without projections. Mesothorax with a_ raised middorsal point and a pair of subdorsal points. Metathorax with two large sub- VOLUME 32, NUMBER 2 91 Figs. 14-20. Vanessa annabella (Field): (14) pupa, dorsal view, 19 mm; (15) pupa, lateral view; (16) pupa, ventral view; (17) adult male, dorsal view, 47—mm expanse; (18) adult male, ventral view; (19) adult female, dorsal view, 49-mm expanse; (20) adult female, ventral view. dorsal white spots, raised anteriorly. Abdomen with two small subdorsal white spots on A-1 bordering those on metathorax. A-2 to A-7 each with one very small middorsal point and a pair of more prominent subdorsal points. Color variable from overall tan to mottled dark browns, sometimes with a greenish golden cast. 992 JOURNAL OF THE LEPIDOPTERISTS SOCIETY A spiracular line, darker brown than ground color, is variable in expression. Mea- surements (average of five specimens): length 19 mm; width 7 mm; depths: thorax 6.25 mm, saddle 5.25 mm, abdomen 7.25 mm. Duration 8-11 days. Adults (Figs. 17-20). Sexual dimorphism subtle, females having a more rounded hindwing than males, especially at M:, Cu, and Cuz. Color pattern same in both sexes: tawny orange with black markings, white subapical forewing spots, and blue pupilled hindwing ocelli. On upperside, forewing cell crossed completely by a black bar. Forewing costal bar, between cell end and apex, orange. Forewing apex pointed, not rounded, at M,, with marginal tawny spot in interspace Rs. Hindwing with four blue pupilled submarginal ocelli in interspaces M:, M2, Ms, and Cu, and often a small solid black ocellus in interspace Rs. Hindwing under- sides mottled principally in various buffs, tans, browns, and grays, with a whitish triangle in interspace M» at cell end. Expanse averages between 40 and 48 mm, females often larger than males. Total developmental time for this species is ca. 30 to 36 days. Natural History Vanessa annabella uses a variety of foodplants in the families Urti- caceae and Malvaceae. Native foodplants most frequently used in southern California are Urtica holosericea Nutt. (Urticaceae), Sida spe- cies, Sidalcea malvaeflora (DC.) Gray, and Sphaeralcea ambigua Gray (Malvaceae). Introduced plants include Malva species, especially M. parviflora L., and Althaea rosea (L.) Cav. (Malvaceae). John F. Em- mel, M.D. (pers. comm.) also reports the use of Urtica urens L. ( Urticaceae ). The eggs are laid singly, usually on the uppersides of the leaves. On nettles (Urtica) the eggs are often attached to the sides of the stinging spines. The hatching larva eats away the top and adjacent walls of the egg and crawls to a suitable place on the leaf uppersurface to construct a shelter. This consists of fine silk webbing tied across a leaf midrib, petiole, or small wrinkle on the leaf margin. The young larva lives under this webbing as it feeds on the leaf and places its frass into the webbing, creating a protective camouflage. In the second instar the larva may enlarge the old nest or construct a new one nearby. When the larva is at the growing tip of a nettle stalk, the nest may incorporate two or more of the tiny new leaves. By the third instar the larva is capable of folding a larger area of the leaf or constructing a deeper nest at the petiole. In the fourth instar the entire leaf may be folded together (on Urtica) or closed about the top edges (on Malwa). Frass is allowed to fall out of the nest but often accumulates in piles in the nest bottom. The fifth instar larval nest is usually larger and may incorporate neigh- boring leaves and stems (Fig. 13). Sometimes leaves of nearby plants which are not foodplants are also tied into the nest even though they are VoLUME 32, NUMBER 2 93 not eaten. On plants with small leaves, such as young Malwa, the larva may tie together many leaves before a nest enclosure is completed. On Urtica holosericea, when a single leaf is used, larvae of V. annabella usually construct nests on the uppersides of the leaves, either by folding over one edge and securing it to the leaf surface or by tying both edges together to form an enclosure. The petiole or nearby midribs may or may not be partially cut to cause the leaf to hang vertically. Less frequently, the larvae will fold the leaf edges underneath so that the undersurface forms the nest interior. Pupation sites are on either the foodplant or nearby objects. When the foodplant is used, a leaf chamber is constructed with firm webbing and the larva suspends itself from the chamber ceiling. Larvae in other loca- tions may secure together any nearby objects to approximate an en- closure or may simply pupate in exposed places, such as from twigs or branches. Pupae often react to disturbances by wiggling laterally. Emerging adults hang from the pupal shell or adjacent perch to ex- pand their wings. A reddish brown meconium is ejected and the adult is ready for flight in an hour. Adults of V. annabella may be encoutered in any life zone from sea level to alpine areas where open sunny places are preferred. Both sexes visit flowers. In the afternoon males tend to congregate on hilltops or other exposed places such as forest openings, glades, meadows, and streamside slopes, especially when patches of dry, bare earth are avail- able for sunning. Many man-made situations are particularly favorable: windbreaks of trees, orchard rows, trails, firebreaks, garden paths, and paved sidewalks and driveways. At these locations, when not occupied in sunning, males will chase after each other and the other vanessid butterflies Vanessa atalanta rubria (Fruhstorfer), V. cardui (L.), and V. virginiensis (Drury), along with unrelated butterflies which con- gregate in the same places. They often bravely chase larger insects and birds and in general will investigate anything that flies through their established area, including falling leaves and objects thrown overhead. These activities ultimately bring the males into contact and subsequent courtship and mating with females, but between these encounters the males spend a great deal of time and energy simply chasing each other. From observations made on hilltops in the vicinity of Ventura, California, during November 1976 when all four Vanessa were present, it was noted that any one species will chase the same or any other species, and two or more individuals may join in the chase. The butterflies may chase each other to a height of ca. 20 m or more before breaking chase and quickly gliding down to land once again on the ground with backs to the 94 JOURNAL OF THE LEPIDOPTERISTS SOCIETY sun and wings spread. The butterflies’ wings frequently come into con- tact during these encounters, but without damaging effects, and the re- sulting noise can be heard nearby. Females, although not congregating in the manner of the males, are likely to be found anywhere, feeding, seeking foodplants, or ovipositing, including hilltop localities when the foodplants or nectar sources occur there also. The flight of V. annabella is composed of glides with the wings held horizontally, interrupted frequently by several fluttering beats. Chasing is mostly vigorous fluttering, and the return dives are composed of gliding and braking. Diapause was not investigated, but if it does occur in this species it is almost certainly during the adult stage, as it is in the other vanessid- nymphalinid butterflies. If adults of V. annabella are unable to survive prolonged or severe frosts, the species probably reinvades the greater part of its northern and eastern range from the milder southwestern areas where breeding is continuous throughout the year. Vanessa annabella is easily attracted to suburban gardens by planting Althaea rosea (Hollyhock) or encouraging Malva parviflora (Cheese- weed) to become established. It is an easy butterfly to raise in captivity, even under poor conditions. Larvae collected in the wild on one food- plant (for example, Urtica holosericea) can be switched to other foodplants (Malva, Althaea) when the former is less easily obtained. The larvae are very often parasitized by tachinid flies, which emerge from the mature butterfly larvae or pupae as mature maggots. DiIscussIon Coolidge (1925) noted that the egg ribs of V. annabella varied in number from 11 to 13, with 11 the most common number. My ob- servations confirm this, but eggs with 10 and 14 ribs were found during the present study. This is in partial disagreement with Field (1971), who stated that the genus Cynthia (in which annabella was placed) had from 14 to 19 egg ribs, and Clench (in Howe, 1975), who gave 14 or 15 as the number of egg ribs in the subgenus Cynthia. The larvae of V. annabella are extremely variable, as are the larvae of V. atalanta rubria and V. cardui, and this variability not only makes a written description difficult but compounds the task of providing re- liable characteristics with which the three species can be separated. In general, the descriptions given for the larvae in this article can be com- pared with those for V. a. rubria and V. cardui in subsequent articles. However, fifth instar larvae of V. annabella can always be distinguished VoLUME 32, NUMBER 2 95 by their smaller head capsule width of 2 mm; in V. a. rubria and V. cardui the fifth instar larval head capsule is nearly 3 mm. Once V. anna- bella is identified, V. a. rubria is distinguished by the numerous white cephalic chalazae, which in V. cardui are black. From observations made on V. annabella larvae collected in various locations and larvae reared under controlled conditions, it was noted that the variations in ground color were at least partly due to environ- mental conditions. The light, grayish-white morphs were more fre- quently encountered on plants exposed to full sunshine, whereas the darker morphs were found mostly on plants in secluded, shaded areas. Darker morphs also resulted when larvae were reared under crowded conditions. Because Malva species are especially successful in disturbed areas and are abundantly available throughout the year as foodplants, V. annabella has probably become much more common since the introduction of these weeds from Europe. This is the situation in the coastal southern California lowlands, where a favorable climate prevails and V. annabella can be found in every month of the year. In his revision of the Vanessa butterflies, Field (1971) resurrected the genus Cynthia for the carye, cardui, and virginiensis species groups. Clench (in Howe, 1975) reunites all the species in Vanessa, treating Cynthia as a subgenus. Emmel & Emmel (1973) used the cases of hybridization between V. a. rubria and Cynthia annabella to demonstrate the “close genetic relationship and probable generic identity” of the two species. With no disrespect to the fine work of Field, I favor a treatment similar to that of Clench, with reservations on the precise placement of annabella. There are 10 known cases of hybridization between V. a. rubria and V. annabella: one specimen reported by Edwards (1877), one by Grinnell (1918), one by Gunder (1930), three by Dimock (1973), one by Emmel & Emmel (1973), one specimen collected by Kirby in the collection of the Natural History Museum of Los Angeles County, and two specimens raised by Henne and Ingham in the Peabody Museum collection. Mr. William D. Field (pers. comm.) discovered upon dis- section the partially crippled specimen designated as “Hybrid #3” in Dimock (1973) to be a female, not a male as erroneously reported. In my opinion these occurrences support, at least, the arrangement of Clench and the generic identity suggested by Emmel & Emmel. Biolog- ically, the examples of hybridization may also demonstrate the presence of an as yet incomplete reproductive isolatory mechanism caused by a recent invasion of V. annabella from South America or a recent invasion of V. a. rubria from Eurasia, or invasions by both species. 96 JoURNAL OF THE LEPIDOPTERISTS SOCIETY ACKNOWLEDGMENTS I wish to thank my father, M. W. Dimock, for printing the photographs which accompany this article; Mr. Julian P. Donahue of the Natural History Museum of Los Angeles County for access to the Museum col- lection; Mr. Christopher Henne of Pearblossom, California, for an account of his hybrids; Mr. William D. Field of the Smithsonian Institution, Washington, D.C., for information on the hybrids; Dr. John F. Emmel, Hemet, California, for reviewing the manuscript; and Mr. Alberto Muy- shondt of El Salvador for recommendations on photographic techniques and whose format for life history observations of El Salvador butterflies is the one I followed for this article. LITERATURE CITED Cootmncr, K. R. 1925. California butterfly notes, III. Bull. Brooklyn Ent. Soc. 20(3): 146-147. Dimock, T. E. 1972. Type locality and habitat—Cynthia annabella. J. Res. on the Lepid. 10: 265-266. 1973. Three natural hybrids of Vanessa atalanta rubria x Cynthia annabella (Nymphalidae). J. Lepid. Soc. 27: 274-278. Dyar, H. G. 1889. Preparatory stages of Pyrameis carye Hiibner. Canadian Ent. 21: 237-238. Epwarps, H. 1877. Pacific Coast Lepidoptera, No. 22. Notes on some diurnal Lepidoptera, with descriptions of new varieties. Proc. Calif. Acad. of Scis. 7: 163-174. EMMEL, T. C. & J. F. Emmeu. 1973. The butterflies of southern California. Nat. Hist. Mus. of Los Angeles County, Sci. Ser. 26: 1-148. Fretp, W. D. 1971. Butterflies of the genus Vanessa and of the resurrected gen- era Bassaris and Cynthia (Lepidoptera: Nymphalidae). Smiths. Contribs. to Zool., Number 84: 1—105. GRINNELL, Jr., F. 1918. Some variations in the genus Vanessa (Pyrameis). Psyche 25: 110-115, pl. 4. Gunpver, J. D. 1930. Butterflies of Los Angeles County. Bull. Southern Calif. Acad. of Sci. 29: 39-95. Howe, W. H., coordinating editor. 1975. The Butterflies of North America. Doubleday and Co., Inc., Garden City, L. I., New York. xiii + 633 p. + 97 pl. HucuEnin, J. C. 1921. Life history of Pyrameis caryae in California (Lep., Rhop.). Ent. News 32: 216-217. Journal of the Lepidopterists’ Society 32(2), 1978, 97-102 A NEW SPECIES OF HEMILEUCA FROM THE SOUTHWESTERN UNITED STATES (SATURNIIDAE) Pau M. TusKEs Department of Environmental Toxicology, University of California, Davis. Davis, California 95616 ABSTRACT. JHemileuca griffini Tuskes which occurs in southern Utah and northern Arizona was collected for the first time in 1974. The adult moth is a black and white day flying saturniid which is active during September and October. The larval hostplant is black brush, Coleogyne ramosissima. This species has a unique taxonomic position in that both the adult and larva exhibit morphological characters which are intermediate to the Pseudohazis and Hemileuca groups, thus, a continuum of characters exists between these two previously separated genera. The genus Hemileuca consists of 23 described species, 16 of which have partial or complete distributional patterns north of Mexico. The moths within this genus are large to moderate in size, and exhibit a great deal of hostplant and habitat diversity. Adults are characterized by hav- ing the labial palpi fused to each other forming a small unsegmented bilobed structure; also, the male has bipectinate antennae. Members of Coloradia, the genus most closely related to Hemileuca, have labial palpi which are separate, and males have antennae which are quadripectinate. The last Hemileuca described as a distinct species was chinatiensis (Tinkham), in 1943. The significance of H. chinatiensis as a species with genitalic characters intermediate between Pseudohazis and Hemi- leuca was overlooked by Tinkham; not until Ferguson (1971) was its taxonomic position made clear. Michener (1962) combined the genera Pseudohazis and Hemileuca on the basis of their morphological similarity, but made no mention of chinatiensis. Although Michener included four subgenera within Hemileuca Ferguson chose to abandon the subgeneric names and to consider them as species groups. It is the purpose of this paper to describe a new species of Hemileuca collected for the first time in 1974, and to present additional mor- phological evidence to support the merger of Hemileuca and Pseu- dohazis. The new species of Hemileuca described in this paper is named after Mr. Bruce Griffin, who collected the first specimens. Hemileuca griffini Tuskes, new species Holotype: Male (Figs. la,b). Heap: Eyes dark brown. Frontal and vertex hairs rust red, clypeal hairs dark brown to rust red. Antennae, bipectinate, 0.67 cm long; shaft orange ventrally, dark brown dorsally, pectiniform processes black and finely plumose. THorax: Dorsally clothed with black hairs; long white hairs mixed with tufts of rust red hairs at base of secondaries. Collar, white with rust 98 JOURNAL OF THE LEPIDOPTERISTS SOCIETY red hairs anteriorly and posteriorly. Legs, clothed with black hairs; anterior por- tion of pro- and metathoracic iegs with long rust red hairs. ABDoMEN: Abdominal segments I-VI lustrous black, posterior margin of pleura I-VI lightly fringed with white hairs. Terga I-VI black and sparsely clothed with red hairs. Terga VII and VIII rust red. Forewincs: 2.54 cm long, wings approximately 60% white and 40% black. Veins black. Marginal area black, submarginal area white, but traversed by black veins; postmedial line black and continuous. Distal portion of discal cell with black band extending from postmedial line to costa. Coastal area black. Area between costa and subcostal area adjacent to discal cell white. Antemedial line black and extending from costa, curving out at base of discal cell and continu- ing transversely to hind margin. Basal patch tear-shaped, black and free standing. Basal portion below basal patch clothed with long black and white hairs to hind margin of wing. Ventral surface similar to dorsal. Hrynpwines: 1.87 cm long and approximately 50% white and 50% black. Marginal and submarginal areas similar to those of primaries. Postmedial line diverging in area of the discal cell, forming a circular area with a white center just distal to the discal cell. Basal area black, clothed with long black and white hairs extending almost to the postmedial line along the interior margin of wing. MALE Genirauia: (Fig. 5) Uncus trilobed, dorsal and anterior portion covered with bristles; wide at base, narrowing at apex. Lateral process of transtilla narrow, and not extending past apex of uncus. Transtilla fused, but with shallow groove at apex of medial process. Valves prominent and distinctly winged, apex slightly rounded and not pinnacle-shaped; upper half of ventral margin heavily setose. Four—eight setae *4 as long or longer than juxta, located basolaterally. Entire genitalic structure lightly sclerotized. Allotype: Female (Figs. 2a,b). HEap: Eyes dark brown. Entire head covered with rust red scales. Antennae, bipectinate, 0.70 cm long; shaft orange both dorsally and ventrally pectiniform processes orange to dark brown, and not finely plumose. THorAx: Dorsally clothed with black hairs, long white hairs at base of primaries and secondaries. Long, large tufts of orange hairs at base of secondaries. Orange hairs scattered on posterior portion of thorax. Collar rust red. Legs, similar to those of male, but with more red present on femur of metathoracic leg than on holotype. ABbpoMEN: Abdominal segments I—VII lustrous black with terga I—VII lightly fringed with rust red hairs. Terga of segment VIII rust red. Pleura I—-VIII lustrous black. Forewincs: 2.83 cm long. Similar to those of male, but with the following exceptions: Marginal area more heavily marked with black scales. Black margin continuous around entire wing. Hinpwincs: 2.04 cm long. Similar to those of male but margins more heavily marked with black scales. ParaTyPe VARIATION. The length of the forewing in the 20 males examined averaged 2.54 cm, and ranged from 2.31 to 3.14 cm. The markings on the forewings exhibited little variation, except for the black basal patch. In some individuals the basal patch is slightly more prominant than that of the specimen illustrated (Fig. 1), but in 15 of the 21 paratypes examined it was less developed or almost absent. Al- though the pattern is uniform, the intensity of the scales differ. The wings of what are assumed to be older specimens are cream colored, rather than white, and often partially transparent. The hindwings show more variation than the forewings. Paratypes from Mexican Hat and Bitter Springs Rd. appears similar, but two males from Pierce Ferry Rd. are much darker. The forewings of 16 females examined averaged 2.75 cm, and ranged from 2.67 to 3.13 cm in length. As in the males the VOLUME 32, NUMBER 2 99 forewings showed relatively little variation, but the hindwing differed markedly. In most individuals the hindwings were approximately 50% black and 50% white, while in others they were about 80% black. Types: Howotyre: ¢ ca. 6 mi. S.W. of Mexican Hat, San Juan Co., Utah. Elev. 4800’. Sept. 2, 1974. Bruce Griffin, Collector. ALLOTYPE: @ ca. 0.5 mi. E. of Jct. 89A and 89 on Hwy 89, near Bitter Springs. Coconino Co., Arizona. Elev. 5200’. Collected as 3rd instar larva by B. Griffin and Ken Hansen on Coleogyne ramosissima, May 3, 1975, and reared to maturity on Cercocarpa betuloides by Paul Tuskes, emerged Oct. 12, 1976. Paratypres: Utah: 3 ¢ 2, same data as holotype, 4 ¢@¢ and 3 2@Q same locality as type, Sept. 8, 1976, Kilian Roever; 8 646 and 4 22, Rt. 163, 7 mi. S.W. of Mexican Hat, San Juan Co., Sept. 8, 1974, K. Roever. Arizona: 3 ¢@¢ and 5 Q2@@Q same data as allotype; 2 ¢¢ and 3 29, Pierce Ferry Rd., 32 mi. N.E. Rt. 83, Mohave Co., K. Roever; 2 9 9, Rt. 160, 42 mi. E.N.E. of Keyenta, Navajo Co., Sept. 8, 1974, K. Roever. The types were deposited at the Los Angeles County Museum of Natural History. Paratypes were deposited at the following institutions: American Museum of Nat- ural History, Los Angeles County Museum of Natural History, Dept. of Entomology, University of California, Davis, and the United States National Museum. Larval Description—Last Instar Head: Shiny black with numerous white setae, diameter, 4-5 mm. Clypeus black. Body: Length 45-55 mm, width, 7-8 mm. Ventral surface gray to light brown with an orange cast. Sublateral scoli black; lateral scoli black with slight yellow cast at tips, dorsal scoli of rosette type, yellow with black center. Body with three distinct lateral cream to white bands. Band I from prothoracic segment to caudal segment, passing through sublateral scoli. Band II broken by interseg- mental area and located slightly ventral to lateral scoli. Band III lightly pig- mented broken band, but still obvious, located midway between dorsal and lateral scoli. Segmental area with cream to white colored paniculum, especially common on lateral areas. Secondary setae white. True legs black. Prolegs gray to black. Spiracles orange. Ground color black. Characteristics of H. griffini in relation to other Hemileuca The trilobed uncus of griffini (Fig. 5) is typical of the Pseudohazis group. The only Hemileuca to have a trilobed uncus outside of the Pseudohazis group is H. electra Wright. The uncus of electra is typically bilobed, but apparently some aberrant males have trilobed unci (Fer- guson, 1971). The transtilla of griffini is shallowly grooved at the apex of the medial process, and this characteristic appears to be intermediate between the two subgenera. Fused transtilla are common to all Hemi- leuca with the exception of diana and grotei. Both of these species have bilobed transtilla which are rounded and short compared to the long thin bilobed structure of Pseudohazis. The valves of griffini are prom- inent and distinctly winged, showing much greater development than the typical rounded valves of Pseudohazis but they are not as large as those of most Hemileuca. The genitalia and adult phenotype of griffini show the greatest 100 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Figs. 1-6. 1, Dorsal (la) and ventral (1b) view of male H. griffini (Holotype). 2, Dorsal (2a) and ventral (2b) view of female H. griffini (Allotype). 3, Dorsal view of male H. chinatiensis. t, Dorsal view of female H. chinatiensis. 5, Male genitalia of H. griffini (Holotype). 6, Male genitalia of H. chinatiensis. y) VoLUME 32, NuMBER 2 101 similarity to those of chinatiensis (Figs. 3, 4 & 6). In griffini, the medial process of the transtilla is frequently less sclerotized and not as stout as that of chinatiensis. In addition, the medial process appears narrower at the tip, with a slightly deeper groove than that found in chinatiensis. The apices of the valves are variable: most are rounded, while others have a more prominent constriction similar to, but less developed than, those of chinatiensis. Adult griffini are 25% smaller than those of chinatiensis. Phenotypi- cally griffini males (Figs. la,b) can be distinguished from those of chinatiensis (Fig. 3) in several ways: The basal black patch on the forewing of griffini is usually free standing or almost absent, while in chinatiensis the patch continues uninterrupted to the hind margin of the wing. On the dorsal surface of chinatiensis the area between the costa and radius adjacent to the discal cell is black while in griffini there is a white patch between the costa and radius adjacent to the discal cell. The viens proximal to the post medial line in griffini are narrow and usually black; the veins of chinatiensis are also black, but the scaling diffuses out from the veins giving them the appearance of being 3 to 4 times wider than those of griffini. The black margin of the forewings of chinatiensis is from % to 1% times wider than that of griffini. In general, the wings of griffini are approximately 40% black and 60% white while in chinatiensis they are about 60% black and 40% white. The dorsal portion of the abdomen of griffini may be covered with long rust red and/or black hairs. Thus, the abdomen may vary from black to light orange, with prominent rust red fringe around the anterior portion of each tergum. The last two segments are covered with long red hairs which form a tuft. The abdominal terga of chinatiensis are a uniform rust red in color. These are but a few of the characters which may be used to separate griffini males from those of chinatiensis. Although the males of griffini and chinatiensis are very distinct, the females are similar, with only a few obvious differences. The red abdominal banding, prominent in chinatiensis females (Fig. 4), is less developed or absent in griffini (Fig. 2). In griffini the red hairs are mixed with black hairs and spread randomly over the entire terga but become more abundant near the pleura, giving a diffused red appear- ance to the lateral surface, or the abdomen may be completely black except for the presence of short red hairs at the tip of the abdomen. At present griffini and chinatiensis are though to be allopatric, with the closest population of chinatiensis occurring 350 to 400 miles to the southeast, in western Texas. Examination of last instar griffini larvae indicated that they are similar 102 JOURNAL OF THE LEPIDOPTERISTS SOCIETY in most respects to Hemileuca. That is, secondary setae frequently, but not always, arise from a pinaculum which is either white or cream colored. Of 36 larvae examined, 31 had white pinacula, the remainder were colored, giving them an appearance similar to that of Pseudohazis. Thus, although most individuals appear similar to larvae of chinatiensis, some individuals appear similar to H. hera (Harris). After examination of the larvae of 16 of the 17 species of Hemileuca occurring north of Mexico, this character has been found to be variable only in the larvae of griffini. The two genera, Hemileuca Walker, 1855 and Pseudohazis Grote & Robinson, 1866 were joined by Michener in 1952. Michener based his decision on external morphological characters. Ferguson (1971) showed the intermediate characteristics of chinatiensis in which the genitalia are similar to Hemileuca but the adult phenotype is that of Pseudohazis. With the discovery and description of griffini, a second species with intermediate adult and larval characters has been found. The genitalia of griffini are more similar to genitalia of males of the Pseudohazis group than those of chinatiensis. In addition, the larvae of griffini exhibit mor- phological characters common to both species groups. Thus, a continuum of adult and larval characters exists between the two previously separated genera, Hemileuca and Pseudohazis. ACKNOWLEDGMENTS I would like to thank Mr. Bruce Griffin and Mr. Kenneth Hansen both of Tucson, Arizona, and Mr. Kilian Roever and Mr. Michael Van Bus- kirk of Phoenix, Arizona, for providing distributional data and/or speci- mens. LITERATURE CITED Fercuson, D. C. 1971. The Moths of America North of Mexico. Fasc. 20.2A, Bombycoidea (in part). Classey, London, pp. 101-153. MicHeNER, C. D. 1952. The Saturniidae (Lepidoptera) of the Western Hemi- sphere, Morphology, Phylogeny, and Classification. Bull. Amer. Mus. Nat. Hist. 98(5): 335-502. TinkHaAM, E. R. 1943. Description and biological notes on a new saturniid of the genus Pseudohazis from the Big Bend region of Texas. Can. Ent. 75(9): 159-162. Journal of the Lepidopterists’ Society 32(2), 1978, 103-106 THE STATUS OF OLLIA PARVELLA DYAR: REDESCRIPTION OF IT IN A NEW GENUS (PYRALIDAE) ANDRE BLANCHARD P.O. Box 20304, Houston, Texas 77025 ABSTRACT. Ollia parvella was described from females only. The discovery of a few males shows that it is not a Peoriine but a Phycitine and that it belongs in a new genus: Welderella. When Shaffer (1968) revised the North American Anerastiinae (Auc- torum), he grouped the majority of the species in the subfamily Peoriinae and returned most of the remaining ones to the Phycitinae. A few genera and species remained unplaced and were listed as such at the end of his revision. Blanchard and Ferguson (1975) included three of these unplaced species in the new phycitine genus Rostrolaetilia together with seven new species which were described in the same paper. Ollia parvella Dyar is another species which Shaffer could not place, in this case because no male was available. My wife and I took six specimens of this species (4 males and 2 females) 3 and 5 July 1975 at the Welder Wildlife Foundation Refuge. Through the courtesy of Dr. D. C. Ferguson I was able to borrow from the National Museum two paratypes of the six females which constitute the type series. The comparison of the genitalia of one of my females (slide A.B. 3879) with those of one paratype (slide U.S.N.M. 52945) leaves no doubt that my specimens are conspecific. As Shaffer had suspected, an examination of the male genitalia shows that this species is a Phycitine, although nothing closely related is in- cluded in Heinrich’s revision of this subfamily (1956). Ollia is a synonym of Peoria, not a Phycitine genus. Obviously a new genus is needed. Welderella A. Blanchard, new genus Type species: Ollia parvella Dyar (Figs. 1, 2, 4, 6). Labial palpi porrect, downcurved, extending over three times eye diameter be- yond front, loosely scaled; from beneath seen to be in contact with each other for nearly all their length; second segment two and a half times longer than the third. Maxillary palpi short, squamous. Antennae simple. No ocelli. Forewing smooth, broadest at two thirds distance from base to apex; apex and tornus rounded. Cell about two thirds length of wing. Venation somewhat vari- able, 10 or 11 veins: R;, Rs and R; normally united, but two specimens show on one wing R; separating from R;.; as a faint spur; M. and Ms; stalked for about two fifths their length; Cu: from lower outer angle of cell; Cu. from near the angle. Hind wing: length of cell ill defined (discocellular vein obsolete) but apparently slightly longer than half the length of the wing; Sc and Rs long stalked, Sc separates from Rs as short spur going io costa, Rs continues to near apex; M, straight to 104 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Figs. 1-5. Welderella parvella: 1, male; 2, female; 3, male genitalia; 4, de- nuded labial and maxillary palpi, showing rudimentary tongue; 5, female genitalia. outer margin; M. absent; M; and Cu, stalked for about half their length; Cu, in almost exact prolongation of cubitus. Male genitalia (Figs. 3, 7): Uncus triangular; apex produced, rounded, covered with bristles. Gnathos stout, with very large flanged apical processes; the lobes fusing posteriorly. Aedeagus smooth, moderately stout, about three times as long as maximum width, vesica without cornuti. Tegumen with strong supporting struc- ture forming on each side a wide U, with one branch supporting the gnathos and the other the dorsal processes of the uncus. Vinculum mostly membranous, sup- ported in part by the tegumen and the shallow, wide, well sclerotized saccus. Transtilla incomplete, represented by a pair of irregularly shaped plates. Juxta with anterior margin heavily sclerotized. Valves simple, without clasper; sacculus narrow and short. Female genitalia (Fig. 5): bursa and ductus bursae membranous; ductus bursae rather wide, broadening progressively into pear shaped bursa; signum well sclero- tized, longitudinally infolded; some weak scobinations around it; ductus seminalis from left side of signum; genital opening wide, funnel shaped, sclerotized and scobinate ventrally. VOLUME 32, NUMBER 2 105 mam, 0.5 mm. Figs. 6, 7. Welderella parvella: 6, venation; 7, enlarged part of male genitalia. This genus shares characters with two widely separate groups of Phycitine genera. The male genitalia suggest that it should go near Laetilia: the uncus, the gnathos, the transtilla are quite similar, but the complete absence of ocelli and the longitudinal wing pattern point to a placement near Bandera and Tampa. I take great pleasure in naming this new genus for the Welder Wild- life Foundation, for its staff, whom I have always found ready to help me in every way and for its generous founders, the late Rob Welder and his wife Bessie Welder. Welderella parvella (Dyar) Dyar, 1906, p. 31. Barnes & McDunnough, 1917, p. 149. McDunnough, 1939, p. 36. Kimball, 1965, p. 250. Shaffer, 1968, p. 89. The original description reads: “Costal half of fore wing white with slight darker lines on the veins toward apex. Inner half pale ocherous, shading to gray next to white part. Hind wing whitish.” The Welder Wildlife Refuge females match this description, but the hind wing of the male is whitish only in the one third of it along the inner margin; the other two thirds are blackish gray. Wing expanse: males 12.5—-14 mm., females 13-15.5 mm. Type data: I have not examined the holotype, a female from Brownsville, Texas, June 3 (?), 1904, H.S. Barber, U.S.N.M. type No. 9103; genitalia slide No. 10, Carl Heinrich, Dec. 20, 1932. Specimens examined: Brownsville, Texas, 31 May 1904, 1 2; 8 June 1904, 1 2; (Slide U.S.N.M. No. 52945), both collected by H.S. Barber. Welder Wild- life Foundation Refuge, 3 July 1975, 2 ¢¢4 (slides A.B. 3894, 3854, 3890; these last two slides are all that remains of one of these males), 5 July 1975, 2 4 4, 106 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 2 292 (4 slide A.B. 3857, 2 slide A.B. 3879), all collected by A. & M. E. Blanchard. ACKNOWLEDGMENTS I am indebted to Dr. D. C. Ferguson for arranging the loan of two female paratypes from the National Museum collection and to Dr. J. C. Shaffer for revising the manuscript. LITERATURE CITED Barnes, W. anv J. H. McDunnoucu. 1917. Checklist of the Lepidoptera of Boreal America. BLANCHARD, A. AND D. C. Fercuson. 1975. Rostrolaetilia—A new North Amer- ican genus of the subfamily Phycitinae, with description of seven new species (Pyralidae). Jour. Lepid. Soc. 29: 131-150. Dyar, H. G. 1906. Description of new American moths. J. N. Y. Ent. Soc. 14: 30-31. Hernricu, Cart. 1956. American moths of the subfamily Phycitinae. U.S. Nat. Mus. Bull. 207, viii + 581 p. Kimpati, C. P. 1965. Arthropods of Florida and neighboring land areas, I: Lepidoptera of Florida, an annotated checklist. McDunnoucu, J. 1939. Checklist of the Lepidoptera of Canada and the United States of America, part 2 (Microlepidoptera). Mem. S. Calif. Acad. Sci., vol. 2, no. l. SHAFFER, J. C. 1968. A revision of the Peoriinae and Anerastiinae (Auctorum ) of America north of Mexico. U.S. Nat. Mus. Bull. 280, vi + 124 p. Journal of the Lepidopterists’ Society 32(2), 1978, 107-110 BIONOMIC NOTES ON THE BLOOD-SPOT SKIPPER [HESPERIIDAE: PHOCIDES LILEA SANGUINEA (SCUDDER) | RAyMonp W. NECK Pesquezo Museum of Natural History, 6803 Esther, Austin, Texas 78752 ABSTRACT. Observations on the life cycle of Phocides lilea sanguinea are re- ported. An additional larval foodplant, notes on egg and larval stages, and adult oviposition behavior are described. Phocides lilea sanguinea (Scudder) is a large-sized skipper which exhibits a metallic blue background with a prominent red spot on each dorsal forewing. Basically a tropical species, sanguinea has established breeding populations in the Brownsville, Cameron County, Texas area. This population has been known as Phocides polybius lilea (Reakirt ) but H. A. Freeman (in litt.) prefers treatment of lilea at the species level with subspecific rank being accorded sanguinea. My initial interest in this species arose because of the significance of establishment of per- manent populations on non-native larval foodplants. Field observations have yielded various new bionomic facts concerning this insect. The only plant previously known to support larvae of sanguinea in either the United States (Lipes, 1961) or Mexico (Comstock and Vasquez, 1961; Kendall and McGuire, 1975) is common guava, Psidium guajava (Myrtaceae). My observations in Brownsville have revealed that a con- generic plant, Psidium cattleianum Sabine (strawberry guava) is also utilized as a larval foodplant by sanguinea. Identification of the food- plant was verified using comparative herbarium specimens and _ char- acters given by Bailey (1949: 729). All references below to Psidium refer to P. cattleianum. Newly-laid eggs are a pale but distinct aqua in color. A glistening wet appearance is noticeable for several minutes following oviposition. Within eighteen hours of oviposition, the contents of the egg begin to turn reddish as embryogenesis proceeds. Red coloration appears initially as individual foci which enlarge until the whole egg appears red. Similar egg color changes during development have been reported in Agathymus (Roever, 1964). This red color involves the internal constituents only, as the chorion appears whitish in color. The egg is hemispherical in shape with a diameter of approximately 1.5 mm. A hole encompassing the top one-third of the egg reveals a hatched egg. The egg shell is not eaten by the larva. 108 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Larvae of sanguinea go through a dramatic change in coloration during development. First instar larvae have a bright red body similar in color to the mature egg, but the head is brownish, varying extensively in darkness. Body length is 3.0 to 3.25 mm while the head capsule measures approximately 1.0 in width. Second instar larvae are about 8.0 mm long with a head capsule of 1.5 mm in width. The head of the second instar remains basically brown but is somewhat closer in coloration to the red body color than is the head of the first instar larvae. Larvae of Mexican and South American populations of Phocides were reported to have yellowish intersegmental bands (respectively, Comstock and Vasquez, 1961; Miles Moss, 1949); no such bands were observed on Texas larvae. Larvae of intermediate instars were not observed. Mature larvae are whitish with a slight “bloom” similar to that reported for Phocides pygmalion okeechobee (Worthington) by Srohecker (1938). This “bloom” is a white, powdery exudate which is present on the body of the larva. The body also exhibits a large number of black “pin-prick” marks. The head capsule is reddish-brown with a yellowish “eyespot” on each side. Retreats are formed by the larvae utilizing leaves of the foodplant. Miles Moss (1949) described initial shelters as “small oval, of cutleaf” while older larvae were “content to hide by day between several leaves held together by a few strands of glutinous silk.” The following observa- tions add to the above information: neither Lipes (1961) nor Kendall and McGuire (1975) mention these retreats per se. Immature larvae make two cuts from the leaf margin inward about five mm. This leaf section is then pulled flat over toward the midrib so that a small retreat is formed. This folded-over upper portion is then attached to the lower portion by a plug of silk. The flatness of the retreat in comparison to that formed by Calpodes ethlius (Stoll) on Canna is probably caused by the thick stiff nature of mature Psidiwm leaves as opposed to the thin, pliable leaves of Canna. While most retreats are formed from two- leaves as reported by Comstock and Vasquez (1961), one retreat consisted of three leaves—the apical pair and one of the leaves on the penultimate node. The larva rested on the top leaf or “ceiling” in an upside-down orientation. Mortality is extremely high in the early instars. A check of a single Psidium shrub yielded twenty-seven hatched eggs, but only five living first and second instar (and dead bodies of seven other) larvae in larval folds. Construction of the larval retreat is a task which many sanguinea larvae are not able to complete. Leaves supporting hatched eggs but no VoLUME 32, NUMBER 2 109 larvae occasionally have a single cut similar to the two required to form a retreat. Only a few chrysalids were observed during these studies. The chrysalis is loosely attached to the upper part of the retreat by silk strands. A parasitized chrysalis with many wasp (probably Apanteles) exit holes was found in one retreat. This chrysalis measured 28 mm in length and 8.25 mm at greatest width (second abdominal segment ). Adult female flight behavior in the vicinity of Psidium is quite dis- tinctive. The imago “flits” or “skips” along and above the periphery of the bush. Suddenly the adult will drop to the level of the bush and quickly land on a leaf. Always facing outward toward the tip of the leaf, she quickly lays an egg, and flies upward and around the bush again. If undisturbed, the behavior sequence is repeated. Although both male and female adults were observed flying, only females alighted on any substrate. All but one of these observations in- volved ovipositional landings on Psidium leaves. One female was ob- served to land on a leaf of a bottlebrush shrub, Callistemon citrinus Staph. (= lanceolatus DC.), which is also an introduced member of the Myrta- ceae. After remaining on the leaf for about ten seconds, the adult flew off; no egg was laid. The relatively long period of time spent on the bottlebrush leaf indicates that after initial selection of a prospective plant from a short distance (about 0.5 m) in the air, confirmation of proper ovipositional substrate is required on the leaf surface. The means of confirmation is unknown at this time, but it can be made in a very short time if the substrate possesses the correct phytochemicals. Placement of the eggs is highly predictable as a result of the rigid ovipositional behavioral sequence. Of twenty-seven egg shells found on 23 December 1970 all but one were on the upper surface of the leaf. Terminal leaves tend to be selected for oviposition. Numerical data and probable adaptive significance of this selection will be presented else- where. Adults in July were observed visiting flowers of an introduced orna- mental, lilac-flowered golden dew drop (Verbenaceae: Duranta repens L.). Adults expressed no interest in the flowers of the bottlebrush shrub tentatively selected for oviposition. Knowledge of the seasonal presence of the various life history stages of sanguinea is desirable. Observations of a mature larva revealed no apparent movement or feeding during a ten-day period from Dec. 1970- Jan. 1971. Weather conditions at this time were quite mild; live eggs and immature larvae were present at the same time. In contrast, De- cember 1976 was cold and wet; no eggs or larvae were found on the same bush at that time. 110 JOURNAL OF THE LEPIDOPTERISTS SOCIETY LITERATURE CITED Bartey, L. H. 1949. Manual of cultivated plants. Rev. ed. Macmillan Co., New York, 1116 pp. Comstock, J. A. & L. G. Vasquez. 1961. Estudios do los circlos biologicos en lepidopteros Mexicanos. An. Inst. Bil. Mex. 31: 349-448. KENDALL, R. O. & W. W. McGuire. 1975. Larval foodplants for twenty-one species of skippers (Lepidoptera: Hesperiidae) from Mexico. Bull. Allyn Mus. 27: 7 p. Lives, J. E. 1961. More butterfly records from Brownsville, Texas, including a foodplant of Phocides polybius (Hesp.). J. Lepid. Soc. 15: 114. Mites Moss, A. 1949. Biological notes on some Hesperiidae. Acta Zool. Lilloana 7: 27-48. Roever, K. 1964. Bionomics of Agathymus (Megathymidae). J. Res. Lepid. 3: 103-120. STROHECKER, H. F. 1938. The larval and pupal stages of two tropical American butterflies. Ohio J. Sci. 38: 294-295. Journal of the Pepidonicnss Society 32(2), 1978, 111-115 CLIMATIC REGIMES RESULTING IN UNUSUAL OCCURRENCES OF RHOPALOCERA IN CENTRAL TEXAS IN 1968 RAYMOND W. NECK Pesquezo Museum of Natural History, 6803 Esther, Austin, Texas 78752 ABSTRACT. During 1968 several species of Rhopalocera which normally do not occur in this area (or occur only in small numbers during the latter part of the season ) were abundant in central Texas. Meteorological regimes which influenced this influx of tropical species are discussed. Central Texas is a major ecotonal area between the Nearctic and the very northern fringes of a dilute Neotropical rhopaloceran element which appears as far north as Central Texas under various climatic regimes. Some of these species may occur as far north as Kansas or Nebraska as stragglers, but breeding populations are not established. One such climatic regime occurred in 1968. Discussed below are observations made at the Brackenridge Field Laboratory of the University of Texas at Austin within the corporate limits of the city of Austin. Heliconius charitonius vasquezae Comstock and Brown and Dryas julia moderata (Stichel) (both Heliconiidae) were very common in 1968 as early as June. The existence of numerous fresh specimens and the length of time during which these two species were present (well into the fall months) indicate that breeding colonies of both species had become established. At least one suitable larval foodplant, Passiflora lutea L. ( Passifloraceae ), is present on the grounds of the field laboratory. Noc- turnal roosting aggregations (up to eleven individuals) of H. charitonius were observed at several sites within the eighty-acre area. Different sites were used at various times. Each site was a shrub or tree at the edge of a wooded area. These two heliconians are seen in the Austin area in about half the years, but they normally appear in late summer or fall indicating late season dispersal from areas to the south with perma- nent populations. Sporadic breeding occurs in at least some of these years as indicated by reports for both species in 1966 (Rickard, 1967, 1968 ). Dynamine dyonis (Geyer) occurred commonly from July into the fall. Specimens were normally restricted to wooded areas along a dry arroyo. Breeding occurred as indicated by the fresh condition of most specimens seen throughout the season. D. dionis was initially reported in the Austin area in 1899 (Brues, 1905). A breeding population was established “along the bed of a dried up creek near Austin, Texas.” Brues had not 112 JOURNAL OF THE LEPIDOPTERISTS SOCIETY seen this species before this time (time of initial observations in Austin area by Brues unknown) and did not see it during the three following years, although he collected in the same areas. As his first records were in October, this species probably immigrated into the area in late summer 1899; the progeny of these immigrants were observed by Brues. Brues (1905) records that Mestra amymone (Menetries) was “very common about Austin .. . in former years they were much less numerous.” M. amymone is a permanent resident of the Austin area (Masters, 1970) but was exceptionally abundant in 1968. Adults of Achlyodes thraso tamenund (Edwards ) (Hesperiidae) were commonly seen resting on soil surfaces in open areas in 1968. Kendall (1965) reported this skipper from southern Texas. Records included Goliad, Kleberg, Live Oak and San Patricio Counties (all well south of the Austin area). Hurricane Beulah struck the Texas coast near the mouth of the Rio Grande River on 20 September 1967. Torrential rains covered a large area of south Texas resulting in floods and semi-permanent ponds of water (Grozier, et al., 1968; Baker, 1971). Subsequently, many rhopalo- ceran species not before known from Texas (or the United States) were reported from south Texas, particularly from Cameron and Hidalgo Counties at the southern tip ( Doyle, 1970; Heitzman, 1970; Heitzman and Heitzman, 1972; Kendall, 1970, 1972). The effect of this storm upon the rhopaloceran fauna in south Texas is well-documented and was more than temporary as some species have been found in subsequent seasons (Tilden, 1974). While this storm may have been related to the unusual rhopaloceran occurrences in central Texas in 1968 in an indirect manner, the major cause involves the weather of 1968. The species involved in this temporary northward movement could have occurred in the central Texas area in late 1967 and remained undetected, but their survival of the 1967-68 winter in these latitudes is most unlikely. Coldest temperature at Austin for this winter was 22°F. which is certainly much too cold for these species to survive. Average monthly temperature for winter 1967-68 were generally below normal in southern and central Texas. If these species could survive such cold weather, these taxa would be common in central Texas during many Se€aSons. Torrential rains associated with Hurricane Beulah allowed the develop- ment of larger than normal populations of these species in areas closer to central Texas than is normally the case (southern Texas and/or northern Mexico). The key to the appearance of these species in early 1968 was higher than average rainfall in May 1968 at Austin (8.75” vs. normal VoLUME 32, NUMBER 2 ans 4.22”) and many other localities in central Texas, especially along the Balcones Escarpment (scattered localities actually had lower than normal totals). Rains and cooler than normal temperatures during the summer (U.S. Weather Bureau, 1968, Climatological Data, Texas, p. 73) fostered plant growth and development of certain rhopaloceran populations. Rainfall at Austin in summer 1968 was 18.5% above normal and high temperature was only 98°F. Population movements from south Texas or, more likely, from northern Mexico along the Balcones Escarpment was initiated by large resident populations; survival of these populations was allowed by favorable moisture conditions. None of these forms was seen in central Texas in early 1969 (low temperature during 1968-69 winter was 22°F.). A second northward movement was observed in fall 1968. These move- ments may have involved species more common in northern Mexico at the end of the rainy season. Population development in northern Mexico probably reached levels such that northward population movements ensued. Climatic conditions of central Texas may not have been signifi- cant in the occurrence of the following form as no evidence of repro- duction in these relatively northern areas is known. Rainfall in the Border Country of Texas was up to twice normal during this fall period (Posey, 1968; Wagner, 1969). Colias (Zerene) cesonia (Stoll) is one of the common species in Austin, particularly in spring and fall. This species is normally represented in central Texas by the nominate subspecies which exhibits a reasonably obvious “dog face” even in the female. Several female specimens of C. cesonia collected during October 1968 exhibited a different phenotype, immacusecunda Gunder, with greatly reduced black markings on DFW and DHW. Originally described as a “form 2°” (Gunder, 1928), this form has been treated as an aberration by some authors (Brown, 1965). I believe this form represents a normal (seasonal?) phenotype present in certain populations in Mexico (Neck, pers. obs.). Under certain climatic conditions this form moves northward into Texas. Other observers reported occurrences of tropical or sub-tropical species in the central Texas area in 1968. Biblis hyperia aganisa Boisduval was seen in San Antonio, Bexar County by W. Tyron on 7 October 1968 (Tilden, 1974), Kendall (1972) reviewed the occurrence of the tropical heliconian, Eueides cleobaea zorcaon (Reakirt), in southern Texas as far north as San Antonio in 1968. In an attempt to verify that a particular climatic regime was responsible for the occurrence of the above butterflies in central Texas, weather records for 1899 and 1966 were consulted to determine if peculiar climatic 114 JOURNAL OF THE LEPIDOPTERISTS SOCIETY conditions existed at the time of previous occurrences in central Texas of some of the above species. In 1899, statewide precipitation in Texas was about normal, being 94% of average, but summer precipitation was high, 131% of average (Norquest, 1941). Precipitation records for particular stations for 1899 are not published in readily available form. However, a massive rainstorm occurred in late June 1899 in central Texas. This storm, which produced the worst flood on record for the Brazos River, was centered north of the Austin area, producing over thirty inches of rain in certain areas (Texas State Almanac, A. H. Belo Corp.). If rain from this storm was distributed, albeit in smaller amounts, to the south, then this storm with its attendant rainfall and wind circulation could have been involved in the occurrence of D. dyonis in central Texas in 1899. Ironically the oc- currence of the subtropical butterfly D. dyonis in central Texas in autumn 1899 followed the most severe Texas winter on record. Record low temperatures were recorded at various stations including 12°F. on 13 February at Brownsville at the southern tip of Texas. Central Texas tem- peratures were at or below 0°F. Brues (1905) remarked that “the summer had been very favorable for the development of insects as Hymenoptera and Diptera were more abundant than I have ever seen them in that part of the country.” Occurrence of breeding H. c. vasquezae and D. j. moderata, in central Texas in October 1966 (Rickard, 1967, 1968) probably resulted from heavy rains in August of that year. Following below normal rainfall during the first seven months of 1966, August was excessively wet for both Austin (6.21” vs. normal 2.17”) and San Antonio (4.28” vs. normal 2.25’). Such a large influx of moisture is quite sufficient to cause rapid vegetative growth and large scale movements of various butterfly pop- ulations. Environmental changes over large areas such as southern Texas and northern Mexico provide natural experiments which reveal factors which control the extent and abundance of occurrence of particular species (see earlier report by Gilbert, 1969). Obviously, two major climatic regimes of present-day central Texas prevent the establishment of a fringe Neo- tropical fauna: insufficient rainfall and relative extreme winter cold. LITERATURE CITED Baker, ki. T., Jn. 1971. Relation of ponded floodwater from Hurricane Beulah to ground water in Kleberg, Kennedy, and Willacy Counties, Texas. Texas Water Development Board, 138: 33 pp. Brown, K. S., Jn. 1965. Some comments on Arizona butterflies (Papilionoidea). J. Lepid. Soc. 19: 107-115. VoLUME 32, NUMBER 2 1S) Burs, C. T. 1905. The occurrence of a tropical butterfly in the United States. Entomol. News 16: 11-12. Doyte, J. E., II. 1970. Field notes on three skippers in Texas (Hesperiidae). J. Lepid. Soc. 24: 212. GiutBerT, L. E. 1969. On the ecology of natural dispersal: Dione moneta in Texas (Nymphalidae). J. Lepid. Soc. 23: 177-185. Grozier, R. U., D. C. Haut, A. E. Hutme & E. E. ScHROEDER. 1968. Floods from Hurricane Beulah in south Texas and northeastern Mexico, September—October 1967. Texas Water Development Board Report, 83: 197 pp. Gunper, J. D. 1928. Progeny of female collected in Achiote, Panama. that variant is likely to be genetically homologous with the one reported here. Specimens of this aberration have been deposited in the collections of the Museum of Comparative Zoology, Harvard University; Peabody Museum of Natural History, Yale University; American Museum of Natural History, New York; National Museum of Natural History, Wash- ington, D.C.; and G. B. Small, Panama Canal Zone. Journal of the Lepidopterists’ Society 32(2), 1978, 138-139 A GYNANDROMORPH OF PAPILIO POLYXENES (PAPILIONIDAE ) Cultures of Papilio polyxenes asterius Stoll are maintained in our laboratory for use in ecological studies. In October 1975, a bilateral gynandromorph of this species ap- peared in the second generation of a laboratory culture derived from populations around Brooktondale, Tompkins County, New York. This was the first such specimen observed. Although the external genitalia are male, only the right half of the specimen is male in appearance (Fig. 1). The yellow spots of the inner row of the right forewing Fig. 1. Lab-reared gynandromorph of Papilio polyxenes, dorsal view. are greatly reduced and/or appear as “ghosts” composed of scales intermediate in color between black and yellow. The inner band of the hind wing resembles a normal male except for the last yellow spot (Cu: cell) which is half obliterated by black scales. The hazy blue spots of the hind wing are irregular and appear in cells Cui, Cue, Me, and Ms. The left half of the specimen perfectly resembles a female. The left forewing is 43 mm in length, the right is 41 mm. Instances of gynandromorphism among the swallowtails are rare. Schmid (19 73)-Can. Entomol. 105: 1549-1551) describes natural gynandromorphs of Ornithoptera victoriae Gray and O. priamus L. Skinner (1919, Entomol. News 30: 247) and Cockayne (1935, Trans. Roy. Entomol. Soc. Lond. 83: 509-522) refer to Papilio glaucus L. gynandromorphs. Hybrid crosses between P. polyxenes and other swallowtails in the machaon group have in some cases yielded gynandromorphic individuals (Clark and Sheppard, 1953, Suppl. Entomol. Rec. 65: 1-12; Ae 1964, Bull. Jap. Entomol. Acad. l: 1-10). Edwards (1868-1872, The Butterflies of North America; Philadelphia: VoLUME 32, NUMBER 2 139 Am. Entomol. Soc.) presents a figure of an apparently gynandromorphic P. polyxenes, but offers no data on the specimen. It is not known what caused the butterfly described here to be a gynandromorph. Gardiner (1972, J. Res. Lep. 11: 129-140) notes that the incidence of gynandromor- phism in Pieris brassicae L. cultures is associated with outbreaks of virus and suggests that viral disease may cause such genetic abnormalities. Viral disease is commonly present at low levels in our cultures of P. polyxenes and may account for the appear- ance of this unusual individual. The specimen is located in Lot 1062 of the Entomological Collections at Cornell University. I wish to acknowledge the support of N.S.F. Grant DEB 76-20114 (to Paul P. Feeny ) which covered costs of publication. Witt1uaM S. Biau, Department of Entomology, Cornell University, Ithaca, New York 14853. Journal of the Lepidopterists’ Society 32(2), 1978, 139-140 ELECTROSTRYMON ANGELIA ANGELIA (LYCAENIDAE): THE OLDEST FLORIDA RECORD? The butterflies, Sphingidae and Castniidae from the Strecker collection were trans- ferred from the Field Museum of Natural History to this institution in 1976 on semi- permanent loan in order that they might be more readily utilized by students of these groups. Since that time workers here and elsewhere have made greater use of the Strecker and Reakirt material contained in Strecker’s collection. That collection is, however, an aggravating one by modern standards: there are few labels on individual specimens; rather, the series are labelled with data that in theory apply to all members under the label. This is certainly not the case in all instances. Fortunately, Strecker prepared catalogs to the Papilionidae, Pieridae and Lycaenidae before his death, and in these families the data for individual specimens are recorded. A second problem involves Strecker’s apparently too-eager interpretation of what specimen was what, and from where. Some of the putative Reakirt types in the collection may not be those, and we suspect that Strecker was easy prey for dealers who peddled material mislabelled by locality. The situation with the Strecker col- lection is by no means as bad as that with some other older collections, notably the Ehrmann collection presently housed at Carnegie Museum of Natural History. Nevertheless, the Strecker collection contains some magnificent material—material that is not duplicated in other North American collections. Further, the Strecker collection, with its associated letters, is an historical document. The Lycaenidae, because that family was one for which the catalog was completed, are especially inter- esting. In working through the hairstreaks in the Strecker collection, specimens of Electrostrymon angelia (Hewitson) were found under the label “Thecla hugon Godart’, a synonym of Electrostrymon endymion (Fabricius ). Two of these specimens were placed in the collection after Strecker had compiled the catalog and are indi- vidually labelled “Haiti” and “Port au Prince, Haiti’; both of these butterflies are specimens of the Hispaniolan subspecies boyeri (W. P. Comstock and Huntington). The other specimen is labelled characteristically with an “a”, referring to an entry in the catalog. This specimen is here figured (Fig. 1) and is referable to the Cuban E. a. angelia. The catalog states that the specimen was from “Florida” and that 140 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Fig. 1. Electrostrymon angelia angelia (Hewitson), ¢ upper (left) and under (right) surfaces; FLORIDA, Chas. Dury (Strecker collection). Allyn Museum photos 071477-15/16. Strecker received it from Charles Dury. No date is given, but it is likely that the specimen was taken before 1880. Kimball (1965, Lepidoptera of Florida) does not list Dury among the pioneer Florida collectors, and it is possible that while Strecker received the angelia from Dury, Dury himself may not have collected it. Were it not for the fact that Anderson (1974, J. Lepid. Soc., 28: 354-358) had recorded this species from the Florida Keys, and others have reported it from as far north as the Fort Lauderdale area on the east coast of the state, it would be tempting to dismiss the Strecker specimen as a hoax or a mislabelled specimen. {! suspect, though, that the Dury/Strecker specimen is an authentic one, and quite possibly angelia long has been a member of the Florida fauna, though perhaps not so commonly as in the past few years. It is further likely that specimens of angelia may have been responsible for the long- standing records of E. endymion from Florida. The specimen that Holland (1931, The Butterfly Book: pl. 64, Fig. 32) figured as “endymion” was actually a specimen of E. angelia boyeri (Klots, 1951, Field Guide to the Butterflies ...: 281), and the latter author expressed doubt about the occurrence of endymion in Florida. Riley (1975, Field Guide . . . Butterflies of the West Indies) does not mention endymion from the West Indies, thus strongly suggesting that the species never has occurred in Florida. Lee D. Mitter, Allyn Museum of Entomology, 3701 Bay Shore Road, Sarasota, Florida 33580. Journal of the Lepidopterists’ Society 32(2), 1978, 140-141 OBSERVATIONS ON ERORA LAETA (LYCAENIDAE) IN NEW HAMPSHIRE Erora laeta Edwards, often considered the rarest of eastern butterflies, is eagerly sought by many lepidopterists; too often with negative results. I made my first trip to New Hampshire to capture this species on 21 May, 1977 with fellow collector Reginald Webster. We visited a few areas in Carroll County, near Bartlett, in northern New Hampshire, where R. W. had taken one specimen the previous year. VoLUME 32, NUMBER 2 141 The location seemed typical for /aeta—an abandoned dirt road going through a beech woods. We hiked up the road until we reached a shallow gulley crossing the road. On the other side of the gulley, on damp dirt, was a female E. laeta, which was quickly netted. We then saw another female which was also easily caught. At this point, I was amazed, for we had hoped to find one or two E. laeta in a day of intense collecting, and had caught two in 10 minutes. Continuing along the road, we came to a little trickle of a stream where we caught two more laeta, both females. Upon returning to the first gulley, we caught three more females. Astounded by our luck we decided to try a few more dirt roads in the area. On every one, we found laeta. It was incredible; was this really that rarest of butterflies that we were seeing everywhere we looked? In about three hours of collecting we must have seen over 80 E. laeta, and had collected one or two from every road we tried, ending up with about 12 specimens apiece. When these numbers are com- pared to those reported in previous literature (e.g., Mousley, 1923, The Can. Entomol. 55: 26—29; Field, 1941, Ann. Entomol. Soc. of America 34: 303-316; Smith, 1960, J. Lepid. Soc. 14: 239-240; Roever, 1962, J. Lepid. Soc. 16: 1-4), our sightings seem truly phenomenal. The most specimens previously reported collected at one time were two males and nine females along the slopes of Mount Killington in New Hampshire (Field, op. cit.), and many reports are of individual specimens taken by chance (e.g., Sullivan, 1971, J. Lepid. Soc. 25: 295-296). Of all the individuals that we saw, only two were males, only one of which was captured. Among the females, some were quite worn, while others looked freshly emerged. There seem two likely reasons, not necessarily mutually exclusive, for the dearth of males: 1) the males had emerged earlier in the season and so most had already died, and 2) the males remained up in the trees, and only females came down to drink at the mud. However the male that was caught was freshly emerged, and in most species of butterflies, it is the males that are found “puddling’” (Downes, 1973, J. Lepid. Soc. 27: 89-99). The single male was caught in a field next to the woods. The females were very easy to catch, some not even flying up when the net was clapped over them. This behavior has been noted by previous authors (e.g., Hessel, 1952, J. Lepid. Soc. 6:34), but strongly impressed me when I nearly stepped on one female as I was walking down the road; it flew up from right under my foot. The males, on the other hand, were more restless and difficult to catch, their flight being fast and uneven, with only occasional landings on vegetation. Two possible explanations for the extraordinary abundance of E. laeta that we observed are as follows: It is possible that there was a real population explosion of E. laeta in New Hampshire in 1977 (W. Kiel, who has collected for years in New Hampshire, caught his first E. laeta this spring). It would be interesting to know if other lepidopterists found a similar increase in this species in other areas. An alter- native explanation is that /aeta is really not that rare, but that its behavior on this day was unusual. Perhaps the butterflies normally spend most of their time in the forest canopy, and thus are not accessible to collectors. This day being very sunny, hot (33°C), and humid, perhaps drove them down to the ground to drink. It may be that early emergence and a short flight period make E. laeta seem very rare, but that finding them is really a matter of being in the right place at the right time. At any rate, we were! DEANE Bowers, Dept. of Zoology, University of Massachusetts, Amherst, Massa- chusetts 01003. Journal of the Lepidopterists’ Society 32(2), 1978, 142-144 OBITUARY & WILBUR S. McALPINE (1888-1977 ) Mr. Wilbur S. McAlpine, charter member of The Lepidopterists’ Society, passed away on 30 July 1977, at the Grovecrest Convalescent Home in Pontiac, Michigan, at the age of 88. ‘Mac,’ as he was known by many of his friends, deeply enjoyed nature and the out-of-doors, and was a devoted amateur lepidopterist, spending most of his spare time collecting and studying Michigan moths and butterflies. He was particularly interested in the life history of many local Oakland County species and specialized in the genus Calephelis, the metalmark butterflies. Wilbur was born in Detroit, Michigan, on 30 December 1888, and graduated from Detroit Central High School in January, 1908. He held positions as a draftsman with the U.S. Lake Survey, Michigan Central Railroad, and Detroit Edison, and was also employed as an Assistant Surveyor of Coal Claims in Homer, Alaska during 1906, 1907, 1911 and 1912. He served with the military in the 472nd Engineers during the First World War in 1918. Wilbur became the principal and owner of a mapping, surveying and engineering business under the names of McAlpine Engineers, Inc. and W. S. McAlpine Map Co. from December, 1915, until his retire- ment in 1965, when he sold his business to the employees. They are still conducting the business under the same names. While operating his business, the firm produced complete maps of all Michigan counties, especially detailed maps of Oakland County. McAlpine published an Atlas of Oakland County, and engineered and recorded over two hundred subdivision plats and made numerous farm, residential lot and topographic surveys, which are still in use today. Undoubtedly, many of the Oakland County lepidoptera collected by McAlpine were discovered during his surveying activities. He witnessed the dis- appearance of many of his favorite collecting sites due to the suburban encroachment moving Outward from Detroit. McAlpine will always be remembered by mid-western lepidopterists as the one who described the Swamp Metalmark, C. muticum, and subsequently worked out its life history. In 1971, he culminated his intense interest in the Calephelis with his publica- tion On the revision of the genus, describing 25 new species and 7 new subspecies VoLUME 32, NUMBER 2 143 mostly from Mexico and Central America. McAlpine also worked out the life history of several other Michigan species, including Hyalophora columbia (Smith), Callophrys and Hesperiidae species, which unfortunately were never published. He was one of the first to locate several H. columbia tamarack bogs in southeastern Michigan, and eventually secured numerous cocoons and reared many in his backyard cages. McAlpine was a very determined Michigan butterfly collector who would travel anywhere, anytime to add a new species to his collection. Unfortunately, the many years devoted to his Calephelis project left him with little time for his personal collection and other lepidopteral plans and pursuits. One of his plans that failed to materialize due to his death was the publication of a guidebook of Michigan butter- flies, complete with colored plates of all known species found in the state. This project was a lifelong ambition to stimulate appreciation for and further the knowledge of Michigan butterflies, especially among young people. Two of his friends of long standing, Dr. George W. Rawson and John H. Newman, and this writer to serve as editor, were to collaborate with ‘Mac’. Although this project is still continuing, the extremely high cost of colored plates has made it necessary to re-adjust the original goal. In 1972, McAlpine donated the bulk of his collection, over 12,000 specimens, in- cluding many Calephelis type specimens, to the Smithsonian Institution. Later, he also donated approximately 800 moths and butterflies and 700 miscellaneous insects to the collection at Michigan State University at East Lansing, and a lesser amount to The University of Michigan at Ann Arbor. His collection, rich in Michigan material, included a long series of H. columbia and its cecropia hybrid, Colias interior (Scudder ) and Oeneis chryxus strigulosa (McDunnough ), and miscellaneous Alaskan lepidoptera and other insects. McAlpine was an Honorary Member of The Michigan Entomological Society and held memberships in the former Detroit Entomological Society, The Entomological Society of Canada and The Lepidoptera Research Foundation, Inc. He was affiliated with the Cranbrook Institute of Science in Bloomfield Hills, and was made a Life Member of The Michigan Society of Registered Land Surveyors on 11 February 1970. He travelled widely in the United States and Mexico and examined museum collections and collected Calephelis material; and also visited the British and Paris museums in connection with his metalmark studies. In addition to his interest in lepidoptera, ‘Mac’ was a devoted and active churchman. He particularly enjoyed evangelistic singing as a soloist, and sang in the choir at the First Baptist Church of Birmingham. He and his late wife, the former Minnie Burnett, are survived by one son, Wilbur Burnett, plus 16 nieces and nephews. Mrs. McAlpine died in January, 1975. I wish to express my appreciation to Wilbur's brother-in-law, Mr. Percy C. Burnett of Pontiac, and his minister, Dr. Glenn H. Asquith, Jr. for information in preparing this manuscript. BIBLIOGRAPHY OF WILBUR S. MCALPINE McAtpine, W. S. 1918. A Collection of Lepidoptera from Whitefish Point, Mich- igan. Occ. Papers Mus. Zool. Univ. Mich. 54: 1-26, 1 folding map. 1936. Habitat of Cissia mitchellii in Cass County, Michigan. Bull. Brook- lyn Entomol. Soc. 31: 110, 221. 1937. A Case of Mistaken Identity and Discovery of a New Metalmark (Calephelis) from Michigan. Bull. Brooklyn Entomol. Soc. 32: 43—49, 1 pl. 1938. Life History of Calephelis muticum (McAlpine), Lepidoptera. Bull. Brooklyn Entomol. Soc. 33: 111-121, 1 pl. 1939. A New Metalmark (Calephelis) from Texas (Lepidoptera, Riodin- idae). Bull. Brooklyn Entomol. Soc. 34: 75-80, 1 pl. 144 JOURNAL OF THE LEPIDOPTERISTS SOCIETY _§. P. Huspety AnpD T. E. Puisxe. 1960. The Distribution, Habits, and Life History of Euptychia mitchellii (Satyridae). J. Lepid. Soc. 14(4): 209-236, 3 pl. 1971. A Revision of the Butterfly Genus Calephelis (Riodinidae). J. Res. Lepid. 10(1): 1-125, 8 pl. 1972(1973). Observations on Life History of Oarisma powesheik (Parker ) 1870. J. Res. Lepid. 11(2): 83-93, 2 pl. M. C. Nietsen, Adjunct Curator, Department of Entomology, Michigan State University, East Lansing, Michigan 48824. Journal of the Lepidopterists’ Society 32(2), 1978, 144 BOOK REVIEW FREDERICK WiLLIAM FRoHAWK, by Valezina Bolingbroke, 1977. E. W. Classey Ltd. Park Rd. Faringdon, Oxon., England. 16 p., 3 figures (photographs), cover drawing in color by F.W.F. “< ... 1am going to attempt to give just a brief ‘impression’ of him,” writes the author on page 1 of this little memoir. The author is not an artist nor a writer as was Mr. Frohawk, nor a fellow naturalist, but his own daughter. It is obvious from the start that she felt for him the love and admiration which any father hopes to inspire in his daughter. The narrative is a series of glimpses into the past, rather like im- pressionist paintings seen from a distance, as Ms. Bolingbroke rambles through the English countryside with her readers. One sees an ancient house with moat and drawbridge, a little stone village, an old fashioned garden, a field of cowslips, travellers in a ‘pony trap’—all interspersed with charming and humorous incidents, flashes into the character of her father, his colleagues, his accomplishments and his deep love for - all of nature, from a very small butterfly to a very young human being. Frohawk was both author and illustrator of distinguished books on Ornithology and Lepidopter- ology, but his vast knowledge encompassed many other aspects of nature including wild flowers, reptiles and weather patterns, to name but a few. One is left wanting to know much more than this ‘brief impression’ gives us of F.W.F., as Frohawk was affectionately nicknamed by his friends. Jo Brewer, Editor, The News of the Lepidopterists’ Society, 257 Common St., Dedham, MA 02026. EDITORIAL STAFF OF THE JOURNAL AusTIN P. Puatr, Editor Department of Biological Sciences University of Maryland Baltimore County, 5401 Wilkens Avenue Catonsville, Maryland 21228 U.S.A. Dovuctas C,. Fercuson, Associate Editor THEODORE D. SARGENT, Associate Editor NOTICE TO CONTRIBUTORS Contributions to the Journal may deal with any aspect of the collection and study of Lepidoptera. Contributors should prepare manuscripts according to the following instructions. Abstract: A brief abstract should precede the text of all articles. Text: Manuscripts should be submitted in duplicate, and must be typewritten, entirely double-spaced, employing wide margins, on one side only of white, 8% x 11 inch paper. Titles should be explicit and descriptive of the article’s content, including the family name of the subject, but must be kept as short as possible. The first men- tion of a plant or animal in the text should include the full scientific name, with authors of zoological names. Insect measurements should be given in metric units; times should be given in terms of the 24-hour clock (e.g. 0930, not 9:30 AM). Underline only where italics are intended. References to footnotes should be num- bered consecutively, and the footnotes typed on a separate sheet. Literature Cited: References in the text of articles should be given as, Sheppard (1959) or (Sheppard, 1959, 196la, 1961b) and all must be listed alphabetically under the heading LirerAtTure Crrep, in the following format: SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London. 209 p. 196la. Some contributions to population genetics resulting from the study of the Lepidoptera. Adv. Genet. 10: 165-216. In the case of general notes, references should be given in the text as, Sheppard (1961, Adv. Genet. 10: 165-216) or (Sheppard 1961, Sym. Roy. Entomol. Soc. London 1: 23-30). Illustrations: All photographs and drawings should be mounted on stiff, white backing, arranged in the desired format, allowing (with particular regard to lettering ) for reduction to their final width (usually 4% inches). Illustrations larger than 8% x 11 inches are not acceptable and should be reduced photographically to that size or smaller. The author’s name, figure numbers as cited in the text, and an indication of the article’s title should be printed on the back of each mounted plate. Figures, both line drawings and halftones (photographs), should be numbered consecutively in Arabic numerals. The term “plate” should not be employed. Figure legends must be typewritten, double-spaced, on a separate sheet (not attached to the illustrations), headed EXPLANATION OF FicuREs, with a separate paragraph devoted to each page of illustrations. Tables: Tables should be numbered consecutively in Arabic numerals. Headings for tables should not be capitalized. Tabular material should be kept to a minimum and must be typed on separate sheets, and placed following the main text, with the approximate desired position indicated in the text. Vertical rules should be avoided. Proofs: The edited manuscript and galley proofs will be mailed to the author for correction of printer’s errors. Excessive author’s changes at this time will be charged to authors at the rate of 75¢ per line. A purchase order for reprints will accompany the proofs. Correspondence: Address all matters relating to the Journal to the editor. Short manuscripts such as new state records, current events, and notices should be sent to the editor of the News: Jo Brewer, 257 Common Street, Dedham, Massachusetts 02026 U.S.A. ALLEN PRESS, INC. eRINTED LAWRENCE, KANSAS US. CONTENTS STUDIES ON THE INTERACTIONS OF MORPHO PELEIDES (MORPHIDAE) with LecuMINosAE. Allen M. Young ae | Notes AND DESCRIPTIONS OF EUPTYCHINI (LEPIDOPTERA: SATYRIDAE) FROM THE Mexican Recion. Lee D. Miller LARVAL FoopPpLANT, Lire History NOTEs AND TEMPORAL DISTRIBU- TION FOR SPLENDEUPTYCHIA KENDALLI (SATYRIDAE) FROM Mexico. Roy O. Kendall: Notes ON THE LirE CycLeE AND NATURAL History oF VANESSA ANNABELLA (NYMPHALIDAE). Thomas E. Dimock A New Spectres oF HEMILEUCA FROM THE SOUTHWESTERN UNITED STaTEs (SATURNUDAE). Paul M. Tuskes __... ee | THe STATUS OF OLLIA PARVELLA DyArR: REDESCRIPTION OF IT IN A New Genus (Pyratmae). André Blanchard _ Bionomic NOTES ON THE BLOOD-SPOT SKIPPER | HESPERIIDAE: PHOCIDES LILEA SANGUINEA (SCUDDER)]. Raymond W. Neck __ CiimaTic ReEcIMES RESULTING IN UNUSUAL OCCURRENCES OF RHOPALOCERA IN CENTRAL TExAs IN 1968. Raymond W. Neck A New NAME FoR PAPILIO CERES CRAMER, [1776], Nec Fasrictus, [1775] (NymMpHALmpAE, DANAINAE). Gerardo Lamas _.... Two New PINE-FEEDING SPECIES OF COLEOTECHNITES (GELECHUDAE). Ronald W. Hodges and Robert E. Stevens _...... Lire History AND HaBits OF COLEOTECHNITES EDULICOLA (GELECHI- IDAE) A Pinyon NEEDLE MINER IN THE SouTHWEsST. Robert E. Stevens, J. Wayne Brewer, and Daniel T. Jennings _... Meyrick’s Recorp oF “MECYNA FURNACALIS, GN.” FROM FIJI, WITH A New GENERIC ASSIGNMENT FOR PYRAUSTA HOMALOXANTHA Meyrick (PyYRALIDAE: PyRAUSTINAE). Eugene Munroe and Akira Mutuura “ORANGE” BANDs, A SIMPLE RECESSIVE IN ANARTIA FATIMA (NYM- PHALIDAE). Annette Aiello and Robert E. Silberglied GENERAL NOTES Pieris Hadi oleracea (Pieridae) caught by insectivorous plant. Frances S. Chew i en Screech owl preys on Peridoma plecta (Noctuidae). Dwight G. Smith _. A gynandromorph of Papilio polyxenes (Papilionidae). William S. Blau .. Electrostrymon angelia angelia (Lycaenidae): The oldest Florida record? Lee D. Miller — a Observations on Erora laeta (Lycaenidae) in New Hampshire. Deane Bowers OBITUARY Se a HS Hf se fe Si eo Book REVIEW 65 75 86 88 97 103 107 111 116 118 123 130 a sa oe > te fe — Pies = he ee Sn gee ort. Volume 32 1978 Number 3 JOURNAL of the LEPIDOPTERISTS’ SOCIETY Published quarterly by THE LEPIDOPTERISTS’ SOCIETY Publié par LA SOCIETE DES LEPIDOPTERISTES Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 14 November 1978 THE LEPIDOPTERISTS’ SOCIETY | EXECUTIVE COUNCIL I. F. B. Common, President T. Surrézu, Vice President C. V. Covet, Jr., Ist Vice President Jut1an P. Donanue, Secretary L. A. GozmAny, Vice President RONALD LEUSCHNER, Treasurer Members at large: R. A. ARNOLD J. F. EMMEL C. D. FERRIs E. D. CAsHATT R. R. GATRELLE J. Y. MILLER R. E. STANFORD AE) Paar M. C. NIELSEN The object of the Lepidopterists’ Society, which was formed in May, 1947 and formally constituted in December, 1950, is “to promote the science of lepidopterology in all its branches, ... . to issue a periodical and cther publications on Lepidoptera, to facilitate the exchange of specimens and ideas by both the professional worker and the amateur in the field; to secure cooperation in all measures” directed towards these aims. Membership in the Society is open to all persons interested in the study of Lepidoptera. All members receive the Journal and the News of the Lepidopterists’ Society. Institutions may subscribe to the Journal but may not become members. Prospective members should send to the Treasurer full dues for the current year, together with their full name, address, and special lepidopterological interests. In alternate years a list of members of the Society is issued, with addresses and special interests. There are four numbers in each volume of the Journal, scheduled for February, May, August and November, and six numbers of the News each year. Active members—annual dues $13.00 Student members—annual dues $10.00 Sustaining members—annual dues $20.00 Life members—single sum $250.00 Institutional subscriptions—annual $18.00 Send remittances, payable to The Lepidopterists’ Society, and address changes to: Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 U.S.A. Back issues of the Journal of the Lepidopterists’ Society, the Commemorative Volume, and recent issues of the NEWS are available from the Assistant Treasurer. The Journal is $13 per volume, the Commemorative Volume, $6; and the NEWS, $.25 per issue. Order: Mail to Charles V. Covell, Jr., Memoirs Editor, Department of Biology, Uni- versity of Louisville, Louisville, KY 40208, U.S.A. The Lepidopterists’ Society is a non-profit, scientific organization. The known office of publication is 1041 New Hampshire St., Lawrence, Kansas 66044. Second class postage paid at Lawrence, Kansas, U.S.A. 66044. Cover illustration: Dasychira dorsipennata larva, dorsal and lateral views. From Fascicle 22.2, “Lymantriidae,” by Douglas C. Ferguson, in Moths of America North of Mexico. The drawing was done by E. R. Hodges, Scientific Illustrator, Department of Entomology, Smithsonian Institution. (Reproduced by permission of the author.) J OURNAL OF Tue Leprporprerists’ SOCIETY Volume 32 1978 Number 3 Journal of the Lepidopterists’ Society 32(3), 1978, 145-159 THE INFLUENCE OF ENVIRONMENTAL FACTORS ON ROOSTING IN THE BLACK SWALLOWTAIL, PAPILIO POLYXENES ASTERIUS STOLL (PAPILIONIDAE) JoHN Epwarp Raw ins! AND RoBERT C. LEDERHOUSE? ABSTRACT. Black swallowtails, Papilio polyxenes asterius Stoll, roost singly in west sloping old-fields in central New York. A frenetic search flight, triggered by decreasing radiation regardless of temperature, precedes roosting. Search flights allow swallowtails to check the suitability of various roosts. Roosts selected favor dorsal basking for as long as possible under decreasing solar radiation. Dorsal basking on the roost ceases, and a roosting posture is assumed when a body temperature high enough for flight can no longer be maintained. Selection for efficient roosting is strong since most adult deaths apparently occur when individuals are roosting. Selection for mimetic or cryptic wing patterns is expected to influence the exposed undersurface of the hindwing more strongly than other wing surfaces hidden during roosting. Butterflies are active during daylight hours when ambient conditions of radiation and temperature are high enough to allow behavioral maintenance of body temperatures suitable for flight (Vielmetter, 1958; Watt, 1968; Heinrich, 1972). As radiation and ambient temperature fall during the evening, butterflies seek night roosts where they remain until the next morning. Some species roost in aggregations (Crane, 1957; Urquhart, 1960; Benson and Emmel, 1973; Larsen, 1973; Muyshondt and Muyshondt, 1974; Turner, 1975; Young and Thomason, 1975), but most roost singly. Little is known of roosting behavior in most butterflies in spite of the widely read query concerning roosts in the introduction to Klots’ (1951:7) popular field guide. The popular literature is full of comments about butterflies disappearing into trees and fields during evening hours but no detailed study has ever been reported for a species which roosts singly. 1 Department of Entomology, Cornell University, Ithaca, New York 14853. Section of Neurobiology and Behavior, Cornell University, Ithaca, New York 14853. Current address: Department of Zoology and Physiology, Rutgers University, Newark, New Jersey 07102. 146 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Many butterflies bask in the sun as long as possible in the evening, otten on the actual perch that will be used as the night roost. Such behavior has been termed “vesper warming” by Clench (1966). Basking on the roost has been recorded in the hesperiids Thymelicus lineola Ochsenheimer and Ancyloxypha numitor Fabricius, the lycaenids Everes comyntas Godart and Lycaena phlaeas L., and the nymphaline Phyciodes tharos Drury (Clench, 1966). Powell (1968) records similar behavior for Incisalia iroides (Boisduval) (Lycaenidae). Several authors have suggested that some species choose roosts which are shaded in the setting sun and therefore illuminated by the rising sun earlier the following morning (MacNeill, 1964; Clench, 1966). Such behavior would favor earlier warming by basking the following morning and could conceivably lengthen the period of diurnal activity. Roosting in shadows cast by the setting sun has been observed in Hesperia spp. (MacNeill, 1964) and in the satyrines Euptychia rubricata Edwards and Cercyonis meadii Edwards (Clench, 1966). MacNeill described roosting Hesperia as flying directly away from the sun and roosting on the eastern side of bushes. Clench noted the movements of a Speyeria aphrodite F. (Nymphalidae). By lining a tree up with the setting sun and then flying directly toward the sun, the butterfly arrived at the shaded side of the intervening tree. After repeating this behavior three or four times, it flew off and roosted in a field. It remains unclear if such shade seeking during roosting is widespread among butterflies, let alone the mechansm by which this is accomplished. The environmental cues which release roosting behavior have been inadequately studied. When ambient radiation and temperature are low enough to prevent maintenance of a body temperature suitable for flight in the nymphalid Argynnis paphia (L.), roosting results (Vielmetter, 1958). This does not explain the onset of roosting on warm evenings when ambient temperatures allow sustained and active flight in the absence of direct insolation. Larsen (1973) has suggested that the timing of roosting events may depend more on decreasing radiation from the setting sun than on decreasing ambient temperature. 4 From the preceding it seems that selection of the roost site in a given species may involve one of three different roosting patterns: 1) Roosting in shadows cast by the setting sun, therefore hastening exposure to the rising sun the following morning, 2) roosting and basking in areas having maximal exposure to the setting sun, thereby prolonging the period when body temperatures suitable for flight may be maintained, or 3) roosting in areas not related to the position of the sun. The following study examines in detail the roosting behavior of the VOLUME 32, NUMBER 3 147 black swallowtail, Papilio polyxenes asterius Stoll, and determines what environmental cues direct such behavior. It constitutes the first study of roosting in papilionids [see observations of Gibson and Panchen (1975) on the African Papilio demodocus Esper] and the first detailed study of non-gregarious roosting for any butterfly. MATERIALS AND METHODS The black or parsnip swallowtail, Papilio p. asterius is frequently seen in open clearings and fields throughout eastern North America. The territorial males may be observed for lengthy periods in relatively small areas (Lederhouse, 1978). The darker females are more transient, usually observed flying quickly across these open areas. Roosting was studied in an old field near Brooktondale, southeast of Ithaca, Tompkins County, New York during the summer of 1975 (Fig. 3). This field was bounded on the north and east by corn fields and on the south and west by dense woody growth. The field was divided into a grid consisting of 276 squares each 10 meters on a side. The relative elevation of each corner of every grid square was established by sur- veying. Slopes and aspects (compass headings) for every grid square were determined by treating each square as a planar surface with corner elevations most nearly matching those measured. Plant densities for each square were measured by direct count for large, conspicuous species or by counting within a randomly placed 0.5 m? circle for smaller and more numerous species. Only the most abundant plant species are treated in this study. Plant nomenclature follows Fernald (1950). During each study period, gross solar radiation and ambient tem- perature were recorded using a centrally-located radiometer and hygro- thermograph. Sunset times were established by reference to published readings of the U.S. Naval Observatory for Binghamton, New York (U.S. Printing Office, 1959). Observations of individual behavior were made by persistently fol- lowing one swallowtail during the evening hours until it selected a roost. This was difficult due to the rapid and erratic flight of individuals seeking roosts; more often than not the butterfly was lost from sight. Additional data were obtained by systematically walking over the entire grid during dusk noting the location and microhabitat of individuals already on roosts. Observations of behavior during the middle of the day were taken from concurrent research by one of us (J. E. R.) and used for comparative purposes. Clock times (EDT) were recorded at the onset of flights preceding roosting. Times were also recorded when butterflies were seen to assume 148 JOURNAL OF THE LEPIDOPTERISTS SOCIETY TABLE 1. Behavioral responses and ambient environmental factors associated with roosting in Papilio polyxenes compared with values for those variables associated with other diurnal activity. Roosting Non-roosting 5d est fe! D) (CNY) xt S.D. (N) Flight duration** (seconds ) 96 + 111 (19) 33 = 52 (651) Perch duration (seconds ) (Overnight ) 132 + 208 (807) Perch height* (cm) 58} ae Iles} (8S) 64 + 27 (794) Air temperature 150 cm above the ground when roost posture assumed** ( Centigrade ) PPS AS) as B5) (1510) DAES) a= PLT tl0 Ne) Relative humidity when roost posture assumed** (7%) 76 + 16 (37) 58 + 11 (640) Solar radiation when roost posture assumed** (cal/cm’/min) 16 = .20 (15) Oo = mlian(254)) Asterisks indicate significance level of two-tailed Student’s t-test for differences between roosting and non-roosting mean values: * (.05 >P > .01); ** (P < .01). a roosting posture with folded wings or when they were discovered already in this posture. Identification of individuals at a distance was made pos- sible by marking the marginal and submarginal row of wing spots with felt-tip pens (Lederhouse, 1978). RESULTS Roosting Search Flight Roosting was always preceded by an extremely rapid and erratic flight of longer average duration than flights at other times of the day (Table 1). The shift from non-roosting activity into this search flight was not gradual but occurred abruptly, the individuals accelerating and often disappearing from sight in seconds. In the 8 cases where a single butterfly was observed throughout a search flight, movement was seen to be circular, the buttertly flying in circles often more than 5 m above the ground. During these flights, which avoided shaded areas, individuals often returned to certain spots in the field, flying low and dipping down into the vegetation. Search flights were repeatedly interrupted by short periods of perching, lasting one or two seconds. These were on potential roosts which either were not stable in the wind, were deflected greatly by the butterfly’s weight, did not provide a good gripping surface (smooth grass culms ), were very far from the ground, or were in the deep shade. Butterflies did not select the roost directly but only remained on those perches which were suitable after abandoning many perches which were not. VOLUME 32, NUMBER 3 149 Behavior on the Roost Eventually a suitable roost was located and the butterfly positioned itself so that the frontal plane of its body was perpendicular to the incident radiation from the setting sun, spread its wings, raised its abdomen between the spread anal margins of the hindwings, and then remained motionless (Fig. 1). This basking continued for several minutes, being longer on warm evenings than on cool or hazy ones. Basking ceased if the butterfly came under the advancing shadows of the surrounding vegetation. At the end of basking the abdomen was suddenly lowered, the wings dorsally appressed and swept back such that the hindwings almost totally covered the front wings, and the body brought parallel with the stem or head of the roost plant (Fig. 2). The posture was maintained throughout the night. The following morning, basking occurred as soon as direct sun fell on the roost. Morning basking appeared identical to that occurring in the evening. Butterflies were seen to rotate about the roost so as to be perpendicular to the rising sun. Moming basking was never seen earlier than two hours after sunrise and usually occurred between 0800 and 0900 (EDT) in July and early August. Effects of Ambient Temperature and Radiation on Roosting Mean values of ambient temperature, relative humidity, and solar radiation all differed between roosting and non-roosting situations (Table 1). It was not surprising that the evening hours associated with roosting had significantly lower radiation, lower temperature, and higher relative humidity than earlier in the day. Papilio polyxenes males are incapable of sustained flight with thoracic temperatures below 24 C (Rawlins, unpublished). This value matches that of the average air temperature when the roost posture with folded wings was observed in the evening (23.9 C). Since radiation was very low at these times, body temperatures would rapidly approach ambient levels in quiescent individuals. Several times individuals which had been roosting less than one minute following vigorous flight were observed to be incapable of sustained flight if disturbed. For example, one female disturbed under .09 cal/cm?/min radiation and 23.3 C ambient tem- perature was unable to remain airborne and crashed into the grass after a gliding flight of about 5 m. This suggests that the roost posture is assumed under decreasing radiation when ambient temperatures drop below the range of body temperatures allowing activity. The onset of roost search flight during July and August paralleled the time of meteorological sunset (Fig. 7). This suggests that the onset of al B sa = O ie) WN a7) isu 7p) = ae isa = avy © A = Ay ea — cal en e ea e) 4 o a, VoLUME 32, NUMBER 3 tom TABLE 2. Plant species used as roosts by Papilio polyxenes. Number on heads Number on stems Roost species or apices or culms Total Tragopogon pratensis L.* 4 J. 5 Lonicera morrowi Gray 0 1 Potentilla recta L.* 1! 0 if Solidago altissima L. 0 2 2 Linaria vulgaris Hill 0 1 1 Total dicotyledons = 5 10 Dactylis glomerata L. 2 0 2 Phleum pratense L. 12 4 16 Other grasses 2 b 9 Total grasses 16 RE DT Total roosts 21 16 Sil Percent of total 57% 43% 100% * All roosts were on dead, leafless scapes of the previous season. the search flight is cued by decreasing radiation even though flying or basking on the roost continues until ambient temperatures fall to levels preventing further activity. The occurrence of abbreviated search flights during sudden radiation drops preceding thunderstorms provided additional evidence that de- creasing radiation triggers such search flights. One male patrolling an area high in the field abruptly switched into erratic search flight behavior with the sudden darkening before a thunderstorm even though air tem- perature 1.5 m above the ground was 26.7 C. After being forced down minutes later by heavy rain, this male assumed the roosting posture in an air temperature of ,20.6 C. With the return of direct sun, basking commenced. Description of the Roost Seventy-three percent of the plants selected for roosts were grasses, the remainder being dicotyledonous species (Table 2). Sixty percent of the roosts on dicots were on leafless scapes of dead plants from the previous season. Such plants constituted less than five percent of the total stems within any grid square in the field. = Figs. 1-2. Postures of Papilio polyxenes on the roost. 1. Adult female in dorsal basking posture on culms of grass. 2. Male in folded-wing posture on short head of Phleum pratense, 152 JOURNAL OF THE LEPIDOPTERISTS SOCIETY @-female €-male Fig. 3. Topography of study area showing location of roosts. Each small grid square is 10 m square. Numbers in margins give elevation of isoclines in meters above lowest point in field (northwest corner). Roosts on plant inflorescences or apices were more frequent than roosts on culms or stems. Such heads may provide roosts which are easier to cling to or are less deflected by the butterfly’s weight. Butterflies on roosts are significantly closer to the ground than those perched during daily activity (Table 1). The maximum height of vegeta- tion within a one meter radius of each roost averaged greater than twice the roost height (110 + 17 cm, N = 33). Roosts were invariably in open areas of the vegetation where contact of the roosting butterfly with sur- rounding plants was impossible in all but the heaviest winds. Description of the Roost Sites Roosting occurred primarily on sites which sloped toward the west and were at the higher elevations in the field (Fig. 3). There were no differences between the roosting sites of males and those of females. The distribution of slopes for the ten meter grid squares in which roosting occurred differed significantly from that of slopes for all grid squares in the field (Kolmogoroy-Smirnov Test; .05 > P > .01) (Siegel, 1954) (Fig. 4). The average slope of 53 roost sites was 4.2 + 2.5 (S.D.) degrees above the horizontal. Similarly, the distribution of aspect values toward which roost sites sloped differed significantly from that for all grid VoLUME 32, NUMBER 3 153 SOF a ALL GRID SQUARES Fig. 4 | GRID SQUARES WITH ROOSTS 258 20 % 15 woe S74" \4i-5 J at ae in degrees above cb hae O (n ALL GRID SQUARES 25 Fig.5 O || GRID SQUARES WITH ROOSTS <240 240249 250259 26026? 270279 280289 290299 300309 310319 7320 Aspect in compass degrees % Figs. 4-5. Frequency distributions of slope (4) and aspect (5) values for all grid squares in study area compared with distributions for only those grid squares containing roosts. sites in the field (P < .01) (Fig. 5) even though the field predominantly sloped toward the west. The density distribution of yellow goat’s-beard (Tragopogon pratensis L.) and rough-fruited cinquefoil (Potentilla recta L.) for roosting sites 154 50 40 80 70 60 50 40 30 20 JOURNAL 0 les 4-7 22 Uf Stems of Tragopogon and Potentilla , m? 0) l= 20: 20-4040 Stems of Solidago and Aster / mi I ALL GRID SQUARES | GRID SQUARES WITH ROOSTS % % OF THE LEPIDOPTERISTS SOCIETY 60 0-39 40-79 80-119 7119 Grass Culms / m?* 40 30 20 10 0-19 20-39 40-59 CO79 77 Culms of Phleum and Dactylis | m 50 40 0 1-19 20-39 40-79 >79 Asclepias Plants / 100 m? Fig. 6. Frequency distributions of various plant densities for all grid squares in study area compared with distributions for only those grid squares containing roosts. differed significantly from that for all sites in the field, the density being higher for roosting sites (Kolmogorov-Smirnov Test; .05 > P > .O1) (Fig. 6). The density distribution of goldenrod (Solidago spp.) and New England aster (Aster novae-angliae L.) also differed significantly (P < .01). Roosts did not occur where Solidago and Aster were at highest density; these species usually form dense stands of plants which contact each other greatly in the wind. VoLUME 32, NUMBER 3 155 On the other hand, density distributions of all grass species, of just orchard grass (Dactylis glomerata L.) and timothy (Phleum pratense L.) taken together, and of common milkweed (Asclepias syriaca L.) did not differ significantly (P > .05) between roosting sites and all field sites. Survivorship on the Roost Roosting individuals were quiescent and could easily be approached and caught by hand. Such inability to escape suggested that predation during roosting might be a major, if not the chief, cause of mortality in black swallowtail adults. Twenty-six roosts were visited the following morning before basking occurred. Two male butterflies were missing from their roosts and were never seen again in the study area. Probability of death per night on the roost was estimated as being 2/26 = 0.077 for all butterflies regardless of sex. Thus, an adult could be expected to live through an average of 8.7 nights assuming no other sources of adult mortality were considered [(1—.077)®:* = 0.5) ]. If only males were considered the probability of death on the roost per night was estimated by 2/21 = 0.095. The average life expectancy of males was through 6.9 nights ignoring other sources of mortality. DISCUSSION Black swallowtails favor roosting sites which will allow them to con- tinue basking for as long as possible under decreasing radiation. Selection of relatively open roost sites which slope toward the west favors the maintenance of body temperatures suitable for continued activity for as long as possible. There is no indication of any behavior which would favor warming the following morning. The presence of such behavior in other butterfly species needs more study. The only unique behavior associated with roosting is the roost search flight. It is this flight which provides the mechanism by which a suitable roost is selected. Repetitive brief perches during this flight allow the butterfly to test and reject perches which are unstable, in the shade, or are disturbed by other vegetation. Frenetic activity during the search flight may provide enough body heat metabolically to allow flight to continue under conditions which would prevent further activity if the butterfly were motionless for a short period. Since the search flight occurs in the sun, roosts which are finally selected tend to be on sites having the longest and most direct exposure to the setting sun. Roosting individuals on level sites are more susceptible 156 JouRNAL OF THE LEPIDOPTERISTS SOCIETY 1900 aw! | 1830 = Y 1800 °o 1730 ae vaeeees two hours before sunset 0 O-onset of a roost search flight 1700 June July July July August August August Sept. 25 ©) 15 JAS) ©) 15 25 5 Fig. 7. Comparison of the time of onset of roost search flights with time of sunset. The solid line is a least-squares regression line of onset times with respect to date. to being shaded by plants between them and the sun than are those on sloping sites facing the setting sun. The onset of the erratic roost search flight is apparently triggered by decreasing radiation regardless of temperature. The shift from dorsal basking on the roost to a roosting posture with folded wings appears to occur when ambient temperature and radiation are such that body tem- peratures suitable for flight can no longer be maintained. There appears to be no endogenous rhythm controlling roosting. Such behavior is just as Clearly associated with the onset of unfavorable conditions for flignt at other times of the day. Basking on the roost is similar to basking under more favorable con- ditions. There is no difference between morning and evening, making Clench’s (1966) categories of “vesper” and “matutinal warming” difficult VOLUME 32, NUMBER 3 157 to define for P. polyxenes. Basking on the roost may lengthen the period in the evening when escape from predators or selection of a different roost are still possible should a predator appear or the roost become unsuitable. Selection for roosting behavior and roost preferences appears to be strong in view of the high mortality recorded in this study. Those individuals which fail to secure a steady, protected roost risk dislodgement and predation while cold and helpless at ground level. In a concurrent study, Lederhouse (1978) estimated the probability of permanent disappearance per day due to dispersal and death for males after the first night of adult life to be 0.106. This value for males through the first night of imaginal life was measured as 0.532. The present estimate of male mortality on the roost (0.0953 is therefore about equal to that which Lederhouse ascribes to all causes of death plus dispersal because all marked butterflies in our study were at least two days old. It is con- ceivable that the large loss through the first night of adult life may be attributed to failure to find a suitable roost during that first evening or to considerable dispersal during the first day. Death on the roost may be the chief source of mortality in this popula- tion, since not one instance of diurnal predation has been seen in over 700 hours of close observation, notwithstanding reported cases of diurnal predation by passerine birds (Erickson, 1973). Predation by ants may explain the preference shown for dead dicot scapes as roosts since ants may not frequent such plants during foraging. If death on the roost is the major source of mortality in adult black swallowtails, then selection for mimetic or cryptic coloration patterns would probably operate most strongly on that portion of the butterfly which is exposed during roosting, the undersurface of the hindwings. We cannot help but note that the undersurface of the hindwings of members of the supposed Battus philenor mimetic complex are more similar in coloration than any other part of the wings. This complex includes B. philenor (L.), Papilio troilus L., P. glaucus L. (dark females), P. polyxenes F., and Limenitis arthemis astyanax F. (Brower, 1958). We suggest that selective advantages gained by cryptic or mimetic coloration patterns in many butterfly species may be greatest during roosting and that the exposed undersurface of the wing is more affected by such selection than is the hidden uppersurface. In conclusion, black swallowtails select roosts which maximize exposure to the setting sun and provide stable, secure sites preventing dislodge- ment during the night. A roost search flight triggered by decreasing radiation is the only behavior unique to roosting and provides a 158 JOURNAL OF THE LEPIDOPTERISTS SOCIETY mechanism by which suitable roosts may be selected. Selection for efficient roosting is strong since death on the roost is likely to be the major source of mortality in adults. ACKNOWLEDGMENTS We wish to express our appreciation to Owen Sholes, Dr. Paul Feeny, and Dr. John G. Franclemont for comments on the manuscript. We thank Dr. Mark Scriber for suggestions on the implications of roost mortality on selection for wing coloration. LITERATURE CITED Benson, W. W. & T. C. Emme. 1973. Demography of gregariously roosting populations of the nymphaline butterfly Marpesia berania in Costa Rica. Ecology 54: 326-335. Brower, J. V. Z. 1958. Experimental studies of mimicry in some North American butterflies. Part II. Battus philenor and Papilio troilus, P. polyxenes and P. glaucus. Evolution 12: 123-136. Ciencu, H. K. 1966. Behavioral thermoregulation in butterflies. Ecology 47: 1021-1034. Crane, J. 1957. Imaginal behavior in butterflies of the family Heliconiidae: changing social patterns and irrelevant actions. Zoologica (New York) 42: 135-145. Erickson, J. M. 1973. Bird predation on Papilio polyxenes F. (Papilionidae). J. Lepid. Soc. 27: 16 FERNALD, M. L. 1971. Gray’s Manual of Botany. Eighth ed. D. Van Nostrand, New York. 1632 p. Gipson, D. O. & A. L. Pancuen. 1975. Roosting behavior of the butterfly Papilio demodocus Esp. on the Kenya coast. Entomol. Rec. 87: 156-158. Hernricu, B. 1972. Thoracic temperatures of butterflies in the field near the equator. Comp. Biochem. Physiol. 42A: 459-468. Kiors, A. B. 1951. A Field Guide to the Butterflies of North America, East of the Great Plains. Houghton Mifflin, Boston. 349 p. Larsen, T. B. 1973. Communal roosting among butterflies in the Lebanon. Entomol. Scand. 4: 299-301. LepErRHOUsSE, R. C. 1978. ‘Territorial behavior and reproductive ecology of the black swallowtail butterfly, Papilio polyxenes asterius Stoll. Cornell University Ph.D. Thesis. Ithaca, New York. MacNemz, C. D. 1964. The skippers of the genus Hesperia in western North America, with special reference to California (Lepidoptera: Hesperiidae). U. Calif. Publ. Entomol., 35. 221 p. | Muysnonpt, A. & A. MuysHonptr, Jr. 1974. Gregarious seasonal roosting of Smyrna karwinskii adults in E] Salvador (Nymphalidae). J. Lepid. Soc. 28: 224-229, PoweELx, J. A. 1968. A study of area occupation and mating behavior in Incisalia iroides (Lepidoptera: Lycaenidae). New York Entomol. Soc. 76; 47-57. SieceEL, S. 1956. Nonparametric Statistics for the Behavioral Sciences. McGraw- Hill, New York. 312 p. Turner, J. R. G. 1975. Communal roosting in relation to warning colour in two heliconiine butterflies (Nymphalidae). J. Lepid. Soc. 29: 221-226. U.S. Pare. Orr. 1959. Sunrise and Sunset at Binghamton, New York. Nautical Almanac Office, U.S. Naval Observatory, Wash., D.C. VOLUME 32, NUMBER 3 159 Urounart, F. A. 1960. The Monarch Butterfly. Univ. of Toronto Press. 361 p. VIELMETTER, W. 1958. Physiologie des verhaltens zur Sonnenstrahlung bei dem Tagfalter Argynnis paphia L. I. Untersuchungen im Freiland. J. Insect Physiol. 2: 13-37. Watt, W. B. 1968. Adaptive significance of pigment polymorphisms in Colias butterflies. I. Variation in melanin pigment in relation to thermoregulation. Evolution 22: 437-458. Younc, A. M. & J. H. THomason. 1975. Notes on communal roosting of Heliconius charitonius (Nymphalidae) in Costa Rica. J. Lepid. Soc. 29: 243-255. ANNOUNCEMENT The Xerces Society offers modest research grants to support scientific work which has some likelihood of contributing to the preservation of terrestrial arthropod popula- tions as biological entities. XS is an international, non-profit organization dedicated to the conservation of terrestrial arthropod populations and their habitats. Awards will usually be a few hundred dollars but may be larger if funds are available. Knowledge- able young workers and those without formal professional affiliation are encouraged to apply. For details write to Dr. FRANcIE CuEw, Xerces Grants Committee Chairman, Department of Biology, Tufts University, Medford, Massachusetts 02155, U.S.A. Journal of the Lepidopterists’ Society 32(3), 1978, 160-174 NOTES ON THE LIFE CYCLE AND NATURAL HISTORY OF BUTTERFLIES OF EL SALVADOR. IIC. SMYRNA BLOMFILDIA AND S. KARWINSKII (NYMPHALIDAE: COLOBURINI) ALBERT MuysSHONDT, JR. AND ALBERTO MuysHONDT 101 Avenida Norte #322, San Salvador, E] Salvador ABSTRACT. Descriptions and photographs of the life histories of Smyrna blomfildia and S. karwinskii are presented, and the larval foodplants ( Urticaceae) and the comparative behavioral characteristics of the two species are recorded and dis- cussed. The present taxonomic placement of S. blomfildia and S. karwinskii is questioned, and an alternate interpretation is expressed based on the differing degrees of morphism between the two species. The adaptiveness of polymorphism is explained relative to human-disturbed habitats in E] Salvador, noting that the monomorphic S. karwinskii is the most evolved species, but that S. blomfildia has a flexibility to Overcome adverse conditions because of certain polymorphic characteristics. The peculiar phenomenon of a divergent evolutionary trend between the early stages (larvae and pupae) and an advergent trend between the adults of both species is noted. This is the second of three papers on the Coloburini (Gynaeciini) of El] Salvador. Classically, Colobura dirce L. and Historis odius Fab. have been included with Smyrna in the Coloburini, based, no doubt, on similarities in adult characteristics. Whether or not these form a natural complex of related species is left for others to determine, who can compare the overall characteristics of Smyrna sp., Colobura dirce, and Historis odius. We present here a description of the early stages of Smyrna blomfildia Fab. and S. karwinskii Hibn., records of the larval foodplants, and an account of the behavior observed in both the immatures and adults. We have reared S. blomfildia and S. karwinskii using the same techniques described for the Catonephelini, Charaxini, and Hamadryadini (Muy- shondt, 1973a, 1974b; Muyshondt & Muyshondt, 1975a) with consistently uniform results. Specimens preserved in alcohol have been sent to the American Museum of Natural History, New York City. Life Cycle Smyrna blomfildia Smyrna karwinskii Egg. Almost spherical with flattened Same as S. blomfildia in all respects. base, light green with 10 whitish vertical ribs which fade around the micropyle. Ca. |! mm diameter. Hatches in 5 days. VOLUME 32, NUMBER 3 Ist instar larva. Head roundish, naked, shiny black. Body naked, cylindrical, brownish-green, with transverse rows of white, shallow, small warts on each seg- ment. Legs and tips of prolegs dark brown. Ca. 2 mm when hatched, growing to 4.5 mm in 2 days. 2nd instar larva. Head shiny black, with short, knobby, divergent horns, one on each epicranium, and 8 small, white con- ical projections across head capsule, under epicranial horns. Body brown with trans- verse rows of tiny, forked spines im- planted on white chalaza. Grows to 8 mm in 2 days. 3rd instar larva. Head reddish with thick, short horns (ca. head length) armed with secondary spines placed in the fol- lowing order: a basal row of 3, 1 pointing inwards, 2 anterad; a second row, with 1 pointing caudad, 1 anterad and 1 laterad. Horns terminate distally in a club with 5 short spines. Around base of each horn are 7 spines; around the ocelli are 5 smaller spines. Body predominantly dark brown with some light spots among the spines, which are placed in the following positions: Ist thoracic segment (T-1) with 1 spine subdorsally, 1 subspiracular spine and 1 pedal spine. T-2 and T-3 with 1 subdorsal scolus with a rosette of 5 spines near tip and 1 vertically; 1 supra- spiracular scolus with rosette of 4 spines and 1 vertical; 1 subspiracular spine and 1 pedal spine. Abdominal segments from A-l1 to A-7 have, in addition, 1 dorsal scolus with 2 lateral spines and 1 distal, and behind the subspiracular spine 1 scolus with rosette of lateral spines and 1 distal spine. A-l1, A-2 and A-7 have 1 small ventral spine in line with prolegs. A-8 has an additional scolus caudad with a 6-spined rosette. A-9 with only 1 sub- dorsal scolus directed posterad with rosette of 5 lateral spines and one distal. A-10 with anal shield and 2 lateral groups of small spines directed posterad. Grows to 15 mm in 2 days. 4th instar larva. Same as 3rd instar but body shows various color morphs: 1 dorsally brown with cream dots, rest of body cream also, where black spiracula 161 Same as S. blomfildia, but light green with transverse rows of white warts. Same as S. blomfildia, but body lighter color and lacking dorsal spines in all but 8th abdominal segment. Head light brown with longer and thinner horns than S. blomfildia (1% head length). Lateral spines of head much reduced. Horn terminals more clubbed. Body color basically brown with double transverse rows of whitish dots on each segment and a broken stripe of light color subspiracularly. Spiracula black. Ventral surface dirty light gray, prolegs beige. Body with whitish scoli armed with con- colored black-tipped spines, placed in the following order: T-1 with subdorsal group of 3 small spines, then 1 small supraspiracular spine, 1 small subspirac- ular spine and 2 small pedal spines. T-2 and T-3: 1 subdorsal scolus, short, with rosette of 6 lateral spines and 1 distal spine; 1 supraspiracular scolus with rosette of 5 lateral spines and 1 distal: 1 subspiracular spine and 2 pedal spines. Abdominal segments A-1 and A-7 have a subdorsal scolus with a rosette of 4 lateral spines and 1 distal; supraspiracular scolus with rosette of 4 lateral spines and 1 distal; subspiracular scolus with 4 lateral spines and 1 distal, then two small pedal spines. A-8 has in addition a heavier dorsal scolus armed with 8 spines. A-9 has only 1 supraspiracular scolus directed posterad, with 6 spines. Anal plate on A-10 surrounded by 6 small spines. Grows to 15 mm in 2 days. Head dirty yellow, with some brown markings frontally. Slender horns, slightly bent in some individuals, colored light gray. All head and horn spines white with 162 stand out and all scoli and spines whitish. Another morph mostly black dorsally, with double row of whitish dots along meson. From supraspiracular to ventral area cream colored. Dorsal scoli white, subdorsal and supraspiracular scoli black, the rest whitish. Other morph mostly greenish-white with black stripes covering dorsal and subdorsal area, but much broken by stripes and dots of greenish- white. All scoli greenish-white with light spines. Still another morph similar to the preceding one but with subdorsal and supraspiracular scoli black. The rest whit- ish. Grows to 28 mm in 2 to 3 days. 5th instar larva. Same as 4th instar, growing to 41 mm in 3 to 4 days. Prepupa. No noticeable change. Hangs from anal prolegs, body incurved ven- trally. Lasts 1 day. Pupa. From light brown to very dark brown, abdominal segments darker than the rest, with rows of lighter, shallow warts: 1 supraspiracularly, 1 subspirac- ularly. Spiracula inconspicuously brown. One black spot at either side between wingcase and thorax. Abdomen rounded with no sharp angles; slightly incurved ventrally, shallow depression at the tho- racic union dorsally. Thorax slightly keeled to rounded head. Pointed cre- master dark brown. 25 mm long, 10 mm laterally and dorsoventrally at widest points. Adults emerge between 8-11 days. JOURNAL OF THE LEPIDOPTERISTS SOCIETY black tips. Body ground color light gray with a darker thin mesal stripe, and trans- verse rows of cream colored spots at seg- ment unions. All scoli as in 3rd instar, implanted now on bright yellow chalaza; scoli and spines white with black tips. Grows to 29-30 mm in 2 to 3 days. Drastic change in color. Now mostly brown with black markings on head, be- tween dark horns and down to ocelli lat- erally. The whole body with black ir- regular marblings. All scoli shorter than in prior instars; brown colored as well as spines, legs and prolegs. Grows to 40—42 mm in 3 to 4 days. Same as S. blomfildia. 1 day. Lighter brown and thinner than S. blom- fildia. Abdomen more humped dorsally and with 3 rows of conical spines: 1 prominent subdorsally, 1 of decreasing size supraspiracularly, and 1 still smaller subspiracularly. Thoracic dorsal keel sharply angled midways. Black spots be- tween wingcase and thorax, as S. blom- fildia. Black pointed cremaster set at an angle in relation to body plan. 28 mm long, 11 mm dorsoventrally, 10 mm lat- erally at widest points. Lasts 9-11 days. Adults. Both species show a marked sexual dimorphism in the coloration, males being brighter than females. Wing shape is the same in both sexes. Males. Dorsally forewing golden brown in basal and discal areas with lighter golden- orange band slanting from midcostal margin to tornus. (S. blomfildia tends to be lighter than S. karwinskii.) The rest velvety black, except a subapical row of three, light yellow spots parallel to the light band. Hindwing in S. karwinskii golden-brown mostly, with a black edge along outer margin, 3 mm wide, near outer angle, very thin from there down along the very edge of the wing to anal angle. A submarginal row of faint black dots along the thin portion of the black edge. In S. blomfildia hindwing is a lighter golden-orange and black edge along outer angle becomes submarginal along VoLUME 32, NUMBER 3 163 Figs. 1-11. Smyrna blomfildia: 1, egg, recently deposited, 1 mm; 2 (photo upside- down), lst instar larva, 4.5 mm (note “perch” on central vein); 3, 2nd instar larva, 8 mm (note new “perch” being constructed at edge of leaf); 4, 3rd instar larva, 15 mm; 5, 4th instar larva, 28 mm; 6-10, 5th instar larvae, various morphs, 41 mm; 11, 5th instar larva, close-up of head. outer margin down to Mz where it becomes thin, ending between Cu: and Cuz where it is substituted by a marginal thin edge running between the two, small toothed projections on anal angle, with a whitish dot in the interior one. Inner fold in both species fulvous gray. Ventrally forewing presents basally some black drawings, on light yellow basic color, which is devoid of markings from midcostal margin to tornus, except for a brownish gray border along inner margin, more so in S. karwinskii. From midcostal to subapical costal margin down to mid-outer margin to tornus there is a black zone 164 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Figs. 12-17. Smyrna karwinskii: 12, egg, ready to hatch, 1 mm; 13, Ist instar larva, 3.5 mm; 14, 3rd instar larva, 15 mm; 15, 4th instar larva (note absence of dorsal spines and slender horns ), 29-30 mm; 16, 5th instar larva (note reduced scoli), 40-42 mm; 17, 5th instar larva, close-up of head. limited distally by a diffuse replica of the dorsal subapical dots, which ventrally merge with each other. Apically a gray zone mottled by faint black markings, more con- trasting in S. blomfildia than in S. karwinskii. Hindwings show a complicated pattern of sinuous lines, circles and triangles of dark brown, light brown and whitish color, all darker in S. blomfildia than in S. karwinskii. VoLUME 32, NUMBER 3 165 Figs. 18-20. Smyrna blomfildia: 18, dorsal view of pupa; 19, side view of pupa; 20, ventral view of pupa. Figs. 21-23. Smyrna karwinskii: 21, dorsal view of pupa; 22, side view of pupa; 23, ventral view of pupa. 166 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Figs. 24-27. Smyrna blomfildia: 24, dorsal view of male; 25, ventral view of male (metric scale); 26, dorsal view of female; 27, ventral view of female. Both species present a submargnal row of 4 “eyes” along outer margin, the two at the extremes twice as large as the two interior ones. Both species also have a black spot on the anal angles. S. blomfildia in addition has a second black spot on the first toothed projection. Females. Both species dorsally have the same pattern as the males, but the golden- brown or orange is replaced by dull brown, separated from the black apical area by a light yellow band. The rest as in males and so is the underside of the wings. Body is concolorous to the respective wing coloration. Antennae black, ending with an orange tip, larger in S. blomfildia. Palpi cream colored, proboscis tanned brown. Wing span 70-80 mm. S. blomfildia usually larger than S. karwinskii, and females larger than males, more markedly so in S. blomfildia. Total time from egg to adult, 25-30 days. Natural History Females of both species of Smyrna deposit their solitary eggs on the undersides of leaves of various Urticaceae. In contrast, other species of Coloburini oviposit on Moraceae. We have found the eggs of both Smyrna spp. on Urticastrum mexicanum (leb.) Kuntze, Urera caracasana (Jacquin) Grisenbach, and U. baccifera (L.) Gaudichaud. The fast flying females land on the undersurfaces of leaves for ovi- VoLUME 32, NUMBER 3 167 ie Figs. 28-31. Smyrna karwinskii: 28, dorsal view of male (note absence of indenta- tions in anal angles); 29, ventral view of male; 30, dorsal view of female; 31, ventral view of female. position. This is in striking contrast to the rest of the Coloburini we have been able to study, which oviposit mostly on the upper surface of leaves. Female Smyrna quickly deposit one egg, then move to another plant close by where the process is repeated. It is not uncommon for the same plant to be visited again after a period of time. We have observed that females deposit more than five eggs in sequence before moving away. The plants used are generally less than 3 m tall. The rather small eggs are quite hard to locate, due in part to the stinging properties of the plants, but also to the light green color of the eggs, which makes them inconspicuous against the color of the leaves. After 5 days the larva hatches and consumes most of the eggshell, leaving traces of the wall. It proceeds to feed on the undersurface of the leaf, where it con- structs a resting perch very close to its feeding place, perpendicular to the surface. This is a notable deviation from the usual method of the other perch-making species we know, which always move to the edge of a leaf to feed, and which construct their resting perch by prolongation of a vein. In Smyrna spp., second instar larvae do eventually construct the 168 JOURNAL OF THE LEPIDOPTERISTS SOCIETY perch at the edge of the leaf. From then on, the larvae remain on the undersurface of leaves until pupation. They pupate on the same plant, either under leaves or on petioles, after hanging by the anal prolegs for one day with the thoracic segments incurved ventrally. The pupa hangs loosely from the pointed cremaster, swinging freely with the faintest movement of the plant. The colors of the wings of the pharate adult do not show through the wingcase of the pupal skin when the adult insect is ready to emerge, due to the dark color of the pupa. After emergence, a rusty-colored meconium is ejected while the insect expands its wings. The imago takes a long period of time before its first flight. Both species show marked sexual dimorphism. At the same time, the two species are very similar to one another dorsally. The principal differ- ence is that S. karwinskii has a rounded anal angle on the hindwing, where S. blomfildia has a small toothed projection. We have never seen imagos of either species visiting flowers, but we have seen them feeding on sap flowing from tree wounds, fermenting fruits, mud puddles, etc., where they spend long periods of time, with their wings folded dorsally. Smyrna karwinskii is noteworthy for its seasonal gregarious roosts in high mountains during the dry season (Muyshondt & Muyshondt, Jr., 1974). This phenomenon also occurs in Mexico (R. Wind, Chiapas, pers. comm.; Beutelspacher, 1975). We have never found eggs or larvae of S. karwinskii in the high mountains during the dry or the rainy season, even where the foodplants occur locally. It seems that they move down to lower levels to breed, usually in close proximity to S. blomfildia. The latter very seldom is found at altitudes over 1600 m and does not have the communal roosting behavior of S. karwinskii. It is not uncommon to collect eggs and larvae of both species on the same plant. The adults of both species behave similarly: they have the same fast rustling flight, they perch on tree trunks with their heads pointing down, and the males have a strong territorial defensive attitude, chasing in- truding butterflies of the same or different species. To date we have not found cases of parasitism in these species, but we have very often witnessed predation by Hemiptera (Reduviidae, mostly) which impale the larvae, leaving only the sagging skin. Another cause of severe larval mortality in the fields is a disease causing the larva’s body to burst, releasing a foul-smelling dark fluid. The foodplants, known locally under one vernacular name common to the three species, “Chichicaste,” are plants often used as hedges around coffee plantations, because of the severe stinging caused by the leaves which deters trespassers. Urera baccifera grows to a height of 7 m when left alone. The trunks VOLUME 32, NUMBER 3 169 and older branches are covered with short, wide spines, the younger branches and the leaves with stinging hairs. The leaves are large, coarse, round-cordate and roughly dentate. The small greenish flowers grow in cymes, producing small, translucent, globose fruits with a dark seed inside. These fruits are much sought after by farm children, who pick them by beating the shrub with a stick while catching the rain of falling fruits with a “sombrero.” Thus, hundreds are collected before disposing of them in situ, much to their pleasure. The juicy fruits have a sweetish- refreshing taste and alleviate thirst readily. Urera caracasana grows to about 4 m and is also used in fences. The leaves are smaller, and of variable shapes; more or less elongate, cordate at the base and acute at the apex, with close dentation at the edges. The fruits are red when mature. Both species are used in popular medicine against venereal diseases. Urticastrum mexicanum is a shrub up to 4 m tall, with ovate, crenate leaves. The fruits are achenes. All of these plants have caused painful accidents to tourists unaware of the severe stinging properties of the otherwise handsome leaves. DISCUSSION Smyrna blomfildia is the type-species of the genus Smyrna Hubner, based on the butterfly originally named Papilio blomfildia by Fabricius in 1781 (Hemming, 1967). As he very often did, Htibner misspelled the specific name as “blomfildii,” and this erroneous spelling was subsequently used by several authors, among them Herrich-Schaffer (1864) and Miller (1866), with an additional error: “blomfieldii.” Other authors named the species S. bella Godart and S. pluto Westwood (Seitz, 1921). The only other reports of the early-stages of this genus that we are aware of are by Miiller, who gave a short description of a probable 4th instar larva preserved in alcohol, some rough descriptions based on that of Miiller’s (Seitz, 1921; Hayward, 1964), and a vague comparison be- tween the larvae of Colobura and Smyrna by Brown & Heineman (1972). We believe that ours is the first complete description of the early stages, with photographic illustrations of both Smyrna blomfildia and karwinskii. The genus Smyrna has been placed in various unrelated groups based mostly on the external characters of the perfect insects by many early authors. Doubleday, Westwood and Hewitson (1849) and Boisduval (1870), placed it close to Agrias. Herrich-Schaffer (1864) put this genus in his “familie XI,” together with the related genera Gynaecia and Callizona, and many unrelated ones, among them Euptoieta, Eunica, Pyrrhogyra, Ageronia, Peridromia, Amphichlora, etc. Schatz & Rober (1892), placed Smyrna in their “Gynaecia-Gruppe,” as part of their 170 JOURNAL OF THE LEPIDOPTERISTS SOCIETY larger “Eunice-Gruppe,” together with Callizona and Gynaecia (Colo- bura), probably following Herrich-Schaffer. Today, many authors follow Seitz (1921), who was probably influenced by Reuter (1898) in placing Smyrna in his “Grupe Gynaeciidae,” together with Historis, Coea, Pycina, Megistanis, Gynaecia and Callizona, as an intermediate group between his “Gruppe Epicaliidi” including many genera (Catonephele, Epiphile, Temenis, Pseudonica, Pyrrhogyra, etc.) covered by one of us (Muyshondt, 1973a, b, c, d; 1974a) in previous papers and his “gruppe Hypolimnadidi,” with Hypolimnas misippus L. Other modern authors include this genus and related genera ( Historis, Coea, Colobura, etc.) in the Limenitidini which we believe is erroneous. Different opinions arise, no doubt because of superficial similarities of the adults, which could very well be due to convergent evolution rather than close relationship. Examples are well known of convergence in color, pattern, and shape between unrelated species actually belonging to different families: i.e. Danaidae, Ithomiidae, Heliconiidae and Pieridae. These often form Miillerian and Batesian mimicry complexes, as pointed out by many biologists (Brower, 1972; Brown & Benson, 1974). The very poor knowledge of the immatures of most tropical butterflies has led to errors in the association of species. Descriptions of the im- mature stages are necessary for a more accurate systematic arrangement of the neotropical Lepidoptera. Comparison of the eggs of the two species of Smyrna indicates that they are very closely related. Larvae and pupae are also very similar, although the larvae of S. karwinskii lack the dorsal row of scoli present in S. blomfildia. In contrast, the eggs, larvae and pupae of Smyrna differ considerably from those of Colubura dirce. The immature stages of Historis odius and Coea acheronta resemble each other closely, but have nothing in common with Colobura, and only the larval head shape resembles Smyrna. There are so many drastic differences between the characteristics of the early stages of the species of Coloburini studied by us [Colobura dirce L. (Muyshondt, Jr. & Muyshondt, 1976), Historis odius (Fab.) and Coea acheronta (Fab.), (Ms. in prep.) ], that we question the correctness of the taxonomy of the group. However, we feel that although Smyrna is an aberrant genus in the Coloburini, it represents a link between the Nymphalini and the other genera now included in the Coloburini. By the same token it is also evident that none of these species can be placed in the Charaxinae, as was done by Boisduval (1870), who placed Smyrna between Agrias and Prepona, and said that the larvae of VoLUME 32, NUMBER 3 fal Aganisthos (=Historis) and Prepona, “sont tout-a-fait semblables” (ex- actly alike). This is absolutely incorrect! Prepona and Archeoprepona do resemble each other in the shape of the eggs, larvae and pupae (Muyshondt, 1973e; Muyshondt, 1976), but neither stage resembles even remotely the early-stages of Historis (Ms in prep.). To include Smyrna with the Limenitidini (in which Adelpha belongs) as most modern authors do, is also incorrect, as they have nothing in common with the Coloburini during their early stages. With Limenitidini there are certain imaginal resemblances, but these are not strong enough to place them together. It is noteworthy that the larvae of both species of Smyrna construct a resting perch with frass pellets. Other larvae which use this defensive strategy construct their perch at the edge of the leaf on which they live: some of them pile a barrier of excreta mixed with pieces of dry leaf tissue at the base of the perch (Adelpha spp.); some fasten leaf cuttings with silk which hang from the perch ( Zaretis, Prepona, Archeoprepona); many others leave the perch bare (Biblis, Mestra, Catonephele, Epiphile, Nica, Temenis, Pyrrhogyra, Diaethria, Catagramma, Cyclogramma, Hamadryas, Colobura, Historis, Coea, Apatura, Marpesia). As far as we have been able to ascertain only the two species of Smyrna construct a perch on the underside of a leaf, very close to where the eggshell was consumed. During the 2nd instar, a new perch is sometimes made at the edge of the leaf; the other species mentioned also do this. We interpret this behavior to result from protection afforded to the small larvae by the strong urticating properties of the foodplants, a factor which by itself might deter at least some predators. After the 2nd instar the larvae abandon their perch and wander about the plant on the underside of the leaves. Perhaps the urticating properties of the plant afford continued protection. It is to be noted that the profusion of spines displayed by the larvae of Smyrna spp. from the 3rd instar on do not have urticating properties. Even the ventral prothoracic gland (adenosma), is not readily extruded as in Colobura dirce and other species provided with this apparent means of defense. Thus, it seems that the larvae of Smyrna rely on the protection granted by the plant itself, rather than on the protection they could derive from their own spines and odoriferous gland. One thing puzzles us: although the adults of the two species of Smyrna are strikingly alike, why is it that the larvae and even pupae of the two species show important differences, such as the unequal number of rows of scolii in the larvae, and the different shape of the pupae? We have seen various larvae and pupae of species belonging to the same genus, the adults having a common shape but with very disparate 172 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY — coloration, such as Siproeta stelenes (Young & Muyshondt, 1973) and V. epaphus (Young, 1972), Heliconius petiveranus and H. charitonius; Anartia fatima and A. jatrophae. Still others show differences not only in color, but in the shape of the wings of the adult, as do Catonephele numilia and C. nyctimus. Yet the larvae and pupae, except for minor discrepancies, if any, have the same characteristics, indicating that they undoubtedly both belong to the same genus. We have seen, on the other hand, species placed in the same genus which have very basic differences during their early stages, for example Hamadryas februa, H. guatemalena and H. amphinome; and Anaea eurypile, A. morvus and A. pithyusa, suggesting that they may belong to different but related genera of one sub- family or family (Muyshondt & Muyshondt, Jr., 1975a, b, c; Muyshondt, 1974b, 1975a, b). For these reasons, regardless of the striking re- semblances in adult coloration and shape between Smyrna blomfildia and S. karwinskii, we suggest they might belong to different genera. They would then form another case of evolutionary convergence, perhaps of Millerian mimicry. While S. blomfildia seems to be in the process of finding its optimum larval characteristics, as suggested by the strking polymorphism in larval coloration, S. karwinskii apparently has already achieved stability as it has only one morph. We consider S. karwinskii as the most evolved of the two, because of the uniformity of characteristics maintained during its whole life cycle, and thence the model of the two. S. blomfildia we consider to be the youngest, still an evolving species. This evolutionary phase seems to have momentarily given S. blomfildia an advantageous flexibility to overcome adverse conditions which are reflected in a more abundant population than its more stable relative, S. karwinskii, at least under the conditions in E] Salvador where the habitats are continuously and severely affected by human influences, due to the high population density. It would be interesting to read an explanation of the present phenom- enon, where two species evolve divergently during their early stages, yet seem to evolve advergently during their adult stage. Most of the work of which we are aware on the evolution of butterflies has concentrated on their adult stage, disregarding almost completely their early stages, which perhaps would throw new light on the problem. ACKNOWLEDGMENTS We wish to express our gratitude to all the people who helped us with their guidance, provided reference material, and revised our manuscript to make it presentable, especially Drs. A. B. Klots, A. B. H. Rydon, J. G. Sternburg, and G. L. Godfrey. This paper would not have been possible VoLUME 32, NUMBER 3 Nike without the great help received from Marilyn and Pierre Muyshondt, who did much of the fieldwork and observations. LITERATURE CITED BEUTELSPACHER, C. R. 1975. Notas sobre el suborden Rhopalocera (Lepidoptera ) de las Minas, Veracruz. Rev. Soc. Mex. Lep. A. C. 1(1): 11-20. BotspuvaAL, J. 1870. Considerations sur des Lépidoptéres envoyés du Guatemala a M. de l’Orza. Rennes. Brower, L. P. & J. van ZANDT Brower. 1972. Parallelism, convergence, divergence and the evolution of mimicry. Trans. Conn. Acad. Arts Sci. 44: 59-67. Brown, F. M. & B. Heineman. 1972. Jamaica and its butterflies. E. W. Classey, Ltd. London. 478 p. Brown, K. S. & W. W. Benson. 1974. Adaptive polymorphism associated with multiple Miillerian mimicry in Heliconius erato (Lepid.-Nymph.). Biotropica 6: 205-228. Dovusepay, E., J. O. Westwoop & W. C. Hewrrson. 1849. The genera of diurnal Lepidoptera. Vol. 2. Longman, London. Haywarp, K.J. 1964. Genera et species animalium Argentinorum. Vol. 3. Insecta, Lepidoptera, Familiae Nymphalidearum. Universitas Nationalis Tucumanensis, Fundatio Michaelis Lillo. Kraft Ltd., Buenos Aires. Hemminc, F. 1967. The generic names of the butterflies and their type species. (Lepidoptera: Rhopalocera). Bull. British Mus. (Nat. Hist.), Entomology Suppl. 9. 509 p. HerricH-ScHAFFER, G. A. 1864. Prodromus Systematis Lepidopterorum. Korres. Blatt Zool.-min. Ver. Regensburg, 18. Mt.ier, W. 1886. Siidamerikanische Nymphalidenraupen. Versuch eines natiir- lichen Systems der Nymphaliden. Zool. Jahrb. Zeitschr. Syst., Geogr., Biol. Thiere 1: 417-678. MuysHonpr, A. 1973a. Notes on the life cycle and natural history of butterflies of E] Salvador. IA. Catonephele numilia Esite (Nymphalidae-Catonephelinae ). J. N. Y. Entomol. Soc. 51: 164-174. 1973b. Notes on the life cycle and natural history of butterflies of El Salvador. IIA. Epiphile adrasta adrasta (Nymphalidae-Catonephelinae). J. N. Y. Entomol. Soc. 51: 214-223. 1973c. Notes on the life cycle and natural history of butterflies of F] Salvador. IIIA. Temenis laothoe liberia (Nymphalidae-Catonephelinae). J. N. Y. Entomol. Soc. 51: 224-233. 1973d. Notes on the life cycle and natural history of butterflies of El Salvador. IVA. Pseudonica flavilla canthara (Nymphalidae-Catonephelinae ). J. N. Y. Entomol. Soc. 51: 234-242. 1973e. Notes on the life cycle and natural history of butterflies of El Salvador. I. Prepona omphale octavia (Nymphalidae). J. Lepid. Soc. 27: 210-219. 1974a. Notes on the life cycle and natural history of butterflies of E] Salvador. VA. Pyrrhogyra hypsenor (Nymphalidae-Catonephelinae). J. N. Y. Entomol. Soc. 52: 163-172. 1974b. Notes on the life cycle and natural history of butterflies of El Salvador. IV. Anaea (Memphis) eurypyle confusa (Nymphalidae). J. Lepid. Soc. 28: 306-314. 1975a. Notes on the life cycle and natural history of butterflies of El Salvador. V. Anaea (Memphis) morvus boisduvali (Nymphalidae). J. Lepid. Soc. 29: 32-39. 1975b. Notes on the life cycle and natural history of butterflies of El 174 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Salvador. VI. Anaea (Memphis) pithyusa (Nymphalidae). J. Lepid. Soc. 29: 168-176. . 1976. Notes on the life cycle and natural history of butterflies of E] Salvador. VII. Archaeoprepona demophon centralis (Nymphalidae). J. Lepid. Soc. 30: 23-32. Muysuonpt, A. & A. MuysHonpr, Jr. 1974. Gregarious seasonal roosting of Smyrna karwinskii adults in El Salvador (Nymphalidae). J. Lepid. Soc. 28: 224-229, . 1975a. Notes on the life cycle and natural history of butterflies of E] Salvador. IB. Hamadryas februa (Nymphalidae-Hemadryadinae). J. N. Y. Entomol. Soc. 53: 157-169. 1975b. Notes on the life cycle and natural history of butterflies of El Salvador. IIB. Hamadryas guatemalena (Nymphalidae-Hamadryadinae). J. N. Y. Entomol. Soc. 53: 170-180. 1975c. Notes on the life cycle and natural history of butterflies of El Salvador. IIIB. Hamadryas amphinome (Nymphalidae-Hamadryadinae). J. N. Y. Entomol. Soc. 53: 181-191. Muysuonptr, A., Jr. & A. MuysHonpr. 1976. Notes on the life cycle and natural history of butterflies of El Salvador. IC. Colobura dirce L. (Nymphalidae- Coloburinae). J. N. Y. Entomol. Soc. 54: 22-33. Reuter, E. 1896. Uber die palpen der Rhopaloceren. Acta Soc. Fennic. Helsinfors. ScHatz, E. & J. Roper. 1892. Die Familien und Gattungen der Tagfalter, in Staudinger, O. & E. Schatz, Exotische Schmetterlinge, 2. Seitz, A. 1921. Macrolepidoptera of the World. Vol. 5. Stuttgart. STANDLEY, P.C. 1922. Trees and shrubs of Mexico (Fagaceae-Fabaceae). Contrib. U.S. Natl. Herb. 23, Part 2. 296 p. Younc, A.M. 1972. The ecology and ethology of the tropical nymphaline butterfly, Victorina epaphus. I. Life cycle and natural history. J. Lepid. Soc. 26: 155-170. Younc, A. M. & A. MuysHonpr. 1973. Ecological studies of the butterfly Victorina stelenes (Lepidoptera-Nymphalidae) in Costa Rica and EI Salvador. Stud. Neotrop. Fauna 8: 155-176. Journal of the Lepidopterists’ Society 32(3), 1978, 175-177 SCELIODES LAISALIS (PYRALIDAE): DESCRIPTION OF THE MATURE LARVA AND NOTE ON ITS FEEDING HABIT E. O. OGUNWOLU Entomology Division, National Cereals Research Institute, P.M.B. 5042, Ibadan, Nigeria ABSTRACT. Mature laboratory-reared and field-collected larvae of Sceliodes laisalis Wlk. (Pyralidae) are examined and described and the chaetotaxy illustrated. A brief note on the feeding habit of this species, an economic pest of Solanum fruits in southwestern Nigeria, is given. Taxonomy of immature insects has long been neglected in Nigeria; mostly, general descriptions of immature stages are contained in publica- tions on the biology of some species. Where a complex of species is involved, most taxonomic studies and identifications were of adults. In contributing to the growth of taxonomy of immatures, I here describe the larva of Sceliodes laisalis Wlk. and illustrate its chaetotaxy. Its feeding habit is also described. Sceliodes laisalis, widely distributed in southwestern Nigeria (Akinlo- sotu, 1977), is an economic pest of the garden egg fruit, Solanum macro- carpon L. and S. melongena L. (Solanaceae). Larvae reduce the quality and quantity of the fruits and the seeds. My descriptions are based on 15 field collected specimens and 25 larvae reared in the laboratory. In the following description, names of setae follow Hinton (1946). To describe the feeding habit at Ibadan, I ex- amined fruits of ten S. macrocarpon and S. melongena cultivars and those of Capsicum annuum L.., C. frutescens L., and Lycopersicum esculentum Mill. for Sceliodes attack. Description of Larva General. Length 16.0-22.4 mm, mean = 17.6 mm; width 3.0-4.1 mm, mean = 3.3 mm. Head yellowish brown often with dark brown maculation; ocellar area deep brown, a fuscous band occurs from post-occiput narrowing towards but not reaching the ocellar area (Fig. 1B). Prothoracic shield pale with brown spots and deep brown areas postero-medially; mesothorax suffused with brown; thoracic legs brown. Abdomen pink dorsally excepting the deep brown anal shield; venter pale. Setal pinacula light brown. Spiracles circular, yellowish with brown rim; prothoracic and 8th abdominal spiracles each larger than the others. Head (Fig. 1A). Width 1.1-1.5 mm, mean = 1.2 mm; subspherical in frontal view. Adfrontal suture reaching the acutely angled vertex. Labrum shallow and emarginate, mandible with 5 teeth, all but the distal one pointed (Fig. 5), mesal surface with 4 ridges. Each of the first, second and the sixth ocellus larger than the others; the second ocellus closer to the first than to the third. Posterior seta P2 about % the length of P1; 176 JOURNAL OF THE LEPIDOPTERISTS SOCIETY SDI ————————— Figs. 1-5. Chaetotaxy of Sceliodes laisalis larva. 1, head, frontal view (A) and lateral view (B); 2, thoracic segments; 3, first, sixth through ninth abdominal seg- ments; 4, anal shield; 5, right mandible. this inserted about mid-length of the head. Seta P1 closer to adfrontal seta AF1 than to AF2. Distance from AF2—AFa puncture one-third to one-half that from AF2-AF1. fa punctures well below frontal setae F1; distance between these one-half to three- fourths that between clypeal setae C2. A2, A3, and LI setae obtuse angled. Vertical setae V1 slightly antero-ventrad from V2; this closer to V3 than to V1. Ocellar seta Ol close to the 3rd ocellus, O2 seta ventrad from the Ist ocellus. Distance from genal seta G1-Ga puncture less than 4% that between Ol—Oa puncture. VOLUME 32, NUMBER 3 AT Thorax (Fig. 2). Seta MxD1 outside prothoracic shield. On this seta XD2 dorsad from and closer to SD1 than to XD1. On meso- and metathorax each of dorsal and subdorsal setae on the same pinaculum; lateral seta L3 postero-dorsad from LI; distance from MV2—MV3 about one-third that from MV3-V1. On mesothorax MSDI1 and MSD2 on the same pinaculum but on separate pinacula on metathorax. Pro- thoracic coxae close but not touching, mesothoracic coxae about the coxal width apart, and the metathoracic coxae generally more than the coxal width apart. Abdomen (Fig. 3). D1 and D2 pinacula equal in size and closer on the 8th than other segments. On the 9th segment, dorsal setae D2 on the same pinaculum; D1, a thin seta, is slightly closer to SD1 or equidistant between D2 and SD1. Seta MDI antero-ventrad from D1 on segments 1-8; lateral pinaculum larger on the 8th than on the other segments. Subventral setae. 1: 3: 2: 1:1 on segments 1, 2, 7-9, respectively. Distance between V1 pinacula on the 7th segment more than 2 that of the 9th segment. Anal shield (Fig. 4). Rounded posteriorly, SD1 setae slightly to well above D1 setae. Prolegs on segments 3-6 with crochets, in biordinal mesoseries, respectively numbering 16-22, mean = 19; 17-22, mean = 20; 17-23, mean = 22; and 18-24, mean = 22; and on anal proleg, 14-17, mean = 16. Description of Feeding Habit In feeding, larvae tunnel within the fruit which shows no external sign of damage while larvae are small. The mature larvae exit through holes made on the fruit. These holes serve as entry sites for decay organisms, whose activities eventually cause fruit rot. The cultivars of S. macrocarpon and S. melongena sampled (n = 5 for each) were all attacked by Sceliodes larvae. Two to five larvae were found within each fruit. I found no Sceliodes larvae in fruits of Capsicum annuum, C. frutescens, and Lycopersicum esculentum and neither did Akinlosotu (1977), but Davis (1964) recorded these as hosts of Sceliodes cordalis (Dbld.) larvae. LITERATURE CITED AxINLosotu, T. A. 1977. A check list of insects associated with local vegetables in Southwestern Nigeria. Univ. Ife. Inst. Agr. Res. Bull. 8 (In press). Davis, J. J. 1964. The egg fruit caterpillar. Queensld. Agr. J. 90: 76-78. Hinton, H. E. 1946. On the homology and nomenclature of the setae of lepidop- terous larvae, with some notes on the phylogeny of the Lepidoptera. Trans. Roy. Entomol. Soc. 97: 1-37. Journal of the Lepidopterists’ Society 32(3), 1978, 178-190 MIGRATION AND RE-MIGRATION OF BUTTERFLIES THROUGH NORTH PENINSULAR FLORIDA: QUANTIFICATION WITH MALAISE TRAPS? THoMAS J. WALKER Department of Entomology and Nematology, University of Florida, Gainesville, Florida 32611 ABSTRACT. Malaise traps with single linear barriers perpendicular or parallel to the axis of the Florida peninsula were operated from 18 Sept. 1975 to 17 Sept. 1976 near Gainesville; insects intercepted by the two surfaces of each barrier were captured separately allowing them to be scored as flying northward, southward, eastward, or westward. During the fall, significantly more individuals were caught flying south- ward than northward for eight species of butterflies: Urbanus proteus (Linnaeus); Phoebis sennae (Linnaeus); Precis coenia (Hubner); Panoquina ocola (Edwards); Agraulis vanillae (Linnaeus); Lerema accius (Smith); Urbanus dorantes (Stoll); and Eurema lisa Boisduval and Le Conte. Estimated net numbers flying southward across each meter ranged from 3956 (U. proteus) to 33 (E. lisa). During the spring significantly more individuals were caught flying northward than southward for two species: P. coenia, A. vanillae. Estimated net numbers were 150 and 10 per m, respectively. Malaise traps can continuously and effectively monitor insect migration within the boundary layer. Long-distance flights by insects are frequent and of theoretical and practical interest (Williams, 1958; Johnson, 1969; Dingle, 1972). Such movements are difficult to study because quantification requires identi- fying flying insects and determining their directions of movement as well as counting them. Most long-distance flights of insects may occur at night or at high altitudes making detailed observations impractical, al- though mass flights above 10 m can be studied with radar (Schaefer, 1976; Riley, 1975). Direct visual observation is useful for large insects that fly low in daylight. Butterflies have been the most frequent subjects of such observations (Arbogast, 1966; Baker, 1968b; Balciunas and Knopf, 1977). Since direct observation is difficult and time consuming, the resulting data are generally skimpy and likely to be biased by choice of observation times. Malaise traps (Southwood, 1966) can complement direct observation of flights of low flying insects by continuously sampling without the presence or bias of an observer. Appropriately modified, a Malaise trap can separate insects flying in one direction when intercepted from those flying in another direction. I used four such traps to monitor insect flights within 2 m of the ground for one year near Gainesville, Florida. ‘Florida Agricultural Experiment Station Journal Series No. 457. VOLUME 32, NUMBER 3 179 Fig. 1 (above). N-—S trap at Green Acres site, looking NNW. Fig. 2 (below). Heads at one end of trap, showing how insects from the two sides of the barrier were kept separate. METHODS Special Malaise traps (Fig. 1) were constructed and are now com- mercially available.?, Each trap had a 2.6 X 6 m central barrier. Insects flew into the trap by either of two 2 X 6 m openings that faced in opposite 2D. A. Focks & Co., P. O. Box 12852, University Station, Gainesville, Fla. 32604. 180 JOURNAL OF THE LEPIDOPTERISTS SOCIETY directions. Upon striking the central barrier they sometimes worked their ways toward either end, through a truncated funnel, and into a receptacle where they were killed by vapors from pieces of dichlorvos- impregnated plastic.? Traps were made so that insects entering through one opening remained separated from those entering through the other opening (Fig. 2). The receptacles were emptied daily or two or three times per week depending on the numbers caught. Insects caught within the trap but not present in the receptacles were killed and added to the appropriate batch. Mating status of samples of female migrants was determined by dis- secting for spermatophores. Traps were set at two sites, 15 km apart: (1) Green Acres Farm, Agronomy Dept., U. of Fla., 18 Sept—18 Oct. 1975 (4 traps); 19 Oct.—2 Nov. 1975 (2 traps); 10 Apr.—6 June 1976 (2 traps); (2) Archer Road Laboratory, Entomology and Nematology Dept., U. of Fla. 19 Oct. 1975- 17 Sept. 1976 (2 traps). The first site was an open field with no buildings or woods within 100 m (Fig. 1). The second was a lawn-like area with buildings 50 m to the west and east. Traps were set in pairs with one member of each pair oriented WSW-ENE (perpendicular to the axis of the Florida peninsula and to the predicted track of migrants—henceforth called a N-S trap) and the other (an E-W trap), 30 m away, NNW-SSE (parallel to the axis of the Florida peninsula and at right angles to the N-S trap). The insect-catching devices (heads) of the traps were improved during the first month of the study by changing the receptacles from translucent polypropylene jars to transparent bags (Fig. 2). Even with improved heads, a trap captured only a small portion of the insects that flew over the 6-meter line defined by its barrier. The efficiency of traps was estimated from counts of individuals captured versus individuals evading capture during observation periods (4, 5, 12, and 26 Oct., 1975). RESULTS For eight species of butterflies the N-S traps caught significantly more individuals flying southward than flying northward in the fall (Table 1). For two of the eight, N-S traps caught significantly more individuals flying northward than flying southward in the spring. Estimating net numbers moving southward (or northward) each week or each season requires not only counts of individuals caught but also an estimate of trapping efficiency. During the four observation periods to determine efficiency of N-S traps with improved heads, 28 of 314 Urbanus “e.g. 3 4 6 cm pieces of No Pest or Stable Strip, Shell Chemical Co. 181 VOLUME 32, NUMBER 3 d) pireMyynos uvy} prweMYyHOU SurATZ o1OW APUKOTFIUSIS ; d) piemy}iou UeYy} pleMyY MOS SuIATF o10UL APULOTFIUSIS ¢ ‘OL6I “349g LT—SNY 9G “GLET “AON 9G—1d2S 8T z *(9x0} 99S) esn Ul 919M Spvoy PeAOIdUIT 1O [eUISLIO Jey}oyYM pu Suljerodo sde1} jo Joquinu jUNODDB OUT 9xe} AsYy} ovsnvoeq peddei} siaquinu YM ApoazF10d 93vTJa1100 yOU Op sazeUIT}s9 ssey], “UOTOeIIp aysoddo ur surAou asoy} JO ssaoxe UI oul] ANA-AASMA W-T ®& ssoi0e UOTOIIP 9UO UL SUTAOU JequINnU pojJeUIT}s| ‘(S0'0 > “(S0';0 > 0 0 0 0 0 0 (Se) el G DsSYy A 0 0 0 0 0 0 (IPF) OV CT SaqjuDLOp *() 0 0 (3) 0 @ 0 0 (PL) eVG 6 SMIID “'T (7) SG G (Ol) 6 Sil 0 0 (ABIL) &B 0 anjupa “Vv 0 0 I I 0 0 (Os) av I D1090 ‘d I 0 (7) 0 6 (¢) SG 0 (S98) ers OT apuuas “J i i (OST) PF SGT (Z) G S (GIE) arg eee ‘el “O[[IAsouresy rvou ‘9/6 3deg LT—-Gy6T ‘deg QT ‘suosves INOJ sulinp (sesoyyuered ut ) ueWeDe|dsIp jou poezeuII}so pue pleMmyjynos puke preMy}10U SuTATF podde.y spenprArpur Fo sIequInN] ‘[ FIV], 182 JOURNAL OF THE LEPIDOPTERISTS SOCIETY proteus (8.9%) were captured. The efficiency for single observation periods varied from 2 to 12%: 3 of 28 (11%), 19 of 159 (12%), 2 of 88 (2%), 4 of 39 (10%). The lowest efficiency (2%) occurred when the wind was mainly from the north and the skippers were flying higher than usual: 70% flew over the trap without first hitting the barrier com- pared to 38-43% for the other three periods. So few of the other species were flying that no reliable estimate of capture efficiency was obtained. During the four observation periods 2 of 8 Precis coenia were captured, 0 of 7 Phoebis sennae, and 0 of 3 Agraulis vanillae. None of these ratios differ significantly from the 8.9% observed for U. proteus (chi square, P > 0.05). To simplify calculation of estimates of net displacement (Table 1 and Fig. 3) while keeping well within the limits suggested by the data on trapping efficiency, I assumed that each trap with improved heads captured 10% of the individuals flying over its 6-m line. For N-S traps with original heads, I assumed a 2.5% efficiency, since when operated simultaneously with N-S traps with improved heads their catches were approximately one-fourth as great (e.g. 115 compared to 469 U. proteus ). Conversion to improved heads was completed 17 Oct. 1975. E-W traps caught approximately the same number of insects flying eastward as flying westward. The only significant exception (P < 0.05) was for U. proteus; 432 were caught in eastward flight versus 177 in west- ward flight (=2.4:1) during 18 Sept._17 Nov. Such a bias would be ex- pected if the average track of southbound migrants was east of SSE (158°), the orientation of the central barrier of E-W traps. Balciunas and Knopf (1977) determined that the mean track was 147°—i.e., 11° east of SSE. Beginning 2 Nov. counts of individuals caught in each end of all traps were recorded separately. The north and south ends of E—W traps showed approximately the same biases as the north and south sides of N-S traps. For example, of the 144 U. proteus trapped 2-17 Nov. in one E-W trap, 137 were caught in the south end. (Data from E-W traps were never used in estimating net displacement northward or southward.) The fall flights lasted for six weeks or longer (Fig. 3). The continuous nature of the fall > Fig. 3. (Top to bottom) U. proteus, P. sennae, P. coenia, A. vanillae. Weekly occurrence and net displacement northward or southward 18 Sept. 1975 through 17 Sept. 1976. Downward bars show net displacement southward; upward bars show net displacement northward. Solid bars indicate a significant (P < 0.05) inequality in numbers caught flying northward and southward. The lengths of the bars show the estimated numbers of individuals flying southward (or northward) across 1 m perpendicular to the axis of the Florida peninsula in excess of those flying in the other direction. Estimates were made from the numbers caught in 1 or 2 N-S traps. Traps VoLUME 32, NUMBER 3 183 400, Long-Tailed Skipper (n=2505) (northward) (southward) ® go. Cloudiess Sulphur (n=259) ® z SS ea @ @ E ~~ ® E 3s 100 Cc go. Buckeye (n= 302) ke za LJ = uJ O 6 I ‘AON ‘pO ydas ‘Sny Ajnf{ oun ACN [lidy (panuyu0on) °% AIaVy, 212 JOURNAL OF THE LEPIDOPTERISTS SOCIETY strong flying and wide ranging species which may or may not have used the area for reproduction. All species recorded during this study were previously seen by me in other similar habitat and in other vegetation types in southern Arizona. No species not recorded on the study area were seen in adjacent areas. I thus consider the species list relatively complete. The butterflies of the study area were representative of the Southeastern Arizona desert scrub habitat with considerable influence from the foothill canyon habitat as outlined by Brown (1965). All species seen on the study area were recorded in one or both of the above habitats by Brown (1965) except V. annabella and E. amyntula. The Santa Rita Range butterfly fauna belong to 10 families (if Apaturidae and Heliconiidae are separated from Nymphalidae). Nym- phalidae and Hesperiidae were represented by the most species (8 each) followed by Lycaenidae (7) and Pieridae (6). Overall, the familial com- position was more diverse than in many temperate communities (e.g., Emmel and Emmel 1963b) which often lack representatives of families with southern distribution (Heliconiidae) or with restricted larval food habits (Apaturidae and Libytheidae). Apaturidae was by far the most abundant family in terms of individuals, due to the high abundance and long flight period of A. leilia. Lycaenidae and Libytheidae were next in individual abundance followed by Riodinidae and Pieridae. The two families with most species were represented by very few individuals. This distribution of individuals by family contrasts sharply with that in the Sierra Nevada of California where individual and species distribution was similar (Emmel and Emmel 1963b). Phenology Activity by adult butterflies was observed in all months except November 1970 and January and February 1971. The general pattern of activity was similar during the two years (Fig. 1). During spring and early summer there was a plateau in number of species which then increased rapidly to a peak in late summer before decreasing to a small number of species in fall. In both years, there were peaks of abundance in both spring and late summer with a period of nearly no activity between (Fig. 1). In 1970, the spring peak was much larger, and the summer peak | began earlier and lasted longer than in 1971. Phenology of the fauna as a whole appeared largely related to rainfall. Precipitation for winter 1969-70 was near average, and the spring popula- tion in 1970 was probably typical for the area. The very dry winter of VOLUME 32, NUMBER 3 ANS 00 (0) oe) Oo e) p (e) 20 NO. OF INDIVIDUALS NO. OF SPECIES 4 1970 4- |971 RAINFALL (CM) M A M J J A S O N D MONTH Fig. 1. Rainfall pattern (1 May—5 Sept.) and butterfly activity on the Santa Rita Experimental Range, Arizona, in 1970 and 1971. 214 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 1970-71 was probably responsible for the reduced diversity and abun- dance in spring 1971. Differences in the overall summer flight period can be directly attributed to differences between the summer rainfall patterns (Fig. 1). The first heavy rain of 1970 was on 29 June and was followed by another similar rain less than a week later. These were sufficient to stimulate the summer growth of vegetation. Butterfly diversity and abundance increased rapidly after these rains but diversity decreased rapidly during the below average dryness of August. Abundance increased in September due mainly to the emergence of a large brood of L. bachmanii. Both diversity and abun- dance decreased rapidly in late September and early October. The 1971 rainy season began on 2 July with a relatively light rain. The next rain did not occur until 14 July after which there were several addi- tional, closely spaced rains (Fig. 1). The vegetation responded minimally to the first rain but much of the new growth dried considerably between rains. The second and subsequent rains initiated and maintained the usual spectacular summer growth of vegetation. There was no emergence or immigration of butterflies with the first rain. The second rain, however, was followed by a rapid increase in species number and abundance similar to the increase after the first rain of 1970. The timing of the autumn de- crease was similar to 1970. The 3 principal spring species showed a peak in about late May or early June with continued emergence into early fall as noted above. Several patterns were evident among the rainy season species. Certain resident species showed an immediate emergence within a few days of the first rain. These included A. leilia, D. chara, P. catullus and possibly E. funeralis. Other species including the immigrant D. gilippus, P. sennae and possibly C. caesonia and the resident E. nicippee occurred in peak numbers 2-4 weeks after the beginning of the rains. The second brood of A. palmeri was timed similarly. Additional immigrants, V. cardui and B. philenor, showed peak abundance about 2 months after the beginning of the rains. In both years, however, L. bachmanii reached peak abun- dance in mid-September. In all other instances where sufficient records exist, the summer peak in abundance was later in 1971 than in 1970 similar to the overall pattern for all species combined as discussed above. Nearly all species had greater peak abundance in 1970 than in 1971. The relationships of the various species within a larval food plant guild are complex. Adults of the principal Prosopis feeders, A. palmeri and H. ceraunus, were spring fliers with the latter flying slightly later than the former although A. palmeri had a relatively large brood after the rains in 1970. Another Prosopis feeder, M. leda, emerged only after the rains VOLUME 32, NUMBER 3 AUS began. Of the two species which feed as larvae on Cassia, E. nicippee was resident and P. sennae was an immigrant. Both showed peak abundance from early August to mid-September. There was almost complete overlap of peak flight activity in 1970 but P. sennae showed peak abundance earlier than E. nicippee in 1971. The Celtis feeding guild, represented by A. leilia and L. bachmanii, showed overall peak abundance in mid- September. The two years differed with little overlap in peak flight period in 1970 and almost complete overlap in 1971. The few notes taken on adult resource use indicate use of several flowering species. The spring species were most often seen visiting Acacia and Prosopis flowers. In summer, flowers of Mirabilis multiflora (used principally by P. catullus), Ipomoea coccinea (B. philenor), Hymenoclea salsola (D. gilippus, L. bachmanii) and Zinnia pumila (D. chara, E. claudia) were visited. Temporal partitioning may be partly responsible for increased diversity of some butterfly populations (Clench 1967). Replacement of one set of species by another within a season allows increased diversity without increasing competition for adult resources. It must be recognized, how- ever, that the observed phenology of adult activity is often closely integrated with the phenology of immature stages which is largely beyond the scope of the present discussion. The numerical data in Table 2 were analyzed using the methods of MacArthur (1964) and Ricklefs (1966). Season length was calculated using the information-theoretical measure (H’) for the combined abun- dance of all species (total season) and for each species that was observed in 4 or more 10 day periods (specific season). The specific seasons were averaged and divided into the total season resulting in a figure which indicates species turnover through the total flight season. Use of H’ places greater weight on periods of abundance and less on periods of rarity and more accurately reflects the length of the flight period than extreme dates. Total season length was about 11 ten-day periods in both years (11.4 in 1970, 10.7 in 1971) and average specific season was over 5 ten-day periods (5.31 in 1970, 5.26 in 1971). There appeared, therefore, to be two turn- Overs among the species present. The turnover in 1970 was obvious with the rainy season species clearly replacing the spring species. The 1971 replacement was less clear but apparently involved the replacement of early rainy season species by those of the later parts of rainy season. While this may have had some effect in 1970, it was largely marked by the large spring broods and longer rainy season of that year. In general, the phenological patterns were similar to those found by 216 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Brown (1965). The major differences were that Brown did not find spring broods of A. palmeri, L. marina and H. ceraunus, although he found L. bachmanii and D. chara in spring, two species which I did not observe. In southwestern New Mexico, Ferris (1976) also found phenology to be relatively similar to southern Arizona. He noted a spring brood of all of the above 5 species. Voltinism Voltinism varied both inter- and intraspecifically (Table 2). Two (D. chara, P. catullus) of the 9 common residents were univoltine during both years. E. nicippe and E. funeralis appeared to be so in 1971 but were at least bivoltine in 1970. Eight rare species were encountered frequently enough to determine voltinism. In 1970, M. leda, P. communis and C. hippalus were univoltine whereas S. melinus, E. amyntula and N. iole were bivoltine. In 1971, L. eufala was univoltine and H. isola was bivoltine. Thus 33% of the resident fauna was univoltine in 1970 and 46% in 1971. Univoltine species, in all cases, appeared in summer after the beginning of the rains. Univoltinism in other long summer faunas is principally a spring and early summer phenomenon (Shapiro 1975). The mid- to late summer univoltinism in southern Arizona is undoubtedly related to the greatly increased suitability of the habitat following the summer rains. The remainder of the Santa Rita fauna was at least bivoltine. Three species (A. palmeri, L. marina, H. ceraunus ) had large spring and smaller summer broods, especially in 1971. The summer populations were repre- sented by scattered individuals observed over a period of 2-3 months. In 1970, A. palmeri may have had 4 broods. These 3 species may be spring univoltines in years when the summer rains fail. Four rare species (S. melinus, E. amyntula and E. funeralis in 1970 and H. isola in 1971) had a distinct brood in early June and at least one other of about the same size following the beginning of the rains. S. melinus and H. isola appeared to have but one post-rain brood; the others possibly 2 or 3. In both years, small numbers of A. leilia occurred before the rains. There were 3 distinct peaks possibly indicating 3 additional broods after the rains in 1970, but in 1971 there was but one large peak in mid and late September. Fresh individuals were noted throughout the flight season of both years indicating continued emergence. Ikmergence was limited to after the rains in L. bachmanii, E. nicippe and N. iole. L. bachmanii had a small brood immediately after the be- ginning of the rains, a large brood in September and possibly another one or two smaller broods still later in the season. E. nicippee appeared VOLUME 32, NUMBER 3 J2AT 2-576 1970 2.0 e (971 0.8 0.4 0:2 A M J J A S O N MONTH Fig. 2. Seasonal changes in species diversity (H’) and equitability (J’) of butterflies on the Santa Rita Experimental Range, Arizona. to have at least 2 broods in 1970 but probably only one in 1971 unless the October records were of an additional brood. N. iole had at least 2 broods in 1970 but was all but absent in 1971. Two immigrant species (D. gilippus, P. philenor) occurred in small numbers before the rains and reached a definite peak in numbers in late summer. P. sennae also showed a peak in abundance after the rains. Other immigrant species occurred irregularly in low numbers after the rains. Unworn individuals of these species occurred throughout the season. 218 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Species Diversity Diversity of a population can be examined in several ways. The method most often used for butterflies is species counts for each of several seasons. This diversity measure for the Santa Rita fauna was examined previously (Fig. 1). Another measure of diversity based on information theory (H’) accounts for both species numbers and relative abundance. This measure has not been widely used for insect populations (Janzen and Schoener 1968) mainly because of the wide range of detectability of the component species (see Shapiro 1975). In this study, I made a concerted effort to obtain accurate counts in a relatively narrow transect and none of the species encountered were particularly secretive. I believe that no species was grossly under- or over-represented in my counts and that the numbers in Table 2 reflect the true relative abundance. Therefore, I used these values to calculate H’ for each 10 day period of the two seasons. The ratio of H’ to maximum diversity possible if each species were equally abun- dant is another component of diversity termed equitability (J’, see Pielou, 1966). These were also calculated for each 10 day period. Diversity, measured by H’, showed a seasonal pattern which closely paralleled species counts (Fig. 2). Linear regression analysis showed that the number of species explained over 80% of the variation in H’ (r = .899, N = 31). In contrast, equitability fluctuated widely, showed no correlation (r = .005) with species counts and was most constant during the rainy season (Table 2). Considering each season as a whole, H’ and J’ were slightly greater (2.34 and 0.67) in 1970 than in 1971 (2.20 and 0.63). The average J’ for Santa Rita butterflies of about 0.65 is slightly lower than the average of 0.74 calculated for insect populations containing a larger number of orders and thus greater trophic diversity (see Austin and Tomoff, in press). CONCLUSIONS The complexity of the phenology of the southern Arizona butterfly fauna was first indicated by Brown (1965). He noted that most lowland species were rainy season fliers or had a spring brood and additional broods during the rainy season. He further recognized the wide annual fluctuations with populations dependent on the local precipitation. Several factors seem apparent in the flight patterns of the species present. Strict seasonal phenology with only the magnitude affected by rainfall was exhibited by the 3 principal spring species and by L. bach- manii in fall. Spring flight by the Prosopis feeders is timed for the larvae to take advantage of the fresh herbage or flower buds and for the adults’ VOLUME 32, NUMBER 3 219 nectar source which appears to be mainly Prosopis and Acacia. The second brood of A. palmeri appears timed to the new herbage growth by Prosopis following the summer rains. Flowering by Prosopis following the rains is limited and this may account for the very small numbers of L. marina and H. ceraunus at this season. The flight seasons of the remaining species are nearly limited to after the beginning of the summer rains. This suggests that rainfall itself and/or the resultant increase in humidity combined with the warm summer temperatures act to break diapause of the resident species. The rapidity of which adults appear after the first rain may relate to the stage of the life cycle at which diapause occurs or to a cumulative effect of successive rains. The enforcement of diapause by heat and aridity in desert regions has been previously noted ( Wiltshire, 1956). Non-resident individuals of wide ranging species may be attracted by increased vegeta- tion growth and flowering by many plants. Adults of 12 species of butter- flies were observed feeding at the flowers of 2 plant species, Mirabilis and Zinnia, which flower abundantly and nearly exclusively after the rains begin. The nearly consistent lateness of the 1971 season compared to 1970 (Table 1) is further indication that rainfall is the important ultimate factor in the phenology of many species. The seasonality of the Santa Rita Range butterfly fauna differs con- siderably from other temperate zone faunas previously examined. The general trend in most populations is for a rather rapid increase in species number to a mid summer peak and then a decrease into autumn (Shapiro, 1975). Only the tidal marsh in California showed a peak in species numbers in late summer or fall. No temperate butterfly community showed the strict dependence on summer rains. These differences in phenology (and voltinism) can be largely attributed to climatic differ- ences; the areas studied by Shapiro (1975) were largely Mediterranean or Mediterranean-montane in climate. The Arizona community was similar to others, however, in that peak populations corresponded to peak vegetation growth. Seasonal phenology in southern Arizona shows certain similarities to that in the Neotropics where Ebert (1969) found peak activity during the rainy season. ACKNOWLEDGMENTS This study was conducted while I was funded to study birds by the US/IBP Desert Biome Program under National Science Foundation Grant GB15886 at the University of Arizona. I thank Donald Thomas and Robert Ricklefs for critical comments on an early draft of the manuscript. i) bo — JOURNAL OF THE LEPIDOPTERISTS SOCIETY LITERATURE CITED Austin, G. T., & C. S. Tomorr. Relative abundance in bird populations. Amer. Natur., in press. Brown, K. S., Jr. 1965. Some comments on Arizona butterflies (Papilionoidae). J. Lepid. Soc. 19: 107-115. Ciencu, H. K. 1967. Temporal dissociation and population regulation in certain Hesperine butterflies. Ecology 48: 1000-1006. Epert, H. 1969. On the frequency of butterflies in eastern Brazil, with a list of the butterfly fauna of Pocos de Caldas, Minas Gerais. J. Lepid. Soc. 23, supple- ment no. 3. EMMEL, T. C., & J. F. EmmMen. 1962. Ecological studies of Rhopalocera at Donner Pass, California. I. Butterfly associations and distributional factors. J. Lepid. Soc. 16: 23-44. & 1963a. Ecological studies of Rhopalocera at Donner Pass, Cali- fornia. II. Meteorological influences of flight activity. J. Lepid. Soc. 17: 7—20. & 1963b. Composition and relative abundance in a temperate zone butterfly fauna. J. Res. Lepid. 1: 97-108. Humpurey, R. R. 1968. The desert grassland. Univ. of Arizona Press, Tucson. Lowe, C. H. (ed.). The vertebrates of Arizona. Univ. of Arizona Press, Tucson. MacArruur, R. H. 1964. Environmental factors affecting bird species diversity. Amer. Natur. 98: 387-397. PreLou, E. C. 1966. The measurement of diversity in different types of biological collections. J. Theoret. Biol. 13: 131-144. RickteFrs, R. E. 1966. The temporal component of diversity among species of birds. Evolution 20: 235-242. SHAprro, A. M. 1975. The temporal component of butterfly species diversity. Pp. 181-195 in M. L. Cody and J. M. Diamond (eds.) Ecology and Evolution of Communities. Belkaap Press, Cambridge, Mass. Wixtsuire, E. P. 1956. Notes on the diapause of Lepidoptera in hot arid sub- tropical climates. J. Lepid. Soc. 10: 201-203. Journal of the Lepidopterists’ Society 32(3), 1978, 221-223 A NEW HINDWING ABERRATION OF CATOCALA MICRONYMPHA GUENEE FROM KENTUCKY CHARLES V. COVELL, JR.? Department of Biology, University of Louisville, Louisville, Kentucky 40208 ABSTRACT. An aberration of the underwing Catocala micronympha Guenée with forewings like “form gisela” and entirely black hindwings is described from Oldham County, Kentucky. It is informally named “form sargenti” in honor of Dr. T. D. Sargent. Catocala micronympha Guenée was very common in Kentucky in June and July 1977, and a large number of specimens representing the various forewing variations was collected in several counties. On the night of 15 June I took at blacklight a single male of what appears to be the first known individual with forewings of form “gisela” Meyer, but with hind- wings completely black on the upperside except for the yellowish-white terminal line, fringe, and apical patch (Fig. 1). The underside of the hindwing has a diffuse remnant of the median yellow band below the costal margin (Fig. 3). Wingspan is 4.4 cm. A typical form “gisela” from Bernheim Forest, Kentucky, also taken on 15 June 1977, is shown for contrast (Figs. 2, 4). The locality from which this specimen was taken was the University of Louisville’s research farm, the Horner Bird and Wildlife Sanctuary, about 20 miles (32 km) southeast of Louisville in Oldham County, near the hamlet of Brownsboro. No Catocala aberrations of any kind had been collected or seen there in over 12 years of fairly heavy collecting until this collection was made. No achromatic hindwing aberrations of C. micro- nympha were reported in Barnes and McDunnough (1918), Forbes (1954), or Sargent (1976). Personal communication with Drs. T. D. Sargent and D. C. Ferguson leads me to conclude that this specimen is unique. Sargent (1976) states that, “. . . hindwing polymorphisms are virtually unknown . . .” in Catocala (p. 77), while the forewings in some species (such as micronympha) are highly polymorphic. Single examples of hind- wing aberrations are also rare. He further states (p. 111) that, “In the Catocala the most prominent “sports” are those involving substantial alterations of the normally invariable hindwings of a species.” Those eastern North American underwings for which there are named aberrations involving all-black or nearly all-black hindwings, where contrasting light- colored bands are typical, include the following (with descriptions from 1 Univ. of Louisville Contributions in Biology No. 190 (New Series). bo bo bo JoURNAL OF THE LEPIDOPTERISTS SOCIETY Figs. 1-4. Catocala micronympha Guenée. 1. aberration “sargenti” Covell, upper- side; 2. form “gisela” Meyer, upperside; 3. aberration “sargenti,”’ underside; 4. form “gisela,” underside. Sargent): C. muliercula Guenée, ab. “peramens” Hulst (“HW almost entirely black”); C. ilia (Cramer), ab. “normani’” Bartsch (“blackish FW from base to pm line, and extended black on HW”); C. unijuga Walker, ab. “fletcheri” Beutenmuller (“HW entirely black”); C. grynea (Cramer), ab. “constans” Hulst (“HW almost totally black”); and C. habilis Grote, ab. “depressans” Sargent, named by him on the basis of a single specimen with nearly all-black hindwings which escaped his killing jar (Sargent, 1976, p. 111, 113; Plate V, 2). While Latinized names for aberrations have no standing in zoological nomenclature, Dr. Sargent coined “depressans” and 3 melanic form names in his book as convenient “handles” for such forms. I am therefore fol- lowing his example and name the new aberrant form Catocala micro- nympha, aberration “sargenti,” in honor of Dr. Theodore D. Sargent in recognition of his contributions to the study of North American Catocala. The specimen on which this name is based is now in my possession, but will be deposited in the U.S. National Museum of Natural History at a later date, Note added in proof: On 6 July 1978, Loran D. Gibson collected a second male of this new form at light at Otter Creek Park, Meade County, Kentucky. The specimen was in worn condition, and is in the collection of the University of Louisville. VoLUME 32, NUMBER 3 225 LITERATURE CITED Barnes, W. J. & J. McDunnoucH. 1918. Illustrations of the North American species of the genus Catocala. Mem. Amer. Mus. Nat. Hist. 3 (1), 47 p., 22 pls. Forses, W.T. M. 1954. Lepidoptera of New York and neighboring states. Part III, Noctuidae. Cornell Univ. Agr. Exp. Sta. Mem. 329, 433 p. SarcGENT, T. D. 1976. Legion of Night: The Underwing Moths. Univ. of Mass. Press, Amherst. xii + 222 p., 8 pls. Journal of the Lepidopterists’ Society 32(3), 1978, 223 CONFIRMATION OF THE OCCURRENCE OF AN ALBINISTIC FEMALE FORM OF PHOEBIS PHILEA (PIERIDAE) IN EXTREME SOUTHERN TEXAS Phoebis philea (Johansson) is a large pierid butterfly common in tropical America. Individuals from Mexico enter southern Texas often (annually according to Howe, 1975, The Butterflies of North America, Doubleday, Garden City, N.Y., 633 p.). Males of this species are easily recognized by the striking contrast of yellow and orange portions of the dorsal forewings and hindwings. Females have the marginal dark markings typical of females of the genus, with the yellow and orange wing portions somewhat less contrasting. An albinistic form of philea was named “obsoleta” by Niepelt (1920, Int. Entomol. Zeit. 14: 17); this form corresponds to albinistic female forms in other species of Phoebis. On 12 August 1961 I collected one “obsoleta” in Brownsville, Cameron County, Texas. The dorsal wing surfaces were quite faded with scales totally lacking in isolated areas, particularly on the forewing discal cell area. Scales still present tend to be lightly greenish white. The ventral wing surfaces were also faded, but orange scales remain in sufficient numbers to provide the general color. One previous report of “obsoleta” from the extreme southern tip of Texas is known. Stallings and Turner (1946, Entomol. News 57: 44) reported a specimen collected in the Lower Rio Grande Valley. H. A. Freeman, who collected this first specimen, has kindly provided the data as follows: 23 August 1944 at a roadside park between Pharr and Hidalgo, Hidalgo County. My second specimen is of interest because local lepidopterists probably are unfamiliar with this form. Occurrence of philea in southern Texas is seasonal, with most specimens being reported from September to November (McGuire & Richard, 1974, An Annotated Checklist of the Butterflies of Bentsen—Rio Grande Valley State Park and Vicinity, Texas Parks & Wildlife Department, Mission, Texas, 21 p.). The worn condition of my specimen indicates that it arrived here after long-distance migration from some- where in northern Mexico. Substantial numbers of philea were found at least as far north as central Texas in late summer 1971 following an unusual climatic regime (Neck, unpub. data); no “obsoleta” were seen at this time by local collectors. RAYMOND W. Neck, Pesquezo Museum of Natural History, 6803 Esther, Austin, Texas 78752. 294 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Journal of the Lepidopterists’ Society 32(3), 1978, 224 A NEW WEEDY HOST FOR THE BUCKEYE, PRECIS COENIA (NYMPHALIDAE) The Buckeye, Precis coenia Hbn., is an opportunistic, oligophagous species hitherto recorded from several genera of the plant families Verbenaceae, Plantaginaceae, and Scrophulariaceae in North America. In the Sacramento Valley of lowland central Cali- fornia it feeds on two species of Lippia (Verbenaceae) and one of Plantago (Plantag- inaceae) (Shapiro 1974, J. Res. Lepid. 13: 120) while in the nearby Vaca Hills it occurs on Diplacus (Scrophulariaceae ) and at mid-elevations on the Sierran west slope on Penstemon azureus Benth. (Scrophulariaceae). In the San Francisco Bay area it has been found on garden snapdragons (Antirrhinum), an introduced scroph. On 17 September 1977 an infestation of fifth-instar larvae, probably from a single colonization event, was found on large mats of fluellin, Kickxia spuria (L.) Dumort, growing in cracks in an abandoned roadway at Davis, Yolo Co., California. This Mediterranean scroph is closely related to toadflax (Linaria) and to snapdragon and occurs sporad- ically in lowland California as a pavement and roadside weed. In captivity the larvae ate leaves, buds, flowers and fruit freely. No recorded hosts of any plant family could be found within 30 m of the infested plants. Adult P. coenia were present. Fluellin is a prostrate perennial plant with cordate, dull green, pubescent leaves less than 1 cm long. The flowers, which are open mostly in the morning, are snapdragon- like, with a long spur; they are purple and bright yellow and about 1 cm long. The stems, which may be 50 cm long, form tangled mats up to a meter across. Artuur M. SHapiro, Department of Zoology, University of California, Davis, Cali- fornia 95616, Journal of the Lepidopterists’ Society 32(3), 1978, 224 A SECOND LOCALITY FOR EULYTHIS MELLINATA (GEOMETRIDAE) IN NORTH AMERICA While identifying some Nova Scotian moths for James Edsall of Halifax, a specimen of Eulythis was examined which resembled no species known to occur in the province. A check on the identity of the specimen at the Nova Scotia Museum showed it repre- sents a Palearctic species, Eulythis mellinate F. (South, 1972, The Moths of the British © Isles, Warne, London, 379 p.), a new provincial record and the second locality in North America where this moth has been collected. Sheppard (1975, Ann. Entomol. Soc. Québec 20: 7) recorded this species from Laval, Québec, under the name associata Borkh, Eulythis mellinata is a widespread Palearctic species. In Europe and Britain the larvae feed on red and black currant ( Ribes rubrum L. & R. nigrum L.). Mr. Sheppard informs me that this species has probably become established on Mountain (Alpine) currant (Ribes alpinum L.) hedges in the vicinity of his home at Laval, Québec. The Nova Scotian specimen is a fresh female and was collected on 31 July 1972 at light in Armdale, Halifax, Nova Scotia. Other captures have been recorded in North VoLUME 32, NUMBER 3 DNS) Fig. 1. Eulythis mellinata F. Female from Armdale, Halifax, Nova Scotia. 31 July 1972. J. Edsall. 3.5x. America at Laval (Isle Jesus), Québec on 10 July 1967 (1 male), 24 June 1973 (1 female), 1 July 1973 (1 male) (Sheppard 1975, Ann. Entomol. Soc. Québec 20: 7), 28 June 1974 (1 male), 7 July 1974 (1 female), 29 June 1975 (1 female), 18 June 1976 (1 male) and 24 June 1976 (1 male) (Sheppard, 1977, pers. comm. ). The introduction of Eulythis mellinata in Nova Scotia was almost certainly recent as the specimen was collected in an area which has been intensively collected for the last 30 years, yet this is the only specimen which has been taken to date. The occur- rence of the moth in two widely separated localities in eastern Canada indicates well- established populations, and its occurrence in other eastern North American localities should therefore be expected. A photograph of the adult has been included to aid in identification. KENNETH NEL, Department of Biology, Dalhousie University, Halifax, Nova Scotia. Journal of the Lepidopterists’ Society 32(3), 1978, 225-226 OCCURRENCE OF THYMELICUS LINEOLA (HESPERIIDAE) IN NEWFOUNDLAND The recent rapid spread of the European Skipper, Thymelicus lineola (Ochsen- heimer) in North America, particularly in the northeastern part of the continent, evi- 226 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY dently has excited considerable interest (Burns 1966, Can. Entomol. 98: 859-866; Straley 1969, J. Lepid. Soc. 23: 76; Patterson 1971, J. Lepid. Soc. 25: 222). As far as Canada is concerned it is now listed (Gregory 1975, Lyman Entomol. Mus., McGill Univ., Ste-Anne de Bellevue, Québec, p. 11) as occurring in the provinces of Québec, New Brunswick, Nova Scotia, and British Columbia, as well as Ontario, where it was first noted on this continent in 1910 (Saunders 1916, Ottawa Nat. 30: 116). The butterfly was certainly present in great numbers in one spot in northeastern Nova Scotia (Cape Breton Island) on 26 July 1977 where I found it on grassy waste- land adjacent to an abandoned coal mine at Sydney Mines. This is 3 km north of Sydney whence the ferry sails for Newfoundland, a voyage of 160 km across the Cabot Strait. Having arrived in Newfoundland, I found T. lineola in the western part of the island, on 28 July 1977. The locality was an open grassy area a few metres wide between woodland and Highway 430, 15 km north of Deer Lake. About a dozen of the butterflies (all males) were observed, most being fresh specimens. Three speci- mens were collected and have been deposited in the Can. Nat. Coll., Ottawa. Holland (1969, J. Lepid. Soc. 23: 33-42) collected in the Deer Lake area in 1965 at the same time of year and did not report seeing this species; indeed it does not appear to have been previously reported from Newfoundland. However, the insect has certainly reached the island now, presumably by traversing the Cabot Strait from Nova Scotia in the very recent past. It is perhaps possible that this species used the ferry for the crossing. W. J. D. Eseruir, P.O. Box 370, 17 Division St., Colborne, Ontario, Canada KOK IMO. Journal of the Lepidopterists’ Society 32(3), 1978, 226-228 A PROBABLE NATURAL HYBRID OF PAPILIO EURYMEDON AND P. RUTULUS (PAPILIONIDAE) FROM IDAHO Natural interspecific hybrids seem to be as rare among swallowtails as they are among butterflies in general. In the field the best evidence for hybridization comes usually from intermediacy of such characters as wing shape and color patterns. On 18 May 1976, David H. Wagner and I encountered impressive swarms of Papilio eurymedon Lucas and P. rutulus Lucas visiting muddy spots at the edge of the town of Lowell, Idaho Co., Idaho. All of the individuals were males. Flying at the same place but much less common were P. multicaudatus Kirby, P. zelicaon Lucas, Pieris napi Linné, Anthocharis sara Boisduval, Euphydryas chalcedona Doubleday and Hewitson, and Celastrina pseudargiolus (Boisduval and LeConte). Some of the male swallowtail “clumps” on the moist soil included over 50 butterflies. They were probably seeking sodium (cf. Arms et al. 1974, Science 185: 372-374). Obviously the situation here was ideal for observing variations, and we examined the crowded butter- flies carefully in the hope of finding aberrant forms. The differences between P. eurymedon and P. rutulus were immediately visible as they flew up and settled, often spreading their wings as they crawled over the moist earth. The gray-white ground color of the former contrasted with the bright clear yellow of the latter. Also the much broader black stripes and reduction of ground color of P. eurymedon quickly separated it from P. rutulus. In one group of swallowtails we noticed a perplexing individual that did not fit either P. eurymedon or P. rutulus. Its ground color was whitish lemon-yellow and the bo bo “I VoLUME 32, NUMBER 3 Fig. 1. Male swallowtails from Lowell, Idaho. Above, dorsal view; below, ventral view (magnifications vary slightly). A. Papilio eurymedon. B. Probable P. eurymedon x rutulus. C. P. rutulus. stripes were of intermediate width. The odd specimen was captured and is illustrated together with examples of the two associated species (Fig. 1). The specimen is inter- mediate in the position, extent, and shape of practically every stripe and spot. This evidence supports the conclusion that the odd specimen is a natural hybrid between P. eurymedon and P. rutulus. 228 JOURNAL OF THE LEPIDOPTERISTS SOCIETY We were especially impressed by the precise intermediacy of the individual. Generally speaking hybrid butterflies show such intermediacy, but in some cases they may resemble one parent more than the other. In the genus Limenitis, F: hybrids of L. archippus Cramer and L. astyanax (Fabricius) are intermediate, but backcrosses yield both hybrid-like and parent-like morphs (Platt 1975, Evolution 29: 120-141). In swallowtails at least, even F: hybrids may sometimes show one-sided intermediacy. For example, Clarke and Sheppard (1957, Lepid. News 11: 201-205) bred female P. glaucus L., the eastern North American counterpart of P. rutulus, with male P. eurymedon, and found that the glaucus wing pattern seemed generally dominant to that of P. eurymedon (cf. their Fig. 2 with Fig. 1 of the present paper). The F: progeny of laboratory crosses of P. polyxenes and P. xuthus were like the former parent in 11 out of 14 characters (Remington 1959, J. Lepid. Soc. 13: 151-164). Thus it is possible that all individuals of P. ewrymedon X rutulus found in nature in the future will not be so conspicuously and precisely intermediate as the one figured here. WARREN HERB WAGNER, JR., Department of Botany, University of Michigan, Ann Arbor, Michigan 48109. Journal of the Lepidopterists’ Society 32(3), 1978, 228-231 NOTES ON SOME MOSAIC PIERIS (PIERIDAE) Mosaic specimens occur in many if not all species of butterflies and moths, and are of scientific interest in that they can provide clues to the sequence of events occurring in embryonic or post-embryonic development. The origins of several types of mosaics are discussed by Ford (1945, Butterflies, Collins, London, Ch. 9). The mosaic nature may be sexual (various kinds of gynandromorphs) or homeotic (involving the produc- tion of a normal feature or pattern in an inappropriate location) or neither. The checkered white, Pieris protodice Bdv. & LeC., is one of the most abundant and widespread North American butterflies and shows a conspicuous sexual dimorphism. There are apparently no published reports of gynandromorphs or other sexual mosaics although a bilateral non-sexual aberration inherited in a Mendelian manner has been reported (Shapiro 1970, Wasmann J. Biol. 28: 245-257). I have never seen a mosaic in any institutional or private collection. Figure 1 shows the first such specimen I have turned up in thirteen years of research on this species, including mass laboratory culture through over 30 generations and repeated field sampling in several states. It was collected in a sample of 10 taken 16 October 1977 at Rancho Cordova, Sacramento Co., California. It is a very unusual gynandromorph for a number of reasons. The entire body and three wings are apparently male. The right forewing appears about 40% female, with the inner margin, discal cell and apex mostly female. There are three black spots near the margin in the interspaces where no black normally occurs in either sex. The female characters are confined to the upper surface. Ventrally the forewings are symmetrical and both male. Thus the assumed chromosomal accident must have occurred in a cell all of whose progeny were fated to positions on the surface giving rise to the dorsal lamina and its scales. I have on hand a similar mosaic gynandromorph of Colias eurytheme Bdvy. (figured by Shapiro 1973, J. Res. Lepid. 12: 94) in which the apex of the left forewing is female above, and the rest of the animal male. In this case the sexes do not differ ventrally in the forewing apical area, and it cannot be said with certainty whether 229 SV NS SO AS « VoLUME 32, NUMBER 3 2 (ae) ene | es aS meee a oO S 83 SS ee 24 See Sg ae ate o 6 | fae} IO) a . (Se) (se) 6S Org as ~S— ©) ec aie] E'S Ze a a 5 Zxae! Cho ox eS =— (avy & | | 3 = = [e) Zz Biges 1; gynandromorph (center 230 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Fig. 2. Mosaic Pieris rapae from California. A: bred, Davis, Yolo Co., April 1972; B: wild, Davis, March 1972; C: Southport, Yolo Co., March 1973; D: Sacramento Co., April 1973; E: Suisun Marsh, Solano Co., March 1974. the specimen is aberrant on both surfaces. The same applies to the Colias philodice Latr. figured by Emmel (1964, J. Res. Lepid. 3: 63) as a mosaic gynandromorph. In this case, however, it is unlikely that the specimen is gynandrous at all. The “female” pattern is chaotic and more likely represents patches of male ground color within the black border, a not uncommon occurrence. Figure 2 illustrates some mosaics of Pieris rapae L. from northern and central Cali- fornia. Two of these are clearly sexual. Specimen A is a bilateral gynandromorph bred ex ovo at Davis. Specimen B is a rather worn, field-collected specimen. These are the only California gynandromorphs I have seen in this species, and it is of more VOLUME 32, NUMBER 3 231 than passing interest that B was collected in the same field in the same week as the mother of A. Both specimens show some streaky mosaicism, with patches of male scales on the female side and conversely. Specimens C through E have abnormal, asymmetrical black markings which are confined to the upper surface. (Mosaics of this sort are very rare on the ventral surface. I have only one rapae, a female, with an abnormal black marking confined to the ventral surface and this is a “ray” similar to and perhaps homologous with the Mendelian character found in the protodice group (Shapiro 1973, Wasmann J. Biol. 31; 301-311).) Im specimen E the shape of the spot suggests homeosis, but it is located in an inappropriate interspace; moreover its position corresponds to an obvious crimp in the dorsal lamina, of the sort to be expected from a deformation of the pupal wing-case. Such injuries may occur when an unhardened pupa slips in its silken girdle. In Colias they routinely result in melanization of the area around the injury; if the adult is able to eclose, the resulting pattern is grossly abnormal (compare female figured by Shapiro 1970, Entomol. News 81:50/data document p. 5). Braun (1939, Biol. Bull. 76: 226-240) showed that as pigment precursor spread out- ward across the wing from the body, deposition took place in those scales which were sufficiently chitinized at that time. Control of pattern thus depends on the rate of scale maturation, which may be accelerated around injuries—contributing to mosaics such as these. Since injuries are most likely on the dorsal surfaces, especially of the forewings, it is not surprising that mosaicism is commonest there. The black streaking near the costa of specimen D could have arisen in several ways, but that near the hindwing apex of C suggests an injury resulting in pigment deposition in the corre- sponding scales as a wave of melanin precursor moved across the wing. All of the specimens figured are in the collection of the University of California at Davis. ArTHur M., SHaprro, Department of Zoology, University of California, Davis, Cali- fornia 95616. Journal of the Lepidopterists’ Society 32(3), 1978, 231-233 A MALE-LETHAL GENETIC FACTOR IN PHYCIODES THAROS (NYMPHALIDAE) During the course of four years of rearing studies using Phyciodes tharos Drury, a total of 17 broods was reared of stock from Upper Tyrone Township, Fayette Co., Pennsylvania. Of these, 15 were derived from wild-collected females and the remaining 2 from wild-laid egg patches found on leaves of the foodplant, Aster simplex. Exact egg-hatch data were kept on 16 of the broods, viability and sex ratio data on 13. Of the 17 broods, 3 showed almost total male inviability. In 2 of these 3, most mortality appeared to be during embryonic development, whereas in the third there was normal embryonic viability but about 50% mortality between the first and fourth larval instars. An additional brood (77-63) reared from a wild female collected in Rochester Mills, Indiana Co., Pennsylvania, showed greatly reduced embryonic viability and an almost total absence of male adults (Table 1). Female progeny from two of the abnormal Fayette Co. broods (74-4, 76-2) were mated to males from normal broods of the same population. Each of these isofemale lines (A and B) showed a tendency toward lowered egg fertility (P < .001, Wilcox on 932 JOURNAL OF THE LEPIDOPTERISTS SOCIETY TaBLeE 1. Egg fertility, embryonic viability, adult sex ratio, and prepupal and pupal mortality in broods of Phyciodes tharos from wild-collected eggs and females. “Male- lethal” broods indicated by asterisks. Broo No. of Proportion Proportion Total Proportion Proportion No. Eggs Fertile Hatched Adults Males Mortality 73-1 199 1.000 1.000 U5) 0.493 0.053 73-2 167 1.000 1.000 140 0.464 0.021 13-3* Si 0.978 0.724 207 0.010 0.019 73-4 638 1.000 1.000 73-29 229 1.000 1.000 56 0.428 0.036 73-30 21 0.523 0.381 74-1 162 1.000 1.000 74-2 472 0.998 1.000 74-3 243 0.992 0.984 74-4* 45 1.000 1.000 ~23 0.000 0.000 75-5 161 0.578 0.012 75-7 220 1.000 1.000 Paley 0.524 0.042 75-8 258 0.992 0.976 126 0.484 0.008 75-54 289 0.990 0.996 160 0.519 0.025 75-66 MB) 1.000 0.985 UT 0.416 0.130 15-67 IAS, 1.000 1.000 21 0.762 0.000 76-2* WZ, 1.000 0.444 118 0.000 0.025 77-63* 484 0.983 0.736 174 0.023 0.172 TABLE 2. Egg fertility, embryonic viability, adult sex ratio, and prepupal and pupal mortality in “male-lethal” isofemale lines of Phyciodes tharos. Broo No. of Proportion Proportion Total Proportion Proportion No. Eggs Fertile Hatched Adults Males Mortality Isofemale Line A 74-15 131 0.191 0.333 74-16 417 0.858 0.640 Isofemale Line B 76-10 106 0.066 0.714 76-11 746 1.000 0.614 126 0.000 0.103 76-13 611 0.988 0.778 9 0.076 0.025 76-14 84 0.060 0.800 76-15 226 0.279 0.714 24 0.083 0.000 76-16 1048 0.995 0.715 150 0.000 0.033 two-sample test) and sharply reduced embryonic viability (P < .001) compared with the normal broods, Four broods were reared through to adults in Line B. Each of these showed almost total male inviability (P = .001) (Table 2). In none of the “male-lethal” broods was there an unusually high incidence of mortality during the prepupal and pupal stages. The male lethal crisis appears to occur during embryonic or larval devel- opment. VOLUME 32, NUMBER 3 230 Robinson (1971, Lepidoptera Genetics, Pergamon, New York, 687 p.) has discussed male-deficient broods in several species of Lepidoptera. In Abraxas grossularia L. (Geometridae), a karytotypic aberration in females gives a tendency to produce nearly unisex but normally viable broods. In Hypolimnas misippus L. (Nymphalidae), females from some small island populations produce all female broods with reduced embryonic viability. Here a dominant sex-linked gene has been postulated. Owen (1966, Heredity 21: 443-451) has investigated East African populations of Acraea encedon L.. ( Acraeidae ), some of which contained only 0.6 to 6.2% males. Eggs produced by wild-collected females showed normal viability. Parthenogenesis was ruled out, and the genetic basis of the unisexual broods remains unknown. The present case in P. tharos appears to be similar to that in H. misippus. Pre- sumably, in both cases the disadvantage of heavy selection against male progeny is offset by some selective advantage to the females carrying the tendency toward uni- sexual broods. Cuar.es G. Oxiver, R. D. 1, Box 78, Scottdale, Pennsylvania 15683. Journal of the Lepidopterists’ Society 32(3), 1978, 233-234 OVIPOSITION BEHAVIOR OF COLONIZED HYALOPHORA GLOVERI GLOVERI (SATURNIIDAE) Efficient collection of eggs is an important aspect of maintaining small colonies of giant silkworm moths as breeding stock. This can be accomplished by establishing an oviposition profile for the species being reared and collecting eggs only during the period of peak oviposition. Experience in rearing many species of Nearctic giant silkworm moths has shown that most eggs are deposited during the first few nights after mating. Oviposition profiles reported for Hyalophora cecropia (Linnaeus ) (Taschenberg & Roelofs 1970, Ann. Entomol. Soc. Amer. 63; 107-111) and Callosamia promethea (Drury) (Miller & Cooper, 1977, J. Lepid. Soc. 31: 282-283) are specific examples of this pattern. This paper reports oviposition data for a small breeding-stock colony of Hyalophora gloveri gloveri (Strecker) maintained on wild black cherry (Prunus serotina) in Frederick County, Maryland. Because of the small size of the colony (<12 individuals) observations were limited to five individuals. Five female moths, each of which mated on the first night after emergence, were placed in brown paper bags (lunch size) on the first night after mating; and were transferred to new paver bags each night thereafter until death. After a period of time sufficient to allow all eggs to hatch, the bags were opened to record the number of eggs deposited and the number hatched. The average longevity of the females after mating was 6.6 days; two individuals lived for 6 days and three lived for 7 days. The females deposited a total of 776 eggs during the study. The maximum number of eggs deposited by a single female was 198; the minimum number was 114. The average number of eggs deposited per female was 155.2. Percent hatch was moderate for eggs deposited during the first 4 nights after mating, the average ranging from 60.7% to 78.4%. The total number of larvae produced per female ranged from 60 to 137; the average being 103.2. The 234 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 120 100 Z [_] ces peposiTeD O = 2 BE Larvae propucen = ai Lu — => Cé =e = © 60 ica) OS (Sp) Liu bh S 40 CXS Lu = = Fz am rr et A) 0 ] 2 3 4 5 6 7 NIGHTS AFTER MATING Fig. 1. Oviposition profile for colonized Hyalophora gloveri gloveri females. oviposition profile shown in Fig. 1 indicates that H. gloveri gloveri follows the general pattern reported for other species; and that the optimum time for collecting eggs to maintain small colonies for breeding stock is during the first night after mating. THoMas A, Mitier, U.S. Army Medical Bioengineering Research and Development Laboratory, Fort Detrick, Frederick, Maryland 21701. (This research was not supported by government funds; the opinions contained herein are those os the author and should not be construed as official or reflecting the views of the Department of the Army.) VoLUME 32, NUMBER 3 | 235 Journal of the pepitentents: Society S2(3); 1978; 23 THE MURRAY O. GLENN COLLECTION OF MICROLEPIDOPTERA The collection of microlepidoptera assembled from 1931-1976 by Murray O. Glenn has contributed significantly to the classification and ecology of the Lepidoptera of the midwestern prairies in the USA. It includes ca. 30,000 carefully prepared moths representing 1325 identified species, 949 of which Glenn collected in the prairies, wooded bluffs, and bottomlands near the Illinois River and its tributaries in Putnam and Marshall counties in north-central Illinois. Significantly the area is now the type- locality for 17 species and at least one additional species that currently is being described. The species that have been named from the Glenn collection with Putnam County as the designated type-locality include the following: OLETHREUTIDAE. Polychrosis sambuci Clarke, Endothenia microptera Clarke, Exartema comandranum Clarke, Eucosma uta Clarke, Epiblema naomi Clarke, and Epinotia atristriga Clarke. COSMOPTERIGIDAE. Teladoma incana Hodges. MOMPHIDAE. Batrachedra illusor Hodges and Chedra inquisitor Hodges. WALSHIIDAE. Periploca cata Hodges, Aeaea venifica Hodges, Sorhagenia baucidis Hodges, and Perimede maniola Hodges. GELECHIIDAE. Chionodes asema Clarke and Dichomeris glenni Clarke. OECOPHORIDAE. Agonopterix dimorphaella Clarke. PTEROPHORIDAE. Oidaematophorus glenni Cashatt. The holotypes for all the above named species are at the U.S. National Museum of Natural History (USNM) except for Oidaematophorus glenni which is in the collection of the Illinois Natural History Survey (INHS). Glenn, in 1977, donated his private collection to the INHS and the USNM, the former institution receiving the identified specimens (ca. 20,000) including numerous paratypes and the latter all unidentified moths (ca. 10,000). (His collection of macro- lepidoptera was given to the INHS in 1969. ) Glenn, as a collector, “. . . . seemed to have a genius for coming un with rare and desirable things.” (Klots, pers. comm.). In addition to collecting, he succeeded in ascertaining natural foodplant associations for many of the species that he encountered. Much of this information is presently available only by examining the collection itself but should prove extremely useful to future studies if coupled with the 46-year compilation of flight records associated with the specimens and environmental changes induced by natural or artificial factors. GrorcE L. GopFrey, Section of Faunistic Survey and Insect Identification, Illinois Natural History Survey, Urbana, Illinois 61801. _ 1 All specimens collected by Glenn in this defined region are labelled ‘Putnam Co., IL.” However, it should be noted that while Glenn did most of his collecting in Putnam County he occasionally forayed into the adjoining portions of Marshall County. 236 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Journal of the Lepidopterists’ Society 32(3), 1978, 236 A NEW RECORD FOR CALYCOPIS CECROPS (LYCAENIDAE) IN COLORADO BY AIRCRAFT-INTRODUCTION A new butterfly record for Colorado, Calycopis cecrops (Fabricius), was collected by Howard Bone, in the rear cargo pit of a United Airlines 727 at Stapleton Inter- national Airport, Denver, on 12 July 1977. The flight was a charter from the east coast arriving in Denver around 1030 MDST. The cargo pit doors of such aircraft are left in the open position the entire time the aircraft is on the ground, and this can easily be from 30 minutes to over an hour, which is ample time for any insect to fly into the pit. The mentioning by others (Eliot 1977, J. Lepid. Soc. 31: 75; Riotte 1977, J. Lepid. Soc. 31: 182) that Lepidoptera can be transported by both military and commercial airliners is confirmed by this record. Another possibility of an aircraft-introduced species is that of Tmolus azia (Hewitson). This species has been collected in the state twice. The first specimen was collected by Jim Eff on 16 July 1957, Chatauqua Mesa, Boulder Co., Colorado. The second specimen was collected by Marc Epstein on 26 July 1975, Magnolia Rd., Boulder Co., Colorado. These records are both equidistant from the airport. The facts that 1) there are no other records from the state, that 2) they were collected about the same time of year, and that 3) there were no other records during the 18 vear interval, suggests the possibility these two specimens were introduced by aircraft. This unusual occurrence of stowaways on aircraft could provide a very logical explanation for the introduction of butterflies at great distances from their normal ranges. A butterfly could easily fly into the pressurized, air-conditioned cargo pit of an aircraft, and be flown across the country or overseas in a matter of a few hours. The normal range of Calycopis cecrops is from eastern Kansas through southern Ohio to southern New Jersey and southward to Florida and Texas. The closest distance to Colorado within this range is approximately 500 air miles (805 km), but this specimen traveled some 1500 air miles (2414 km) from the east coast inside an airplane. The normal range of Tmolus azia in North America is southern Arizona and southern Texas, which is approximately 600 air miles (966 km) from Colorado. The specimens of Calycopis cecrops and Tmolus azia (collected by Marc Epstein ) are deposited in the collection of the Denver Museum of Natural History. MicHaEL G. PocuEe, Devartment of Zoological Collections, Denver Museum of Natural History, City Park, Denver, Colorado 80205. Journal of the Lepidopterists’ Society 32(3), 1978, 236-238 NEW FOODPLANT AND OVIPOSITION RECORDS FOR THE EASTERN BLACK SWALLOWTAIL, PAPILIO POLYXENES ON AN INTRODUCED AND A NATIVE UMBELLIFER Papilio polyxenes (Fabr.) is one of the most common Papilionidae in open fields of the eastern United States. Its larvae are considered to prefer plants of the Umbellif- erae, although in laboratory no-choice situations they will eat several species of Rutaceae, Some individuals can also survive when fed on the cucumber (or mountain magnolia) tree, Magnolia acuminata L. (Scriber and Feeny, in prep. ). VOLUME 32, NUMBER 3 ear In Greene County, Ohio, and Ithaca, New York, the preferred foodplant for Papilio polyxenes (Fabr.) appears to be the introduced wild carrot, Daucus carota (L.), as it likely is for most of the northeastern United States. In the eastern United States, a variety of species of Umbelliferae have been reported as foodplants (Scudder, 1889, The butterflies of eastern United States and Canada, 2: Forbes, 1960, Cornell University Agr. Expt. Sta. Memoir #371; Teitz, 1972, An index to the described life histories of Macrolepidoptera of the continental United States and Canada, Vol. 1; Tyler, 1975, the Swallowtail Butterflies of North America, Naturegraph). In addition to carrot, those plants upon which polyxenes larvae have been observed naturally in New York (J.M.S.) are wild parsnip, Pastinaca sativa L., poison hemlock Conium maculatum L., angelica Angelica atropurpurea L., and goutweed, Aegopodium podo- graria L. In 1976, polyxenes larvae in Ohio were found (M.D.F.) upon bulb-bearing water hemlock, Cicuta bulbifera L., angelica, Angelica atropurpurea, and wild parsnip, Pastinaca sativa in addition to wild and cultivated carrot. Here we have two separate observations of polyxenes on plants which should be reported due to the apparent lack of any previous natural observations and also due to their particular ecological significance. Our first observation is that of a female ovipositing on a characteristically woodland native plant species, Cryptotaenia canadensis (L.), Honewort (Fernald, 1950, Gray’s Manual of Botany, 8th ed.). The observation (J.M.S.) was made on 15 July 1977 at 10:00 hrs along a wooded creek at the end of Carlsbrook Drive in the township of Beavercreek, Ohio (Greene County). Conditions were favorable for oviposition, with the temperature approximately 30°C and the humidity also very high. Although these seem to be the ideal conditions for oviposition of most eastern swallowtails, solar radiation was probably a very important contributing factor as well, especially with the potential thermoregulation of body temperature in adults (R. C. Lederhouse, pers. comm. ). Although P. polyxenes larvae eat Cryptotaenia in laboratory no-choice conditions (Erickson, 1975, Psyche 81: 109-130; Scriber, 1975, Comparative nutritional ecology of herbivorous insects; Generalized and specialized feeding strategies in the Papilion- idae and Saturniidae Ph.D. Thesis Cornell University, Ithaca, N.Y.), the adults are rarely seen flying in forested areas where the foodplant occurs. Reasons for use of this particular wooded patch in Ohio are uncertain. This female may have drifted into the area more by chance than choice since the wooded habitat was a rather narrow strip in an otherwise open area of residential lawns and first year successional habitats. Inside the woods the polyxenes female hovered and circled several Crypto- taenia plants before depositing one egg in an immature flower head. The female did not investigate the other Umbelliferae (Sanicula, Heracleum, Osmorhiza) nearby, and instead flew off across the lawns out of sight. Although larvae ate and survived upon Heracleum maximum (Bartr.) plants from this same wooded location, post- flowering Heracleum plants in mid-July were less suitable for larval growth than were the mid-May plants (Finke, 1977, Factors controlling the seasonal foodplant utilization by the specialized herbivore, Papilio polyxenes. (Lepidoptera: Papilionidae) M.S. thesis, Wright State Univ., Dayton, Ohio). We do not know whether the pre- flowering Heracleum plants would have been more attractive for oviposition by polyxenes in May. P. polyxenes larvae from Greene County, Ohio refused to eat Sanicula gregaria Bickni., and others died after several days of eating Osmorhiza longistylis (Torr.) and O. claytoni (Michx.) (Finke, ibid.). Utilization of other woodland umbellifer species, Taenidia integerrima and Thaspium barbinode by Papilio joanae Heitzman may have contributed to reproductive isolation by habitat and the relatively new species status of this polyxenes relative (Heitzman, 1973, J. Res. Lepid. 12: 1-10). The significance of habitat as an isolating mechanism for adults and larvae of the two species is an interesting aspect of their ecology which needs further investigation. Our second observation was made in Ithaca, New York (Tompkins County) on 938 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 5 June 1977. Eight Papilio polyxenes larvae were observed feeding upon one plant of the introduced Lovage, Levisticum officinale (Doch). It is uncertain if these larvae (all of which were molting from the 3rd to 4th instar) were derived from one or several adults. It is extremely unlikely that they wandered onto the Levisticum from another plant, as there were no other umbellifers nearby. Larvae were collected and reared through to pupation on Levisticum. Adults have been preserved as voucher specimens (at the Univ. of Wisconsin, Madison). The 1973 and 1974 Ithaca populations of P. polyxenes produced larvae which consumed Levisticum officinale and grew at rates comparable to those for larvae on 35 other Umbelliferae in laboratory no-choice situations (Scriber, ibid.). Papilio polyxenes larvae obtained from an adult caught in Costa Rica by Paul Feeny all refused to eat or else ate and died in a similar no-choice situation (Scriber, pers. obs. ). Some polyxenes larvae from a Costa Rican female x Ithaca male mating were, however, able to eat Levisticum and grew to the 2nd and 3rd instar before the culture was lost to virus (J.M.S. and R. C. Lederhouse). The genetic basis for this feeding ability remains undetermined. Some recent studies with Costa Rican polyxenes larvae indicate a marginal and variable ability to utilize Levisticum does exist in this Central American population (Wm. Blau, pers. comm. ). The Levisticum officinale—Papilio polyxenes interface would appear to offer a good system for investigation of the evolutionary dichotomy underlying differences in ovipositional and larval feeding stimulants. The closely related P. machaon in Sweden, for instance, will oviposit freely on Levisticum, but 100% of the larvae die feeding on it (Wilkund, 1975, Oecologia 18: 185-197). Wilkund (ibid) suggests that the rareness and relatively recent introduction of the plant into Sweden may partially account for nonavoidance of the plant by ovipositing adults. The differences in larval feeding success between polyxenes of New York and Costa Rican populations may also be a function of the amount of time the plant and insect have been in contact. Levisticum does not, to our knowledge, occur in Costa Rica (Standley, P. C., 1938, Flora of Costa Rica, Field Mus. Natur. History, Chicago Botanical Series, Vol. 18 (no. 420); and Wm. Blau, pers. comm. ). In summary, we would like to emphasize the fact that foodplant utilization by P. polyxenes populations appears variable, depending upon local habitat factors and plant phenology in any particular year as well as regional or geographic host- plant preferences which may have evolved over a longer period of time. More field observations and laboratory studies could clarify many unknown or puzzling aspects of the coevolution of the Papilionidae and their hostplants. J. Mark ScriperR AND Mark FINKE, Department of Entomology, University of Wis- consin, Madison, Wisconsin 53706. VOLUME 32, NUMBER 3 ZOO Journal of the Lepidopterists’ Society 32(3), 1978, 239-240 BOOK REVIEW Tue British BUTTERFLIES, THEIR ORIGIN AND ESTABLISHMENT, by R. L. H. Dennis, 1977. E. W. Classey Ltd., Park Road, Faringdon, Oxon., England SN7 7DR. 318 pp., 20 figs., 15 tables. $17.50 US. Here is a book that should be of interest to all students of biogeography. It is well done but many general collectors of butterflies will find it hard going. First, it is not a book to help determine what species you have from the British Isles. It is a book that lives up to its second title. Dennis has divided his book into four sharply separate sections. North Americans may find the first part, “Geomorphological Frame- work” a bit puzzling at first. The table on p. 7 (the first page of the first chapter) sets forth the nomenclature used for the Pleistocene in northwestern Europe, including the British Isles. Add a third column to this, naming the equivalent North American terms, and you will be allright. The other sections in sequence are “Recent Rhopalocera geography and habitat adjustments,’ “Subspecies and subspeciation,” and lastly “The arrival sequence and establishment of the British Rhopalocera.” In addition, there are four appendices containing useful information, particularly for those of us on this side of the ocean who lack intimate knowledge of the British fauna and flora. The author has done an unusually good job assembling a wealth of data about the Pleistocene in the British Isles. He retells this in detail. It is best to have a good scale map of Great Britain and Ireland at hand unless you are intimately familiar with the geography of them. Maps would have helped in this section, but I suppose cost would have been prohibitive. In the second part of the book Dennis treats two subjects: zoogeography and adaptation to the environment. There is considerable redundancy, but it is not obtrusive. It may be helpful. Here is clearly demonstrated why the British Isles is the ideal place for such a study at this time. The region is essentially a closed system for butterflies with few migrants and substantial sea barriers. Collectors have been active for two centuries or more and their data are available. The region is small enough—a total area considerably less than the State of Montana. The amount of information about the area is greater than that for any com- parable area in either Canada or the United States. This gives the zoogeographer an ample working sample. The geography side is equally well-reported. The ideas of geological mapping and stratigraphy and the foundation of modern geology are British inventions of the 18th century. Detailed large scale mapping is available for the entire United Kingdom. All of these are needed before such a task as Dennis set for himself can be confidently attacked. The second half of the book, Sections C and D, contains an able discussion of subspeciation as evident in the British Isles. Dennis’s interpretation of subspeciation, as related to geography, flora and modern and past climates, can be duplicated no- where on the American continents. The last section is the interpretation of the data presented in the first three sections. Here Dennis had two earlier similar studies to use, and to agree or disagree with. He did all three. B. P. Beirne wrote several times on the subject and summed up his knowledge in The Origin and History of British Macrolepidoptera found in the Trans. Roy. Entomol. Soc. Lond., 98, 1947. E. B. Ford set forth his views in detail in Butterflies published by Collins, London. The latest edition of this delightful book was released in 1957. Dennis had several advantages over either of these able authors. He had a large number of precise radiocarbon dates and the results of the very recent and extensive paleontological studies of Pleistocene and Recent (Flandrian) insects. Needless to say, Tables 14 and 15, setting forth the ideas of the three writers, show progressive changes in opinion. It will be a very long 240 JOURNAL OF THE LEPIDOPTERISTS SOCIETY time before a comparable work can be written for any part of the New World. This is a beautiful example of what can be done when data are available. The book is an important guide for those who would engage upon detailed zoogeographic studies. F. Martin Brown, 6715 S. Marksheffel Rd., Colorado Springs, Colo. 80911. Journal of the Lepidopterists’ Society 32(3), 1978, 240 OBITUARY JAMES H. BAKER (1910-1978) Mr. James “Jim” Huffman Baker, charter member of the Lepidopterists’ Society, died April 14, 1978 at St. Luke’s Hospital in Boise, Idaho after a long illness. He was 67. Jim was born Aug. 14, 1910 in Baker, Oregon, the son of Deering F. and Bernice Huffman Baker. He graduated from Baker High School in 1928, was employed by the Citizen’s National Bank in Baker, and then ran the family grocery, Baker's Super- market, for over 35 years. Jim was a man of many activities. In addition to his lifelong interest in insects of many orders, he bowled, traveled, was interested in general nature study, collected rocks, and was an antique dealer and a gem worker. He published several scientific papers, and his extensive collecting disclosed several insects that were subsequently named, including Ewphydryas anicia bakeri Stallings and Turner, and Celastrina argiolus bakeri (Clench). He worked closely with both the American Museum of Natural History and the Smithsonian Institution. He was also a member of the Coleopterists’ Society. | Jim will be greatly missed by all of his many friends and colleagues who have enjoyed his company and his family’s hospitality. He is survived by his wife, Ilah; a son, James Michael Baker; a daughter, Judith Ann Haswell of Pullman, Washington; and two granddaughters. J. W. Titpen, 125 Cedar Lane, San Jose, California 95100. EDITORIAL STAFF OF THE JOURNAL AusTIN P. Puiattr, Editor Department of Biological Sciences University of Maryland Baltimore County, 5401 Wilkens Avenue Catonsville, Maryland 21228 U.S.A. Frances S. Coew, Managing Editor Dovctas C. Fercuson, Associate Editor THEODORE D. SARGENT, Associate Editor NOTICE TO CONTRIBUTORS Contributions to the Journal may deal with any aspect of the collection and study of Lepidoptera. Contributors should prepare manuscripts according to the following instructions. Abstract: A brief abstract should precede the text of all articles. Text: Manuscripts should be submitted in duplicate, and must be typewritten, entirely double-spaced, employing wide margins, on one side only of white, 8% xX 11 inch paper. Titles should be explicit and descriptive of the article’s content, including the family name of the subject, but must be kept as short as possible. The first men- tion of a plant or animal in the text should include the full scientific name, with authors of zoological names. Insect measurements should be given in metric units; times should be given in terms of the 24-hour clock (e.g. 0930, not 9:30 AM). Underline only where italics are intended. References to footnotes should be num- bered consecutively, and the footnotes typed on a separate sheet. Literature Cited: References in the text of articles should be given as, Sheppard (1959) or (Sheppard, 1959, 196la, 1961b) and all must be listed alphabetically under the heading LrreRATURE CrreED, in the following format: SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London. 209 p. 196la. Some contributions to population genetics resulting from the study of the Lepidoptera. Adv. Genet. 10: 165-216. In the case of general notes, references should be given in the text as, Sheppard (1961, Adv. Genet. 10: 165-216) or (Sheppard 1961, Sym. Roy. Entomol. Soc. London 1: 23-30). Illustrations: All photographs and drawings should be mounted on stiff, white backing, arranged in the desired format, allowing (with particular regard to lettering ) for reduction to their final width (usually 4% inches). Illustrations larger than 8% x 11 inches are not acceptable and should be reduced photographically to that size or smaller. The author’s name, figure numbers as cited in the text, and an indication of the article’s title should be printed on the back of each mounted plate. Figures, both line drawings and halftones (photographs), should be numbered consecutively in Arabic numerals. The term “plate” should not be employed. Figure legends must be typewritten, double-spaced, on a separate sheet (not attached to the illustrations), headed EXPLANATION OF FicuRES, with a separate paragraph devoted to each page of illustrations. Tables: Tables should be numbered consecutively in Arabic numerals. Headings for tables should not be capitalized. Tabular material should be kept to a minimum and must be typed on separate sheets, and placed following the main text, with the approximate desired position indicated in the text. Vertical rules should be avoided. Proofs: The edited manuscript and galley proofs will be mailed to the author for correction of printer’s errors. Excessive author’s changes at this time will be charged to authors at the rate of 75¢ per line. A purchase order for reprints will accompany the proofs. Correspondence: Address all matters relating to the Journal to the editor. Short manuscripts such as new state records, current events, and notices should be sent to the editor of the News: Jo Brewer, 257 Common Street, Dedham, Massachusetts 02026 U.S.A. ALLEN PRESS, INC. et LAWRENCE, KANSAS UusS.m CONTENTS Tue INFLUENCE OF ENVIRONMENTAL FACTORS ON ROOSTING IN THE BLACK SWALLOWTAIL, PAPILIO POLYXENES ASTERIUS STOLL (PaPILIONDAE). John Edward Rawlins & Robert C. Leder- house) 2 NoTeEs ON THE LiFE CYCLE AND NATURAL HIsTOoRY OF BUTTERFLIES or Ex SALvaporE. IIC. SMYRNA BLOMFILDIA AND S. KARWINSKII (NYMPHALIDAE: CoLosurini). Albert Muyshondt, Jr. & Alberto Muyshondt 0 ee SCELIODES LAISALIS (PYRALIDAE): DESCRIPTION OF THE MATURE LARVA AND NOTE ON ITs FEEDING Hasit. E. O. Ogunwolu ___ MIGRATION AND RE-MIGRATION OF BUTTERFLIES THROUGH NORTH PENINSULAR FLORIDA: QUANTIFICATION WITH MALAISE TRAPS. Thomas J. Walker Hysrios BETWEEN CALLOSAMIA AND SAMIA (SATURNUDAE). Richard S. Peigler EE RHOPALOCERA OF WEST VircINIA. Bastiaan M. Drees & Linda Butler 2 a PHENOLOGY AND DIVERSITY OF A BUTTERFLY POPULATION IN SOUTHERN Arizona. George T. Austin a) A New Hinpwinc ABERRATION OF CATOCALA MICRONYMPHA GUENEE FROM Kentucky. Charles V. Covell, Jr... GENERAL NOTES Confirmation of the occurrence of an albinistic female form of Phoebis philea (Pieridae) in extreme southern Texas. Raymond W. Neck _.. A new weedy host for the Buckeye, Precis coenia (Nymphalidae). Arthur M. Shapiro 2 A second locality for Eulythis mellinata (Geometridae) in North America. Kenneth Neil 2 Occurrence of Thymelicus lineola (Hesperiidae) in Newfoundland. W. J. D. Eberlie A probable natural hybrid of Papilio eurymedon and P. rutulus (Papilion-— idae) from Idaho. Warren Herb Wagner, J. -.-2------2----22----2-eeneeeneeee Notes on some mosaic Pieris (Pieridae). Arthur M. Shapiro ....-...-----.- A epee genetic factor in Phyciodes tharos (Nymphalidae). Charles G. Oltwer on I Oviposition behavior of colonized Hyalophora gloveri gloveri ( Saturniidae). Thomas A. Miller 2.0500 OE) The Murray O. Glenn Collection of Microlepidoptera. George L. Godfrey A new record for Calycopis cecrops (Lycaenidae) in Colorado by aircraft- introduction. Michael G. Pogue —....)\ New foodplant and oviposition records for the eastem Black Swallowtail, Papilio polyxenes on an introduced and a native Umbellifer. J. Mark Scriber & Mark Fink Nores AND News BOOK “REVTew nn DR ALA Osrrganen ok 145 160 175 178 19] 198 207 221 Volume 32 1978 Number 4 JOURNAL of the LEPIDOPTERISTS’ SOCIETY Published quarterly by THE LEPIDOPTERISTS’ SOCIETY Publié par LA SOCIETE DES LEPIDOPTERISTES Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 28 February 1979 THE LEPIDOPTERISTS’ SOCIETY EXECUTIVE COUNCIL : I. F. B. Common, President T. Surr6zu, Vice President C. V. CovE.LL, Jr., Ist Vice President JuLt1an P. DonanuE, Secretary L. A. GozmMAny, Vice President RONALD LEUSCHNER, Treasurer Members at large: R. A. ARNOLD J. F. EMMEL C. D. FErris E. D. CAsHATT R. R. GATRELLE J. Y. MILLER R. E. STANFORD Al Be Prarr M. C. NIELSEN The object of the Lepidopterists’ Society, which was formed in May, 1947 and formally constituted in December, 1950, is “to promote the science of lepidopterology in all its branches, . . . . to issue a periodical and other publications on Lepidoptera, to facilitate the exchange of specimens and ideas by both the professional worker and the amateur in the field; to secure cooperation in all measures” directed towards these aims. Membership in the Society is open to all persons interested in the study of Lepidoptera. All members receive the Journai and the News of the Lepidopterists’ | Society. Institutions may subscribe to the Journal but may not become members. Prospective members should send to the Treasurer full dues for the current year, together with their full name, address, and special lepidopterological interests. In 4 alternate years a list of members of the Society is issued, with addresses and special interests. There are four numbers in each volume of the Journal, scheduled for February, May, August and November, and six numbers of the News each year. Active members—annual dues $13.00 Student members-—annual dues $10.00 Sustaining members—annual dues $20.00 Life members—single sum $250.00 Institutional subscriptions—annual $18.00 Send remittances, payable to The Lepidopterists’ Society, and address changes to: Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 U.S.A. Back issues of the Journal of the Lepidopterists’ Society, the Commemorative Volume, and recent issues of the NEWS are available from the Assistant Treasurer. The Journal is $13 per volume, the Com:nemorative Volume, $6; and the NEWS, $.25 per issue. Order: Mail to Charles V. Covell, Jr., Memoirs Editor, Department of Biology, Uni- oa versity of Louisville, Louisville, KY 40208, U.S.A. 2 The Lepidopterists’ Society is a non-profit, scientific organization. The known office of publication is 1041 New Hampshire St., Lawrence, Kansas 66044. Second class postage paid at Lawrence, Kansas, U.S.A. 66044. Cover illustration: Dasychira dorsipennata larva, dorsal and lateral views. From Fascicle 22.2, “Lymantriidae,” by Douglas C. Ferguson, in Moths of America North of Mexico, The drawing was done by E. R. Hodges, Scientific Illustrator, Department of Entomology, Smithsonian Institution. (Reproduced by permission of the author. ) \ JOURNAL OF Tae LeEpiIpopreERIStTs’ SOCIETY Volume 32 1978 Number 4 Journal of the Lepidopterists’ Society 32(4), 1978, 241-250 THE ZALE SETIPES SPECIES COMPLEX (LEPIDOPTERA: NOCTUIDAE) EE Topp Systematic Entomology Laboratory, IIBIII, Federal Research, Sci. & Educ. Admin., U.S. Dept. of Agriculture’ ABSTRACT. The setipes complex of the noctuid genus Zale Hiibner is revised. The identity of two species, confused for more than 100 years, is clarified. Letis incipiens Walker is removed from the synonymy of Zale setipes (Guenée) and ele- vated to a subspecies of Z. peruncta (Guenée). Z. discisigna discisignata Draudt is cited as new synonym of Z. setipes (Guenée) and Z. setipes 2 f£. postmedialis Draudt, Homoptera aemona Druce (in part), Zale notipennis Draudt are new synonyms of Z. peruncta (Guenée). The first United States record of typical Z. peruncta {Guenée ) is listed. The noctuid genus Zale Hubner, as currently recognized, is composed of a large number of moderately large moths, many with a rather similar pattern of cryptic wing maculation. The pattern of maculation is usually composed of numerous irregular transverse or oblique lines, the moths presumably resembling the bark of trees on which the moths may rest. They vary in color from nearly black to pale yellow brown or light gray. A few species, especially some from tropical America, have areas of pale green scaling on the wings, but that color usually fades very rapidly after death to yellow or yellow brown. Identification of species has been difficult in the past and many misidentifications have occurred. Two closely related species, Zale setipes (Guenée) and Z. peruncta (Guenée) have been confused, misidentified and misnamed since 1869. The purpose of this paper is to indicate the proper appli- cation of the names, to describe and illustrate the characters that dis- tinguish the species, to detail the specific geographic distributions and to record Z. peruncta (Guenée) from the United States (Texas). 1¢/o U.S. National Museum, Washington, D.C. 20560. 242 JOURNAL OF THE LEPIDOPTERISTS SOCIETY The two species are very similar in maculation and both species are sexually dimorphic, both males and females of each species more closely resembling the same sex of the other species than the opposite sex of their own species. The dimorphism is expressed in differences in the shape of the forewing and in the pattern of maculation. The males have a narrower forewing, somewhat produced apically, the termen nearly straight or even excavate before tornus, therefore differing from the females and the other species of Zale which have broad forewings and a rounded termen. The pattern of maculation of the males ap- proaches that of some species of Metria Hiibner (= Safia Guenée). The females resemble females of other Zale species in wing shape and maculation. Examples of setipes are consistently larger and have more pale scaling in the postmedial area of the forewing than do examples of peruncta. Excellent characters for specific separation exist in the male genitalia and in the shape of the sternal plate of the eighth ab- dominal segment of the female. In 1965 the author studied the types and syntypes of Metria and Zale in the collection of the British Museum (Natural History) in order to correctly identify species from the Antilles and to obtain information necessary for possible future generic revisions. Nearly 80 slides of geni- talia, mainly of types, were prepared and the errors in the application of the names, Zale setipes (Guenée) and Zale discisigna (Walker), were discovered. The types of all the names relating to the setipes complex, including the lectotype of Z. peruncta (Guenée) which was sent to me at that time from Paris, were studied. | History Guenée described Xylis setipes (1852, p. 7, Noctuélites Pl. 15, Fig. 6) from a single male from Nova Friburgo, Brazil and Homoptera peruncta (1852, p. 9) from 2 specimens without locality. Guenée suggested that one specimen of peruncta was a male lacking antennae, but this seems unlikely since he placed the male of setipes in a separate genus, Xylis Guenée, while placing peruncta in Homoptera Boisduval with other typical Zale species. The lectotype of peruncta is a female specimen from the Paris Museum selected by Viette (1951, p. 161). The colored illustration of the type of setipes accompanying the original description is excellent. For a number of years the relationship of setipes and peruncta and the sexual dimorphism in the complex were not recognized. During that period Walker described males of peruncta as Homoptera ustipennis ({1858| 1857, p. 1071) and Letis incipiens (1858, p. 1266). In 1869 (p. VoLuME 32, NUMBER 4 243 157) Herrich-Schaffer identified specimens of peruncta from Cuba (true setipes is not known from the Antilles) as Xylis setipes Guenée and the trivial name has since been misapplied by all authors to date. In the collections of the U.S. National Museum and the British Museum (Natural History) the name was likewise misapplied. Moschler (1890, p. 202) listed females in his treatment of “setipes,” but no discussion of sexual dimorphism was included. He did, however, wonder why the females he studied were only 4245 mm in expanse whereas the size given for setipes in the original description was 55 mm. Butler (1879, p. 41) recognized that Walker’s ustipennis was related to setipes in the statement: “H. ustipennis, a Xylis.” There is no indication as to his specific concept of setipes. The sexual dimorphism of the complex had still not been recognized by Druce (1889, p. 341). He utilized setipes in the same sense as Herrich-Schiaffer and listed ustipennis as a separate species from Panama; both names were placed in Xylis. He did not refer to Homoptera peruncta Guenée. Females of both species of the complex obviously were present in his series of the new species, Homoptera aemona, because he stated: “The specimens from Guatemala are rather larger and are paler in colour than those from the Volcan de Chiriqui. Our figure is taken from one of these latter.” In an unex- plained action Hampson (1898, p. 250) placed setipes in Polydesma Boisduval and used ustipennis as a form of that combination for ex- amples of peruncta from St. Lucia and Grenada. Hampson (1913, pp. 208-210, text figs. 54 and 55) treated both species and provided keys to and illustrations of the males. He placed the generic names Homop- tera Guenée and Xylis Guenée in the synonymy of Zale Hiibner. The two species of the setipes complex (as subgenus Xylis) were separated from the other species of Zale in the key because the hind tibiae of male are fringed with long hair and the hindwing with termen some- what excurved at middle, the costa lobed [expanded] at the base. The type of setipes now in the British Museum (Natural History) was not available to Hampson as it was not received by that institution until 1928. Unfortunately, Hampson apparently did not check the original description and illustration of setipes and continued to use the name incorrectly for peruncta which he placed along with all other names of the complex in the synonymy of setipes as identified by him. Hampson made another error in treating true setipes by calling it Zale discisigna (Walker). He thought the worn, damaged female holotype of Ho- moptera discisigna Walker ( [1858] 1857, p. 1066) represented the female sex of the large species of the complex. This error was perpetuated in collections and in the literature. Homoptera discisigna Walker does 244 JoURNAL OF THE LEPIDOPTERISTS SOCIETY resemble setipes somewhat and cons:dering the condition of the type, the error is partially understandable, but discisigna is considerably smaller and is not even congeneric. The type of discisigna had been studied before by other workers and the species placed in Peteroma Schaus or Barcita Moschler in collections. Dognin apparently was con- fused by the use of Homoptera discisigna Walker for different species in collections and must have written to William Schaus about the problem. He attached a note from Schaus’ reply of June 22, 1922 on a male of peruncta from Tucuman, Argentina now in the U.S. National Museum. Schaus informed him that he thought Walker had described two species named discisigna and that he believed that the one refer- rable to Peteroma was described in “Characters of Lep. Het.” Schaus obviously was wrong; no other description by Walker with the trivial name discisigna has been located. It seems likely that Schaus’s belief in a second description probably developed because discisigna was originally described as a Homoptera and because Hampson applied the name to a Zale species. SYSTEMATICS Zale setipes (Guenee ) (Figures 1-2, 9 and 12) Xylis setipes Guenée, 1852, p. 7; 1858, Pl. 15 ( Noctuélites), Fig. 6—Walker, 1857, p. 1052.—Druce, 1889, p. 341 (in part ).—M6schler, 1890, p. 202 (in part). Zale setipes (Guenée), Draudt, 1940, Pl. 70, row b (setipes @ ). Xylis ustipennis, Druce not Walker, 1889, p. 342 (in part).—Hampson, 1913, p. 208 (synonym of discisigna, Hmpsn. ). Homoptera aemona Druce, 1889, p. 344 (in part )—Hampson, 1913, p. 208 (synonym of discisigna, Hmpsn. ). Homoptera discisigna, Druce not Walker, 1890, p. 345 (in part ).—Hampson, 1913, p. 208. Zale discisigna, Hampson not Walker, 1913, p. 208, Fig. 54——Haimbach, 1928, p. 216.—Draudt, 1940, p. 454 (in part). Zale discisigna ab. discisignata Strand, 1917, p. 43 (= discisigna ab. 1 of Hampson. An infrasubspecific name, excluded. ). Zale discisigna discisignata Draudt, 1940, p. 455 (= discisigna ab. 1 of Hampson and ab. discisignata Strand.) [New synonymy. ] Diagnosis. Length of forewing, male, 24 to 27 mm, average 24.8 mm; female, 23 to 27 mm, average 24.2 mm. Pattern of maculation as illustrated (Figs. 1 and 2). Ground color of male paler than female and males of Z. peruncta (Guenée); trans- verse lines in medial area of forewing distinctly marked; hindwing of male with dark subterminal shade between veins M, and Cw, reaching termen only at vein M3. Maculation of hindwing of female variable, with (Fig. 2) or without blue-white postmedial spots, ground color sometimes paler than females of peruncta, but usually about the same darkness. Male genitalia as illustrated (Fig. 9), apical process of ventral margin of valve rather sigmoid in shape, longer than the thin, rather rec- tangular apical process of costa of valve. Base of uncus with triangular (apex slightly VoLuME 32, NuMBER 4 245 6 Adults of Zale setipes complex. Fig. 1, setipes, ¢, Chiriqui, Panama; 2, setipes, 2, “Cent. Amer.”; 3, peruncta peruncta, ¢, Juan Vinas, Costa Rica; 4, p. peruncta, ©, Orizaba, Mexico: 5, p. incipiens, 6, Cuba; 6, p. incipiens, 2, Convento, Dominican Republic. curved distad) lateral flanges: flanges present also on tegumen, bilobed, the depres- sion between lobes variable in depth and caudal lobe sharp pointed (Fig. 9) or shorter and rounded (Noctuidae genitalia slide No. 5033 of holotype). Female genitalia (Fig. 12) with a pair of large rectangular sternal plates present below ostium. Types. The Houoryre, ¢, of Xylis setipes Guenée from Nova Friburgo, Brazil, Noctuidae genitalia slide No. 5033 and the Hotoryre of Zale discisigna discisignata Draudt, a 2 from Volcan de Atitlan, [Guatemala], Noctuidae genitalia slide No. 5087, are in the British Museum (Natural History), London, England. Distribution. The species is known to occur from Mexico to Brazil, but is not known from the West Indies. Specimens from the following localities have been examined. MEXICO: Jalapa; Orizaba. GUATEMALA: Volcan de Atitlan; Chejel: Cayuga: “Guatemala.” COSTA RICA: Tuis; Juan Vinas. PANAMA: Chiriqui. COLOMBIA: Pacho, Ost-Cordill. ECUADOR: Jatunyacu, Oriente; Abitagua, Ori- 246 JouRNAL OF THE LEPIDOPTERISTS SOCIETY ente. BRAZIL: Castro, Parana; Rio Janeiro; Petropolis; “Casa Br.’; Ponte Nova, Rio Xingu, Amazonas; Nova Friburgo. Discussion. The larger size, paler ground color and distinctive male and female genitalia distinguish this species from Z. peruncta (Guenée). Zale peruncta peruncta (Guenée) (Figures 3-4, 8 and 11) Homoptera peruncta Guenée, 1852, p. 9.—Walker, 1857, p. 1069.—Viette, 1951, p. 161 (Lectotype designation. ). Xylis setipes, Gundlach not Guenée, 1881, p. 358 (in part).—Druce not Guenée, 1889, p. 341 (in part )—Mo6schler not Guenée, 1890, p. 202 (in part). Polydesma setipes, Hampson not Guenée, 1898, p. 250. Zale setipes, Hampson not Guenée, 1913, p. 209, fig. 55 (peruncta (Guen.), ustipennis (Wlk.), incipiens (Wlk.) and aemona (Druce) as synonyms ).—Wolcott not Guenée, 1923, p. 169; 1936, p. 432; 1951, p. 603.—Haimbach not Guenée, 1928, p. 216.—Draudt not Guenée, 1940, p. 455 (in part ).—Schaus not Guenée, 1940, p. 229 (in part ).—Biezanko, Ruffinelli and Carbonell not Guenée, 1957, p. 50. Zale setipes ab. postmedialis Strand, 1917, p. 43 (= setipes, ab. 1 of Hampson. An infrasubspecific name, excluded. ). Zale setipes 2 f. postmedialis Draudt, 1940, p. 455, pl. 70, row b (= setipes, ab. 1 of Hampson and ab. postmedialis Strand). [New synonymy. ] Homoptera ustipennis Walker, 1857, p. 1071. [New synonymy. ] Xylis ustipennis (Walker), Butler, 1879, p. 41.—Druce, 1889, p. 342 (in part). Polydesma setipes £. ustipennis (Walker), Hampson, 1898, p. 250. Homoptera aemona Druce, 1889, p. 344, pl. 31, fig. 3 (in part). [New synonymy. ] Zale discisigna, Draudt not Walker, 1940, pl. 70, row 6 ( é, discisigna). Zale notipennis (sic) Draudt, 1940, p. 455 (misspelling of ustipennis Wlk. ? As synonym of setipes, Draudt). [New synonymy. ] Diagnosis. Length of forewing, male, 19 to 23 mm, average 20.4 mm; female, 19 to 23 mm, average 21.7 mm. Pattern of maculation of male (Fig. 3) similar to that of setipes, but ground color, particularly median part of forewing darker; dark subterminal shade of hindwing between veins M and Cu reaching termen for most shades width. Female marked and colored as in setipes, sometimes slightly darker, hindwing maculation variable as in setipes. Male genitalia with process from costa of valve longer than process from ventral margin, the latter slightly clavate or mitten- shaped, both processes (Fig. 8) quite different than in setipes. Flanges at base of uncus in typical subspecies nearly rectangular. Flanges of tegumen thornlike, apices slightly recurved. Sternal plate of female genitalia ovoid, caudal margin variable, usually terminating in two short bluntly pointed processes with a prominent narrow medial emargination (Fig. 11), occasionally emargination reduced in length, an ex- treme example with median caudal lobe that is very weakly emarginate (lectotype of Homoptera aemona Druce). Types. The Lecroryre, 2, of Homoptera peruncta Guenée, locality unknown, is in the Muséum National, Paris, France. The Hotoryre of Homoptera ustipennis Walker, 4, locality unknown, Noctuidae genitalia slide No. 5091, and the syNTyPEs of Homoptera aemona Druce and Zale setipes £. postmedialis Draudt are in the British Museum (Natural History), London, England. Druce had examples of both this species and true setipes in his original series of aemona from Mexico, Guatemala, and Panama but did not indicate the number of examples either in total or from the respective countries. He illustrated a specimen, a 9, from Volcan de Chiriqui, Pan- ama. There are three specimens in the British Museum (Natural History) from that locality. One labeled Homoptera aemona Druce, Type @, Noctuidae genitalia slide No. 5090 has been selected and is presently designated as LECTOTYPE. The name VoLuME 32, NuMBER 4 QAT woe Male and female genitalia of Zale setipes complex. Fig. 7, peruncta incipiens, 4; 8, p. peruncta, 6 aedeagus not shown; 9, setipes, ¢ aedeagus not shown; 10, p. incipiens, 2; 11, p. peruncta, 2; 12, setipes, 2. postmedialis was proposed for “ab. 1” of Hampson who did not indicate number of specimens or locality. The specimen labeled as type, a female, Noctuidae genitalia slide No. 5088 from Grenada has been selected and is now designated LECTOTYPE. Distribution. The typical subspecies occurs from southern Texas to Argentina on the continent and in the Antilles from Grenada to Puerto Rico. The specimen from Texas was collected on 27 November 1973 by A. and M. E. Blanchard. It represents a new record for the United States. I have examined specimens from the following localities. TEXAS: Santa Ana Refuge, Hidalgo Co. MEXICO: Jalapa; Misantla; Orizaba; Cordoba; San Cristobal las Casas, Chiapas. COSTA RICA: Tuis; Juan Vinas. PANAMA: Chiriquii COLOMBIA: Sta. Marta. VENEZUELA: Aroa. ECUADOR: Abitagua, Oriente. BRAZIL: St. Catherines [Santa Catarina]; Alta da Serra, Sao Paulo; Rio Janeiro; Theresopolis.s PARAGUAY: Sapucay; “Paraguay.” ARGENTINA: Tucuman. GRENADA: Grand Etang; “Grenada.” ST. VINCENT: Montreal District. ST. LUCIA: 1.5 mi S. Mt. Gimie; “St. Lucia.” DOMINICA: Clarke Hall; Grand Savanne; Pont Casse. VIRGIN ISLANDS: Gallows Point, St. John. PUERTO RICO: 4 mi SE. Ciales; Ciales. 248 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Discussion. This species is smaller than setipes and the ground color is slightly darker, especially the median area of the forewing and the subterminal spot between veins Ms and Cus. The two apical processes of the valve of the male gentitalia and the sternal plate of the female genitalia are differently shaped than those structures in setipes. This subspecies may be separated from the other subspecies by area of occurrence and by characters of the male and female genitalia discussed in the diagnosis of the atypical subspecies. The species of the setipes complex of Zale do not appear to be common in most areas judging from the number of examples in collections and from my personal collecting experience. Only at Grand Etang, Grenada, to my knowledge, has a species of the complex been collected in large numbers (personal light trap collecting). Time of year, weather conditions, time of night, collecting locality, and collecting technique may in part explain the reduced captures elsewhere, but I believe some other factor is responsible. A few days later on St. Vincent and then on St. Lucia collecting with traps in apparently similar ecological locations resulted in only a few captured specimens. In a three year survey of Dominica, 1964-1966, eight different collectors collected only four examples of peruncta. It is true that traps were not utilized there and the species are also known to be only temporarily attracted to light, settling soon on the plants some distance away. However, I personally collected other species of Zale there in large numbers by collecting on such plants, the specimens logated by their glowing, light reflecting eyes. Zale peruncta incipiens (Walker), new status (Figures 5-6, 7 and 10) Letis incipiens Walker, 1858, p. 1266.—Hampson, 1913, p. 209 (synonym of setipes, Hampson ).—Schaus, 1940, p. 229 (synonym of setipes, Schaus ). Zale incipiens (Walker), Draudt, 1940, p. 455 (synonym of setipes, Draudt). Xylis setipes, Herrich-Schaeffer not Guenée, 1869, p. 157.—Gundlach not Guenée, 1881, p. 358 (in part); 1891, p. 195.—Druce not Guenée, 1889, p. 341 (in part )—Moschler not Guenée, 1890, p. 351.—Ragués not Guenée, 1914, p. 141. Xylis sctipes (sic), Anonymous not Guenée, 1895, p. 73 (misspelling of setipes). Zale setipes, Hampson not Guenée, 1913, p. 209 (in part)—Schaus not Guenée, 1940, p. 229 (in part). Diagnosis. Length of forewing, male, 18.5 to 20.5 mm, average 19.5 mm; female, 20.0 to 22.0 mm, average 21.2 mm. It seems likely that the range in size will probably approach that of the typical subspecies when more material is available for study. Only five pairs have been examined. The pattern of maculation appears to be essentially identical to that of peruncta peruncta and similarly variable. The male and female genitalia differ consistently from those of the typical subspecies. The apical processes of the valve of the male genitalia (Fig. 7) are more slender than in typical peruncta, the process of the costa distinctly sinuous. Flanges at base of uncus thornlike, each with apex bent caudad. Flanges of tegumen much larger, not thornlike in shape as in typical peruncta, apex variable in shape, up-curved and blunt (Fig. 7) or sharp-pointed and caudally directed (holotype). Female genitalia with sternal plate of eighth abdominal segment smaller than in typical subspecies, the caudal lobes larger in proportion to plate size (Fig. 10). Type. The Hororypr, 2, from St. Domingo, Noctuidae genitalia slide No. 5089 is in the British Museum (Natural History), London, England. Distribution. Known only from Cuba and Dominican Republic. The specimens studied are labeled as follows. CUBA: Santiago; Cayamas; “Cuba.” DOMINICAN REPUBLIC: St. Domingo; San Francisco Mts., St. Domingo; Hotel Montana, 10 km NE Jarabacoa, La Vega Proy.; 1.3 km S Loma de Cabrera, Dajabon Prov.; Convento, 12 km S Constanza. VoLuME 32, NuMBER 4 249 Discussion. The true status of this entity is not known. It has been placed as a subspecies of peruncta because of the geographic isolation and to express the close relationship of the two entities compared to setipes. At the present time incipiens and typical peruncta occur on the neighboring islands of Hispaniola and Puerto Rico respectively. The former population probably representing an old invasion from Central America, the latter a more recent invasion from northern South America through the Lesser Antilles. LITERATURE CITED AnonyMous. 1895. Catalogo numérico del museo zool6gico Cubano (Museo Gund- lach). Alvarez, Habana. 112 pp. BrEZANKO, C. M. DE, A. RUFFINELLI, & C. S. CARBONELL. 1957. Lepidoptera del Unieuay. Revta. fac. Agron. Mioncaidee! No. 46; 1-152. Butter, A. G. 1879. On the Lepidoptera fof the Amazon, collected bye Dr. James W. H. Trail, during the years 1873 to 1875. Trans. Sauomall Soc. London, 1879, pp. 19-76. Draupt, M. 1939-1940. Noctuidae: Catocalinae. In Seitz, Die Gross-schmetter- linge der Erde, 7: 417-461, pls. 57-71. Stuttgart: Kernen. [pp. 417-428, 1939; pp. 429-461, 1940.] Druce, H. 1881-1900. Lepidoptera: Heterocera, Vol. 1. In Godman & Salvin, Biologia Centrali-Americana, Zoologia, Insecta. i—xxxii, 1-490, pls. 1-41. Lon- don: Taylor & Francis. [pp. i-xxxii, 1900; pp. 1-24, 1881; pp. 25-32, 1883; pp. 33-112, 1884: pp. 113-160, 1885; pp. 161-200, 1886; pp. 201-256, 1887; pp. 257-344, 1889; pp. 345-440, 1890; pp. 441-490, 1891.] GuENEE, A. 1852. Histoire Naturelle des Insectes. Species Général des Lépidop- téres. V. 7 (Noctuélites, V. 3): 442 pp. Librairie Encyclopédique de Roret, Paris. 1858. Ibid., 58 Pls. [Noctuélites, 1-24; Deltoides et Pyralites, 1-10; Uranides, 1; Phalénites, 1-22 and Siculides, 1.] GunpuacH, J. 1881. Contribucion a la Entomologia Cubana. Pt. 1 Lepiddpteros. Montiel, Habana. 445 pp. . 1891. Apuntes para la fauna Puerto-Riquenma. Pt. 7 Lepiddpteros. An. Soc. Espan. Hist. Nat., 20: 109-207, 323-384. Harmpacu, F. 1928. A list of the species and descriptions of new forms of the American genus Zale and a new form of Safia (Lepidoptera: Noctuidae, Cato- calinae). Trans. Amer. Entomol. Soc., 54: 215-231. Hampson, G. F. 1898. The Moths of the Lesser Antilles. Trans. Entomol. Soc. London, 1898, Pt. III, pp. 241-260, Pl. XVII. . 1913. Catalogue of the Lepidoptera Phalaenae in the British Museum. V. 13: i-xiv, 1-609, pls. 222-239. Taylor & Francis, London. HerricH-ScHaAEFFER, G. A. W. 1869. Die Schmetterlinge der Insel Cuba. Noctuina (part). Corresp.-Blatt zool.-min. Ver. Regensburg, 23(10): 153-160. Moscuter, H. B. 1890. Die Lepidopteren-fauna von Portorico. Abhandl. Senckenb. Nat. Ges., 16: 69-360, 1 Pl. Racugs, P. V. 1914. Museo Cubano “Gundlach” Catalogo General. Zoologia. 156 pp. Scuaus, W. 1940. Insects of Porto Rico and the Virgin Islands—Moths of the family Noctuidae. Scientific Survey of Porto Rico and the Virgin Islands, V. 12, pt. 2, pp. 177-290. New York Academy of Science, New York. STRAND, E. 1917. Neue Aberrationen der Noctuiden-Subfamilien Hadeninae, Eras- triinae, Catocalinae, Mominae, und Phytometrinae. Arch. Naturgesch., Abt. A 82, Heft 2 (1916), pp. 28-50. VieTtTE, P. 1951. Sur quelques Noctuelles decrites par Guenée (1852-1854). Bull. Mens. Soc. Linn. Lyon, 20(7): 159-162. 250 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Waker, F. [1858] 1857. List of the specimens of lepidopterous insects in the collection of the British Museum. Pt. 13, Noctuidae, pp. 983-1236. 1858. Ibid., Pt. 14, Noctuidae, pp. 1237-1519. Woxcort, G. N. 1923. Insectae Portoricensis (A revised annotated check-list of the insects of Porto Rico with descriptions of some new species.). Journ. Dept. Agric. Porto Rico, 7(1): 1-312. . 1936. Insectae Borinquenses (A revised annotated check-list of the insects of Puerto Rico.). Journ. Agri. Univ. Puerto Rico, 20(1): 1-600, 190 text figs. . 1951. The insects of Puerto Rico. Lepidoptera. Journ. Agric. Univ. Puerto Rico, 32(3) (1948): 537-748, 62 text figs. Journal of the Lepidopterists’ Society 32(4), 1978, 251-255 A LOST AND MISPLACED TAXON (LEPIDOPTERA: TORTRICIDAE ) J. F. Gates CLARKE Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560 ABSTRACT. The rediscovery of “Antithesia montana” Bartlett-Calvert is recorded. _ Adult female genitalia are figured, and its assignment to the Tortricidae is established. In 1893 (p. 831, Pl. 1, Fig. 4) Wm. Bartlett-Calvert described and figured Antithesia montana from Lolco, Araucania, Chile, without indi- cation of its family connections. As far as I am able to ascertain, it was not until 1922 (p. 163) that the species was mentioned again, this time by Meyrick, when he placed the species in the Oecophoridae, in the genus Hypercallia. Obviously, Meyrick never saw the species, but based his placement on misinterpretation of the badly illustrated an- tennae, as the long labial palpi of Hypercallia. In January 1974, Dr. Oliver S. Flint, Jr., Department of Entomology, Smithsonian Institution, rediscovered this species in Argentina, at Pu- cara, on the Rio Honthue so now it will be possible to establish more accurately its taxonomic position. Bartlett-Calvert’s type has disappeared; at least I have not been able to locate it. Dr. Ariel Camousseight, Chief, Seccion Entomologia, Museo Nacional de Historia Natural, Santiago informed me that the type is not in that museum, and Dr. Klaus Sattler, of the British Museum (Natural History), where some of Bartlett-Calvert’s material was de- posited, informed me that montana is not represented in that collection. “Lolco, Araucania” is given as the type locality. “Araucania . . . was the name of a former region of Chile . . . now mainly comprised in the provinces of Arauco and Valdivia.” Lolco, however, is now in the province of Malleco. The specimen before me, collected by Dr. Flint, came from Pucara, just over the border between Argentina and Chile. Pucara is situated approximately 225 km south of Lolco and 25 km west of San Martin de los Andes. The original description in Spanish is as follows: “Las alas ante- riores, por encima con la mitad basilar amarilla; la mitad esterna i una parte de la base negruzca, o con reflejos de luz rojizo-negruzco; en el centro de la mitad esterna hai una mancha redonda amarilla, encerrada por un circulo negro; las posteriores de un color moreno-negruzco bril- bo Ol bo JOURNAL OF THE LEPIDOPTERISTS SOCIETY 2 _ Figs. 1-2. Proeulia montana (Bartlett-Calvert). 1, reproduction of the original figure; 2, from Argentina, Pucara. VotuMe 32, NuMBER 4 na 8 liante; por de bajo, todas las alas son morenas inclinandose a negro; la cabeza de color amarillo; el torax i abdomen moreno oscuro; las franjas en las alas anteriores son negruzcas i en las posteriores moreno-claras.” A free translation of the above follows: “The forewings above from center to base yellow; the outer half and part of the base dark brown, or with reflections of reddish brown; in the center of the outer half there is a round yellow spot contained by a black circle; the hindwings of a shining tawny dark brown; the under- sides of all the wings are brown inclining to black; the head yellow; the thorax and abdomen dark brown; the fringes of the forewing are dark brown and in the hindwings clearly brown.” The description fits the specimen in hand and needs no emendation. The female genitalia are described below for the first time. No male is available. Genus Proeulia Clarke Proeulia Clarke, 1962, Proc. Biol. Soc. Washington, 75, 293-294 (Type species.— Eulia robinsoni Aurivillius, in Skottsberg, The Natural History of Juan Fernandez and Easter Island, 3: part 2, 266, Pl. 11, fig. 17). Proeulia montana ( Bartlett-Calvert), new combination Antithesia montana Bartlett-Calvert, 1893, Santiago de Chile, Univ. Anales, 84:831, Pl. 1, Fig. 4. Hypercallia montana (Calvert), Meyrick, 1922, In Wytsman, Genera Insectorum, 180: 163 Male genitalia unknown. Female genitalia (USNM 24331). Ostium very broad, strongly sclerotized inwardly. Anirum not differentiated from the strongly sclerotized, very short ductus bursae. Bursa copulatrix membranous without ventral sclerotized process. Ductus seminalis from latero-ventral surface of bursa copulatrix. Type. Lost. Type locality. Chile, Malleco, Lolco. Distribution. Chile, Argentina. Foodplant. Unknown. Remarks. Although this species lacks one feature characteristic of the genus Proeulia, the sclerotized process from the ventral surface of the bursa copulatrix, I do not hesitate to place montana in this genus. As pointed out by Obraztsov (1964), “Only in the description of the wing venation are some modifications necessary.” He points out that veins 6 and 7 of the hindwing are sometimes slightly separate, as opposed to being stalked, as originally described; also that veins 3 and 4 of hindwing “are either connate or slightly separate at origin” the latter condition found in montana. Obraztsov also points out that the peculiar process from the ventral surface of the bursa copulatrix is reduced in some species, and speculates that it might dis- appear in some taxa. In the case of montana this process is absent, as predicted by Obraztsov. Since the identity of this species appears to be beyond doubt, the designation of a neotype is not necessary. 254 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Fig. 3. Proeulia montana (Bartlett-Calvert), ventral view of female genitalia. VoLuUME 32, NuMBER 4 USS, ACKNOWLEDGMENTS The photographs were made by Victor E. Krantz, and the drawing of the genitalia was made by Mr. George Venable, both on the staff of the Smithsonian Institution. LITERATURE CITED BARTLETT-CALVERT, W. 1893. Nuevos Lepiddépteros de Chile. Santiago de Chile, Univ. Anales, 84: 813-834, Pls. 1, 2. Meyrick, E. 1922. In Wytsman, Genera Insectorum. Lepidoptera Heterocera. Oecophoridae, 180: 1-200, Pl. 1-6. Osraztsov, N. S. 1964. Neotropical Microlepidoptera, V, Synopsis of the species of the genus Proeulia from central Chile (Lepidoptera: Tortricidae). Proc. U.S. Nat. Mus., 116( No. 3501), p. 183-194, Pl. 1-9. Journal of the Lepidopterists’ Society 32(4), 1978, 256-260 LARISA SUBSOLANA, A NEW GENUS AND SPECIES OF MOTH FROM EASTERN NORTH AMERICA (OLETHREUTIDAE) WILLIAM E. MILLER U.S. Dept. of Agriculture, Forest Service, North Central Forest Experiment Station, Folwell Ave., St. Paul, Minnesota 55108 ABSTRACT. Larisa Miller, new genus, is proposed for Larisa subsolana Miller, new species. Larisa is intermediate between the subfamilies Laspeyresiinae and Eucosminae but is tentatively placed in the former. Larisa subsolana is described from more than 130 adult specimens representing a geographic range from Texas and Florida north to Michigan, Ontario, and Massachusetts. Capture dates in Florida range from March 14 to September 27; elsewhere April 10 to August 7. I have noticed the species discussed here in museums and private collections for a decade. It was sometimes identified as Epinotia, Gyp- sonoma, or Laspeyresia, genera which represent two olethreutid sub- families. Detailed study eventually showed that the species cannot be placed in any existing genus. This report is based on more than 130 adult specimens. The letter n denotes number of observations or speci- mens underlying a particular statement. Larisa Miller, new genus Male and female. Head: Maxillary palpus with two developed segments (3 n); labial palpus slightly upturned, second segment expanding apically; antenna 2/5 xX forewing length; scaling of front and crown dense, bushy. Thorax: Smooth-scaled; metathoracic legs unmodified. Forewing: Smooth-scaled; slightly broader toward termen; costal fold absent; costa slightly and uniformly curved from base to apex; apex acute; termen convex; dorsum curved; 12 veins, all separate, upper internal vein of cell arising between veins 10 and 11, vein 11 arising near middle of cell (Fig. 1,5). Hindwing: Costa convex near middle; apex acute; termen concave; dorsum straight between veins 1b and 3; pecten normal; veins 3 and 4 stalked to almost connate; vein 5 straight or slightly bent at base toward 4; veins 6 and 7 stalked (Fig. 1, 5 n). Abdomen: Smooth-scaled; eighth segment of male with a pair of lateral scale tufts; eighth tergite of female with scales as well as setae. Male genitalia: Uncus well developed, sclerotized and bifid; gnathos fused ven- trally across middle; hami long, finger-like; valva simple with rudimentary clasper, a tuft of fine setae on base of sacculus; one to several long, slender setae may be present on latero-ventral surface of cucullus; aedeagus sleeve-like, short, tapered; deciduous cornuti present; dorsal plate of anellus not developed. Female genitalia: Papillae anales simple; posterior apophyses slightly longer to slightly shorter than anterior apophyses; sterigma shield-shaped with short finger-like projections beside ostium, ostium on anterior margin; ductus bursae short, enlarged near middle, sclero- tized except for a short distance beyond enlargement, convoluted at junction with corpus bursae; dual thorn-like signa. Type-species. Larisa subsolana, new species. Comments. Larisa keys to Laspeyresiinae or Eucosminae (Heinrich 1923, Obraztsov 1958) depending on character variability and interpretation. It is an intermediate genus but is tentatively placed in Laspeyresiinae. In the male, the VoLUME 32, NUMBER 4 ASST Figs. 1-6. Larisa subsolana, new species. 1, Venation of fore- and hindwing. 2, Fore- and hindwing of specimen from Devil’s Den State Park, Arkansas. Length of forewing 5.0 mm. 3, Male genitalia of specimen from 3 km E Palmdale, Florida. 4, Enlargement of aedeagus of preceding male. 5, Female genitalia of specimen from preceding locality. 6, Enlargement of sterigma and associated structures of preceding female. 958 JOURNAL OF THE LEPIDOPTERISTS SOCIETY rudimentary clasper and well developed uncus are characteristic of Eucosminae. Within Laspeyresiinae, Larisa most resembles Laspeyresia and Hemimene or Pammene (Heinrich 1926, Obraztsov 1960) but differs from both by its convex forewing termen, long setae on outer surface of cucullus, setal tufts on sacculus, well developed hami, and in previously enumerated details of forewing or hindwing venation. Larisa is feminine gender and a patronym for Larisa K. Miller, my volunteer assistant. Larisa subsolana Miller, new species Male. Length of forewing 3.8-5.8 mm (71 n). Head: Labial palpus brown, scales white-tipped, length of second segment 1 x eye diameter and 2.9-4.0 x length of apical segment as estimated from scaled and descaled specimens (21 n); front and crown light brown; antenna brown. Thorax: Brown dorsally, including tegula, scales white-tipped; shining white ventrally; pro- and mesothoracic legs brown on outer side, scales white-tipped, tarsi white-banded, shining white on inner side; metathoracic legs shining white. Forewing (Fig. 2): Length 2.6-2.9 x width (5 n); ground color of upper side brown; basal patch sharply delineated; middle crossband grayish brown grading apically to darker brown, a thin brown line cen- trally from costa to dorsum; distal third grayish brown, tinged in costal half with rust; fringe brown; underside light brown, mottled with white in costal area. Hindwing (Fig. 2): Widest membranous part 1.1-1.4 x that of forewing (5 n); upperside, underside, and fringe light brown. Abdomen: Grayish brown dorsally, paler ventrally, including genital scaling. Genitalia (Figs. 3-4): Width of valval neck 0.43-0.74 greatest width of cucullus (24 n), the individual values showing a normal frequency distribution; 8-17 deciduous cornuti or empty cornutus sockets (2m): Female. As described for male except forewing length 4.1-6.3 mm (61 n) and brown genital scaling. Genitalia (Figs. 5-6, 20 n): Sterigma with short finger-like projections lateral to ostium bursae; posterior apophyses slightly longer to slightly shorter than anterior apophyses. Types. HoLtotype ¢: ARKANSAS, Devil’s Den State Park, Washington Co., June 26, 1966 (R. W. Hodges), No. 72093 in National Museum of Natural History. ALLOTYPE 2: ARKANSAS, same data as holotype except May 30, 1966, in National Museum of Natural History. Paratypres, 10 specimens: ARKANSAS, same data as holotype except 22 May 1966, ¢ genitalia slide USNM Tor 2, wing slide WEM 5; same data as holotype except 20 May 1966, ¢ genitalia slide LKM 1219766; 13 km SE Ethel, Arkansas Co., 9 July 1969 (R. L. Brown); MISSISSIPPI, Clinton, Hinds Co., 14 July 1974 (Bryant Mather), No. 73267, 2 genitalia slide LKM 403772; MICHIGAN, East Lansing, Ingham Co., 15 July 1968 (J. P. Donahue), ¢ genitalia slide JAB 34; T4N, R2W, Sec. 35, Ingham Co., 12 June 1966 (J. P. Donahue), ¢ genitalia slide KAK 73; ALABAMA, 21 km SW Greensboro, 23 April 1976 (J. B. Heppner); FLORIDA, 3 km E Palmdale, 4 May 1974 (J. B. Heppner), @ genitalia slide JBH 455; same data as preceding except ¢@ genitalia slide JBH 454; NEW YORK, Ithaca, 2 July 1976 (J. G. Franclemont), 2 genitalia slide RLB 645. Para- types are in National Museum of Natural History; California Insect Survey, Univer- sity of California, Berkeley; Florida State Collection of Arthropods; Cornell Uni- versity; University of Minnesota, Twin Cities; and collections of Richard L. Brown, John B. Heppner, and Bryant Mather. Specimens not designated as paratypes are in the above repositories, also University of Michigan and Field Museum of Natural History. Geographic distribution. Present records for the species occur from Texas and Florida north to Michigan, Ontario, and Massachusetts (Fig. 7). Biology. Available biological information is based on adults captured in flight. The hostplant is unknown. There is probably more than one generation a year. Capture dates in Florida range from March 14 to September 27 (20 n); elsewhere, VoLuME 32, NUMBER 4 259 Fig. 7. Distribution of records for Larisa subsolana. April 10 to August 7 (116 n). If the largest sample of moths from one locality (46 n, Devil’s Den State Park, Arkansas, 20 May—-22 July 1966, R. W. Hodges) represents one generation, the flight period is longer than that of many olethreutids and suggests a protected place of development insulated by shade, soil, or woody tissue. This sample also shows protandry typical of olethreutids (the median capture date of males preceding that of females by 12 days) and a male/female ratio of 0.92, essentially unity. ACKNOWLEDGMENTS I thank the following for specimen loans and other assistance: Jerry A. Powell, University of California, Berkeley; Don R. Davis, National Museum of Natural History; Roland L. Fischer, Michigan State Uni- versity; Thomas E. Moore, University of Michigan; Bryant Mather, Clinton, Mississippi; Charles P. Kimball, West Barnstable, Massachu- setts; and Henry Dybas, Field Museum of Natural History. I am espe- 260 JOURNAL OF THE LEPIDOPTERISTS SOCIETY cially indebted to Richard L. Brown, Cornell University, and John B. Heppner, University of Florida. After more than 50 Larisa specimens had come to my attention, I prepared a draft of this paper and asked several workers including Brown and Heppner to review it. Unknown to me, both were also independently studying the insect and each generously forwarded research information and more than 60 additional specimens to me. LITERATURE CITED Hernricu, C. 1923. Revision of the North American moths of the subfamily Eucosminae of the family Olethreutidae. U.S. Natl. Mus. Bull. No. 123, 298 pp. 1926. Revision of the North American moths of the subfamilies Laspey- resiinae and Olethreutinae. U.S. Natl. Mus. Bull. No. 132, 216 pp. Opraztsov, N. 1958. Die Gattungen der Palaearktischen Tortricidae. II. Die Unterfamilie Olethreutinae. Tijdsch. Entom. 101: 229-261. . 1960. Die Gattungen der Palaearktischen Tortricidae. II. Die Unter- familie Olethreutinae. 3. Teil. Tijdsch. Entomol. 103: 111-143. Journal of the Lepidopterists’ Society 32(4), 1978, 261-272 NOTES ON MEXICAN ACTINOTE (NYMPHALIDAE: ACRAEINAE) AND THEIR RELATIVES, WITH DESCRIPTION OF A NEW SUBSPECIES JACQUELINE Y. MILLER AND LEE D. MILLER Allyn Museum of Entomology, 3701 Bay Shore Road, Sarasota, Florida 33580 ABSTRACT. The application of names described for South American species to their Central American, and especially Mexican, counterparts has led to great con- fusion in the literature. Actinote stratonice oaxaca is described from Oaxaca, Mexico; this insect had been reported previously as the nominate subspecies. Three species of the Actinote thalia group, A. calderoni, lapitha and thalia are illustrated, redescribed and discussed. A key for the separation of the three species is provided. The first two species are recorded from Mexico—calderoni had been previously misidentified as lapitha from there. One name, A. lapitha zilchi Franz and Schroder, is synony- mized to calderoni. Collecting in Mexico over the past forty years has yielded many butterfly species that were previously unknown from there, not a few of which were totally unexpected. These unexpected taxa have created many problems, usually for one of two reasons: 1) the butterfly was an already described Central or South American species and was de- scribed as new from Mexico because of a lack of comparative material or an ignorance of the pertinent literature; or 2) many species (espe- cially those figured and described in Seitz) incorrectly have been as- cribed to the Mexican fauna, again because of a lack of adequate comparative material. Both situations are well demonstrated in the Nymphalidae: Acraeinae. The Hoffmann (1940) catalog lists only four species of this subfamily within the borders of Mexico, but recent collecting has uncovered one that has been misidentified in collections and in correspondence—the one with which the first-mentioned species had been confused and an undescribed subspecies of a well-known South American insect. In the hope of unravelling the confusion in this small subfamily (within the Mexican borders), we offer these notes. Actinote stratonice oaxaca J. Miller and L. Miller, new subspecies Figs. 1-6 Male. Head, thorax and appendages black; abdomen black with a reddish-brown midsternal line. Upper surface of wings similar to that of the nominate subspecies, but paler, and with the following differences: forewing totally black anteriad of cell (partially reddened in other subspecies ); dark marking at end of forewing cell much smaller than in other subspecies and black area at base of forewing cell and along inner margin more restricted than in other populations. Under surface pattern also paler than in s. stratonice with forewing differences as noted for upper surface bo o>) bo JouRNAL OF THE LEPIDOPTERISTS SOCIETY Figs. 1-4. Actinote stratonice oaxaca J. Miller and L. Miller, new subspecies. 1-2, Holotype ¢, upper (1) and under (2) surfaces; MEXICO: OAXACA: Can- delaria Loxicha, 550 m. 3-4, Paratype 2, upper (3) and under (4) surfaces; same locality. Both specimens in Allyn Museum of Entomology. and in addition the pale orange patches in the forewing cell area darkened basad, blending to tawny-yellow distad; black areas admixed with pale yellow scales, espe- cially in base hindwing. ¢ genitalia as illustrated, differing from those of other stratonice in only minor respects. Length of forewing of Holotype ¢ 24.4 mm, those of eleven of the @ Paratypes range from 26 to 31 mm. Female. Differs from the 2 of nominate stratonice in the same manner as does the @, but additionally the basal black areas of the forewing encompass the proximal third of the cell and below it along the inner margin, and all of the black areas below are completely suffused with yellow scaling. One @ Paratype has a hindwing supernumerary vein off Rs on the right side. @ genitalia as illustrated and com- paring well with those of other subspecies. Lengths of forewings of 12 2 Para- types range from 28.2 to 38 mm. Specimens examined. Described from 25 specimens, 13 males and 12 females, from the state of Oaxaca, Mexico. Types. Hotorype ¢: MEXICO: OAXACA: Candelaria Loxicha, 550 m, 8.ix. 1969 (KE. C. Welling). Pararypres: all MEXICO: OAXACA: same locality as Holotype, 14 15.xi.1967, 16 15.ix.1968, 12 21.vii.1970, 14 19 27.viii.1970, 19 21.vii.l1973 (all E. C. Welling M.); El Portillo del Rayo, Candelaria Loxicha, 14 32 17.xi.1967 (all E. C. Welling M.), 64 49 18.vii.1976 (all de la Maza family); Rio Molina, Mpio. Suchistepec, 2200 m., 19 10.x.1967 (E. C. Welling M.); San Jose Pacifico, Mpio. Rio Hondo, 2400 m., 14 9.x.1967 (E. C. Welling M.); Puente VoLUME 32, NuMBER 4 263 Figs. 5-6. Actinote stratonice oaxaca J. Miller and L. Miller, new subspecies. 5, 6 genitalia of Paratype; MEXICO: OAXACA: Candelaria Loxicha; preparation M-3630 (Jacqueline Y. Miller). 6, 2 genitalia of Paratype; same locality; prepara- tion M-3602 (Jacqueline Y. Miller). del Guajolote, Jalatengi, 19 21.iii.1975 (E. Fernandez), 14 21.xi.1975 (de la Maza family); Dos de Mayo, 1¢ 18.v.1976 (de la Maza family). Disposition of type series: Holotype ¢, two ¢ and four @ Paratypes in the collection of the Allyn Museum of Entomology; two ¢ and three @ Paratypes in the collection of E. C. Welling M. and eight ¢ and five 2 Paratypes in the de la Maza collection. Remarks. The subspecific name refers to the state from whence the new sub- species came. A. stratonice is recorded in Seitz from the Sierra Madre de Santa Marta of Colombia and southward through the mountains of Venezuela and into Ecuador. The present subspecies was first recorded in the literature from Mexico by de la Maza R. and de la Maza E. (1975), but at the time they did not recognize it as a separate subspecies. The subspecies oaxaca is characterized by its overall dull coloration and by the suffusion of the under surface with pale yellow scales. In none of the other subspecies of stratonice do these characters appear. The disjunct distribution of stratonice, with a Mexican subspecies widely separated from its nearest relatives, is indeed intriguing. 264 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Figs. 7-10. Actinote calderoni Schaus. 7-8, 4, upper (7) and under (8) sur- faces; MEXICO: CHIAPAS: Mapastepec. 9-10, 2, upper (9) and under (10) surfaces; same locality. Both specimens in Allyn Museum of Entomology. The Identity of the Mexican Actinote thalia Group Species Pale Mexican specimens of thalia group Actinote in most collections usually have been identified as A. lapitha (Staudinger). Comparison of most Mexican material with Staudinger’s description and subsequent literature citations suggest that these specimens could not be true lapitha. Accordingly, we searched museum and private collections and the literature for documented material that might shed light on the correct name for the Mexican insects. The results were surprising, and to avoid future confusion the following redescriptions are offered to aid in the identification of these butterflies. Actinote calderoni Schaus, 1920 Figs. 7-12 Actinote calderoni Schaus, 1920: 434 (TL—Anteos, El Salvador). = Actinote lapitha zilchi Franz and Schréder, 1954: 80; fig. 2 (TL—Km. 30, Son- sonate Rd., La Libertad, El] Salvador). [New Synonymy. ] Male. Head, thorax and first two abdominal segments dark brown covered with a few fuscous and tawny dorsal hairs; remaining abdominal segments naked VoLUME 32, NUMBER 4 265 Figs. 11-12. Actinote calderoni Schaus. 11, @ genitalia; MEXICO: CHIAPAS: Mapastepec; preparation M-3633 (Jacqueline Y. Miller). 12, 2 sterigma; same lo- cality; preparation M-3601 (Jacqueline Y. Miller). dorsad; thorax and first two abdominal segments covered with a few tawny ventral scales; last abdominal segments ventrally naked; pleural line buff. Palpi tawny with a few black scales along inner surface only. Antennae and legs black. Upper surface of wings very thinly scaled with white, giving a dusky, off-white appearance; forewing smoky at apex and marginally to Cuz-2A; veins slightly dark- ened and interneural spaces with single smoky stripes. Under surface of wings similar, but a faint, dark comma-shaped marking lies from M;-Cu: to Cui-Cuz of the forewing and numerous long, dark hairs lie on the veins, especially of the hindwing. 6 genitalia as illustrated. Forewing lengths of the ¢ examples examined range from 21.5 to 24.5 mm. Female. Similar to ¢, but paler, especially the forewing smoky markings above and below. @ genitalia as illustrated, generally characterized by having the sterigma more sculptured than in related species. Lengths of forewings of 2 specimens at hand range from 25 to 30.5 mm. Types and specimens examined. We have examined 15 specimens, seven males and eight females, from the following localities: EL SALVADOR: Anteos, 1¢ (Hotoryee, USNM), 12 (possible Paratype, CM). GUATEMALA: Tiquisate, 28.vi.1947, 12 (AMNH). MEXICO: CHIAPAS: Mapastepec, various dates, 1939- 1959, 24 59 (AME), 3é6 12 (AMNH); San Jeronimo, 600 m., 17.vii.1975, 192 (E. C. Welling M.). The records from Mexico and Guatemala are apparently the first for either country. The present insect has been masquerading in Mex- ican collections for years as A. lapitha (Staudinger), a species that is abundantly distinct. Not only have workers on Mexican butterflies been confused on the identity of calderoni, but also those in other parts of Central America. The fact that Schaus’ (1920) description appeared in an entomologically obscure journal has resulted in the paper never being cited previously by workers on Actinote. Were the “Fifty-Year Rule” still in effect in 266 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Figs. 13-16. Actinote lapitha (Staudinger). 13-14, ¢, upper (13) and under (14) surfaces; MEXICO: CHIAPAS: Tierra Blanca, Mpio. La Trinitaria, 1500 m. (E. C. Welling M. collection). 15-16, 2, upper (15) and under (16) surfaces; no locality data (AMNH collection ). the International Code of Zoological Nomenclature, the name calderoni could be ignored with impunity, but happily this rule was repealed a few years ago, so we must return to the oldest name. Franz and Schroder (1954) had obviously not seen the Schaus description, but their excellent figure of the type of A. lapitha zilchi is referable to specimens of calderoni that we have seen (including the Holotype), and since the two taxa were described from within 100 km. of one another, it is evident that they represent the same species. Actinote lapitha (Staudinger), 1888 Figs. 13-18 Acraea lapitha Staudinger, 1888: 82 (TL—“Chiriqui” ). Male. Head, thorax and abdomen clothed with black dorsal hairs; head, palpi and most of thorax also clothed with black ventral hairs; small patch of tawny scales on meso- and metathoracic preepisterna and an additional such patch on metathoracic epimeron; abdomen clothed with admixed fuscous, tawny and buff scales; pleural line tawny and buff only. Antennae and legs black. Ground color of forewing above translucent and tawny with margins outlined in dull gray-brown, especially at apex; prominent gray-brown transverse marking from end cell to Cu.-2A, interspersed with VoLUME 32, NuMBER 4 267 Figs. 17-18. Actinote lapitha (Staudinger). 17, ¢ genitalia; MEXICO: CHI- APAS: Tierra Blanca, Mpio. La Trinitaria; preparation M-3416 (Jacqueline Y. Mil- ler). 18, 2 sterigma; no data; preparation M-1657 (Jacqueline Y. Miller). tawny scaling from end cell to M;-Cu:; veins and interneural striping prominent, gray-brown. Hindwing above also translucent tawny, dull gray-brown at costa and along margin; veins heavily darkened with gray-brown and interneural markings of same color short and heavy. Forewing below similar to upper surface, but margins and apex not so dark and with an additional dull gray-brown bar across middle of cell. Hindwing below as above with a faint gray-brown cell end marking from costa to base of M; and more prominent veinal and interneural blackish-brown striping. Fringes of both wings blackish on both surfaces. ¢ genitalia as illustrated. Lengths of forewings of the three ¢ specimens examined range from 22.8 to 23.6 mm. Female: Similar to ¢, but thorax sparsely clothed with tawny scales, markings of all wings paler, but extra-discal band of forewing more prominent and base of cell of same wing somewhat overscaled with fulvous. 2 genitalia as illustrated; sterigma somewhat heavier than that of the next species and not quite so ornamented as in calderoni. Lengths of forewings of the two 2 examples before us 20.2 and 23.1 mm. Types and specimens examined. We have seen two females and three males of this insect. PANAMA: Jicaron Island, 14—15.i.1902, 1¢ (BMNH). COSTA RICA: Puerto Golfito, 4.vii.1965, 1¢ (Gordon B. Small, Jr. collection). MEXICO: CHI- APAS: Tierra Blanca, Mpio. La Trinitaria, 1500 m., 15.ix.1972, 1¢ (E. C. Welling M.) No Data, 22 (AMNH). Evidently the Costa Rican record is a new, but not unexpected one. Hoffmann (1940: 672) lists lapitha from “Tierra caliente de la costa del Pacifico de Chiapas,” no true specimens of that species are in the Hoff- mann collection in the AMNH. All of the specimens in Hoffmann’s material were calderoni, and one of these bore a determination label in Hoffmann’s hand of “Actinote lapitha Staudinger.” Since Hoffmann obviously confused lapitha with calderoni, we feel that Mr. Welling’s specimen of the former is the first authentic record from Mexico. A. lapitha was described from the Chiriqui region of Panama, but the type specimen is apparently no longer extant, perhaps having been 68 JouRNAL OF THE LEPIDOPTERISTS SOCIETY Figs. 19-22. Actinote subhyalina (Staudinger). 19-20, ¢, upper (19) and under (20) surfaces; PERU: Rio Cachiyacu, Iquitos (BMNH). 21-22, Lectotype @ (see designation in text), upper (21) and under (22) surfaces; PERU: Yurimaguas (ZMHU). destroyed during World War II (H. J. Hannemann, pers. comm.). We have been unable to locate an authentic Chiriqui specimen in more recent collections; hence, we do not designate a Lectotype here. Actinote subhyalina (Staudinger), 1888 Figs. 19-24 Acraea subhyalina Staudinger, 1888: 81; pl. 32 (TL—Yurimaguas, Peru). Description. Sexes similar; Head, thorax and abdomen above and below blackish-brown clothed with fuscous to buff scales. Palpi blackish-brown; legs brown and antennae dull reddish-brown. Forewing above buff, translucent and laved with fulvous toward base (darkest in @); apex and margin dull brown; veins darkened; transverse markings: a jagged blackish-brown one across cell about %4 distance from base to end and a second one from costa across end cell to anal margin (not prominent below Cu; in ¢). Hindwing above buff, translucent, with darker veins; margin outlined in dull brown; interneural striping in anal region; prominent dark brown bar from costa to Cu;-Cuz and a dull brown mark across distal end of cell. Under surface of wings as above, but dull brown markings slightly overscaled with buff, hindwing distal band more diffuse and an additional dull brown stripe in hindwing cell Sc+Ri-Rs. Lengths of forewings of all specimens examined ranged from 20 to 22 mm. Genitalia of ¢ and 9° as illustrated. Specimens examined. We have been able to examine only a single ¢ and six ? specimens. PERU: Rio Cachiyacu, Iquitos, [18]93, Stuart, 14 39 (BMNH); Yurimaguas, 19 (ZMHU); No data, 22 (CM). VoLUME 32, NUMBER 4 969 Figs. 23-24. Actinote subhyalina (Staudinger). 23, 4 genitalia; PERU: Rio Cachiyacu, Iquitos; preparation M-3403 (Jacqueline Y. Miller). 24, @ genitalia of Lectotype; PERU: Yurimaguas; preparation M-3411 (Jacqueline Y. Miller). Staudinger (1886: 81) described A. subhyalina from 12 2 specimens taken at Yurimaguas. Certainly the figured 2° from Berlin was one of Staudinger’s syntypes, and the two specimens from CM may have been, but in the case of the latter two specimens this cannot be ascertained with precision. One of the CM specimens bears the label “Acraea subhyalina/ from Dr. O. Staudinger/1885,” a date that was three years before the description; the second specimen bears a number (Stauding- ers?) only, “386.” The specimen received from the ZMHU is defi- nitely from the type locality, was in the Staudinger collection and has been labelled as “Origen” by Staudinger or someone subsequent to him. It is the logical candidate for designation as the Lectotype of the name, and we have so labelled it, affixing a red, partially printed, partially handwritten (italics) label to it: “Lectotype/ Acraea/ subhyalina/ Staudinger, 1888/ designated by Jacqueline Y. Miller/ & Lee D. Miller, HOLT. DIscUusSION The impetus for this project was a series of seven specimens in the Allyn Museum collection from Mapastepec, Chiapas, Mexico. Exami- nation of these specimens revealed that while they were closely related to A. lapitha (the name associated with them), they were abundantly distinct. Letters for additional specimens brought two from the AMNH that were in agreement with the original description of lapitha (but without data) and five more of the odd one, four of which were from the 270 JoURNAL OF THE LEPIDOPTERISTS SOCIETY Hoffmann collection, taken by Dr. Escalante. A later trip to CM yielded one more specimen and the first clue to the identity of the Mexican material that we had. The additional label on the CM specimen stated that it was “Actinote calderoni Schaus,” but at the time we were unable to find the original description or any reference to it, and we still as- sumed that the name might have been a manuscript name only. Re- checking the series and the type collection at the USNM yielded not only the type specimen of calderoni, but also finally the reference to the original description of this elusive name. The confusion did not end there, though. The two true lapitha from the AMNH had a fulvous basal flush on the forewing, thus resembling subhyalina. This led us to wonder if the specimen figured by Staudinger (1888) was subhyalina or lapitha. Letters to the BMNH and the ZMHU brought additional specimens of Staudinger’s insects and some very helpful information. All three species are rare in collections, especially the males. Since we encountered such difficulty in making determinations in the thalia group, we present the following key to aid other workers to the species treated here. 1. Forewing above with dark transverse band at end cell from costa to near anal angle: 2-222. Sone nce ee 2. 1’. Forewing with no such band; Mexico to El Salvador calderoni Schaus. 2. Hindwing above with prominent dark discal markings at end of cell; Peru PE pk lial RE 8 cca are ed TEE Ly ail al Shah subhyalina (Staudinger). 2’. Hindwing above without prominent marking at end cell; Mexico to Panama eg dont GM Nu | aetna Di eth A es lapitha (Staudinger). Essentially the key characters for the separation of the species in this complex are the dark bars across the cells of both wings. A. subhyalina shows these bars on the upper surfaces of both wings, A. lapitha has only the one on the forewing and A. calderoni has neither. Seemingly the orange flush at the base of the upper forewing should be diagnostic, but whereas it is prominent in most subhyalina, it also appears in some female lapitha, hence it is diagnostic of neither. | The male genitalia (Figs. 11, 17, 23) are similar, but subtly different, in all three species. The valvae are elongated and slightly curved dorsad in calderoni, whereas they are squarecut posteriad in both lapitha and subhyalina. The saccus of subhyalina is much more elongate than is that of either of the other two species. Most of the differences between females of these species lie in the sterigmal region. The posterior margin of the lamella postvaginalis is U-shaped and narrow in lapitha, U-shaped and expanded in subhyalina and even more enlarged and W -shaped in calderoni. The entire opening VoLUME 32, NuMBER 4 TEAL of the ostium bursae is darkly sclerotized in a narrow ring in lapitha, a somewhat broader darkly sclerotized ring in subhyalina but only darkly sclerotized in four separate areas around the opening in calderoni, but the area around the ostium itself is lightly sclerotized. The genital capsule is larger in calderoni than in the other species. The tarsi, as is true of all members of the thalia group, are asym- metrical and comparable one with another. Geographically A. subhyalina can be immediately separated from lapitha and calderoni, none of the three species being found in Colombia or Ecuador, as far as we know. It would not be surprising to see subhyalina from at least the latter country, and there might be sym- patry between subhyalina and lapitha in Colombia. A. lapitha, as re- corded here, has a much more extensive range than previously believed, and calderoni is not restricted to El Salvador. Either of these species, or both, may well be found in Honduras and Nicaragua. We hope that this paper will encourage others to try to fill in the distributional blanks for this interesting group. ACKNOWLEDGMENTS We are deeply indebted to the following for loan of material and for access to their collections (abbreviations) of Actinote: Dr. F. H. Rindge, American Museum of Natural History (AMNH); Mr. H. K. Clench, Carnegie Museum of Natural History (CM); Mr. W. D. Field, National Museum of Natural History (USNM); Mr. P. R. Ackery, British Museum (Natural History) (BMNH); Dr. H. J. Hannemann, Zoologische Museum der Humboldt Universitat (ZMHU); Dr. T. Escalante and Messrs. R. de la Maza R. and J. de la Maza E., Mexico, D. F., Mexico; Mr. E. C. Welling M., Merida, Yucatan, Mexico and Mr. G. B. Small, Jr., Balboa, Canal Zone. The Allyn Museum of Entomology is abbreviated AME in part of the paper. Mr. Field and Dr. J. F. G. Clarke of the USNM were instrumental in obtaining a copy of the Schaus paper for us. We also thank Mr. S. R. Steinhauser for pointing out the Franz and Schroder paper to us and to Mr. H. W. Dybas of the Field Museum for obtaining a copy of it for our use. Special thanks are due Mr. A. C. Allyn of this institution for taking the photographs contained herein and for reading the manuscript. LITERATURE CITED Franz, E. & H. Scurover. 1954. Tagfalter (Lep. Rhopalocera) aus El Salvador. Senckenbergische Biol. 35: 75-87; figs. bo ~l bo JOURNAL OF THE LEPIDOPTERISTS SOCIETY HorrMAnn, C. C. 1940. Catalogo sistematico y zoogeografica de los Lepidopteros mexicanos. la. Parte. Papilionoidea. An. Inst. Biol. 11: 639-739. Jorpan, K. 1924. Acraeidae. In Seitz, A., Macrolepidoptera of the World 5: 358— 374. DE LA Maza R., R. & J. DE LA Maza E. 1975. Adiciones al Catalogo de Lepidop- teros mexicanos. Rev. Soc. Mexicana Lepid. 1: 62-63; figs. Scuaus, W. 1920. Descriptions of two new species of butterflies from tropical America. J. Washington Acad. Sci. 10: 434-435. STAUDINGER, O. 1888. In Staudinger, O. and E. Schatz, Exot. Schmett. (1): 81- oo. Pio: Journal of the Lepidopterists’ Society 32(4), 1978, 273-276 PAPILIO ARISTODEMUS (PAPILIONIDAE) IN THE BAHAMAS Harry K. CLENCH Carnegie Museum of Natural History, Pittsburgh, Pennsylvania 15213 ABSTRACT. Two subspecies of Papilio aristodemus Esper, both new, are de- scribed from the Bahamas: driophilus (TL: Cutlass Bay, near Dolphin Head, Cat Island), known from Cat, South Andros, and North Andros islands; and bjorndalae (TL: Man of War Bay, Great Inagua Island), known only from Inagua and strikingly different from any known subspecies, though apparently derived from driophilus. Papilio aristodemus is an Antillean swallowtail with a strong tendency to vary geographically. Nominate aristodemus Esper 1794 occurs on Hispaniola; the subspecies temenes Godart 1819 is found on Cuba and on Little Cayman in the Cayman Islands (Carpenter & Lewis 1943); subspecies ponceanus Schaus 1911 is known only from southeastern Florida, particularly Key Largo. An old record of the species for Puerto Rico (cf. Comstock 1944: 535), subspecies unknown, is not substantiated by more recent captures. This species recently has been discovered in the Bahamas (Clench, 1977). I first found it on South Andros Island in early June 1974. A year later, in early June 1975, I took it also at the southern end of Cat Island, and in 1976 I collected a specimen on North Andros. The populations on these islands are not absolutely identical, but they are close enough to be referred to the same subspecies, described below as driophilus. Miss Karen Bjorndal, a graduate student at the University of Florida, Gainesville, spent over a year on Great Inagua Island, from April 1975 to August 1976, studying the energy budget and nutritional ecology of the Green Turtle, Chelonia midas. While there she also made a collec- tion of butterflies, which she has generously donated to Carnegie Mu- seum of Natural History. In her collection are two specimens of a striking new subspecies of aristodemus, in several ways the most distinct of all. It is a pleasure to name it in honor of Miss Bjorndal. Papilio aristodemus driophilus, new subspecies Papilio aristodemus ponceanus: Clench 1977:190. Description. Much closer to ponceanus (Florida) than to either a. aristodemus (Hispaniola) or temenes (Cuba). This is shown particularly by its sharing with ponceanus such traits as the thin median yellow-ocher band, and the complete, only slightly curved, subterminal row of yellow-ocher lunules, both on the forewing above. From ponceanus, however, it differs in these ways: (1) On the hindwing upperside the subterminal yellow, orange, or red-orange spot 274 JOURNAL OF THE LEPIDOPTERISTS SOCIETY in Cu-2A is completely separated by a black bar from the terminal yellow or orange distad. In ponceanus these two pale areas are connected by a narrow isthmus along Cuz. (2) On the forewing above, the cell is rather densely and evenly sprinkled with pale (greenish) scales. In ponceanus the sprinkling is extremely sparse and tends to be limited to the basal and costal parts of the cell. (3) The projections of the hindwing termen at the vein-ends (including the tail) are longer than they usually are in ponceanus (but the latter is inclined to be variable in this respect). (4) On the forewing above, the segment of the median band in M:-Mb is broadly in contact with the next posterior segment, in M2-M;. In ponceanus the M:-M2 seg- ment is smaller, and posteriorly separated from the next one by a fuscous gap or (rarely) touches the next segment at a point only: the two segments are never in broad contact. (5) On the hindwing upperside the median pale band is somewhat broader than in ponceanus. In driophilus the segment in the cell is consistently wider than the fuscous in the cell just distad; in ponceanus the pale band is here subequal to the fuscous in width, or it is somewhat narrower. Length of forewing. Male, one only, 47.0 mm; female, 48.0-50.0 mm, mean (of 4), 49.1 mm. Measurements are of the type series only. Types. Ho.otype, 2, Cutlass Bay, near Dolphin Head, southern Cat Island, Bahamas, 6.vi.1975, leg. H. Clench, sta. 259 b; C. M. Acc. 27783. Paratypess, 13 3@, as follows: 22, same data as holotype; 1¢ 12, the same except 4.vi, sta. 257 b. Holotype and paratypes, C. M. Ent. type series no. 680. Remarks. In addition to the type series I have examined 4¢ 22 from South Andros Island, Bahamas: ca. 2 mi S Driggs Hill, 2-8.vi.1974, leg. H. Clench. The forewing length of this series is as follows: males, 43.5-49.0 mm, mean (of 4), 45.0 mm; females, 47.5--51.0 mm, mean (of 2), 49.2 mm. These specimens agree closely enough with the Cat Island series, notably in all points mentioned in the above description, that I believe them correctly referred to the subspecies driophilus. The agreement, however, is not perfect and the two island samples show a few, mostly statistical, differences: (a) On the hindwing above, the subtornal pale spot (as in (1) above) is dark orange, with little or no pale edging, in all the Cat Island specimens; it is light orange, more or less heavily edged laterally with yellow, in all but 1¢ from South Andros (in which it is dark orange). (In ponceanus; dark orange with slight lateral yellow. ) (b) On the hindwing above, a small rusty spot in the base of cell M3-Cu: is present in 12 12 (40%) of the Cat Island series, but is totally absent from the South Andros series. (In ponceanus: 56%). (c) The discal cell on the forewing underside is filled with smooth, pale yellow- ocher in all Cat Island specimens; in all South Andros specimens the cell has periph- eral fuscous and faint distal longitudinal fuscous streaks. (In ponceanus: as on South Andros, but the fuscous is even heavier. ) (d) On the forewing underside, the subapical transverse fuscous bar from costa (just distad of, and parallel to, the conspicuous pale bar on the forewing upperside ) extends inward to cross cell Rs-M: in 1¢ (17%) from South Andros, in 42 (80%) from Cat Island. In the remaining individuals it does not reach that interspace (In ponceanus: 22%.) On South Andros driophilus flew in dense scrub, usually 1-2 m above the ground, only briefly and occasionally pausing to feed at the flowers of shrubs in that height range. The butterflies were mostly in the scrub itself and they entered roadways or other open areas only to cross from VoLUME 32, NUMBER 4 UTS one part of the scrub to another. These habits they shared fully with P. andraemon bonhotei Sharpe 1900, which flew with driophilus, and the two were virtually indistinguishable on the wing. On Cat Island the habits of driophilus were similar except that individuals were seen more often in open areas, especially at the flowers of ornamental vines and shrubs around the hotel where I stayed. On 28 September 1976 I took a single male driophilus just north of Nicolls Town, North Andros, a new record for that island. I saw no others and am at a loss to explain the late capture date. The butterfly is quite fresh and was found flying in a somewhat overgrown old field. It, too, is referable to the new subspecies, although differing in a few respects (e.g., the median pale band on the hindwing above is thicker than in any other driophilus seen except one of the female paratypes from Cat Island; and its distal edge is straight [as in bjorndalae], not convex near Rs and M,). With regard to traits (a) through (d) above: (a) the subtornal pale spot is dark orange with slight lateral yellow (as in ponceanus); (b) it has no rusty spot in M,;—-Cu,; (c) on the forewing below it agrees with South Andros specimens in the discal cell coloration; (d) also on the forewing below, the subapical fuscous bar extends inward only to R;. Papilio aristodemus bjorndalae, new subspecies Description. Differs in two major traits from all previously known subspecies of aristodemus: (1) a large patch of rusty red is present on the hindwing upperside between M: and the inner margin, and between the cell-end and the diffuse, faint band of sprinkled blue scaling that basally edges the subterminal row of pale spots; and (2) on the hindwing, both above and below, the subterminal pale spots posterior to Mz are distally displaced and reduced in size, so that the row is essentially parallel to the termen throughout and the component spots are of similar thickness and more quadrate (less lunular). The latter trait is particularly conspicuous on the underside. The rusty red patch varies in the two specimens at hand, but I can- not tell whether the variation is sexual or individual. In the female the patch is large, the component spots contiguous, and there is even a minute extra dot of the same color in Mi-M2; in the male the component spot in Cui-Cu: is wanting, and those in Me-M;-Cu; are thin and short, separated by fuscous along the veins. The median pale band of the forewing upperside is thin, as in subspecies pon- ceanus, driophilus, and aristodemus, and slightly or not at all broken at Mb, as in driophilus; on the forewing upperside the subterminal row of pale spots is lightly curved (as in all subspecies except nominate aristodemus, in which it is strongly curved, almost angulate, near Cui), and continues strongly costad to Ri, as in pon- ceanus (in a. aristodemus it stops at M2; in temenes at about M,; in driophilus at R; or Ru, the segment in R.-R;s being often weak or wanting). On the hindwing upperside, in Cu-inner margin, the subterminal pale bar is connected to the pale terminal area by a narrow isthmus along Cu, as in ponceanus (in all other sub- species the two pale areas are usually completely separated by intervening fuscous). The median pale band on the hindwing upperside is thin, about as in ponceanus or even thinner, and about half as thick as that in driophilus, and its distal edge is straight, not convex near Rs, as it is in ponceanus and driophilus. This median band 276 JOURNAL OF THE LEPIDOPTERISTS SOCIETY posteriorly curves distad at the inner margin and runs along the margin almost to the blue bar, as in driophilus (in the others it intersects the inner margin at a high —often right—angle and does not run distad). Basad of the subterminal pale spots is a band of sprinkled blue scales, strongest in Cu:-2A but extending, weaker, costad to M, or Rs, essentially as in driophilus and ponceanus (in nominate aristodemus it is absent except for the segment in Cuz-2A, and in temenes it is usually so). Both specimens are smaller than any other aristodemus I have seen. Length of forewing. Male, 40.0 mm; female, 44.0 mm. Types. Ho.orypee, 2, Man of War Bay, Great Inagua Island, Bahamas, 4.x.1975, leg. Karen Bjorndal. C. M. Acc. 29104. Paratype, ¢, Calf Pond, northwestern Great Inagua, 18.v.1976, leg. Karen Bjorndal. C. M. Acc. 29104. Holotype and paratype, C. M. Ent. type series no. 690. Remarks. This subspecies apparently was derived from driophilus of the central Bahamas, although it has departed from it to an unusual and striking degree. The large rusty red patch on the hindwing above gives it a distinctive appearance, but the patch is foreshadowed by the small, obscure, rusty red spot that appears in M;-Cu; in some driophilus and ponceanus (see character (b) in the Remarks under driophilus above). Miss Bjorndal comments (in litt.): “From September to December [1975] and from May to August [1976] swallowtails were flying on Inagua. I was unable to distinguish which species [aristodemus or andraemon|. They were commonly seen in open scrub, dense scrub, coppice, coastal areas and [in the residential area of | Matthew Town.” In April 1977 on Little Inagua Island I repeatedly saw, but was unable to capture, a swallowtail in the short, narrow strip of low forest on the western coast, about a mile south of Northwest Point. Like Miss Bjorndal, I was unable to tell which of the two species it might have been. LITERATURE CITED Carpenter, G. D. H., & C. B. Lewis. 1943. A collection of Lepidoptera (Rho- palocera) from the Cayman Islands. Ann. Carnegie Mus. 29: 371-396, ill. Crencu, H. K. 1977. A list of the butterflies of Andros, Bahamas. Ann. Carnegie Mus. 46: 173-194, ill. Comstock, W. P. 1944. Insects of Porto Rico and the Virgin Islands, Rhopalocera or butterflies. Scient. Survey Porto Rico and the Virgin Islands (New York Acad, Sci.) 12(4): 421-622, ill. Journal of the Lepidopterists’ Society 32(4), 1978, 277-281 THE NAMES OF CERTAIN HOLARCTIC HAIRSTREAK GENERA (LYCAENIDAE) Harry K. CLENCH Carnegie Museum of Natural History, Pittsburgh, Pennsylvania 15213 ABSTRACT. The palearctic genus Strymonidia Tutt 1908 and the nearctic genus Euristr'ymon Clench 1961, both in current use, are both synonymized to Fixsenia Tutt 1907, on the basis of male genital structure. The currently used palearctic genus Nordmannia Tutt 1907, and many other generic names not in general use, must be synonymized to Satyrium Scudder 1876, on similar structural evidence. Both Fixsenia and Satyrium are shown to be holarctic, and both are unusually variable in external facies. All species known to belong to both genera are listed. The correct generic names to be applied to some of the palearctic hairstreaks has long been a problem. The species concerned are those listed by Higgins & Riley (1970: 235-238) and Higgins (1975: 109-112) for Europe under the genera Nordmannia and Strymonidia, together with numerous related species in the central and eastern Palearctic. The usage of Higgins & Riley, and Higgins, is representative of that current in the Palearctic literature. Study indicates, however, that neither of these generic names is tenable, and that assignment of the species (i.e., which species are congeneric with which) must be modified. Malicky (1969) has shown that the long unused generic name Fixsenia Tutt must be revived for the widespread species pruni Linnaeus in addi- tion to the (Asiatic) type species, herzi Fixsen. Some time ago I pointed out (Clench 1961: 212) that pruni was congeneric with certain North American hairstreaks to which I gave the new generic name Euristrymon. In consequence of Malicky’s discovery, Euristrymon must therefore fall to Fixsenia. Malicky also concluded that other Old World hairstreaks (he specif- ically cites spini, ilicis, w-album, and acaciae) are congeneric among themselves, and collectively differ from Fixsenia in the presence of a serrated ventral keel at the distal end of the penis in the male genitalia, as well as in certain larval and pupal characters. Malicky assigned all these species to the genus Strymonidia Tutt, a name which has long been in general use for many of them. The name Strymonidia, however, has as its generic type another little known species from Asia, thalia Leech, which Malicky was not able to study. I have seen no thalia either, but from published illustrations it seemed quite similar to herzi, the type of Fixsenia, and I was curious about its genitalic structure. In correspondence with Dr. Malicky I asked him about it. He was able to borrow a specimen and sent me a drawing of the genitalic 278 JOURNAL OF THE LEPIDOPTERISTS SOCIETY preparation he made. This showed clearly that thalia is indeed con- generic with herzi. Therefore the generic name Strymonidia is also a synonym of Fixsenia and cannot be applied to the species Malicky cites. The species Malicky included in Strymonidia, and additional Old World species as well, are themselves congeneric with a large number of Nearctic species now united in the genus Satyrium Scudder. This name, Satyrium, has priority over any other and must therefore be applied to these palearctic species. The discovery that the numerous species of Satyrium in both Old and New Worlds are all congeneric has created a large generic synonymy, particularly involving the Palearctic species. The apparent reason is that despite their close structural similarity they have diversified to a con- siderable degree in appearance. A number of species, among them the type of the genus, combine the un-hairstreak attributes of incon- spicuous or absent scent pad, absence of tails and tornal lobe, and essentially spotted patterns. They much resemble blues, and many were originally so described. Bethune-Baker (1892), in fact, made a special point of demonstrating that several of them were actually hairstreaks instead of blues. The external structural diversity of Satyrium and Fixsenia, in male scent pad, in wing shape, in tails, and in pattern, finds a remarkable parallel in the New World hairstreak genus Strymon Hiibner 1818 (i.e., as delimited in Clench 1961: 215ff). In Strymon an even more striking genitalic homogeneity is associated with similar variation in external structures. Apparently in both these genera extensive adaptive radiation has occurred, although the environmental significance of most of the affected external traits remains largely obscure. The male genitalia of the European species of both these genera are well figured by Higgins (1975). The formal synonymy and nomenclatorial data for both Fixsenia and Satyrium, together with their characters and known included species, are as follows: Fixsenia Tutt Fixsenia Tutt 1907, Nat. Hist. British Butts. 2: 142, type species by original designa- tion, Thecla herzi Fixsen 1887; Hemming 1967: 193; Malicky 1969: 38, 61. = Leechia Tutt 1907, l.c., type species by original designation, Thecla thalia Leech 1893; Hemming 1967: 249. Junior homonym of Leechia South 1901. See Strymonidia. - Strymonidia Tutt 1908, op. cit.: 483, replacement name for Leechia Tutt 1907, q.v., with the same type species; Hemming 1967: 419. New subjective synonym. = Euristrymon Clench 1961; 212, type species by original designation, Papilio favonius J. &. Smith 1797; Cowan 1970: 10; dos Passos 1970, J. Lepid. Soc. 24: 33. New subjective synonym. = Thecla, Nordmannia, etc., of authors, in part. VoLUME 32, NuMBER 4 279 The generic characters of Fixsenia are: hindwings usually tailed, tornal lobe usually present, if slight, but no tornal cleft. Male genitalia: uncus lobes low and transverse, the lateral border short; vinculum with dorsal part wide, but without posterior shoulder process, abruptly narrowing to the strap-like ventral part; anterior border of vinculum without corematal processes; saccus present but short, rarely if ever longer than its width at middle; valvae contiguous to more or less their middle, then divergent, the mesial edges not dentate, and with no terminal spine; penis apically upcurved and flared, wih two terminal cornuti of about equal diameter, but with no terminal ventral keel. In the Palearctic the last character, the absence of a ventral penial keel, is sufficient to separate Fixsenia from Satyrium, the only other genus with which it may be confused; but the remaining characters are necessary to discriminate it from other New World Strymonine genera. The known members are: Palearctic species: herzi Fixsen 1887; thalia Leech 1893; pruni Linnaeus 1758. Nearctic species: favonius J. E. Smith 1797; ontario Edwards 1868; polingi Barnes & Benjamin 1926. Satyrium Scudder = Argus Gerhard 1850, Versuch. Mon. europ. Schmett. (1):4, type species by mono- typy, Lycaena ledereri Boisduval 1848; Hemming 1967: 56. Junior homonym of Argus Bohadsch 1761. Satyrium Scudder 1876, Bull. Buffalo Soc. Nat. Sci. 3: 106, type species by original designation, Lycaena fuliginosa Edwards 1861; Comstock & Huntington 1958, J. New York Ent. Soc. 66: 116; Ziegler 1960, J. Lepid. Soc. 14: 20; Hemming 1967: 403; dos Passos 1970, op. cit.: 28. = Callipsyche Scudder 1876, l.c., type species by original designation, Thecla behrii Edwards 1870; Comstock & Huntington 1958, op. cit.: 105; Hemming 1967: 91. = Neolycaena de Niceville 1890, Butts. India, Burmah and Ceylon 3: 15, 64, type species by original designation, Lycaena sinensis Alphéraky 1881; Hemming 1967: 308. = Edwardsia Tutt 1907, l.c., type species by original designation, Papilio w-album Knoch 1782; Hemming 1967: 156. Junior homonym of Edwardsia Costa 1838. See Chattendenia. = Felderia Tutt 1907, l.c., type species by original designation, Thecla w-album Knoch 1782; Hemming 1967: 156. Junior homonym of Edwardsia Costa 1838. See Chattendenia. = Felderia Tutt 1907, l.c., type species by original designation, Thecla w-album Knoch var. eximia Fixsen 1887; Hemming 1967: 193. Junior homonym of Fel- deria Walsingham 1887. See Thecliolia. = Klugia Tutt 1907, l.c., type species by original designation, Papilio spini [Denis & Schiffermiiller] 1775; Hemming 1967: 242. Junior homonym of Klugia Robineau- Desvoidy 1863. See Tuttiola. = Kollaria Tutt 1907, l.c., type species by original designation, Thecla sassanides Kollar [1849]; Hemming 1967: 242. Junior homonym of Kollaria Pictet 1841. See Superflua. = Erschoffia Tutt 1907, l.c., type species by original designation, Thecla lunulata Erschoff 1874; Hemming 1967: 169. Junior homonym of Erschoffia Swinhoe 1900. See Pseudothecla. = Bakeria Tutt 1907, l.c., type species by original designation, Lycaena ledereri Bois- duval 1848; Hemming 1967: 72. Junior homonym of Bakeria Kieffer 1905, but never replaced. 280 JOURNAL OF THE LEPIDOPTERISTS SOCIETY — Nordmannia Tutt 1907, l.c., type species by original designation, Lycaena myrtale Klug 1834; Hemming 1967: 315. — Chattendenia Tutt 1908, op. cit.: 483; Hemming 1967: 109. Replacement name for Edwardsia Tutt 1907, q.v., with the same type species. = Thecliolia Strand 1910, Entomol. Rundsch. 27: 162; Hemming 1967: 439. Replace- ment name for Felderia Tutt 1907, q.v., with the same type species. = Tuttiola Strand 1910, l.c.; Hemming 1967: 451. Replacement name for Klugia Tutt 1907, g.v., with the same type species. = Superflua Strand 1910, l.c.; Hemming 1967: 420. Replacement name for Kollaria Tutt 1907, qg.v., with the same type species. = Pseudothecla Strand 1910, l.c.; Hemming 1967: 386. Replacement name for Erschoffia Tutt 1907, q.v., with the same type species. = Thecliola Waterhouse 1912, Index Zool. 2: 299; Hemming 1967: 439. Incorrect Subsequent Spelling of Thecliolia, q.v. = Necovatia Verity 1951, Rev. franc. Lépid., Suppl.: 183, type species by original designation, Papilio acaciae Fabricius 1787 [proposed as a subgenus of Stry- monidia Tutt 1908, vide supra]. Note: This generic name was overlooked by Hemming (1967), Cowan (1968, 1970), and the Zoological Record. I thank Lt. Col. J. N. Eliot for calling it to my attention. = Thecla, Strymon, Strymonidia, etc., of authors (in part). I have examined the male genitalia of the following species and find them all congeneric: fuliginosum, type species of Satyrium behrii, type species of Callipsyche w-album, type species of Edwardsia, Chattendenia spini, type species of Klugia, Tuttiola sassanides, type species of Kollaria, Superflua lunulatum, type species of Erschoffia, Pseudothecla ledereri, type species of Bakeria myrtale, type species of Nordmannia acaciae, type species of Necovatia I have not seen the species sinensis (type species of Neolycaena), but pretiosum Staudinger is extremely closely related, perhaps conspe- cific, and its genitalia, along with those of some other Satyrium, were figured by Bethune Baker (1892). His figures show that pretiosum, and hence most likely sinensis, is a Satyrium. I have not seen eximium Fixsen (type species of Felderia, Thecliolia), but from illustrations it seems to be a close relative of w-album and I conclude, therefore, that it is probably congeneric. The generic characters of Satyrium are: male genitalia: distal end of penis with ventral serrated keel; penis with two terminal cornuti, one of which is usually dentate; distally divergent valvae; tips of valvae without a mesial hair-like fringe. In the Palearctic the presence of a serrated penial keel will discrim- inate it from Fixsenia, or any other Strymonine. In the New World, however, several other genera share this keel, from which the additional characters will separate it. The known species belonging to Satyrium are as follows. I include only those whose genitalia I have examined either directly or in published illustrations, except for the species fol- VoLUME 32, NuMBER 4 281 lowed by “(?),” which are included provisionally, on the basis of ex- ternal facies alone. Palearctic species: eximium Fixsen 1887 (?); w-album Knoch 1782; spini [Denis & Schiffermiiller] 1775; latior Fixsen 1887; sassanides Kollar 1849: lunulatum Erschoff 1874; pretiosum Staudinger 1886; sinensis Alpheéraky 1881 (?); ledereri Boisduval 1848; myrtale Klug 1834; teng- stroemii Erschoff 1874; ilicis Esper 1779; acaciae Fabricius 1787. Nearctic species: fuliginosum Edwards 1861; behrii Edwards 1870; auretorum Boisduval 1852; saepium Boisduval 1852; tetra Edwards 1870 (= adenostomatis Hy. Edwards 1877); liparops Le Conte 1833; kingi Klots & Clench 1952; calanus Hiibner 1809; caryaevorum McDunnough 1942; edwardsii Scudder 1870; sylvinum Boisduval 1852; californicum Edwards 1862; acadicum Edwards 1862. It is worth noting that at one time it was believed (e.g., Riley 1958: 285) that myrtale was congeneric with Erora Scudder 1872. The latter genus, however, is representative of a wholly New World, primarily tropical, group with no palearctic members at all. This group differs considerably from Satyrium, not only in the absence of a serrated penial keel, but also in the high lateral margins of the uncus lobes, the single, always interior, cornutus, and in addition a terminal penial tooth that is either external and part of the shaft or else internal, on the vesica, and eversible. The vesica, moreover, is usually scobinate. LITERATURE CITED BETHUNE-BAKER, G. T. 1892. Notes on Lycaena (recte Thecla) rhymnus, teng- stroemii, and pretiosa. Trans. Entomol. Soc. London 1892: 27-31, ill. Ciencu, H. K. 1961. Tribe Theclini, pp. 177-220, in P. R. and A. H. Ehrlich, How to know the butterflies. W.C. Brown, Dubuque, Iowa. Cowan, C. F. 1968. Annotationes Rhopalocerologicae. Author, Berkhamsted, England. 20 pp. 1970. Annotationes Rhopalocerologicae 1970. Berkhamsted, England: author; 70 pp. Hemminc, F. 1967. The generic names of the butterflies and their type-species (Lepidoptera: Rhopalocera). Bull. Br. Mus. Nat. Hist. (Entomol.), Suppl. 9: 509 pp. Hiccrns, L. G. 1975. The classification of European butterflies. Collins, London. 320 pp., ill. & N. D. Rmey. 1970. A field guide to the butterflies of Britain and Europe. Collins, London. 380 pp., ill. Mauicxy, H. 1969. Uebersicht ueber praimaginalstadien, bionomie und O6kologie der mitteleuropaischen Lycaenidae (Lepidoptera). Mitt. Entomol. Ges. Basel (N.F.) 19: 25-91, ill. Ritty, N. D. 1958. The genera of holarctic Theclinae: A tentative revision. Proc. Tenth Int. Congress Entomol. 1: 281-288. Journal of the Lepidopterists’ Society 32(4), 1978, 282-288 OVER-WINTERING BEHAVIOR IN EUPHYDRYAS PHAETON (NYMPHALIDAE) M. DEANE BOWERS Department of Zoology, University of Massachusetts, Amherst, Massachusetts 01003 ABSTRACT. The behavior of the pre-diapause larvae of Euphydryas phaeton (Nymphalidae) associated with over-wintering is described. Observations are based on studies of five wild populations in central Massachusetts over a three-year period, as well as on larvae reared in the laboratory. The functional significance of this larval behavior is discussed. Most Lepidoptera over-winter as either an egg or pupa; some nym- phalid butterflies (e.g. the Mourning Cloak, Nymphalis antiopa L., and Compton’s Tortoiseshell, N. vau-album Boisduval and Leconte), and certain groups of moths [e.g. the Lithophanini (Schweitzer, 1977) | overwinter as adults. In only a relatively few cases do Lepidoptera spend the winter in the larval stage. The Viceroy, Limenitis archippus Cramer (Nymphalidae) for example, is multivoltine and may enter a facultative diapause in the third instar during late summer and fall (Hong and Platt, 1975). Unusually, however, the Baltimore Checker- spot, Euphydryas phaeton Drury (Nymphalidae), and other members of the genus Euphydryas, are univoltine and exhibit an obligatory dia- pause in the fourth instar. Although some descriptive work has been done on the development of the early, pre-diapause instars of E. phaeton, little has been reported of the actual behavior of the larvae during over-wintering. Observa- tions on several wild colonies of E. phaeton and on larvae reared. in the laboratory have revealed some interesting behavioral aspects of the over-wintering process in this species. METHODS Observations were made on five colonies of E. phaeton in Hampshire, Franklin, and Hamden Counties in central Massachusetts from 1974—_ 1977. The site of each colony was wet and marshy, typical habitat for E. phaeton and its primary foodplant, Turtlehead (Chelone glabra L.., Scrophulariaceae). The five sites differed in elevation, area, amount of Turtlehead present, and size of the E. phaeton population; but these differences were not factors in the present study. At each of the colonies, I determined the absolute numbers of egg masses, and (later) larval webs. At the three largest colonies, heights of the pre-hibernation webs above the ground were measured. All VoLUME 32, NuMBER 4 283 colonies were usually observed at least once a week. In addition I reared larvae from eggs in the laboratory at 22-25°C under continuous light. Developmental times for each instar, dates of larvae entering the pre-hibernation web and ceasing feeding, and time to the third molt were recorded for these larvae. OBSERVATIONS Euphydryas phaeton females usually oviposit two to three large masses of from 100-600 eggs each (Scudder, 1889; Bowers, pers. obs.) from late June through July in Massachusetts. The eggs hatch in about 20 days. When the larvae hatch they may construct a small web on the leaf next to the egg shell but move to the growing tip of the plant within 24 hours and begin web construction and feeding. This feeding web is extended down the length of a stalk of Turtlehead as the larvae devour the leaves. The larvae are gregarious and develop to the end of the third instar in approximately three weeks. About mid-August in central Massachusetts, the larvae stop feeding, and thicken and compact a section of the web (this date may vary widely, depending on the hatch date of a particular cohort of larvae, elevation, and current climatic conditions). Upon entering this “pre- hibernation web” larvae become quiescent and molt to the fourth instar in about five days (from 3-7 days in the laboratory) (Edwards, 1875; Scudder, 1889; Bowers, pers. obs.). Edwards (1875) reported that larvae enter this web about 15 July in West Virginia, but at higher elevations in Massachusetts this web entrance can occur as late as the second week in September. Previous authors (e.g. Scudder, 1889; Clark, 1927) have referred to this strengthened and compacted web as the “hibernating web.” How- ever, larvae do not spend the winter in this web at all, but leave it and move into the litter; this web is thus better referred to as the “pre-hibernation web,” which is the term used throughout this paper. This designation also serves to differentiate it from the feeding web which encloses plant parts on which the larvae are feeding. The pre-hibernation web may be constructed at the base of the Turtlehead stalk on which the larvae have been feeding, but is some- times found a short distance away. This web, as well as the feeding web constructed throughout the development of the pre-hibernation larvae, includes the stalk and leaves of Turtlehead as well as adjacent plants such as ferns, grasses and herbs (Figs. 1-3). The pre-hibernation webs are always constructed above ground level; most are found at a height of more than 50 cm. The mean height for 43 pre-hibernation 284 JOURNAL OF THE LEPIDOPTERISTS SOCIETY VoLUME 32, NUMBER 4 285 webs measured in the summer of 1976 was 73 cm (range: 30-127 cm; standard deviation: + 25.50). I have never observed a pre-hibernation web directly on the ground. Once the larvae enter the pre-hibernation web they do not feed at all. There may be some movement and occasionally a larva or two will leave and wander over the web surface. If the larvae are disturbed by movement of the web, they will move around a bit, but are usually inactive. An exception to this occurs when the web is damaged by storms or predators. In order to observe larval reactions to such damage, I broke open several webs. When the web is broken, the larvae begin moving around almost immediately and one or two larvae begin to crawl out of the web and over the surface. Other larvae soon follow. Within a day or two the web has been repaired and the larvae return to their inactive state. Thus, although the larvae are quiescent and not feeding, they can still react to stimuli such as disruption of the web. The larvae do not spend the winter inside this web as has been suggested by previous authors (e.g. Edwards, 1875; Scudder, 1889; Clark, 1927). Rather, about the end of October, the larvae move out of the web en masse to the base of a plant on which the pre-hibernation web was constructed. Here they form a large contiguous aggregation among the dead grass and litter on the ground. This movement occurred during the weeks 27 October-4 November 1975, 22 October-29 October 1976, and 20 October-27 October 1977. While in this aggregation, in- dividual larvae are active on warm days, moving from the primary mass as far as 25 cm. On cool, cloudy days or early in the morning, the larvae are usually found in a tight group, but individuals will quickly move around and out of the mass when disturbed. After approximately a week (from two days to two weeks; different for each aggregation of larvae), groups of about 10 to 100 individuals from this large mass move distances of 5-100 cm away and roll up leaves and bits of debris and fasten them with silk. It is here that they spend the winter. From 1974 to 1977 over 100 pre-hibernation webs were examined and in all webs the larvae moved into the litter. All the larvae remain near the pre-hibernation web and thus close to the Turtlehead stalk on which < Figs. 1-3. Development of the feeding web of E. phaeton on C. glabra: 1, beginning web formation at top of plant; 2, further development of web with more leaves enclosed; 3, extension of web to encompass several stalks of C. glabra and adjacent plants. 286 JOURNAL OF THE LEPIDOPTERISTS SOCIETY they originally fed. Because Turtlehead is perennial this behavior en- sures an adequate food supply for larvae emerging the following spring. On warm, sunny days during the winter, larvae may become active, and groups may shift position by a few cm, rolling up new leaves in which to spend the remainder of the winter. When the larvae are dug up from under the snow in January and brought to temperatures of 20-25°C, they quickly become active and crawl around the container in which they are confined. Usually they will not feed at this time. Clark (1927) brought hibernating larvae indoors in February and tried feeding them forced shoots of Lonicera japonica Thunb. (Caprifoliaceae ), an alternate foodplant for this species (Clark, 1927; Scudder, 1889). After wandering around for a week, they began to feed, but all eventually died before reaching pupation. If larvae are brought indoors in this manner, allowed to be active and then returned to outside conditions, they resume their resting state with no apparent ill effects, emerging from diapause in the normal manner in the spring. I attempted to break diapause in E. phaeton by subjecting several egg masses and their emerging larvae to 24 hours of light throughout development and hibernation. Although little is known about breaking diapause in univoltine insects, Beck (1968) suggests this method. These lab-reared larvae formed a pre-hibernation web in the same way as larvae in the field, yet were occasionally active. The larvae were given fresh food when plants seemed wilted or were consumed, and _ they fed occasionally through the end of October, with approximately 40% mortality. Between 25 October to 1 November there was a fourth molt (which does not occur in natural populations) that was synchronized within any single group of larvae but not among the groups. After this time, most of the larvae were inactive, and by mid-January all the larvae had died. Although attempts to break diapause in other insects with an obligatory diapause have been successful (e.g. Beck, 1968), this attempt was not. None of the larvae survived and none exceeded the size attained by hibernating larvae in the field. Thus diapause in this univoltine insect is truly obligatory and probably under genetic rather than photoperiodic control; the latter is probably the case in most multivoltine species (Beck, 1968; Hong and Platt, 1975). DISCUSSION Karly authors (e.g. Scudder (1889) and Edwards (1884)) believed that the larvae of E. phaeton spent the winter in the pre-hibernation web (which they called the “hibernation web”). Observation of these VoLUME 32, NUMBER 4 287 webs throughout the fall in Massachusetts shows that it would not be feasible for larvae to remain in the web: although the pre-hibernation web has many more layers of silk and is smaller and more compact than the feeding web, wind and rain begin to damage it by the end of October. Before winter has advanced very far there is little left of the web, and the larvae cannot repair the web during the winter. One of the harshest winter microclimates is just above the snow level where most of the pre-hibernation webs are found. Air temperature fluctuations here are much more dramatic than those at the soil surface (MacKinney, 1929; Mail, 1932) where larvae aggregate. Vegetation and litter reduce these temperature fluctuations, and snow provides further insulation (MacKinney, 1929; Mail, 1932). In Massachusetts there is snow cover for most of the winter and thus the larvae are quite well protected. Wind is also an important agent above the snow or ground level, but would have little impact on the larvae under snow or litter cover. Thus, regardless of snow cover, by moving out of the pre-hibernation web into the litter, larvae escape extreme tem- perature fluctuations and the desiccating effects of wind characteristic of winter conditions. The question, then, is why should larvae construct a pre-hibernation web at all; why not move into the litter in August when feeding stops? Perhaps the groups of hibernating larvae would be easy prey for ground predators such as spiders and beetles which are abundant at the end of the summer. By the end of October, however, most of these pred- ators are absent or inactive and the larvae could safely move into the litter. In summary, pre-hibernation larvae of E. phaeton exhibit three major behavioral sequences: first, construction of the feeding web during the first three instars; second, abandonment of this web and construction of a small, compact pre-hibernation web in which the larvae remain quiescent; third, departure from this web and movement of smaller groups into the litter where they form an over-wintering site by rolling up leaves and bits of debris and fastening them with silk. The last two sequences require appreciable expenditures of energy (for movement and silk-making) while no food is being eaten. This expenditure sug- gests that these behaviors are necessary to ensure larval survival over the fall and winter. ACKNOWLEDGMENTS I am very grateful to Dr. Theodore D. Sargent for his encouragement and editorial comments. This research was supported in part by a Sigma Xi Grant-in-Aid of Research. 288 JoURNAL OF THE LEPIDOPTERISTS SOCIETY LITERATURE CITED Beck, S. D. 1968. Insect photoperiodism. Academic Press, New York. 288 pp. Criark, A. H. 1927. Notes on the melitaeid butterfly Euphydryas phaeton (Drury ) with descriptions of a new subspecies and a new variety. Proc. U.S. Nat. Mus. Wash. 71, article 11, #2683. pp. 1-22. Criark, S. H. & A. P. Puarr. 1969. Influence of photoperiod on development and larval diapause in the Viceroy butterfly, Limenitis archippus. J. Insect Physiol. 15: 1951-1957. Epwarps, W. H. 1875. Notes on butterflies. Canad. Entomol. 7: 150-151. 1884. Description of the preparatory stages of Melitaea chalcedon, Bois., with some notes on larvae of M. phaeton. Papilio 4: 63-70. Hone, J. W. & A. P. Puarr. 1975. Critical photoperiod and day-length threshold differences between northern and southern populations of the butterfly, Limenitis archippus. J. Insect Physiol. 21: 1159-1165. MackKinney, A. L. 1929. Effects of forest litter on soil temperature and soil freez- ing in autumn and winter. Ecology 10: 312-321. Mart, G. A. 1932. Winter temperature gradients as a factor in insect survival. J. Econ. Entomol. 25: 1049-1053. ScuppER, S. 1889. The Butterflies of the Eastern United States and Canada. W. H. Wheeler, Cambridge, Mass. ScHweitzER, D. F. 1977. Life history strategies of the Lithophanini (Lepidoptera: Noctuidae, Cuculliinae), the winter moths. Ph.D. dissertation. Univ. of Mass., Amherst, Mass. 304 pp. Journal of the Lepidopterists’ Society 32(4), 1978, 289-303 INTER-SPECIFIC HYBRIDIZATION INVOLVING LIMENITIS ARCHIPPUS AND ITS CONGENERIC SPECIES (NYMPHALIDAE) AUSTIN P. PLATT Department of Biological Sciences, University of Maryland Baltimore County, 5401 Wilkens Avenue, Catonsville, Maryland 21228 GrorGE W. RAWSON 10405 Amherst Avenue, Silver Spring, Maryland 20902 GEORGE BALOGH 3607 No. 98th Street, Milwaukee, Wisconsin 53222 ABSTRACT. The occurrence of 43 natural hybrids involving Limenitis archippus and its congeneric species (L. arthemis-astyanax, L. lorquini, and L. weidemeyezrii ) is reviewed. Nine of these hybrid records are reported for the first time. Data based on laboratory crosses are given in order to document the purported wild hybrid specimens. Reasons underlying the observed natural hybridization are suggested and their evolutionary implications are discussed. Species of the North American genus Limenitis readily undergo inter- specific hybridization both in nature and in the laboratory (Edwards, 1882: Scudder, 1889; Field, 1904, 1914; Newcomb, 1907; Gunder, 1934; Remington, 1958, 1968; Platt, 1975). The occurrence of 22 natural hybrids and the laboratory documentation of them in crosses involving either 1) L. arthemis arthemis Drury or 2) L. arthemis astyanax Fabri- cius X L. archippus Cramer have been reviewed and discussed by Monroe (1953); Grey (1968); Shapiro and Biggs (1968); Platt and Greenfield (1971), and Greenfield and Platt (1974). Since then, John- son (1974) and Arbogast (1976) have reported two other wild-collected L. arthemis astyanax X L. archippus hybrid specimens. Likewise, the natural occurrence of four L. lorquini Boisduval x L. archippus hybrids (Gage, 1970; Perkins and Gage, 1970) and five L. weidemeyerii Ed- wards X L. archippus hybrids (Cross, 1936, 1937; Simpson and Pettus, 1976) have also been recorded. Lab-bred equivalents of these wild hybrids are shown in Fig. 1. In this paper we shall review these past records and will report records of nine other naturally occurring Limenitis hybrids involving L. archippus, a species broadly sympatric with other members of the genus. We will also present new information obtained from laboratory bo CO i) JouRNAL OF THE LEPIDOPTERISTS SOCIETY DORSAL VENTRAL lem Fig. 1. Representative lab-bred F, male hybrid specimens. 1) Form “arthechip- pus, brood 893, No. 17, May 7, 1977; 2) form “weidechippus,” brood 576, No. 7, Sept. 4, 1971; 3) lorquini 9 x archippus 6 (unnamed hybrid), brood 987, No. 1, Sept. 8, 1978. These, and all other lab-bred specimens, were reared on either Salix babylonica L. or Prunus serotina Ehrh. VoLUME 32, NuMBER 4 291 UNITED STATES $000", 10000" contours “EH. Frowschnar ond BLE. Woodion. Jr Fig. 2. Distribution map of 43 known wild, inter-specific F; hybrids involving cross-breeding with L. archippus collected from “prior to 1872” through 1976. All records for which the sex is known are males. These hybrids are widely distributed geographically. Touching symbols represent two (or more) specimens from the same locality. Most hybrids have been collected late in the season ( August-November ). Complete data for these hybrids are given in Tables 1, 1A, and 2. (Map reproduced with the permission of the Missouri Botanical Garden, St. Louis, Mo. ). crosses recently made by Platt, followed by a brief discussion of the relationships between L. archippus and its close relatives. Tables 1, [A and 2 summarize the collection data for all 43 records of wild hybrids involving L. archippus and its congeneric species. The geographic distributions of the various hybrid forms have been plotted in Fig. 2. The new hybrid locality records given in Table 1 are those from Maine (hyb. form “arthechippus” Scudder), Wisconsin, Tlinois, Michigan, New Jersey, Virginia, and Florida (hyb. form “rubidus” Strecker). The specimens from Idaho, Illinois, Michigan, New Jersey, Virginia, and Florida probably represent state records for these hybrids. The Florida record represents the first report of a natural hybrid between L. a. astyanax Fabricius and the sub-species L. archippus floridensis Strecker. The live specimen was observed circling a shrub willow along the edge of Mud Lake by G.W.R. while he was lunching JOURNAL OF THE LEPIDOPTERISTS SOCIETY (LO6T ) quroomen []94% se “S[oqe] SBLTT UOrs -Ud00V pUP ‘[[OD soureg sey Quo ‘sfeqey ‘snyy UATyoorg reaq YI0q *7[09 “WN’S'D "UIUOO *si0od “Uasyel N‘O'W "UIUOD ‘sIod ‘UOAIT, ‘—D “AY SSOP “G ysoreg ysoreg “5 (STL) Bese "uuLoO ‘siod “7 ‘q ‘[ sa}ou pue 010g ‘"T X xea[dui0o xpupfiysp-stmaywd sijvuaul)? 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Y[OFION sosnyoesse jy ayouve[ puvla1ounss AA erueAlAsuue dg a syslog elueAlAsuueg [PI0L ouIeg \sey Aueqry YIOX MIN uapAIqd suryduio J, yIOX MAN yepnog — eqoPUR]Y pve s[V ailysoyy aysduepyT MON svoyunpessed JOosqousg JUIRTY [eert}UOJ “OTA ulseg Aensnevoyeyy — oaqand drysuao J, AVUnODy 93VYS IO A}I[VIO'T IO QOUTAOIg (‘s1oyyne oy} JO uOrsstutiod Aq poonpordoay) ‘a[qu[rVAv JOU SPAX WOHRULLOFUL FEY} O}BOIPUL SOYSeCT “OST tt STLOL ‘playueerg pue Yq wWoIF polzrpour ‘spriqdy xvuDAZsp-sray PD siuaUYT WYSUvI-p[IM A[snorAdid QZ JO sploovdy “WT ATAV$ 295 VcLUME 32, NuMBER 4 "(‘uruto9 ‘sod ‘aspury "H ‘A “Id ‘6S6T ‘S10IM) “H'N’W'V 94} Ur are suoutIoods om} oY J, ‘sesueyIy “OD UoTUy) ‘opeiog [y Ww evuULTY “YA ‘SAY AQ pivdureq v Ul LCE ‘9% ‘SNY v{Ndoo UT poazdaT[00 o10M snddiyouw "7T XK xpuvfysp ‘p'T & Ve ‘eITq Aq Ajp}UID -9l 9pvUl U99q BALY{ Sasso1d [VOOAdTOVI JBYIQ ‘“ssoId P snuay.iW “DT K J snddiyoiw “7 ue woi (PIGL) Ploy Aq Posvor o19M snddwoay.p seul JYSIA ; *‘SoTPUL UI9q DAVY 9}eP 0} PoszaT[OO susuTIOeds [[e ‘UMOU SI Sv IezZOSUT , GL [F701 II 140d “4 “f WI yAo0q a{ PT OLGI “GS Ides oruoyuy ues Iexog Sexo], uosuyol “y PPusolg “VA Py Z C9G6I “Pp ‘Ides SG a o® Anne Eee TeN ospuly *H “A 1° HNWY ?T SE6T “T 3des AUD asoy ryseng sesue yy yuurs *[ (€G6T ) 9or1u0/y Q SIOIIOS “Y Omi SPBI “Ides O][FAsmo'T UOSIoFfo[ Ayonjquay (FOBT) PIP od qoorf OT = puvysy suoT — ylOX MON so}0u UOI}9T[OO [X9G H 2}eq drysumoy, A}UNOD) 2381S pure 901n0sg 10/2} 10}09][0D ‘ON IO A}T[eoO'T IO 9OUTAOLY ‘ponunuoy ‘WI AIdVy, 296 JouRNAL OF THE LEPIDOPTERISTS SOCIETY DORSAL VENTRAL | lcm Fig. 3. 1) Wild-caught and 2) lab-bred male specimens of L. arthemis astyanax x L. archippus floridensis F1 hybrids (form “rubidus”). Data for the lab-bred speci- men are brood 986, No. 1, Sept. 19, 1977 (Data for wild-caught specimen are given in Table 1). with members of the Florida Audubon Society in the northeastern portion of the Ocala National Forest. He recognized the specimen as an “off-color” Limenitis and investigated it further. The insect then settled on the willow shrub. Since he had no net, George made a “desperate strike” at it with his cap, knocking the butterfly to the ground and collecting it. The specimen is illustrated in Fig. 3, along with a similar, single specimen recently reared by A.P.P. in the laboratory from a hand-paired cross between a Maryland astyanax ? X a floridensis é (from a stock obtained just east of the Everglades [near Home- VoLUME 32, NuMBER 4 297 DORSAL VENTRAL lcm Fig. 4. Small, weak, and faded lab-bred F, female hybrid specimens, dorsal and ventral views. 1) Form “weidechippus,” brood 576, No. 1, Aug. 28, 1971; 2) form “rubidus,’ brood 648, No. 4, July 26, 1973. stead] in southern Florida). Astyanax evidently is relatively uncommon in central Florida. Near his home at New Smyrna Beach (Volusia Co.), G.W.R. has seen only two specimens of astyanax during the past 20 years. Also, F. Rutkowski (pers. comm.) recently collected a ¢ astyanax 1.5 miles north of Shamrock (Dixie Co.) in Florida, and he mentioned other records from as far south as Dade Co. (Kimball, 1965). Possibly such scarcity of one (or both) species accounts in part for such inter- specific hybridization (Simpson and Pettus, loc. cit.). So far as is known, all of the wild hybrids thus far collected have been males, although five broods containing small, faded, weak (and often malformed) F, females (Fig. 4) were reared at UMBC in June and July, 1973 by crossing inter-specific strains having different geo- graphic origins (Maryland L. astyanax 22 X Vermont L. archippus DORSAL JouRNAL OF THE LEPIDOPTERISTS SOCIETY VENTRAL VoLUME 32, NUMBER 4 299 é 4). Among these broods, there were a total of 39 (27%) F, females among 143 hybrid “rubidus” progeny. Two of the larger broods, in- volving sibling female astyanax and the same male archippus parent, yielded 1:1 sex ratios. Earlier crosses reported by Platt (1975) showed that such inter- specific crosses, in which strains from the same (or closely adjacent) northeastern geographic origins were crossed, gave rise to complete adult heterogametic (female) inviability. However, robust females of the “arthechippus” and “rubidus” hybrid phenotypes (Fig. 5) also have been obtained by backcrossing F, hybrid males to females of the three parental forms (arthemis, astyanax, and archippus, respectively). Such backcrosses often have low viability, but sometimes yield relatively large numbers of progeny (Platt, loc. cit.). Although fertile crosses have been obtained in all possible reciprocal combinations, crosses using L. arthemis, astyanax, lorquini, or weidemeyerii 22 X L. archippus 3 4 have been the easiest to effect in the laboratory using hand-pairing. The fact that such a pairing also has been seen in the wild (Klots, 1959; Table 1A) suggests that these inter-specific hybrid crosses may occur most often in this direction in nature, as well. Table 1 indicates that eight of the nine previously unreported arthemis- astyanax X archippus hybrids, like most of those reported earlier, were collected during the late summer and fall months (August-November). Only one (from New Jersey) was collected in June, a time suggesting that it most likely arose from an over-wintering larva. Thus, these new records, as well as the previous ones, support the contention that the ecological and behavioral barriers normally preventing inter-specific hybridization in Limenitis tend to break down later in the season (Green- field and Platt, loc. cit.). This break down may well be correlated with the onset of facultative larval diapause in Limenitis which occurs during the third instar. Thus, it seems as if those individuals most often selecting mates of the wrong species are the very ones which seem to be “genetically mal-adapted” to their environment (that is, they are the ones which are not diapausing at that time of year when they are <& Fig. 5. Representative robust, lab-bred hybrid-type backcross females. 1) L. arthemis 2 xX F, hybrid “arthechippus” 8 (“arthechippus-like” morph), brood 63c, No. 4, July 30, 1968; 2) L. archippus 2 xX F; hybrid “arthechippus” ¢ (“arthe- chippus-like” morph), brood 915, No. 45, June 13, 1977; 3) L. arthemis 2 xX F; hybrid “rubidus’ 8 (“proserpina-like” morph), brood 95B, No. 4, Dec. 26, 1968; 4) L. archippus 2 & F; hybrid “rubidus” ¢ (“rubidus-like” morph), brood 757, No. 10, Sept. 15, 1975. Such backcross females also occur in parent-type morphs (see Pia LO): JoURNAL OF THE LEPIDOPTERISTS SOCIETY 300 ‘T]OO ‘sted ‘Y oney[g suoje LECT *9ECT “S801D SaAAIMOS ‘GC'Q OT QE6T Bor10 IOAUNG ‘IU UdATS JON OpR10[OD sqis poutnso.id ‘Yysorz ‘U99S T ‘WYSNeo T (9L6T ) sn}jog pue uosduiis uosdullg "5 ‘YP 2G C/6T ‘6 eunf SuT[[OD HOT IOUWILIO'T OpeIO[OD (9L6T ) ‘snyY “TOULOWA “SO sn}jog pue uosduiis MD dD TI PEST GZ ‘sny SuT][OD HO IOUILIO'T OpeI0[OD (ssoIn _snddiysapiam,,) snddryouw “JT X wwahawapian “Ty ‘[[Oo ‘sted wed Yeo ‘UIUUOD “sted ‘ase ‘Weysulssen Ys PT TLOI (LPO Ayou'y ‘Iu epy oyep] ee “sted A, PUelPM “UULOD ‘sied ‘ase ‘AIOMOSUOW “Of LT TL6OI F 3des ‘Iu “‘puelyorny uojueg UOSUTYSE AA uigeesee Ry, PURTYOY "UIULOD “sted ‘ase ‘AIQUIOSUOW ‘Of PPT 696T ‘9 A4[nL ‘Iu ‘puelyony uojwueg UOSUIYSe AA OLET 98e5 pue Mgousi2d cA, PULTE SUIYIOT ‘OLE “sed SHED) VA "Bl =O CO6L ‘F ADL ‘Iu “puepyory uojwueg UO}SUTYSE AA, (wIoF poweuuN) snadiyoiw “JT X tumb.o) “TJ $9}0U UOT}O9T[OO xoS 9 31eq drysuMo y, Aqunoy 9381S pure 9010s 10/2 10}99T[OD ‘ON 10 A}ITV00'T ‘snddiyoiv “JT X (ApaAoedse. ‘nlahawapian pue twinbio)) sijuawty JO soloeds pepueq Ulo}soA\ OA\} OU} SUIA[OAUL SpliqAYy pyIA\ oUIu JO SpIOO9Y “FZ AIAVY;, VoLUME 32, NuMBER 4 301 supposed to be doing so). Such hybridization, perhaps, represents a “last chance” effort to reproduce. The similar rare natural hybrids reported between both of the western banded Limenitis (L. lorquini and L. weidemeyerii) and L. archippus are listed in Table 2. The eight known dates of siting or capture sug- gest once again that either 1) the hybrids emerged early enough in the year (June and July dates) so that it is a relative certainty that the larvae from which they arose over-wintered in hibernacula, or 2) the specimens were collected in the late summer or fall months. Gage’s (pers. comm.) collection dates for the small “hybrid swarm” near Rich- land, Washington (Table 2) suggest that the four lorquini x archippus hybrids, in fact, represent progeny from at least three different matings. The same may be said for the four “weidechippus” records from Colo- rado, as well. All of these wild western hybrids are males, and they closely resemble hybrid from “arthechippus” in possessing a_ partial postmedial white band dorsally (Fig. 1). Two crosses between Colo- rado weidemeyerii 22 X Massachusetts archippus 6 ¢ have been made by laboratory hand-pairings to date, yielding 22 ¢¢ and nine 2° (seven of the latter being malformed). All of these F,’s, although showing some phenotypic variability, are referrable to hyb. form “weide- chippus’ (Platt, unpub. data). During the past summer two crosses between Oregon L. lorquini °° X Maryland L. archippus ¢é were carried out. All 16 F, progeny were males. An additional cross involving an F, hybrid, arthemis-lorquini 2 (Massachusetts = Oregon stocks, respectively) x Maryland archippus ¢, yielded 21 male hybrid-like progeny. Thus, L. archippus, which is broadly sympatric with its congeners, will occasionally hybridize with all of the other allopatric species of Limenitis in nature. However, such crosses evidently are rare, leading to the supposition that morphological, behavioral, visual, and possibly pheromonal cues, as well as habitat isolating mechanisms, normally operate to prevent such inter-specific hybridization. These barriers against gene exchange between the viceroy and its close relatives some- times tend to break down, usually when one or both species are rare, and often toward the end of the breeding season, at times when the majority of developing Limenitis larvae are entering diapause. Laboratory data show that inter-specific strains having different geo- graphic origins may be genetically more compatible than similar strains from the same locality, as judged by either the presence or absence of adult females in the F, generation. Thus, genetic incompatibility be- tween the viceroy and its congeneric species is viewed as being of local 302 JOURNAL OF THE LEPIDOPTERISTS SOCIETY origin, suggesting that archippus may have arisen from a banded an- cestral species by the process of sympatric speciation. (The senior author would appreciate hearing from members of the Society who may have knowledge of other records of wild Limenitis hybrids. ) ACKNOWLEDGMENTS We are grateful to those individuals who provided us with informa- tion regarding the collection of hybrid specimens as noted in Table 1. We thank Dr. A. Maizels for sending the floridensis stock and Mr. P. J. Kean for making the inter-specific hand-pairing involving this strain. Mr. S. J. Harrison and Mr. T. Williams have assisted with the insect rearing and preservation. We thank Dr. W. D. Field, Dr. D. C. Ferguson of the U.S.N.M., and Dr. K. Bagdonas of the University of Wyoming for the information relating to hyb. form “weidechippus’. Mr. E. Gage kindly provided data and color photos of the lorquini xX archippus hybrids. LITERATURE CITED Arsocast, R. T. 1976. Capture of a hybrid Limenitis arthemis astyanax x L. archippus (Nymphalidae) in southern Georgia. J. Lepid. Soc. 30: 4. Cross, F.C. 1936. (Notitle). Hobbies 41: 112. 1937. Butterflies of Colorado. Proc. Colo. Mus. Nat. Hist. 16: 3-28. Epwarps, W. H. 1882. Descriptions of new species of butterflies found in the United States. Papilio 2: 45-49. Frecp, W. L. W. 1904. Problems in the genus Basilarchia. Psyche 11: 1-6. . 1914. Hybrid butterflies of the genus Basilarchia. Psyche 21: 115-117. Gace, E. V. 1970. A record of a naturally occurring Limenitis hybrid (Nymphal- idae). J. Lepid. Soc. 24; 270. GREENFIELD, J. C., Jn. & A. P. PLatr. 1974. Report of the capture of an additional hybrid between Limenitis arthemis astyanax and L. archippus (Nymphalidae). J. Lepid. Soc. 28: 72—75. Grey, L. P. 1968. (No title). In No. Amer. Ann. Summary, News Lepid. Soc., No. a, p, 29) Gunver, J. D. 1934. A check list revision of the genus Basilarchia Scud. (Lepid.: Rhopalocera ). Canad. Entomol. 66: 39-48. Jounson, K. 1974. An aberrant interspecific hybrid of Limenitis (Nymphalidae ) from Wisconsin. J. Lepid. Soc. 28: 163-165. KimBati, C. P. 1965. The Lepidoptera of Florida, in Arthropods of Florida and neighboring land areas. Fla. Dept. Agric., Gainesville, Fla. Vol. 1: 363 pp. Kiors, A. B. 1959. A mixed mating of two species of Limenitis Fabricius (Lepidop- tera, Nymphalidae). J. N.Y. Entomol. Soc. 67: 20. Monroe, B. L. 1953. Research Associate, Florida State Collection of Arthropods, Division of Plant Indusiry, Florida Department of Agriculture and Consumer Services. Journal of the Lepidopterists’ Society 32(4), 1978, 309 A PARTIALLY ALBINIC ABERRATION OF PHYCIODES THAROS (NYMPHALIDAE ) On 25 July 1977, I took a partially albinic male aberration of Phyciodes tharos Drury (Fig. 1) in Upper Tyrone Township, Fayette Co., Pennsylvania, at an eleva- tion of 1100 ft (335 m). All the normally tawny or brown coloration characteristic of this species was replaced by an extremely pale, orange-tinged cream color. The black markings were not affected. I know of no similar specimens. The aberration has been deposited in the collection of the Peabody Museum of Natural History, Yale University, New Haven, Connecticut. Cuar.es G. Ottver, R. D. 1, Box 78, Scottdale, Pennsylvania 15683. Fig. 1. Phyciodes tharos Drury: A., B. pale (partially albinic) male aberration, dorsal and ventral sides; C., D. typical male from same locality, dorsal and ventral. Journal of the Lepidopterists’ Society 32(4), 1978, 310 A RECORD OF AGRIAS AMYDON (NYMPHALIDAE) FROM COSTA RICA Two female specimens of Agrias amydon Hew. subspecies (?) were taken in Parque Santa Rosa, Guanacaste Province, Costa Rica. This is the first substantiated record of this genus from Costa Rica. Agrias zenodorus smalli Miller & Nicolay has been anticipated from Costa Rica and there exists in the literature a supposed sight record from Turrialba, Cartago Province, Costa Rica (Miller & Nicolay 1971. Bull. Allyn Mus. (1):1-5). It was totally unexpected to find Agrias in a habitat like Guanacaste where there is a strongly marked dry season in contrast to the usual wet forest habitats in other countries where the genus Agrias occurs. I have compared the specimens with A. amydon in the U.S. National Museum, the Carnegie Museum, and the Allyn Museum of Entomology and have found them to be differently marked on the dorsal surface of both fore and hind wings than the comparative material. Description of this A. Amydon subspecies is impossible on the basis of only two female specimens and must await more material. On the basis of the specimens examined in the above mentioned museums and in the literature available to me, A. amydon is recorded only as far north as Colombia. These specimens represent a considerable range extension and a new record for Costa Rica. One specimen has been placed in the collection of the Allyn Museum of Entomology, Sarasota, Florida. Pump J. DeVries, Museo Nacional de Costa Rica, Department de Historia Natural, Apartado 749, San Jose, Costa Rica. Fig. 1. Agrias amydon Hew., subspecies (?), female: dorsal (left), ventral (right). Wingspan = 6 mm. Journal of the Lepidopterists’ Society 32(4), 1978, 310-311 ADDITIONAL FUNCTION OF THE LEPIDOPTERAN PROBOSCIS The most unique part of the lepidopteran body form is the proboscis. This pre- hensile tube functions mainly as a suction device for nutrient procurement. A second, but minor, function has been observed in a variety of unrelated butterflies and moths VoLUME 32, NUMBER 4 311 —fluid “pumping.” Reports describe these individuals as having an extended pro- boscis at a moisture source and simultaneously voiding fluid from the anus (Clench 1957, Lepid. News 11:18-21; Roever 1964, J. Res. Lepid. 3:103-120; Hessel 1966, J. Lepid. Soc. 20:242; Jobe 1977, Entomol. Gaz. 28:8). Interpretations of this be- havior have been speculative generalizations. Personal observations have revealed a third—also minor—function of the lepidopteran proboscis. A female Atalopedes campestris (Boisduval) (Hesperiidae) was found floating with fluttering wings on the water surface of a wading pool in a residential backyard in Austin, Travis Co., Texas. The skipper did not appear to be severely injured, probably due to a relatively short period of partial submergence. Nevertheless, the scales were quite wet. Subsequent observation of this individual revealed that it was rapidly probing the scaly covering of the anterior part of its body by continued manipulation of its proboscis. Most of the probing involved ventral and lateral scales of the thorax. A definite color-lightening effect was observed. This change in coloration would indi- cate a reduction in the amount of water which had a plastering effect upon the scales. Decrease in amounts of water present among the scales could result from two effects of probing by the proboscis. Simple separation of adjacent matted scales by mechanical movement of the proboscis would increase evaporation rates because of an increase in scale surface area exposed to air. Also, water could be and probably was being physically removed by suction via the proboscis. The primary behavioral regime enlisted in this task would involve an activity related to previous function—suction action of the proboscis. Therefore, physical removal of water should be regarded as the initial function upon which natural selection acted. Enhancement of evaporation rate by scale separation initially was an ancillary result of this behavior. Such enhancement could be further increased by selection favoring rapid random movements as opposed to sucking up water at one spot and relying on capillary pressure to maintain a continuous film of water. Although this behavior was observed to function to remove water from scales following partial immersion in water, it could also function to remove moisture from rain (dew?) on butterflies roosting in exposed sites. Removal of water may be important for several reasons. Flying ability may be reduced if water is present in sufficient amounts to appreciably increase the weight of the insect. A matted water/ scale film present on the body surface may interfere with spiracular inhalation or favor development of pathogenic populations. Even if most air were inhaled via abdominal spiracles, removal of thoracic surface water may decrease abdominal and wing surface water via capillary action. RayMonp W. Neck, Pesquezo Museum of Natural History, 6803 Esther, Austin, Texas 78752. JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY INDEX TO VOLUME 32 (New names in boldface ) aberrations, 221, 309 Abraxas grossularia, 233 Achalarus lyciades, 201 Achlyodes thrasa tamenund, 112 Acraea encedon, 233 A. lapitha, 266 A. subhyalina, 268 Acraeidae, 233 Acraeinae, 261 Actinote calderoni, 261, 264 A. lapitha zilchi, 261, 266 A. stratonice oaxaea, 261 A. subhyalina, 268 A. thalia, 261 Adelpha, 171 Aeaea venifica, 235 Aganisthos, 171 Agathymus, 107 Ageronia, 169 Agonopterix dimorphaella, 235 Agraulis vanillae, 178, 211 Agrias, 169, 310 A. amydon, 310 A. zenodorus smalli, 310 Aiello, Annette, 135 Amaryssus astenous, 116 A. minos, 116 Amblyscirtes aesculapius, 28 A. samoset, 28, 200 A. vialis, 200 Amphichlora, 169 Amplypterus gannascus, 56 Anaea andria andria, 200, 211 A. eurypile, 172 A. morvus, 172 A. pithyusa, 172 Anartia amathea, 135 A. fatima, 135, 172 A. jatrophae, 172 Ancyloxypha numitor, 146, 201 Anthocharis midea, 202 Antithesia montana, 251 A. sara, 226 Apanteles, 109, 128 Apatura, 171 Apaturidae, 212 Apodemia palmeri, 209 Archeprepona, 171 Argynnis paphia, 63, 146 Asterocampa celtis celtis, 28, 203 A. clyton clyton, 203 A. leilia, 209 Atalopedes campestris, 200, 311 Atopothoures ovaliger, 55 Atrytone delaware delaware, 200 Atrytonopsis hiana hiana, 200 Auctor hujus operis, 116 Austin, George T., 207 Autochton cellus, 28, 201 Baker, James H., 241 Balogh, George, 289 Bandera, 105 Barcita, 244 Bartlett-Calvert, Wm., 251 Batrachedra illusor, 235 Battus philenor, 28, 157, 201, 209 behavior, 38, 56, 233, 282 Berenbaum, May, 303 Biblis, 171 B. hyperia aganisa, 113 bionomics, 107 Blanchard, André, 55, 103 Blau, William S., 138 Blood-spot skipper, 107 Boloria bellona, 28, 204 B. selene myrina, 204 book reviews, 63, 144, 239 Botys furnaclis, 130 Bowers, Deane, 140, 282 Braconidae, 128 Brephidium exilis, 211 Brewer, J. Wayne, 123 Butler, Linda, 198 Calephelis borealis, 202 C. muticum, 142, 200 Calpodes ethlius, 108 Calydna lusca, 46 C. virginiensis virginiensis, 202 Callaghan, Curtis j., 37 Callizona, 169 Callophrys, 3, 143 C. augustinus croesiodes, 202 C. barryi, 4 C. byrnei, 4 C. gryneus, 3 C. g. castalis, 7 C. g. gryneus, 6, 202 C. g. sweadneri, 6 C. henrici henrici, 24, 28, 202 C. hesseli, 3 C. irus irus, 200 Grlokiees C. nelsoni, 3 C. niphon niphon, 202 VoLUME 32, NUMBER 4 C. polios polios, 202 @.4siva, 3 C. s. juniperaria, 6, 14 C. s. mansfieldi, 14 C. turkingtoni, 3 Callosamia angulifera, 191 C. promethea, 59, 191, 233 C. securifera, 194 Calpodes ethlius, 108, 200 Calycopis cecrops, 202, 236 Calydna lusca, 46 Castniidae, 139 Catocala, 63 . grynea, 222 . habilis, 222 . ilia, 306 . marmorata, 306 . micronympha, 221 . muliercula, 222 . unijuga, 222 Catagramma, 171 Catonephele, 170 C. numilia, 172 C. nyctimus, 172 Catonephelini, 160 Celastrina argiolus, 34 C. a. bakeri, 240 C. ebenina, 20, 203 C. pseudargiolus, 20, 28, 203, 226 Cercyonis meadii, 146 C. pegala pegala, 204 Charaxinae, 170 Charaxini, 160 Chedra inquisitor, 235 Chelonia midas, 273 Chew, Frances S., 129, 159 Chionodes asema, 235 Chlosyne gorgone carlota, 203 C. harrisi harrisi, 200 C. janais, 59 C. lacinia crocale, 211 C. nycteis nycteis, 203 Chrysocharis, 128 Glark J. iF. Gates, 251 Clench, Harry K., 273, 277 climatic regimes, 111 Coea acheronta, 170 Cogia hippalus, 209 Coleotechnites, 118 . edulicola, 118, 123 . milleri, 123 . moreonella, 120 . ponderosae, 118 . ponderosana, 122 . starki, 123 Colias eurytheme, 28, 202, 209, 228 aS SiGie@ PD} GQ) ©} >) C. cesonia, 113, 200, 209 CC intenor 43 202, C. philodice, 28, 202 collection, 235 Colobura dirce, 160 Coloburini, 160, 170 Coloradia, 97 Copacodes aurantiaca, 211 courtship, 41 Cosmopterigidae, 235 Covell, Charles V., Jr., 221 Cryptotainia canadensis, 237, 304 Cyclogramma, 171 Cydimon poeyi, 307 Cyllopsis dospassosi, 75 C. gemma freemani, 84 C. nayarit, 84 C. pephredo, 84 C. pseudopephredo, 84 C. wellingi, 75 Cynthia, 94 C. annabella, 95 C. cardui, 95, 203 C. carye, 95 C. virginiensis, 95, 203 Danaidae, 170, 204 Danainae, 116 Danaus gilippus, 209 D. plexippus, 204, 209 Demons, IRs ling lal 2s) descriptions, 75, 175, 261 DeVries, Philip J., 310 Diaethria, 171 Dichomeris glenni, 235 Dicladocerus, 128 Dimock, Thomas E., 88 Drees, Bastiaan M., 198 Dryas julia moderata, 111 Dymasia chara, 209 Dynamine dyonis, 111 Eberlie, W. J. D., 226 Electrostrymon angelia angelia, 139 E. a. boyeri, 140 E. endymion, 139 Endothenia microptera, 235 Epargyreus clarus clarus, 28, 201 Epiblema naomi, 235 Epimetes, 116 Epinotia, 256 E. atristriga, 235 Epiphile, 170 Erora, 279 E. laeta, 35, 140, 202 Erunniss 24354 6 305 E. baptisiae, 201 313 314 JOURNAL OF THE LEPIDOPTERISTS SOCIETY E. brizo brizo, 201 F. thalia, 279 E. b. lacustra, 28, 61 F. w-album, 277 E. funeralis, 209 foodplant, 3, 20, 86, 236, 303 E. horatius, 201 Gelechiidae, 118, 123, 235 E. icelus, 201 genitalia, 7, 304 E. juvenalis, 25, 28, 201 genetic factor, 231 E. lucilius lucilius, 201 geographic distribution, 3, 259, 291 E. martialis, 201 Geometridae, 224 E. pacuvius, 305 Gilbert, Lawrence E., 54 E. persius, 200, 305 Glaucopsyche lygdamus nittanyensis, 203 E. zarucco zarucco, 200 Godfrey, George L., 235 Euchloe olympia olympia, 202 Goya ovaliger, 55 Eucosma uta, 235 G. stictella, bo Eucosminae, 256 Graphium marcellus, 201 Eueides cleobaea zorcaon, 113 Gynaecia, 169 Eulophidae, 128 Gynaeciidae, 170 Eulythis mellinata, 224 Gynaeciini, 160 Eunica, 169 gynandromorph, 138 Euphaedra ceres, 116 Gypsonoma, 256 Euphydryas anicia bakeri, 240 habitat, 20 E. chalcedona, 226 habitss 1235 sie E. phaeton, 204, 282 hairstreak, 277 Euphyes bimacula, 200 Hamadryas, 171 E. dion dion, 200 H. amphinome, 172 E. conspicua conspicua, 200 H. februa, 172 E. vestris metacomet, 200 H. guatemalena, 172 Euptoieta, 169 Hamadryadini, 160 E. claudia, 204, 209 Hardwick, D. F., 49 Euptychia areolata areolata, 200 Harkenclenus titus mopsus, 202 E. cymela cymela, 28, 904 Heliconiidae, Jia 170, 185, 212 E. gemma gemma, 204 Heliconius charitonius, 172 E. hermes, 28 H. c. vasquezae, 111 E. h. sosybius, 204 H. erato petiverana, 44 E. rubricata, 146 H. ethilla, 43 Euptychiini, 75 H. petiveranus, 172 Eurema, 39 Heliothis fastidiosa, 49 E. lisa, 178, 202 H- glomosa) BO E. nicippee, 202, 209 H. imperspicua, 51 Euristrymon, 277 H. inclara, 51 E. ontario ontario, 200 H. lanul, 51 Everes amyntula, 209 He Ree a2 Y , E. comyntas, 34, 146, 203 EGE, oe ay, Oe tee Lares H. rubiginosa, 52 Exartema comandranum, 235 H. siren, 53 Felderia eximium, 280 H. spectanda, 53 Feniseca tarquinius tarquinius, 202 H. sulmala, 53 Fiji, 130 Hermiargus ceraunus, 209 Finke, Mark, 236 H. isola, 209 lixsenia acaciae, 277 Hemileuca chinatiensis, 97 F. favonius, 279 H. diana, 99 F. herzi, 277, 279 H. electra, 99 F. ilicis, Bag H. griffini, 97 F’. ontario, 279 H. grotei, 99 F. pruni, 277, 279 H. hera hera, 59, 99 F. polingi, 279 Hemimene, 258 F. spini, 277 Hesperia, 146 VOLUME 32, NUMBER 4 . attalus attalus, 200 . leonardus, 201 . metea, 200 . Sassacus sassacus, 201 . columbia, 61 me Hesperiidae, 107, 112, 143, 199, 225, 304, 310 Hesperiinae, 185, 189 Historis odius, 160 Hodges, Ronald W., 118 Homoptera aemona, 242, 243 H. discisigna, 243 H. peruncta, 242 H. ustipennis, 242 Hyalophora, 191 H. cecropia, 194, 233 H. columbia, 143 H. gloveri gloveri, 233 hybridization, 95, 191, 226, 289 hybrids, 191, 226, 289 Hylephila phyleus, 189, 201, 211 Hymenoclea salsola, 215 Hymenoptera, 128 Hypercallia, 251 Hypolimnadidi, 170 Hypolimnas misippus, 170, 233 NG-Z.N., AS Incisalia iroides, 44, 146 insectivorous plant, 129 interactions, 65 inter-specific hybrids, 191, 226, 289 “arthechippus,” 289 eurymedon X rutulus, 226 lorquini < archippus, 289 “rubidus,” 289 “weidechippus,” 289 Ithomiidae, 170 Jennings, Daniel T., 123 Johnson, Kurt, 3 Junonia coenia, 203 Kendall, Roy O., 75, 86, 307 Kohler, Steve, 1, 57 Kricogonia lyside, 59 Laetilia, 105 Lamas. Gerardo, 116 Larisa subsolona, 256 Laspeyresia, 256 Laspeyresiinae, 256 lectotype designations, 49 Lederhouse, Robert C., 145 Leptotes marina, 209 Lerema accius, 178, 200 Lerodea eufala, 200, 211 Lethe anthedon, 28, 204 L. appalachia, 204 L. creola, 200 315 L. eurydice eurydice, 204 L. europa malaya, 64 Letis incipiens, 242 letter to the editor, 54 Leucochimona philemon, 46 Libytheana bachmanii bachmanii, 203 L. b. larvata, 209 Libytheidae, 203, 212 life cycle, 88, 160, 282 life histories, 123 Limenitidini, 170 Limentis, 228, 289, 305 L. archippus, 203, 228, 289 L. a. floridensis, 289 L. arthemis arizonensis, 203 L. a. arthemis, 203, 289 ik, Ga, asvimmabs, 2S, sie 208) CAE 228, 289 L. lorquini, 289 L. weidemeyerii, 289 Lycaeides melissa, 59 L. m. samuelis, 200 Lycaena arota, 44 L. hyllus, 202 L. phlaeas, 146, 202 Lycaenidae, 59, 139, 140, 146, 199, 212, 236, 277 Lycorea pieteri, 116 malaise traps, 178 Marpesia, 171 M. berania, 43 McAlpine, Wilbur S., 142 Meander felsina, 37 Mecyna furnacalis, 130 Megistanis, 170 Mellichamp, T. Lawrence, 20 Mestra, 171 M. amymone, 112 Metria, 242 Mexican Actinote, 261 Meyrick, 130, 251 microlepidoptera, 235 mid-valval flexion, 304 migration, 178 Miller, Jacqueline Y., 261 Miller, Lee D., 63, 64, 75, 139, 261 Miller, Thomas A., 233 Miller, William E., 256 mimicry, 157, 170 Ministrymon leda, 209 Mitoura, 3 Momphidae, 235 Morphidae, 65 Morpho achilles, 73 M. amathonte, 73 MENCUDTISt M. granadensis, 66, 73 316 JOURNAL OF THE LEPIDOPTERISTS SOCIETY M. peleides, 65 P. a. temenes, 273 M. theseus, 73 P. blomfildia, 169 mosaics, 228 P. ceres, 116 Munroe, Eugene, 130 P. cresphontes cresphontes, 201, 211 Mutuura, Akira, 130 P. demodocus, 146 Muyshondt, Alberto, 160 P. eurymedon, 226 Muyshondt, Albert, Jr., 160 P. glaucus, 28, 138; lav) 201226 Najas ceres, 116 P. joanae, 237, 304 Nastra lherminier, 201 P. multicaudatus, 226 Nathalis iole, 200, 209 P. polyxenes asterias, 138, 145, 157, natural history, 88 201, 228, 236, 303 Neck, Raymond W., 107, 111, 224, 310 P. rutulus, 226 Neil, Kenneth, 224 P. troilus, 28, 157, 201 Neilsen, M. C., 142 P. xuthus, 228 Neolycaena sinensis, 280 P. zelicaon, 226 new foodplant, 236, 303 Papilionidae, 1, 138, 139, 145, 226, 236, new genera, 103, 116 Dns, BIO} new generic assignment, 130 Parnassius clodius gallatinus, 1 new record, 236 Peigler, Richard, S., 191 new species, 75, 78, 82, 97, 118, 256 Peridroma plecta, 134 new subspecies, 75, 248, 261, 273 Peridromia, 169 Nica, 171 Perimede maniola, 235 Noctuidae, 134, 241, 306 Periploca cata, 235 Nordmannia, 277 Peteroma, 244 notes and news, 19, 48, 159 phenology, 207 Nymphalidae, 59, 88, 116, 135, 146, 160, Phocides lilea sanguinea, 107 185, 199)919) 924031) 9e8 961, Pl nolbus Ifeau lon 282, 289, 309, 310 P. pygmalion okeechobee, 108 Nymphalini, 170 Phoebus agarithe maxima, 59 Nymphalis antiopa, 203, 282 P. sennae, 178, 209 N. milberti milberti, 203 P. s. eubule, 202 N. vau-album, 200, 282 P. philea, 223 Nymphidium lisimon attenuatum, 46 Pholisora catullus, 201, 209 N. mantus, 46 Phyciodes batesti, 203 obituaries, 142, 240 P. tharos, 28, 146, 203, 231, 309 Oecophoridae, 235, 251 Pieridae, 59, 129, 139, 170, 185, 200, 212, Oeneis alberta, 57 DPN), ONS, O. chryxus strigulosa, 143 Pieris brassicae, 139 Ogunwolu, E. O., 175 P. napi oleracea, 129, 226 Oidaematophorus glenni, 235 P. protodice, 202, 209, 228 Olethreutidae, 235, 256 P. rapae, 202, 230 Oliver, Charles G., 231, 309 P. virginiensis, 28, 35, 202 Ollia parvella, 103 pine feeding species, 118 orange bands, 135 Platt, Austin P., 289, 305 Ornithoptera priamus, 138 Poanes hobomok, 28, 200 O. victoriae, 138 P. massasoit hughi, 200 Ostrinia, 130 P. m. massasoit, 200 oviposition, 233, 236 P. viator, 200 Pammene, 258 P. yehl, 200 Panoquina ocola, 178, 200 je zabulon, 200 Panthiades m-album, 200 Pogue, Michael G., 236 Papilio andraemon, 273 Polites coras, 201 P. aristodemus, 273 P. a. bjorndalae, 275 P. a. driophilus, 273 P. a. ponceanus, 273 . mystic, 201 . origenes origenes, 201 . themistocles, 201 . vibex vibex, 200 ae Bene pias ac! VoLUME 32, NUMBER 4 Polychrosis sambuci, 235 Polydesma, 243 Polygonia comma, 28, 203 P. faunus faunus, 203 P. interrogationis, 28, 203 P. progne, 203 Pompeius verna verna, 200 population structure, 37 Precis coenia, 178, 224 predation, 134 Prepona, 170 proboscis, 310 Problema byssus, 200 Proeulia, 253 P. montana, 253 Pseudohazis, 97 Pseudonica, 170 Pteromalidae, 128 Pterophoridae, 235 Pycina, 170 Byralidae,, po, 103, 175 Pyrausta homaloxantha, 130 Pyraustinae, 130 Pyrginae, 184 Pyrgus centaureae wyandot, 201 P. communis, 201, 209 Pyrrhogyra, 169 Rawson, George W., 289 rediscription, 103 Restinga butterflies, 37 Rhododipsa aden, 49 Rhopalocera, 111, 198 Riodinidae, 37, 200 Robb, Jeff, 56, 59 roosting, 145 Rostralaetilia, 103 Rawlins, John Edward, 146 Safia, 242 Samia cynthia, 191 Satummidae, 59, 97, 191, 233 Satyridae, 57, 86, 200, 204 Satyrium acaciae, 273, 280 . acadica acadica, 200 . acadicum, 281 . adenostomatis, 281 . auretorum, 281 . behrii, 280 . calanus, 281 . californicum, 281 Sue. jalicen 202 . caryaevorus, 200, 281 . edwardsii, 202, 281 . eximium, 281 . fuliginosum, 280 - tlicis, 281 NN RNNRNNN RNNNN . kingi, 281 . latior, 281 . liparops, 281 S. l. strigosa, 202 . lunulatum, 280 . myrtale, 280 . pretiosum, 281 . saepium, 281 . sassanides, 280 . sinensis, 281 . spini, 280 . sylvinum, 281 . tetra, 281 . tengstroemii, 281 . w-album, 280 Sceliodes cordalis, 177 S. laisalis, 175 Schinia approximata, 49 S. Gr, DO S. dolosa, 49 . hanga, 50 . labe, 49 . lora, 49 . neglecta, 52 . obscurata, 52 . pyraliodes, 49 . tanena, 53 ultima, 53 Scott, James A., 304 Scriber, Mark J., 236 Sebethis, 116 Shapiro, Arthur M., 223, 228 Shields, Oakley, 61 Silberglied, Robert E., 135 Siproeta epaphus, 43, 172 S. stelenes, 172 Smith, Dwight G., 134 Smyrna bella, 169 S. blomfildia, 160 S. karwinskii, 160 S. pluto, 169 Sorhagenia baucidis, 235 Speyeria aphrodite, 146, 204 S. atlantis atlantis, 204 S. cybele cybele, 28, 204 S. diana, 204 Sphingidae, 56, 139 species diversity, 218 NNN NNNNNRNRNRNHnRNHNHNN ANNNNnNNNN Splendeuptychia kendalli, 75, 86 S. salvini, 78 Staphylus ceos, 209 S. mazans hayhurstii, 201 Stevens, Robert E., 118, 123 Strymon, 278 S. melinus, 209 S. m. humili, 202 S17 318 Strymonidia thalia, 277 Systematics, 1, 49, 75, 97, 103, 116, 118, 130: 139. 241 25125652615 2738; TT Tampa, 105 Taygetis mermeria excavata, 75 T. m. griseomarginata, 75 Teladoma incana, 235 Temenis, 170 temporal distribution, 86 Thecliolia, 280 Thymelicus lineola, 146, 200, 225 Texola eulauda perse, 211 Thorybes bathyllus, 201 T. confusis, 200 T. pylades, 28, 201 Tilden, J. W., 241 Tmolus azia, 236 Todd) Bok. 244 Tortricidae, 251 Trogus pennator, 303 Tuskes, Paul M., 97 unusual occurrences, 111 Urania fulgens, 307 U. leilus, 307 Urbanus dorantes, 178 U. proteus, 178, 200 Uresiphita, 130 Urticastrum mexicanum, 166 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Vanessa annabella, 88, 211 V. atalanta, 203, 211 V. a. rubria, 93 V. cardui, 93, 209 V. virginensis, 93 voltinism, 216 Wagner, Warren H., Jr., 20, 226 Walker, Thomas J., 178 Wallengrenia egeremet, 201 Walshiidae, 235 Welderella, 103 W. parvella, 105 Wilkinson, Ronald S., 306 Xanthopsamma, 130 X. aurantialis, 132 X. homaloxantha, 132 X. youboialis, 132 Xylis setipes, 242 Young, Allen M., 65 Zagrammosoma multilineatum, 128 Zale discisigna discisigna, 241 Z. notipennis, 241 Z. peruncta, 241 Z. p. ineipiens, 248 Z. setipes, 241 Z. s. £. postmedialis, 246 Zaretis, 171 Zerene cesonia, 113, 209 Zinnia pumila, 215 PLANT INDEX FOR VOLUME 32 Acacia greggii, 209 Acer rubrum, 33 A. saccharum, 33 Adiantum pedatum, 34 Aegopodium podograria, 237 Aesculus octandra, 33 Ailanthus altissima, 192 Althaea rosea, 92 Anethum graveolens, 303 Angelica atropurpurea, 237, 303 Antirrhinum, 224 Arachis hypogea, 67 Aruncus dioius, 20, 26 Asclepias syriaca, 155 Aster novae-angliae, 154 A. simplex, 23] Asteraceae, 32 Astilbe biternatum, 32 Athyrium filix-femina, 34 Bambusa aculeata, 79, 86 Betula lenta, 33 Bidens pilosa, 185 Botrychium virginianum, 34 Callistemon citrinus, 109 Canna, 108 Caprifoliaceae, 29, 285 Capsicum annuum, 175 C. frutescens, 175 Carex plantaginea, 34 Carpinus caroliniana, 33 Celtis pallida, 207 Cercidium microphyllum, 207 Cercocarpa betuloides, 99 Chamaecyparis thyoides, 3, 7 Chelone glabra, 282 Cicuta bulbifera, 237, 303 C. maculata, 303 Cimicifuga racemosa, 34 Coleogyne ramosissima, 97 Conium maculatum, 237, 304 Cornaceae, 29 VoLUME 32, NUMBER 4 Cornus florida, 33 C. racemosa, 29 C. stolonifera, 29 Cryptotaenia canadensis, 237, 303 Cupressaceae, 3 Cynodon dactylon, 86 Dactylis glomerata, 151, 155 Dacus carota, 237, 303 Dioclea, 72 D. wilsoni, 67 Diplacus, 224 Drosera rotundifolia, 129 Duranta repens, 109 Erythrina crista-galli, 67 Euphorbiaceae, 307 Fagus grandifolia, 33 Gramineae, 86 Geranium maculatum, 24 Heracleum maximum, 237 Hydrangea arborescens, 21 Hymenoclea salsola, 215 Impatiens capensis, 34 Inga, 72 Ipomoea coccinea, 215 Juniperus, 3 J. ashei, 4 J. californica, 6 J. communis, 6 J. depeana, 4 J. flaccida, 7 J. horizontalis, 4 . monosperma, 4 . occidentalis, 4 . osteosperma, 4 . pinchotii, 4 . scopulorum, 5 . silicicola, 6 . virginiana, 4 Kickxia spuria, 224 Laportea canadensis, 34 Leguminosae, 65 Levisticum officinale, 238 Lindera benzoin, 33 Linaria, 224 L. vulgaris, 151 Lippia, 224 Cy ay ay ey ey ey Liriodendron tulipifera, 34, 192 Lonicera japonica, 286 L. morrowi, 151 Lycopersicum esculentum, 175 Machaerium floribundum, 65, 72 Magnolia acuminata, 236 Malwa, 92 M. parviflora, 92 Malvaceae, 92 Marcgraviaceae, 45 Medicago sativa, 67 Mirabilis multiflora, 215 Moraceae, 166 Mucuna, 72 Myrtaceae, 107 Norantea brasiliensis, 45 Omphalea diandra, 307 Opuntia fulgida, 209 O. spinosior, 209 Osmorhiza claytoni, 237, 303 O. longistylis, 237, 303 Oxypolis filiformis, 303 Passifloraceae, 111 Passiflora lutea, 111 Pastimaca sativa, 237 Penstemon azureus, 224 Phleum pratense, 151, 155 Pinus contorta, 123 P. edulis, 123 P. jeffreyi, 123 Plantaginaceae, 224 Plantago, 224 Potentilla recta, 151, 153 Prosopis juliflora, 207 Prunus serotina, 233, 290 Psidium cattleianum, 107 P. guajava, 107 Ptilimnium capillaceum, 303 Quercus dumosa, 61 QO. durata, 61 Rhododendron, 36 Ribes alpinum, 224 R. nigrum, 224 R. rubrum, 224 Rutaceae, 236 Salix babylonica, 290 Sanicula gregaria, 237 Saxifragaceae, 32 Scrophulariaceae, 224 Sida, 92 Sidalcea malvaeflora, 92 Sium suave, 303 Solanaceae, 175 Solanum macrocarpon, 175 S. melongena, 175 Solidago altissima, 151, 154 Sphaeralcea ambigua, 92 Spermolepis divaricata, 303 Spiracea, 28 Taenidia integerrima, 237, 303 Thaspium barbinode, 304 Thuja occidentalis, 129 Tiarella cordifolia, 34 Tilia americana, 34 Tragopodon pratensis, 32, 151, 153 319 320 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Trillium, 34 Urtica holosericea, 90 Umbelliferae, 236 U. urens, 92 Urera baccifera, 166 Urticastrum mexicanum, 166 U. caracasana, 166 Verbenaceae, 109, 224 Urticaceae, 92, 160 Vibernum lentago, 29 Zinnia pumila, 215 ERRATA p. 20, abstract, line 7, W. ebenina = C. ebenina p. 35, third line from bottom, Marylan = Maryland p. 134, note title & Vol. 2 Contents on back cover, Peridoma = Peridroma p. 172, line 2, V. epaphus = S. epaphus p. 214, third paragraph, line 7, C. caesonia = C. cesonia p. 224, second paragraph, line 4, E. mellinate = E. mellinata Vol. 3 Contents on back cover, fourth line from bottom, Mark Fink = Mark Finke (The Editor regrets these errors ) Date of Issue (Vol. 32, No. 4): 28 February 1979 EDITORIAL STAFF OF THE JOURNAL AusTIN P. Piatti, Editor Department of Biological Sciences University of Maryland Baltimore County, 5401 Wilkens Avenue Catonsville, Maryland 21228 U.S.A. Frances S. Cuew, Managing Editor Dovuctas C. Fercuson, Associate Editor THEODORE D. SARGENT, Associate Editor NOTICE TO CONTRIBUTORS Contributions to the Journal may deal with any aspect of the collection and study of Lepidoptera. Contributors should prepare manuscripts according to the following instructions. Abstract: A brief abstract should precede the text of all articles. Text: Manuscripts should be submitted in duplicate, and must be typewritten, entirely double-spaced, employing wide margins, on one side only of white, 8% xX 11 inch paper. Titles should be explicit and descriptive of the article’s content, including the family name of the subject, but must be kept as short as possible. The first men- tion of a plant or animal in the text should include the full scientific name, with - authors of zoological names. 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Gates Glarke 2.0 +0 LARISA SUBSOLANA, A New GENUS AND SPECIES OF MOTH FROM EASTERN NortH AMERICA (OLETHREUTIDAE). William E. Miller Notes ON MEXICAN ACTINOTE (NYMPHALIDAE: ACRAEINAE) AND THEIR RELATIVES, WITH DESCRIPTION OF A NEW SupssPEciEs. Jacqueline Y. Miller & Lee D. Miller 0 PAPILIO ARISTODEMUS (PAPILIONIDAE) IN THE BAHAMAS. Harry K. Clench THe NAMEs OF CERTAIN HoLarctic HAIRSTREAK GENERA (LYCAE- NIDAE). Harry K..Clench 0 ee OVER-WINTERING BEHAVIOR IN EUPHYDRY AS PHAETON (NYMPHALIDAE). M. Deane Bowers INTER-SPECIFIC HYBRIDIZATION INVOLVING LIMENITIS ARCHIPPUS AND irs CONGENERIC SPECIES (NYMPHALIDAE). Austin P. Platt, George W. Rawson, & George Balogh GENERAL NOTES Taenidia integerrima, a new foodplant record for Papilio polyxenes (Papilionidae).. M. Berenbaum ‘22.0000 Mid-valval flexion in the left valva of asymmetric genitalia of Erynnis (Hesperiidae). James A. Scott (i000 Catocala ilia (Noctuidae) feeding on decaying fruit in an inner-city en- vironment. Ronald S, Wilkinson. 0.0.0.0. Periodic occurrence of Urania fulgens (Uraniidae) in the United States. Roy O.. Kendall 0000000008 i OS A merely albinic aberration of Phyciodes tharos (Nymphalidae). Charles G. Oliver sl a 0 A A record of Agrias amydon (Nymphalidae) from Costa Rica. Philip J. DOV 148 esc ces ID Oa I A Additional function of the Lepidopteran proboscis. Raymond W. Neck -.. Eprror’s Nore a a a ek be St me Fb en tr ts eo ef oh a Se Pr mS 241 251 256 261 273 277 282 289 Volume 33 1979 Number 1 JOURNAL of the LEPIDOPTERISTS’ SOCIETY Published quarterly by THE LEPIDOPTERISTS’ SOCIETY Publié par LA SOCIETE DES LEPIDOPTERISTES Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS cat 4 June 1979 THE LEPIDOPTERISTS’ SOCIETY EXECUTIVE COUNCIL I. F. B. CoMMoN, President T. SHIROZU, Vice President C. V. COVELL, JR., lst Vice President , JULIAN P. DONAHUE, Secretary L. A. GOZMANY, Vice President RONALD LEUSCHNER, Treasurer Members at large: R. A. ARNOLD J. F. EMMEL C. D. FERRIS E. D. CASHATT R. R. GATRELLE J. Y. MILLER R. E. STANFORD A. P. PLATT M. C. NIELSEN The object of the Lepidopterists’ Society, which was formed in May, 1947 and for- mally constituted in December, 1950, is “to promote the science of lepidopterology in all its branches, ....to issue a periodical and other publications on Lepidoptera, to facilitate the exchange of specimens and ideas by both the professional worker and the amateur in the field; to secure cooperation in all measures” directed towards these aims. Membership in the Society is open to all persons interested in the study of Lepi- doptera. All members receive the Journal and the News of the Lepidopterists’ Society. Institutions may subscribe to the Journal but may not become members. 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Order: Mail to Charles V. Covell, Jr., Memoirs Editor, Department of Biology, Uni- versity of Louisville, Louisville, KY 40208, U.S.A. The Lepidopterists’ Society is a non-profit, scientific organization. The known office of publication is 1041 New Hampshire St., Lawrence, Kansas 66044. Second class postage paid at Lawrence, Kansas, U.S.A. 66044. Cover illustration: Third instar larva of Limenitis archippus Cramer (Nymphalidae) preparing to enter winter diapause. The larva is resting on the lip of its hibernaculum constructed from the basal portion of a chewed tubular willow leaf (Salix babylonica Linnaeus) covered with silk. In the autumn such larvae begin facultative diapause in response to decreasing day-length. Original drawing by Mr. George C. Ford, Jr., Graph- ics Illustrator, Department of Biological Sciences, University of Maryland Baltimore County, 5401 Wilkens Avenue, Catonsville, Maryland 21228. eS ae a ¥ } % 4 JOURNAL OF Tue LeprpopreRiIsts’ SOCIETY Volume 33 1979 Number 1 Journal of the Lepidopterists’ Society 33(1), 1979, 1-5 PAITITIA NEGLECTA, GEN. N., SP. N. FROM PERU (NYMPHALIDAE: ITHOMIINAE )? GERARDO LAMAS Museo de Historia Natural “Javier Prado,” Universidad Nacional Mayor de San Marcos, Apartado 1109, Lima-100, Pert ABSTRACT. Paititia neglecta, gen. n., sp. n., from Peri, San Martin, Juanjui, is described herein. This monotypic genus is considered to be the most primitive member of the tribe Mechanitini of the Ithomiinae. Two specimens, representing a new genus and species of Ithomiinae, lay unnoticed for almost 50 years in the collections of the British Museum (Natural History ), London, and the American Museum of Natural His- tory, New York. The BMNH female was included among a series of un- identified Methona examples in the Rothschild collection, while the AMNH male had been identified (and labelled) as “Xanthocleis ino” by the late R. M. Fox. The third known specimen (a male) was presented to the “Javier Prado” Museum in 1976 by the collector, Mr. José M. Schunke. Paititia Lamas, new genus External diagnostic characters. Large Mechanitini (Fox, 1956), very similar in size and color pattern to Thyridia psidii ssp. and Methona spp. May be distinguished from Thyridia psidii (Linnaeus) by the absence of small red dots on base of forewing above, and the presence cf two separate, white spots on cell Sc-R:-R; of hindwing below. Paititia may be separated from Methona spp. by the long hair patch extending beyond the discal cell apex of the male hindwing above, which in the latter is restricted to the basal half or two-thirds of that cell. In Fox’s key (1940), the males of Paititia will key out to Xanthocleis Boisduval (i.e., Thyridia Hiibner), but may be readily differentiated by the characters given above. The females will key out to Athesis Double- day; however, they can be easily distinguished by the wing shape and 1 The present paper forms part of a D.Sc. dissertation submitted to the Departamento de Zoologia, Instituto de biociéncias, Universidade de S40 Paulo, Sao Paulo, Brasil (Lamas, 1973). As a con- densed version of that thesis will not be published in the near future, I have decided to make available the new genus and species of Ithomiinae described therein. bo JOURNAL OF THE LEPIDOPTERISTS SOCIETY Fics. 1-11. Paititia neglecta, gen. n., sp. n.: 1, palpus; 2, male foreleg; 3, fe- male foreleg; 4, male forewing; 5, male hindwing; 6, female hindwing; 7, male geni- talia, ventral view; 8, male genitalia, lateral view; 9, penis; 10, male eighth tergite, dor- sal view; 11, female genitalia, ventral view. color pattern (wings long and narrow, with yellowish transparent areas in Paititia, short and wide, with reddish-brown translucent areas in Athesis ). Palpi (Fig. 1). Basal segment curved, adpressed to the head; second free, 1.5 times longer than the basal; apical fusiform, very small, one-fifth to one-eighth the length of the basal (longer in male). Antennae. Club yellow, with approximately 15 segments, very slightly widened, VOLUME 33, NUMBER 1 3 barely more so than the pedicel, which is black; as long as the forewing discal cell, that is, slightly longer than one-half the forewing length. Male forelegs (Fig. 2). Reduced; femur plus trochanter as long as the stout articulate coxa; tibia one-fourth longer than femur; tarsus two-fifths the length of the tibia, with slender spines on ventral side. Female forelegs (Fig. 3). Reduced, coxa articulate; femur as long as coxa; tibia equal to femur plus trochanter; tarsus with 4 apparent segments, two-fifths the length of the tibia; basal segment twice the lengh of the remainder; second and third short, apical consisting of the fourth and fifth fused; first segment with two pairs of spurs, the outer ones slightly longer; second and third segments with a pair of spurs each; second, third and apical segments with trichoid sensillae. Venation (Figs. 4-6). Sc and R: on forewing of both sexes running parallel, with- out anastomosing, Sc ending shortly beyond discal cell apex; Ri arising from cell, base of Re anastomosed to Rs-s-s, well beyond cell apex; rs-m: absent or very short, straight; mi-mz long, angular, long Re arising from angle; msz-ms also long, straight. Male hindwing with bifid hum, both arms well developed; Sc and R separate at base, Sc ending beyond discal cell apex; rs-m: straight, slightly more than half as long as mms; mi-mz angular, both arms straight and of equal length, Rc on angle; Me2-ms straight. Hair patch complete, running along upper portion of discal cell, behind radial vein, from base to beyond cell apex. Female hindwing. Similar to male, except that Sc ends scarcely beyond cell apex; rs-mi very short, less than one-third the length of m2-ms; no hair patch. Male genitalia (Figs. 7-10). Eighth tergite with two short and distally widened lateral lobes, without sclerotized claws; saccus short, one-half as long as the valva; tegumen hood-like, not separated by a suture from uncus; uncus stout, posterior area with a strong claw bent downwards; juxta V-shaped, poorly developed; appendices angulares not sclerotized; gnathos lightly developed, its arms being united below the tuba analis by means of a membrane only; valvae symmetrical, long, quite wide, bear- ing two claw-like appendices on their caudal ends, upper claw shorter and stronger than lower; penis long, slender, curved between the anterior and middle thirds; fora- men penis very long, one-third of the total length; gonoporus flared, terminal; vesica with cornuti. Female genitalia (Fig. 11). Vaginal plate irregular, funnel-shaped, with a lateral aliform process on left side; ostium bursae located left of longitudinal axis of abdomen; caudal end of ostium bursae strongly sclerotized and slightly bent left and downwards, remainder of ductus very long; bursa copulatrix with poorly developed signa and with a globular appendix bursae. Type-species: Paititia neglecta Lamas, sp. n. Etymology. The generic name is based on the Quechua word “‘Paititi,” which re- fers to the fabulous kingdom of “El Dorado,” so ardently sought by explorers and ad- venturers in South America, almost ever since the discovery of the New World. It should be treated as being of feminine gender. Paititia neglecta Lamas, new species Male (Fig. 12). Wing margins and transverse bands black, transparent areas with a yellowish tinge. Hindwing below with two white spots on cell Sc + Ri-R; and a row of marginal double white spots on cells M:-M>2 to Cu.-2A. Humeral spot white, costal line pale yellow. Abdomen below, and all other body spots, white. Female (Fig. 13). Similar to male, hindwing white, marginal spots smaller. Types. HoLorypPe @, Juanjui, San Martin, Peru; xi.34 (G. Klug), deposited in the British Museum (Natural History). One PARATYPE ¢, Achinamiza, San Martin, Peri; 14.1.26 (H. Bassler, station F6001); AMNH Acc. 33591, in the American Museum of Natural History, New York. One PARATYPE ¢, Iberia, Madre de Dios, Peri; 27.vi.75 (J. M. Schunke ), in the Museo “Javier Prado,” Lima. Etymology. The specific name is the Latin word for “forgotten.” 4 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Fics. 12-13. Paititia neglecta, gen. n., sp. n.: 12, paratype male, Achinamiza; 13, holotype female. Dorsal left, ventral right. DiIscussION Relationships Paititia is considered to be the most primitive member of the tribe Mechanitini (Lamas, 1973). It indeed seems to represent a link between the Tithoreini and Mechanitini. Paititia neglecta resembles members of the genus Olyras Doubleday (cf. Fox, 1940, 1956), in the shape of the male forelegs, wing venation and genitalia, and even the presence of the white spots on the hindwing cell Sc + Ry-R, below (present in most forms of Olyras). However, Olyras differs by the divided hair patch and by the valvae having only one short claw-like process, the females being even more different. Paititia is very close to Thyridia Hiibner (= Xanthocleis Boisduval), both having almost identical venation, but I believe the differences pre- sented by the male and female genitalia and foretarsi are enough to sep- arate them. VoLUME 33, NUMBER 1 5 Natural history Nothing is known about the life-habits of Paititia neglecta. Two of the localities where it has been found (Juanjui and Iberia) are characterized by rather dry tropical forest. This may represent an example of a primi- tive species which has been displaced to marginal habitats by a more modern and aggressive species, as seems to be the case with Melinaea mnasias ( Brown, 1977). ACKNOWLEDGMENTS Research for the present paper was mainly financed by grants “Bio- légicas 70/671” and “72/849” of the Fundacéo de Amparo a Pesquisa do Estado de Sao Paulo (FAPESP), Brasil. I acknowledge with special thanks the help and hospitality of Mr. R. I. Vane-Wright (British Museum [Natural History] ), Dr. F. H. Rindge (American Museum of Natural His- tory, New York), and the advice of Drs. P. E. Vanzolini and N. Papavero (Museu de Zoologia da USP, Sao Paulo) and K. S. Brown, Jr. ( Universi- dade Estadual de Campinas, Sao Paulo). I am deeply obliged to Mr. J. M. Schunke for having obtained the third known specimen of Paititia neglecta for the “Javier Prado” collections. LITERATURE CITED Brown, K. S., Jr. 1977. Geographical patterns of evolution in Neotropical Lepi- doptera: differentiation of the species of Melinaea and Mechanitis (Nymphali- dae, Ithomiinae). System. Entomol. 2: 161-197. Fox, R. M. 1940. A generic review of the Ithomiinae (Lepidoptera: Nymphali- dae). Trans. Amer. Entomol. Soc. 66: 161-207. 1956. A monograph of the Ithomiidae (Lepidoptera). Part I. Bull. Amer. Mus. Nat. Hist. 111: 1-76. 1967. A monograph of the Ithomiidae (Lepidoptera). Part III. The tribe mechanitini Fox. Mem. Amer. Entomol. Soc. 22: 1-190. Lamas, G. 1973. Taxonomia e evolucdo dos géneros Ituna Doubleday (Danainae) e Paititia, gen. n., Thyridia Hiibner e Methona Doubleday (Ithomiinae) ( Lepidop- tera, Nymphalidae). D.Sc. Thesis to the Departamento de Zoologia, Instituto de Biociéncias, Universidade de Sao Paulo, vii + 225 p. Journal of the Lepidopterists’ Society 33(1), 1979, 6-20 EXPERIMENTAL HYBRIDIZATION BETWEEN PHYCIODES THAROS AND P. BATESII (NYMPHALIDAE ) CHarLES G. OLIVER! R. D. 1, Box 78, Scottdale, Pennsylvania 15683 ABSTRACT. F: hybrids and backcrosses were made between the nymphalid but- terflies Phyciodes tharos and P. batesii. The two species differ in larval, pupal, and adult phenotypic appearance, ecology, and larval diapause response. Genetic in- compatibility was shown by significant hybrid inviability, growth irregularities, and abnormal adult sex ratios and development times. The reciprocal F: hybrids differed greatly in their expression of incompatibility. Hybrid inviability is attributed to break- downs in the genetic mechanisms controlling growth and development. The relationship of Phyciodes tharos Drury to P. batesii Reakirt has been little understood despite the fact that they both occur in compara- tively densely populated areas of the northeastern United States and have been known to be distinct for well over a century. This confusion appears to be due to two causes. First, P. tharos is common to abundant over the entire range of P. batesii. Since P. batesti occurs in widely sepa- rated, small populations, it is probably often overlooked due to its super- ficial resemblance to P. tharos. Second, the biology of P. tharos itself actu- ally is poorly known. Rearing and hybridization studies now in progress in my laboratory indicate that “P. tharos” in the Northeast consists of two entities differing in larval and adult phenotypic appearance and voltinism and showing significant incompatibility when hybridized in the laboratory (Oliver, in prep.). The more southern entity, hereafter referred to as “Type A,” occupies the Transition and Austral Life Zones, whereas the more northern “Type B” is the “P. tharos” of northern New England, northern New York State, and southern Canada. The phenotypic differ- ences of Types A and B have resulted in Type B passing as P. batesii in many collections, although it resembles P. batesii little more than does Type A. Phyciodes batesti is very local in the Northeast. It appears to be restricted to dry sites, often of the barrens type. The Onondaga Co., New York, pop- ulation used in these experiments occurs on dry limestone ledges. In — the Appalachians of Pennsylvania and West Virginia, the localities I have investigated are for the most part shale barrens or rocky riparian slopes. One of the best known localities, along the banks of the Ottawa River near Aylmer, Quebec, was described by McDunnough (1920) as the “lower dry slopes of a small ridge.” Both Types A and B of P. tharos occur in a wide variety of habitats, including those of P. batesii. 7 Adjunct. Assistant Professor, Dept. of Biology, West Virginia University, Morgantown, West Virginia 26506. VoLUME 33, NUMBER 1 t Phyciodes batesii flies in early June in West Virginia and southwestern Pennsylvania and in mid June in central New York State. In West Virginia and southern Pennsylvania this is between the rather discrete first and sec- ond broods of P. tharos Type A, which has a total of three to four broods. In Onondaga Co., New York, however, P. batesii and P. tharos Type B fly together. Thus, the observation of Forbes (1960) that broods of P. tharos and P. batesii alternate at any given locality seems to apply only to the southern portion of the range of P. batesii. The life history of P. batesii was described and figured by McDun- nough (1920). The larva and pupa differ in a number of characters from those of P. tharos (Table 1) and are much more like those of P. campes- tris, which were figured by Comstock (1930) and compared briefly to P. tharos in another paper of mine (Oliver, 1978). First and second instar larvae of both P. campestris and P. batesii live communally within a loose web spun over the feeding area on the foodplant. P. tharos has a similar communal behavior, but no web is spun. McDunnough was able to obtain oviposition by P. batesii on “a species of Aster with heart-shaped leaves,” found wild larvae on this plant, and successfully reared them through to adults. This Aster was probably A. undulatus L., which is very common in P. batesii habitats. Wild-caught P. batesii females from Onondaga Co., New York, refused to oviposit on Aster undulatus in the laboratory, but laid readily on A. simplex ( Willd.) Burgess, a common foodplant of P. tharos. Newly-hatched larvae of both Phyciodes fed readily on A. undulatus when transferred to its leaves. Larvae of both species also accepted A. laevis L. More than 200 unmated, laboratory-reared P. batesii adults were re- leased shortly after eclosion during August into a western Pennsylvania old-field habitat containing abundant Aster simplex (but no A. undula- tus). Several pairs were observed in copulo the next day. In early Octo- ber a group of small larvae was recovered from a clump of A. simplex and reared through to normal adults the following spring, indicating that P. batesii may choose more than one species of Aster as natural foodplants. In the northern Midwest (e.g., Michigan and Wisconsin) P. batesii has a somewhat different biology. Colonies occur in moist areas, and there is sometimes a partial second brood (Nielsen, in litt.). Populations in the Northeast seem to be strictly univoltine. Midwestern P. batesii differ slightly in appearance from those in the Northeast and may possibly rep- resent a separate entity. PROCEDURE Stock of P. batesii used in these experiments was derived from four wild-inseminated females collected 11 June 1976, in Syracuse, Onondaga JOURNAL OF THE LEPIDOPTERISTS SOCIETY Juesqe 10 JuIeF AT[eNsN ‘opqeisze A J9}U99 Yor[q yurodurd *M wAMo1g 10 yorlTg o} eIPSur9ezUyT Jeordeqns = ueipeyy o]vIPoUtIezUT 9] RIPOU19}UT uMOIg AvIS [[e JSOWTY 9} VIPOUIIOUT uo} -elIeA [eyuerted poesoxa 6 6 ‘Q}eIPOULIOUL P P ‘afqeriea AIO 9} VIPSW19zU] UMOIG 9} [OOOUT) 9} ¥IPSULI9}U] 9} PIPOULIO}UT OHM. uMoIg Yysryurd yySsTT esuly ystyurd *M UMOIg spuqsy yuosoid Jusosa10 IaATIS ATUO ATTeNS(-) Prd MOEA Aes ofed Ato jeordeqns < ueipeyy pueq snonulju0d UOJ 0} puay, MOT[PA MeI}S 0} BSURIO O[e Avis y1ed Avis [[e JSOWLY A¥Is Yep pue oY posroyooyD, yoryq Ajueur -wopeid varie SUI ‘AAvoy AIO A ques -qe jsowye suorjoefo1d [esiop ‘OUI]JNO UL pepuNnor s10Wl Yon, UMOIG 9yeTOOOYD AT[P19}LT pue A][esiop pezoLIjse1 “19 ueAd “AAvoy ‘teduoy UMOIG JYSIT SNOIO[OOUOD uMOIg YsTyurd YSTT esul} ysryurd *M UMOIg yu90se10 SUIpNo[O Ueayo ‘asie'T 19}U90 yorlq qurodurd 0/M 10 ‘A UMOIg MOT[EA MeI]S Ye IOAqIS uevipeut < jeordeqns pueq SUIULIOZ JOU ‘poze]OsT esurlo yIeqd UMOIg sarey AuMe} JO Uor}zodoi1d ose] “MA Ati yep pue Avis YS] potoyooyD svole OS5UvIO uado asie] ‘WY SsIT AlPATeTOYy peounouoid AJ@AT}eJeI suondefoid [esiop ‘OUITJNO UI Ie_nsue s1oW Yon, UMOIG JYSI'T Ajjeteyey pue AT[esiop sAIsue}X'y uayoIq “UIe ony UMOIG Yep 10 ABI UMOIG 9}BTODOYO yAIeG yoyed yrep jeuy *), (4Juo 2 2) sjods yep jeulsieurqns [jews ‘9 IO[OO punoIsS AH “CG soyoyed yor[q [eordeqns ‘sa UBIPOUL JSOMOT ‘SOZIS ‘OY “Pf sjods y1ep uvIpeut MAE Iojoo punois zYSIT MA's IO[OO qnyo [euUa\UY *T jeque, “g IO]OO 91n}I4seA Apog ‘¢ IO[OO SOBULIT BUIAA “Z uotssoidxe ureyed yorlg ‘T [esiogd ‘Vv aD ee souvivedde [[e12AQ edng ‘TI UOI}RIOTOO [PBS “F a[nsdeo peey ‘seyojed a} AA “28 sodiiys yYST [esiog “p Io]Oo dij apolaqny, ‘9 IO[OO apo1eqny, “q IojOoo punois Apog “ve BAIL] INV ‘T HSO9Dq *d SOLDY} * J qajyoRINyD ‘spriqdAy ‘yy Iey] pur ‘Vsazpq *g ‘so1py} ‘g Jo souvievodde ordAjousyd jo uostiedwo) ‘T FIavy -VoLUME 33, NUMBER 1 9 Co., N.Y. Stock of P. tharos Type A was derived from four females taken 10 June 1976, in Acme, Westmoreland Co., Pennsylvania, and of Type B from two females taken 11 June 1976, at the Onondaga Co., N.Y. locality. Cultures were maintained at 15 to 28°C and separated into controlled photoperiod (18 h light/24 h or 24 h light/24 h) and natural photoperiod (for Fayette Co., Pa., latitude 40°N) groups. Artificial lighting after sunset was provided by a 100-watt incandescent bulb at a distance of 1 to 2 meters. Matings were made by the hand-pairing method (Clarke, 1952). Fecundated females were provided with cut sprigs of Aster simplex for oviposition. Eggs were left in situ until hatching and the aster sprigs kept fresh in water. Larvae were reared on cut sprigs of A. simplex in water and housed in 10 X 20 cm glass cylinders. The photoperiodically-regulated larval diapause in Phyciodes occurs at the beginning of the third instar. Diapausing larvae were removed from active cultures, placed in groups in 90 mm plastic petri dishes, sealed in airtight containers, and stored in a domestic refrigerator at 0 to 2°C until April or May of the following year. Upon removal from the diapause con- tainers al] larvae were maintained at 15 to 28°C and 18 h light/24 h until pupation, at which time they were transferred to natural light conditions. F, progeny of wild-collected females was used for the hybrid pairings and as parental-type stock for backcrosses; no stock used was inbred. Ob- servations were made on parental population and F,; hybrid phenotypic appearance, interspecific courtship behavior, development periods and adult eclosion patterns, fertility, adult sex ratios, embryonic, pupal, and adult viability, and on backcross embryonic viability. Controls were reared concurrently at all times for comparison with experimental broods. Data on egg fertility, viability, and sex ratios were treated statistically using the Wilcoxon Two-sample Test. Adult fertility was measured by a count of the number of visibly developing eggs divided by the total num- ber of eggs laid after a single mating. Development periods from hatch- ing of the egg to eclosion of the adult were estimated by calculating the 99% confidence intervals for the medians of the distributions (Owen, 1962). Distributions of development times within broods or series of broods were represented by adult eclosion curves, graphs of the number of adults eclosing from pupae each day. RESULTS Interspecific courtship behavior Courtship in both P. tharos and P. batesii is apparently dependent on a variety of stimuli. The presence of a butterfly of appropriate coloration 10 JOURNAL OF THE LEPIDOPTERISTS SOCIETY VoLUME 33, NUMBER 1 Wl and size elicits approach by males. If the approached individual does not leave or, in the case of conspecific males, show aggressive behavior, as attempt at copulation will be made. Females which do not wish to cop- ulate avoid the male’s probing genitalia by dorsal or lateral movement of the abdomen. Males of both P. tharos and P. batesii showed the courtship approach response when presented with females of the other species. After an initial response, however, males of neither species attempted to copu- late. Some stimulus, perhaps olfactory, appeared to terminate courtship. Phenotypic appearance Differences in phenotypic appearance of the fifth instar larvae, pupae, and adults of the parental species and F, hybrids are summarized in Table 1. Typical specimens of adults are shown in Fig. 1. There was wide varia- tion in adult phenotypic appearance among the F, hybrid broods. The artificial illumination levels used to extend daily photophase in these experiments eliminated facultative diapause in P. tharos (see below ), but the induction of the naturally-occurring adult seasonal forms was little affected. The photoperiodic regulation of polyphenism in P. tharos has been described in a previous paper (Oliver, 1976). Univoltine P. batesii do not, of course, show natural seasonal polyphenism. Under the artificial long-day laboratory conditions described above, however, non- diapausing larvae produced late summer and early fall adults which had significantly heavier expression of the dark wing pattern elements on both the ventral and dorsal sides. This artificially induced form did not differ from the naturally occurring phenotype as the seasonal forms of P. tharos differ from each other. F, hybrid adults emerging in September and Octo- ber showed some expression of the short photophase phenotype but not to as great an extent as did the P. tharos controls. Phyciodes tharos and P. batesii differ in the length of time required for full embryonic development. Eggs of P. tharos kept at natural outdoor temperatures during early August hatched after 6 days. Those of P. batesii required an additional day or day and a half. The hatching times of the F, hybrids varied within broods from 6 to 7% days. Fertility, viability, and sex ratio Egg fertility and embryonic viability of control broods was very high (Table 2). Fertilizability of eggs was slightly reduced in the F, hybrid << Fic. 1. Parental-type and F, hybrid adults: Row A—P. tharos; B—P. batesii; C—F;, hybrids P. batesii 2 x P. tharos 6; D—F: hybrids P. tharos 2 x P. batesii ¢. Specimens show, left to right, male dorsal, female dorsal, male ventral, female ventral. JOURNAL OF THE LEPIDOPTERISTS SOCIETY (cO'=d) 860° = 8880 (‘S'N) G00’ += 8160 (c00' = d) GSO’ + FS0'0 (100° > d) O13 + FST‘0 (100° > d) FLI + F6F'0 (100 >d) 81¢ = §Z10 (100° > d) €&3° = 91¢°0 (SN) 8S0° = 6860 O10’ + 0660 SE0° = 866'0 620 = S66'0 a[IMoF/peyoye HY (Ol =d) &z0o (SN) Silo’ (eco > d) 99&° (420° 90 (Oi = a) Bae (SN) 676° (cO’= d) 680° (SN) 613 660° LO} LGO" + Gv60 LI8 + 086'0 6606 + 8670 LOS "+ 6660 6061 + 908'0 VSLE Sasso1oyoeg s= SISO LL6B + L160 C6LV + OLL0 GOGI spuqay + 8860 cEGC + 1860 OLEV + 1860 SOVY S[O1QUOD [R}UIIeg DOM an © 2(2 YFXAN X 6 0G) X & YF AN 2 YAXN X 6(P 0G X & YI XN) P(P vg X 6 Y2 XN) X 6 0G 2 Ya XN X 6(P Y2 XN X 6 0G) 2 (2 09d X 6 YFXN) X 6Y2 AN 2 yz ed X & DG 2 YFXN X 6 0G 2 0d X 6 YAXKN ‘AN ‘osnoeidg :pq ‘A'N ‘osnoeiAg :y7 ‘eg ‘ouloy :Y} Prey /e[H10 4 S239 JO ‘ON spooiq JO ‘ON DUnLTY ‘s[O1] -U09 YA SpriqAy FO UOstedutod 0} LofaI POUROIFIUSIS JO S}SoJ, “UOT}LIAVP pAIVpUL}s SoJBOIpPUL ON[PA [BUIOap PUODIG “spooiq sso1oyOR pue ‘priqAy ty ‘(9q) usazqg “gd “(Y}) somy? “d JO (9[H419F/poyoyey ) AYTIqeIA oruoAIquia pue (ple /afjiey) APpQIeF B89 uvsyy “Z ATAVE$E VOLUME 33, NUMBER | 13 crosses. Embryonic viability was heavily depressed in the P. batesii 2 x P. tharos 6 F, hybrid series, but not demonstrably affected in the recip- rocal cross. Embryonic viability of the P. batesii 2 x N.Y. P. tharos é Fy hybrid series may differ from that of the P. batesii 2 X Pa. P. tharos ¢ series (P = .10). Backcross embryonic viability was drastically reduced. Embryonic viability of both F; hybrids and backcrosses showed great variability among broods (Table 2). Post-larval viability (i.e. during pupation and eclosion and as pupae) was greatly reduced in the P. batesii 2 x P. tharos é F, hybrids and to a lesser extent in the reciprocal cross. The decrease in viability was almost entirely during the pupal stage (Table 3). Adult sex ratios of the P. batesii 2 x P. tharos 6 F, hybrid series did not differ significantly from those of the parental broods (Table 3). In the two P. tharos 2 x P. batesii é broods, however, female adults were en- tirely absent (Brood 76-43) or greatly reduced in numbers (to 16.07% in Brood 76-42). Structural abnormalities Structural abnormalities involving segmental irregularities were rela- tively common in the P. batesii 2 x P. tharos 6 hybrid broods, though absent in the parental controls and reciprocal crosses. Between 0 and 15% (no exact counts made) of the larvae of each hybrid brood showed a lack of development of one side of an abdominal segment. The af- fected segment half was both narrower and shorter than the correspond- ing half, often lacked a tubercle, and resulted in the larval abdomen abruptly bending to one side. Two larvae were segregated and observed throughout development. In these, the semental irregularity persisted through the pupa and into the adult (Fig. 2). Both larvae produced ap- parently otherwise normal male adults. F, hybrid males from the cross P. batesii 2 d) F216 VIG (so0'>d) OF FI (SN) 02°0 c 2 vaXs Yi (100° >d) FILS (SN) S&S (CSN) LST CSN) O1L¥P £66 (100 >d La9E (SN) O9'T 8 P YyrX 6 0g cc'ss8 IGG 99°G L8°S¥P 989 oo'8 991 v sjou0o pq 8F'E6 srv'0 LEO 88° 6F SIS! 6S°7 cL 0 9 sjoquol Yj s}[npe s]npe 9}9[du100ur (2 % ueeur) SUISO[Va eevednd eedndoid spooiq ssOIn /se1eds OTA peyddrp UWOISO[O Oe x9S ‘ou [8}0, peed preg JO "ON ‘s[01}U09 YA spliqAy Jo suostIVdut0d 0} 1oFfo1 QOURSIFIUSIS JO S}SAT, “SassoIOyoRq puke spliqAy ‘yf I1ay} pue “(pq ) usayoq *d “(Y1) SOLDYyy “d 97PIS YOK MON Fo uonednd 19}ye pue Sutnp AjTIqeraAur jo (saseyusoied) 9oueplouy = “¢ ATAV | VoLuME 33, NUMBER 1 Fic. 2. P. batesii 2 x P. tharos 6 F, hybrid male adult showing asymmetrical development of abdominal segment (arrow). 16 JOURNAL OF THE LEPIDOPTERISTS SOCIETY TABLE 5. Development times in days from hatching of egg until eclosion of adult for non-diapausing P. tharos and P. batesii control broods, Fi hybrids, and backcrosses. Medians with 99% confidence. See text for rearing conditions. Brood Date Males Females no. hatched N Min—Max Median N Min—Max Median P. tharos (New York )—Natural photophase 76-8 19-26 June WH 32-53 34-36 134 33-52 38-39 76-9 19-23 Jun 216 32-50 34 190 32-41 36-37 P. tharos (New York )—18 h light/24 h 76-38 13 Aug 84 33—44 36-37 95 37-54 42-44 76-40 13 Aug 90 33-46 34-35 79 35-70 37-42 76410 | 13 Aue Bee 38/1 31 | teams =39 P. batesii—24 h light/24 h 76-1 21-25 Jun Di 38-107 49-56 on 43-116 56-57 76-2 Bilis) |own 54 33-105 49-76 49 40-116 55-76 76-3 24-25 Jun 23 40-97 48-55 34 48-134 51-62 76-4 21-25 Jun 30 38-86 41-64 33 38-89 45-55 P. batesii 2 x P. tharos 6—18 h light/24 h 76-5 13-15 Aug 11 40-61 42-59 8 35-54 35-54 76-6 14-18 Aug 61 39-73 49-53 65 33-56 41-46 76-7 15-18 Aug 15 ATT 51-67 D4 34-65 42-47 76-8 20-21 Aug 4 Silas} 51-58 5 39-56 39-56 76-10 15-21 Aug 40 45-74 51-54 on 35-55 39-42 foal 22-29 Aug 28 44-65 49-57 19 38-57 41-48 76-13 22-23 Aug 3 59-69 — iL 50 — (Fl 4 Jun 4 44-84 = 9 3440 34—40 77-2 30 May-2 Jun 24 40-60 42—A7 18 36-48 36—40 77-3 3-5 Jun 10 34-41 34-41 6 32-36 — P. tharos 2 x P. batesii 6 —18 h light/24 h 77-42 15-18 Aug 188 35-62 41-42 77-43 16-17 Aug AO 36-58 38-40 (P. tharos 2 x P. batesii 6)? x P. tharos 6—18h light/24 h 77-26 19-20 Jul 42, 28—46 29-34 54 29-45 37-38 77-34 28-29 Jul 110 30-54 34-37 97 29-37 30-31 77-38 31 Jul-l Aug 79 29-41 33-35 106 32-44 36-37 P. tharos 9 x (P. batesii 2 x P. tharos 4 ) 6 —18 h light/24 h 7728) 29-31) Tul 18 30-645 e0eso 77-32 24-27 Jul 164 27-66 33-35 97 28-65 31-36 other hand, showed significant incidence of larval diapause when reared during June and July on natural photoperiod. This diapause response was entirely facultative, since there was a complete absence of diapause in larvae reared on 18 h light/24 h. Phyciodes batesii reared on natural photoperiod during June and July showed a 100% incidence of larval diapause. Many of those reared on 24 h VoLUME 33, NUMBER 1 17 NUMBER OF ADULTS ECLOSING SES yp 116 134 DAYS UNTIL ECLOSION Fic. 3. Distributions of times required for development of New York State P. tharos, P. batesii, and Fi hybrid broods from hatching of eggs until eclosion of adults. A—P. tharos hatching in late June, natural photophase, B—P. batesii hatching in late June, 24 h light/24 h; C—F, hybrid P. batesii 2 x P. tharos é hatching in early August, 18 h light/24 h; D—F: hybrid P. tharos 2 x P. batesii & hatching in early August, 18 h light/24 h. light/24 h, however, developed without diapause, indicating that at least part of the culture was composed of facultatively diapausing individuals. There was no incidence of diapause in F, hybrid larvae from the cross P. batesii 2 X P. tharos ¢. The reciprocal hybrid, however, had an inci- dence of diapause intermediate between those of the parental species (Table 4). Survival of the F; hybrid larvae during diapause storage was normal compared with that of the parental species. Development periods and eclosion patterns The median development periods from hatching of the egg to eclosion of the adult were significantly longer for non-diapausing P. batésii than for concurrently reared non-diapausing P. tharos from New York State or Pennsylvania (Table 5). In addition, emergences were very scattered, producing a very different eclosion pattern from that of either population of P. tharos ( Figs. 3A and B). 18 JouRNAL OF THE LEPIDOPTERISTS SOCIETY TaBLE 6. Post-diapause development periods of P. tharos, P. batesii, and their Fi hybrid P. tharos 2 x P. batesii 6. Medians with 99% confidence. All larvae re- moved from storage on same day (4 April 1977). Brood Males Females no. N Min—Max Median N Min—Max Median P. tharos (New York) 76-8 105 41-49 43-44 153 42-56 49-50 P. batesii 76-1 1 46 — 8 44-50 44-50 76-2 5 46-48 — i 49-51 49-51 76-3 14 43-A7 43-46 13 45-54 45-48 76-4 4 45-51 — 5 48-53 — P. tharos @ x P. batesii 3 76-42 = 36 72—107 74-83 F, hybrids of the cross P. batesii 2 X P. tharos é showed eclosion pat- terns intermediate between P. tharos and P. batesii, but male develop- ment periods about the same as those of P. batesii. Females of this cross had shorter development periods and tended to emerge before the males, rather than the normal reverse (Fig. 3C). Male development times of the cross P. tharos ? X P. batesii 6 were significantly shorter, though not as short as those of concurrently reared P. tharos. The spread of eclosion times for male adults was much more like that of P. tharos than P. batesii (Fig. 3D). All P. tharos ? X P. batesii 6 F, adults emerging without diapause were males; all those emerging after diapause, females. Resumption of feeding after diapause of these larvae was much delayed after removal from cold storage, and growth was much slower than that of the parental controls (Table 6). DIscussION It is clear from the results that P. tharos and P. batesii are well-differ- entiated species. There are marked differences in phenotypic appearance, voltinism, development rate, and ecology. The adults show strong be- havioral isolating mechanisms during courtship in the laboratory. The heavy reduction in F, hybrid viability and fertility and in backcross em- bryonic viability indicates a high degree of genetic incompatibility be- tween the species. Phyciodes tharos and P. batesii appear to have achieved rather similar phenotypes (compared to P. tharos and P. campestris, for example) by somewhat different genetic means. This is attested to by the wide range of phenotypic variation in the F; hybrid adults and is especially marked in F, females from the cross P. batesii 2? x P. tharos 6. Expression of the VOLUME 33, NUMBER 1 19 dorsal dark pattern elements ranges from as dark or darker than P. batesii to almost as light as P. tharos (Fig. 1). This indicates that the two species look more alike than they actually are. I have discussed at length in another paper (Oliver, in press) the genetic basis of incompatibility effects involved in hybrid breakdown and surveyed the literature on viability of butterfly hybrids. The hybrid in- compatibility shown between P. tharos and P. batesii may involve differ- ences in the genetic control of hormones that direct growth and develop- ment. Disruption of normal hormonal control leads to formation of inviable embryos, abnormal tissue differentiation patterns, lessened fer- tility, lowered ability to pass from one life cycle stage to another, and ab- normal development rates. In general the genetic incompatibility between P. tharos and P. batesii is fairly similar to that shown between P. tharos and P. campestris mon- tana Behr (Oliver, 1978). Both sets of F; hybrids have similar adult eclosion patterns. However, embryonic viability is reduced much more in the series P. tharos x P. batesii than in P. tharos X P. campestris. On the other hand, there is a much greater deficiency of F; hybrid females in the series P. tharos X P. campestris than in P. tharos x P. batesii. These differences and those in ecology, voltinism, and so on indicate that P. batesii and P. c. montana are physiologically quite distinct, and P. batesii probably should not be regarded as an eastern representative of P. cam- pestris. P. batesii is more specialized than either of the other species. It appears to have evolved from multivoltine stock by lowering of the thresh- old of diapause induction to include all naturally encountered photoperiod conditions. Although capable of feeding on at least several species of Aster, it has become closely associated with (though perhaps not re- stricted to) a single species, A. undulatus, and seems in the Northeast to be found only in the rather narrow habitat range of this aster. Other but- terflies have followed similar courses of evolution. Pieris virginiensis Edw., for example, has evolved univoltinism in apparently the same way (Shapiro, 1971; author's unpub. data) and has become restricted to the narrow habitat of its single foodplant, Dentaria diphylla Michx. ACKNOWLEDGMENTS I greatly appreciate the help of Mr. Edward Jennejohn, Manlius, New York, in obtaining laboratory stock of Phyciodes batesii. Mr. Joseph O. Brenneman provided invaluable photographic assistance. LITERATURE CITED Ciarke, C. A. 1952. Hand pairing of Papilio machaon in February. Entomol. Rec. 64: 98-100. 20 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Comstock, J. A. 1930. Egg, larva, and pupa of Phyciodes campestris Behr. Bull. So. Calif. Acad. Sci. 29: 136. Forses, W.T.M. 1960. Lepidoptera of New York and neighboring states. Part IV. Memoir 371, Cornell U. Agric. Exper. Sta. McDunnovucp, J. 1920. Notes on the life history of Phyciodes batesi Reak. (Lepid. ). Canad. Entomol. 52: 56-59. Outver, C.G. 1976. Photoperiodic regulation of seasonal polyphenism in Phyciodes tharos (Nymphalidae). J. Lepid. Soc. 30: 260-263. 1978. Experimental hybridization between the nymphalid butterflies Phyciodes tharos and P. campestris montana. Evolution 32: 594-601. 1979. Genetic differentiation and hybrid viability within and between some Lepidoptera species. Am. Nat. (in press ). Owen, D. B. 1962. Handbook of statistical tables. Addison-Wesley Publ. Co., Reading, Mass. SHAprro, A. M. 1971. Occurrence of a latent polyphenism in Pieris virginiensis (Lepidoptera: Pieridae). Entomol. News 82: 13-16. Journal of the Lepidopterists’ Society 33(1), 1979, 20 TEMPORARY RANGE EXTENSION AND LARVAL FOODPLANT OF DYNAMINE DYONIS (NYMPHALIDAE) IN TEXAS The northern normal limit of Dynamine dyonis Geyer in Texas is Cameron and Hidalgo counties. The occurrence of D. dyonis north of its usual range was noted on 27 July 1966, when I collected a tattered female on the Salado Creek, three miles southeast of the Northeast Preserve, a city park in San Antonio, Texas. The inunda- tions of Hurricane Beulah in September 1967 produced lush vegetation in south Texas and may have caused the invasion of D. dyonis to extend as far north as Collin Co. (18 Sept. 1968, 2 ¢ 6, leg. Edward Reid). Further evidence of this movement are records by the following collectors in other counties in 1968: Gonzales Co., Hidalgo Co. (both M. A. Rickard); Bell Co., San Patricio Co. (both R. O. and C. A. Kendall); Travis Co. (C. J. Durden); Brazos Co. (J. E. Hafernik). The last known record in 1968 was Bexar Co., 23 Nov., 1 @, leg. J. F. Doyle. The total number of D. dyonis taken by collectors in Texas from 27 July 1966 through 23 Nov. 1968 was 134 (68 64, 66 292). To my knowledge no populations remain in central or northern Texas. On 5 May 1968, I observed a female D. dyonis as it fluttered about a trailing plant in a dry creek bed in the Northeast Preserve. The plant, Tragia ramosa Torrey (Eu- phorbiaceae), and the butterfly were caged and placed outdoors at my home in San Antonio, Texas. Twenty-four eggs were deposited that same day. The first larva emerged on 9 May. Only 6 larvae remained on 19 May because of cannibalism. Adults — which emerged were: (31 May) 3 6 6,1 9; (1 Jume) 1 6,1 @. Larvae were collected at the Northeast Preserve site in 1968 and reared on T. ramosa. These larvae were collected on 19 May and pupated 24 May. One adult ( é ) emerged on 5 June. Larvae also were collected on 14 July and pupated between 17 and 18 July. Adults (1 6, 1 2) emerged on 23 and 25 July. Josepn F, Doyxe III, 11839 Monticeto Lane, Stafford, Texas 77477. Journal of the Lepidopterists’ Society 33(1), 1979, 21-28 NOMENCLATORIAL CHANGES IN EUCOSMINI (TORTRICIDAE) RicHARD L. BROWN Department of Entomology, Comell University, Ithaca, N.Y. 14853 ABSTRACT. The genera Kundrya, Norma, and Erinaea are synonymized with Rhopobota. Female genitalia of R. unipunctana, R. dietziana, and R. finitimana are illustrated. Griselda stagnana and G. myrtillana are transferred to Rhopobota. Epi- blema separationis, formerly a subspecies of E. praesumptiosa, is recognized as a species. Female genitalia of both species are illustrated. Notocelia trimaculana, N. illotana, N. culminana, and N. purpurissatana are distinguished from Epiblema. Problems of identification, classification, and evolutionary relationships of the Olethreutinae have persisted in spite of the economic importance of many species in this subfamily. Heinrich’s revision (1923) of the Eucosmini was based principally upon characteristics of wing venation and male genitalia. In recent years Bentinck and Diakonoff (1968), Diakonoff (1973), and Obraztsov (1958-1968), have recognized the value of the female genitalia in differentiating species as well as defining the genera. This paper presents modifications in the classification of selected Eucosmini genera and species as a result of the examination of the female genitalia and other characters. Changes in Rhopobota Rhopobota Lederer, 1859, Wien. Ent. Monat., 3: 366. Type species: Tortrix naevana Huebner [1814-1817], by monotypy. Although Lederer considered R. naevana a senior synonym of Tortrix unipunctana Haworth [1811], R. naevana is now recognized as a junior subjective synonym of R. unipunc- tana. Norma Heinrich, 1923, U.S. Natl. Mus. Bull. 123: 191. [New Synonymy. ] Type species: Epinotia dietziana Kearfott, 1907, by monotypy. Kundrya Heinrich, 1923, U.S. Natl. Mus. Bull. 123: 192. [New Synonymy.] Type species: Kundrya finitimana Heinrich, 1923, by monotypy. Erinaea Meyrick, 1907, Journ. Bombay Nat. Hist. Soc., 18: 141. [New Synonymy. ] Type species: Erinaea chlorantha Meyrick, 1907, by monotypy; a junior subjec- tive synonym of Teras verditer Hampson, 1891 (Diakonoff, 1950). The genera Norma and Kundrya were considered by Heinrich to be close to Rhopobota. Heinrich distinguished Kundrya by a character of the forewing venation, Ry and R; united. These two veins were described as stalked in Rhopobota and approximate in Norma. Rhopobota was separated from the first two on the basis of the porrect socii which are 22 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Zug en gen ae ea 2 Fics. 1-2. Female genitalia including seventh abdominal segment: (1, left) Rhopobota unipunctana (Bellingham, Washington; USNM 17763); (2, right) Rhopo- bota dietziana (Ethel, Arkansas; R. L. Brown prep. 683). Scale line = 1 mm. apically fused. Meyrick (1907) described Erinaea based upon a species from Ceylon, and did not indicate a relationship with Rhopobota. Unique characters of the female genitalia of all the type species and a reassessment of previous distinctions provide the basis for synonymizing Erinaea, Norma, and Kundrya with Rhopobota. The most distinctive character is the sclerotization of the sides and base of the corpus bursae and distal area of the ductus bursae. A separate sclerotized band around the ductus bursae is located near the colliculum at the inception of the VoLuME 33, NUMBER 1 93 Fic. 3. Female genitalia including seventh abdominal segment of Rhopobota finitimana (Falls Church, Virginia; USNM 17762). Scale line = 1 mm. ductus seminalis. The colliculum is sclerotized ventrally and extends be- yond the lamella antevaginalis. Both the lamella antevaginalis and lamella postvaginalis are well developed and may be fused with or separate from the seventh sternite (Figs. 1, 2, 3; Clarke, 1958, pl. 169). These charac- teristics are also shared by the Asian species, R. eclipticodes (Meyrick) and R. microrrhyncha (Meyrick), as figured by Clarke (1958, pl. 170) and the Palearctic species, R. ustomaculana (Curtis), figured by Bentinck and Diakonoff (1968, fig. 189b, c). The eighth tergite which surrounds the papillae anales possesses scales as well as simple setae in the three North American species, dietziana, finitimana, and unipunctana. Scales may be present or absent on the eighth tergite among the species of the related genera, Epinotia, Ancylis, and Chimoptesis. 24 JOURNAL OF THE LEPIDOPTERISTS SOCIETY The male genitalia of finitimana, dietziana, and unipunctana are simi- lar in having a bifurcate uncus with widely separated arms that are fin- gerlike and weakly sclerotized. The socii are long, porrect, and densely setose, separate in finitimana and dietziana and fused apically in uni- punctana. The gnathos of Rhopobota is reduced and weakly sclerotized medially. The aedeagus is short in finitimana, longer in dietziana and uni- punctana. The male of chlorantha has not been examined. The condition of veins Ry and R; in the forewing represents a transi- tion from approximate in dietziana (n=17) to stalked in unipunctana (n = 37) and united in finitimana (n =7). The length of the stalk varies among the specimens of unipunctana examined. These veins are stalked in the type of chlorantha, as illustrated by Clarke (1958). All three North American species of Rhopobota feed on Ilex ( Aquifolia- ceae). R. unipunctana also feeds on Vaccinium (Ericaceae) and is a pest of cranberry, Vaccinium macrocarpon Aiton (Heinrich, 1923). The lar- vae of unipunctana and dietziana were described by MacKay (1959) and were considered to be closely related. Analysis of photographs and drawings of the genitalia (Bentinck and Diakonoff, 1968; Pierce and Metcalfe, 1922) provides evidence for syn- onymizing Griselda stagnana (Denis and Schiffermueller) and G. myr- tillana (Westwood) with Rhopobota. Obraztsov (1945) included these two species in Griseida Heinrich, based on the presence of a costal fold in the forewing of the male. Powell (1964) provisionally retained these two in Griselda but indicated that they might not be congeneric with the type species, G. radicana (Walsingham), the sole North American spe- cies. The male and female genitalia of G. radicana are similar to those of Epinotia hopkinsonana (Kearfott) and E. subviridis Heinrich, all coni- fer feeders. The presence of the costal fold in stagnana and myrtillana does not justify their separation from the other Rhopobota species which lack one. Secondary sexual characteristics, such as the costal fold of the male, are seldom of generic value, as is emphasized by Diakonoff (1973) in his study of the South Asiatic Olethreutini. My investigations of Epinotia also show the costal fold may be present or absent in closely related spe- cies. Rhopobota stagnana ({Denis and Schiffermueller]). [New Combination.] Tortrix stagnana [Denis and Schiffermueller], 1775, Ankundung eines Systemati- schen Werkes von den Schmetterlingen der Wienergegend, p. 131. Tortrix fractifasciana Haworth, [1811], Lepidoptera Britannica, 3: 466. anaes fractifasciana, Pierce and Metcalfe, 1922, Genitalia British Tortricidae, p. 75, pl. 26. Griselda fractifasciana, Obraztsov, 1945, Zeitschr. Wiener Entomol. Ges., 30: 33-34. The female of R. stagnana has a corpus bursae with sclerotized sides. VOLUME 33, NUMBER 1 25 The sterigma is similar in shape to that of R. dietziana, both lamella post- vaginalis and antevaginalis are well developed. A sclerotized band around the ductus bursae is located near the colliculum. Scales are pres- ent on the eighth tergite, as shown in the specimen figured by Bentinck and Diakonoff (1968). R. stagnana occurs in England and central Europe (Meyrick, 1895), and feeds on the flowers and seeds of Scabiosa columbaria L. (Dipsa- ceae ) in England ( Ford, 1949). Rhopobota myrtillana (Westwood). [New Combination. ] Sericoris myrtillana Westwood, In Humphreys and Westwood, 1845, British Moths, 2: 146, pl. 89, fig. 15. Grapholitha vacciniana Zeller, 1846, Isis von Oken, p. 248. Rhopobota vacciniana, Pierce and Metcalfe, 1922, Genitalia British Tortricidae, p. fospl. 26. Griselda vacciniana, Obraztsov, 1945, Zeitschr. Wiener Entomol. Ges. 30: 34. Griselda myrtillana, Bradley, 1959, Entomol. Gazette, 10: 72, pl. 11. The female has a similar, although weaker, sclerotization of the corpus bursae than that described above. The sclerotized band around the ductus bursae is located near the colliculum. Both the lamella antevag- inalis and postvaginalis appear well developed and separate from the seventh sternite. Scales on the eighth tergite probably are present, but are not evident in the figures. The male has a rudimentary clasper on the valva, similar to R. wnipunctana and ustomaculana. R. myrtillana occurs through north and central Europe and the British Isles. The larvae feed on Vaccinium myrtillana L. (Ericaceae) (Mey- rick, 1895). Changes in Epiblema Huebner Epiblema separationis was described by Heinrich (1923) as a sub- species of praesumptiosa Heinrich but is raised to the species level in this paper. Heinrich characterized separationis by its smaller size, the absence of brown spots on the inner margin of the forewing ocellus and vein 1A, and the more rounded cucullus. E. separationis has a forewing expanse of 9-11 mm (n=10); praesumptiosa has a forewing expanse of 14-17 mm (n=11). However, the maculation of the forewing varies with the brown spots present or absent in each. The most conspicuous differences are found in the female genitalia and seventh abdominal segment (Figs. 4, 5). E. separationis lacks signa on the corpus bursae, whereas praesumptiosa has two well developed signa. The papillae anales of separationis are widened apically but those of praesumptiosa are nearly uniformly wide. The lamella postvaginalis is more setose and emarginate posteriorly in separationis. The sclerotiza- 6 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 4-5. Female genitalia including seventh abdominal segment: (4, left) Epi- blema separationis (San Benito, Texas; USNM 17735); (5, right) Epibelma prae- sumptiosa (Brownsville, Texas; USNM 17708). Scale line = 1 mm. tion of sternite VII is relatively narrower and longer in separationis than in praesumptiosa. Both species lack sclerotization of the ductus bursae; however, the ductus bursae of praesumptiosa is distinct in being striate. Both species possess setae but lack scales on tergite VIII. E. separationis possesses a tri-lobed eversible pouch ventrally, anterior to the papillae anales. This pouch can be everted by forcing alcohol with a syringe into the anterior opening of the seventh segment after it has been separated from the rest of the abdomen. The pouch appears to be glandular under high magnification. VOLUME 33, NUMBER 1 27 The male genitalia of separationis differ from those of praesumptiosa in having shorter setae on the corona of the rounded cucullus, a smaller rudimentary clasper, and a truncate uncus. Male genitalia of both species have been figured by Heinrich (1923). E. separationis appears to be closely related to discretivana (Hein- rich). Both species lack signa and sclerotization of the ductus. E. prae- sumptiosa is thought to be most closely related to numerosana (Zeller ), grossbecki Heinrich, abruptana (Walsingham), deflexana Heinrich, and exacerbatricana Heinrich. This species group shares the derived char- acteristic of long coronal setae on the cucullus. Sclerotization of the duc- tus is also lacking in this group. MacKay (1959) included insidiosana Heinrich in this group. However, insidiosana has short coronal setae on the cucullus and a sclerotized band around the ductus bursae. The distribution of separationis within the United States is limited to southern Texas and Florida. The larvae, described by MacKay (1959), have been reared from galls of Borrichia frutescens (L.) (Compositae ) in both localities. E. praesumptiosa is limited to southern Texas; the host plant has not been identified. Four Notocelia species, trimaculana (Haworth), illotana (Walsing- ham), culminana (Walsingham), and purpurissatana (Heinrich), were included in Epiblema by Heinrich (1923). N. suffusana, a junior subjec- tive synonym of trimaculana, was reassigned to Notocelia from Epiblema by Bentinck and Diakonoff (1968). MacKay (1959) described the ex- ternal feeding larvae of culminana and trimaculana and considered them to be a separate genus but retained them in Epiblema. Notocelia is distinguished from Epiblema by the presence of two non- deciduous cornuti at the apex of the aedeagus and a well-developed lamella antevaginalis of the female. The antevaginal plate is reflexed outward producing a projecting ostium. The ductus bursae of the female is lightly sclerotized from the ostium to the heavily sclerotized band at the inception of the ductus seminalis. ACKNOWLEDGMENTS I wish to acknowledge with my appreciation the following individuals and institutions who have provided specimens used in this study: D. R. Davis, U.S. National Museum of Natural History, Smithsonian Institu- tion; J. G. Franclemont, Cornell University; Museum of Comparative Zoology, Harvard University; W. W. Moss, Academy of Natural Sciences of Philadelphia; E. Munroe, Canadian National Collection; F. H. Rindge, American Museum of Natural History. Drs. Franclemont, William Mil- ler, and Jerry Powell have read parts of the manuscript and made sug- gestions for its improvement. I thank Amy Louise Trabka for the draw- 28 JOURNAL OF THE LEPIDOPTERISTS SOCIETY ings. This study was supported in part by National Science Foundation Grant DEB 77-15808 and a Grant-in-Aid of Research from the Society of Sigma Xi. LITERATURE CITED Bentinck, G. A. G. anp A. Diaxonorr. 1968. De Nederlandse Bladrollers (Tor- tricidae). Mono. Ned. Entomol. Vereen., No. 3. 201 p., 33 pl. Crarke, J. F. G. 1958. Catalogue of the type specimens of Microlepidoptera in the British Museum (Natural History) described by Edward Meyrick. Vol. 3. British Museum (Natural History), London. 600 p. Draxonorr, A. 1950. The type specimens of certain oriental Eucosmidae and Carposinidae. Bull. Brit. Mus. (Nat. Hist.) Ent. Vol. 1(4): 275-300, 8 pl. _ 1973. The South Asiatic Olethreutini (Lepidoptera, Tortricidae). Zool. Mono. Rijksmus. Nat. Hist., No. 1. i—xii, 700 p. Forp, L. T. 1949. 0.99). Allele frequencies were estimated directly from the phenotype frequencies for each locus. The formula for heterozygosity of an individual locus is H = 1-p,? where p equals the frequency of an allele at the locus. An estimate of the genetic distance separating the two populations was made using the method of Nei (1975) which provides an estimate of the mean number of codon differences per structural gene locus. This index may take values from 0, representing populations with no alleles in com- mon, to 1, representing populations with identical frequencies of the same alleles. RESULTS Mark-recapture analysis of population size The calculated estimates of the parameters P,, ¢, and B; for each day are given in Table 1, and the daily estimates of population size ( P,) are shown in Figure 1. These data indicate two distinct peaks in population size: one on 10 July and a second, smaller peak on 14 July. Using a method of graphical estimation (Southwood, 1966) the area under the population estimate curve was calculated and this total, divided by the 32 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY TasBLE 1. Results of analysis of mark—recapture data. No. Proportion marked No. of Standard of animals Survival new Total error of recaptures at risk rate animals population estimation Day A A A A A ho ae (i) (a; ) (M;) (9;) (B;) (P,) (q Vee) ) 1 — 0 158 — — 0.0 2 .136 16.6 .736 98.3 W221) TOS 3 129 24.0 IL JU) 193.3 186.0 117.0 4 nlip2 Bla) 941 330.7 378.3 Iso) 5 sll 75.8 1.68 166.1 682.9 396.4 6 .210 178.0 .296 —8.5 847.6 BE 4 7 .250 60.3 913 —15.9 2AN.2 139.0 8 382 77.0 .410 223.6 201.6 70.5 9 AS) 39.4 Wo 1F —30.9 305.4 202.3 10 Pale eee 136 28.8 MT as) eal ith .194 (a3) — — (65.7) 1.0 12 093 (03) sa nae (32.3) 2.0 13 .066 (01) — — (15),23) 2.0 14 .000 (00) —— — — — average adult lifetime (derived from the mean survival rate ¢;), provided an estimate of total population size of 2,912 insects. No L. eurydice in- dividuals were ever netted in the woods, nor were any L. appalachia caught in the Pothole area. A similar mark-recapture program was begun with L. appalachia at a site in the woods about 100 meters from the L. eurydice population. This particular site was chosen because of the consistently higher densities of butterflies observed there in comparison to the woods in general. How- ever, this study was abandoned because of an extremely low rate of re- captures and a general scarcity of L. appalachia individuals. In the first four days of the study, 29 specimens were netted, marked and released in 2 manhours of search; only one of these marked individuals was ever re- captured. It became apparent that the site represented only a temporary aggregation point for L. appalachia, and not a stable population unit. In- dividual insects tended to move extensively through large areas of the wooded bog basin, and several similar high-density sites were subsequent- ly found. Electrophoretic analysis of protein polymorphisms The frequencies of all alleles for the eight enzyme-synthesizing gene loci for each species are shown in Table 2. In Lethe eurydice, one locus (IDH-II) was represented by a single al- lele in all individuals. Two loci were dominated by single alleles with frequencies greater than 99 percent (GOT and a-GPD). Two loci were VOLUME 33, NUMBER 1 33 900 800 600 A Pop. estimate (P; ) re Seow 7G 9) OMe i213, 14. 15 16 17 18 19 20 21 22 Date of sample (July 1975) Fic. 1. Daily estimates of population size (P, ) for Lethe eurydice at McLean Bogs Reserve, Pothole area. dominated by single alleles with frequencies greater than 95 percent but less than 99 percent (MDH-I and MDH-II). Of the other three poly- morphic loci, one locus had two alleles represented in the population (IDH-I), one locus had three alleles (PHI), and one locus had four al- leles (PGM ). The mean heterozygosity for the population is 0.175. The heterozygosi- ties of individual loci are listed in Table 3. The genotype frequencies at all loci were not signifcantly different from those predicted by the Hardy- Weinberg expression using the x? goodness-of-fit test (Sokal and Rohlf, 1969 ). Lethe appalachia showed a similar pattern of electromorphic variation. 34 JOURNAL OF THE LEPIDOPTERISTS SOCIETY TasLE 2. Results of analysis of allozyme variation. Lethe eurydice Lethe appalachia : Results of Allele Sample Allele Sample G-test Locus Allele frequency size frequency size (p= GOT ar — 320 .012 162 N.S. a id, .988 | b .003 — PHI a .103 340 — 164 <.001 b .679 61 c .240 A421 d — .018 PGM a orl 308 488 162 <.001 - b 046 .265 c me, 204 d .006 043 | MDH-I a .034 320 .006 162 : 036 b .966 .994 . MDH-II a 975 320 1.00 162 | .O1 b .025 — | @-GPD b 993 320 1.00 162 INES: c .007 — IDH-II a .876 322, .988 162 <.001 b me .O12 IDH-II a 1.00 226 1.00 150 N.S. Three loci (MDH-II, a-GPD and IDH-II) were represented in the pop- ulation by single alleles. One locus (MDH-1I) was strongly dominated by an allele with a frequency greater than 99 percent. Two loci (GOT and IDH-I) were dominated by single alleles with frequencies between 95 and 99 percent. The remaining two loci had three (PHI) and four (PGM) alleles present in the population. | TaBLE 3. Estimates of heterozygosity per locus. Locus Lethe eurydice Lethe appalachia GOT .006 024 PHI .470 DUO men PGM 083 .648 MDH-I .066 .012 MDH-II .048 .000 a-GPD 014 .000 IDH-I pad 2 024 IDH-II .OO0 .000 Mean nus 2 VOLUME 33, NUMBER 1 BS The mean heterozygosity of the L. appalachia pupulation was 0.152. Heterozygosities of individual loci can be found in Table 3. No locus showed significant deviation from Hardy-Weinberg expectation using the x” goodness-of-fit test. The two species are highly significantly different (p < .01, G-test) in allele frequencies at four of the eight loci examined, and significantly different (p = .036) at another. These data clearly indicate reproductive isolation in sympatry. Using the genetic distance measure of Nei (1975) the distance separating these species equals 0.145, a figure well within the rather wide range of available estimates of distances separating sibling pairs. (Nei’s measure can range between a maximum distance of 1.00, representing no alleles in common, to a minimum of 0.0, representing total identity. ) A very small number of individuals (six) of Lethe portlandia captured at the McLean Bogs Reserve were analysed at the same eight en- zyme loci. Although the sample size was insignificant for statistical pur- poses, the electromorphs at six of the loci tested represented clearly dif- ferent mobility classes from those present in either L. appalachia or L. eurydice. DIscuUSSION The results of this study fully agree with the conclusion of Cardé, Sha- piro and Clench (1970) and Shapiro and Cardé (1970) that the butter- flies Lethe eurydice and L. appalachia are distinct, although very simi- lar, sibling species. Both the mark-recapture study and the analysis of genetic variation demonstrate that the two are genetically isolated and habitat-segregated, at least in the locality studied. The mark-recapture study indicates that the two species strongly resist crossing over from the open to the wooded habitat, or vice versa. It also suggests that the two populations have contrasting spatial structure. Lethe eurydice occupies a small, isolated, concentrated, uniform patch of acceptable habitat, while L. appalachia occupies a more “fine-grained” habitat, moving extensively between more fragmented and diffuse sites of maximum acceptability. LITERATURE CITED Carpe, R. T., A. M. SHarmo & H. K. Ctencw. 1970. Sibling species in the euryd- ice group of Lethe (Lepidoptera: Satyridae). Psyche 77: 70-103. CueERMock, F. G. 1947. Notes on North American Enodias (Lepidoptera). En- tomol. News 58: 29-35. FreLp, W. L. 1936. New North American Rhopalocera. J. Ent. Zool. (Pomona College) 28: 17-26. Jotty, G.M. 1965. Explicit estimates from capture-recapture data with both death and immigration—stochastic model. Biometrika 52: 225-247. 36 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Ne1, M. 1975. Molecular population genetics and evolution. North Holland Re- search Monographs, Frontiers of Biology, Vol. 40. North Holland Publ. Co., Amsterdam-American Elsevier, N.Y. and Oxford. SELANDER, R. K., M. H. Smiru, S. Y. YANG, W. E. JoHNson & J. B. GENTRY. 1971. Biochemical polymorphism and systematics in the genus Peromyscus. I. Varia- tion in the old-field mouse Peromyscus polionotus. Stud. Genet. VI: 49-90 (Univ. Texas Publ. 7013). SHapiro, A. M. & R. T. Carpe. 1970. Habitat selection and competition among sibling species of Satyrid butterflies. Evolution 24: 48-54. SokaL, R. R. & F. J. Rontr. 1969. Biometry. W. H. Freeman and Co., San Fran- cisco. SoutHwoop, T. R. E. 1966. Ecological methods, with particular reference to the study of insects. Methuen and Co., Ltd., London. Journal of the Lepidopterists’ Society 33(1), 1979, 36 NOTES AND NEWS THE JAMES H. BAKER COLLECTION The James H. Baker collection of insects has been received at the Department of Entomology, Smithsonian Institution. Baker’s material consists of slightly more than 24,400 specimens, primarily Lepidoptera, but also contains many Coleoptera and Diptera. Among the Lepidoptera the collection is especially rich in Geometridae, Baker’s specialty. Most of the specimens in this collection are from eastern Oregon, but Baker enjoyed a wide correspondence and traded considerably; consequently there is a rather liberal sprinkling of moths and butterflies from localities other than Baker’s home state. Baker also collected in such places as Arizona, Idaho, and Nevada, so there is a nice representation of species from those areas. J. F. Gates Ciarke, Dept. of Entomology, U.S.N.M.N.H., Smithsonian Institution, Washington, D.C. 20560. | Journal of the Lepidopterists’ Society 33(1), 1979, 37-41 THE LARVA OF CRYPTOCALA ACADIENSIS (BETHUNE) (NOCTUIDAE)! TrmotHy L. McCase New York State Museum, Albany, New York 12234 ABSTRACT. The mature larva of Cryptocala acadiensis (Beth.) (Lepidoptera: Noctuidae) is described. Apocynum androsaemifolium LL. was found to be an ac- ceptable food plant; additional acceptable and unacceptable food plants are listed. Eggs were laid singly and the minimum developmental time from egg to adult was 116 days in the laboratory. This is a much shorter developmental time than what would be expected in nature as the mature larva is presumed to overwinter. In the type description, Cryptocala acadiensis (Bethune, 1869) was placed in the genus Anarta Ochsenheimer because of its small size and black bordered, yellow hind wings. Benjamin (1921) recognized it as a synonym and the older name for Rhynchagrotis gilvipennis (Grote) and erected Cryptocala for it. Prior to this paper nothing was known of the life history. Mikkola and Jalas (1977) report that Rumex is the host plant of the very close (if actually distinct) species, Noctua (Cryptocala) char- dinyi Boisduval. Cryptocala acadiensis occurs from Labrador south to Massachusetts and west to the Pacific. Its flight period is from July to August (Forbes, 1954). A female of C. acadiensis was taken at ultraviolet light on 16 July 1977 in the Adirondacks, 6 mi east of Indian Lake, 1820 ft, Hamilton Co., New York. The following day 39 eggs were laid singly in a holding jar. The larvae eclosed in seven days and were offered a selection of plants. The first instar larvae initially accepted the blossoms of Hypericum perforatum L.., Sagittaria latifolia Willd., the blossoms and leaves of Apo- cynum androsaemifolium L., and the leaves of Prunus virginiana L., Achil- lea millefolium L., Sambucus canadensis L., and Spiraea latifolia ( Ait.) Borkh., but the limited feeding and continual wandering of the first instar larvae indicated that most of these plants were unacceptable. Only A. androsaemifolium was continuously utilized by the first instar larvae and all later instars were reared to maturity on A. androsaemifolium leaves. Plants refused by the first instar larvae include: Rubus idaeus L., Ame- lanchier laevis Wieg., Pteridium aquilinum (L.) Kuhn, and Vaccinium myrtilloides Michx. The larvae feeding on A. androsaemifolium remained healthy (no dis- ease) but grew slowly. They started pupating on 12 September 1977 and adults began to emerge on 10 November 1977. Presumably, this species would normally overwinter as a mature larva and pupate in the spring. 1 Published by permission of the Director, New York State Museum, Journal Series No. 254. 38 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Fic. 1. Cryptocala acadiensis, Adirondack Mts., New York: photograph of living, ultimate instar larva. The larvae were cultured in tins in total darkness (interrupted only by the addition of fresh leaves every two days) at 22°C (+8°). These artificial conditions speeded development as is the case with many spe- cies which normally feed at night. The early pupation is also typical of many Lepidoptera which have a non-obligatory diapause. The illustrations that accompany the descriptions of the last larval in- star were drawn to scale using an ocular grid. All scale lines represent 0.5 mm. The terminology and abbreviations follow Godfrey (1972) with the exception of coronal punctures. General (Fig. 1). Head 2.06-2.23 mm wide. Total length 26-30 mm. Abdominal prolegs present on third through sixth segments. Head and body smooth. Setae simple, insertions in small, flat, black tubercles. Spiracle A-8 0.28 mm high. Seta D1 0.40 mm long. Coloration (living material). General head and body color light brown; a light middorsal line and a light line just below D2 on all segments; venter below spiracles — light; spiracles also light. _ Head (Fig. 2). Epicranial suture 1.04 mm long. Height of frons 1.40 mm. Ad- frontal punctures (AFa) anterior and second adfrontal seta (Af-2) posterior to apex of frons. Coronal punctures 5 (Ca-5), posterior setae 1 & 2 (Pl & P2), and lateral seta (L.) each arise from a black pigmented spot. Ocellar interspaces between Ocl- Oc2 and Oc2—Oc3 each equal to diameter of Oc2; Oc3—Oc4 one-third diameter of Oc4; Oc4—Oc6 approximately 1.5 times diameter of Oc4; Oc4—Oc5 2.0 times the diameter of Ocd4. Mouthparts. Hypopharyngeal complex (Fig. 3): spinneret subsequal to labial palpus, apex bearing short spinules; stipular seta (S) at anterior dorsal apex of pre- VoLUME 33, NUMBER 1 39 Fics. 2-3. Cryptocala acadiensis, Adirondack Mts., New York: 2, frontal aspect of head; 3, oral aspect of left mandible. AO JOURNAL OF THE LEPIDOPTERISTS SOCIETY Ji Y/’ 4) \ ee iMG Wij i Y <7) ) | Wa Le ye Lib if Wh Wy We Iics. 4-6. Cryptocala acadiensis, Adirondack Mts., New York: 4, left aspect of hypopharyngeal complex; 5, left dorsolateral setal arrangement of prothorax; 6, left dorsolateral setal arrangement of first abdominal segment. VoLUME 33, NUMBER 1 Al mentum; distal region of hypopharynx covered with fine spines; proximolateral region bearing single row of about 8 distinct spines. Mandible (Fig. 3): inner ridges dis- tinct, with prominent basal tooth; sixth outer tooth low, divided into smaller subteeth. Thoracic segments. Prothorax (Fig. 5): cervical shield weakly sclerotized, with two punctures between setae D-1 and XD-1, one puncture between XD-1 and XD-2 (located two-thirds distance from XD-1 to XD-2) and one puncture along posterior margin of shield behind D-2; SD-1 and SD-2 on same pinaculum; L1 and L2 also on same pinaculum; SV-1 and SV-2 separate. Meso- and metathoracic segments with a non-setiferous puncture on same pinaculum as SD-1. Abdominal segments. Ab-] (Fig. 6): two subventral setae (SV-1 & SV-3); Ll dorsal to spiracle. Ab-2—6 with three subventral setae. Ab-7 and Ab-8 with only one setae in subventral group. Crochets: uniordinal, 16-17 per third abdominal proleg, 18—21 per fourth, 20-24 per fifth, 22-26 per sixth. Material examined. Thirteen specimens, 6 miles east of Indian Lake, 1820 ft., lat. 43°45’30” long. 74°10’14”, Hamilton Co., New York, 10 September 1977, from ova of female collected, determined, and reared by T. L. McCabe. The basal mandibular tooth and the spinule-tipped spinneret seem to indicate a relationship to Ochropleura plecta (L.) (see Crumb, 1956, for larval description), but this is not substantiated by the male genitalia (figured by McDunnough, 1928). ACKNOWLEDGMENTS I thank Drs. J. G. Franclemont and G. L. Godfrey for reviewing this manuscript. Mr. Stanley J. Smith verified the plant determinations and voucher plant specimens are deposited in the New York State Museum. Funds have been provided by the New York State Museum, Albany, New York. LITERATURE CITED BENJAMIN, F. H. 1921. A study of the noctuid moths of the genera Lampra Hbn. and Cryptocala, gen. nov. Bull. South. Calif. Acad. Sci. 20: 133. BETHUNE, C. J. S. 1869. A new species of Anarta from Nova Scotia. Trans. Nova Scotia Inst. Nat. Sci. 2(3): 84. Crus, S. FE. 1956. The larvae of the Phalaenidae. U.S. Dept. Agric. Tech. Bull. 1135: 99. Forses, W. T. M. 1954. Lepidoptera of New York and neighboring states, Cornell Univ. Agri. Exp. Sta. Mem. 329: 71. Goprrey, G. L. 1972. A review and reclassification of larvae of the subfamily Hadeninae (Lepidoptera, Noctuidae) of American North of Mexico. U.S. Dept. Agric. Tech. Bull. 1450, 265 pp. MrKxkoLa, K. & I. Jauas. 1977. Yokkoset 1 [Noctuidae of Finland]. Otava, Hel- singssa. 256 pp. McDunnoucu, J. H. 1928. A generic revision of North American Agrotid moths. Nat. Mus. Canada Bull. 55, 78 pp. Journal of the Lepidopterists’ Society 33(1), 1979, 42-49 COURTSHIP BEHAVIOR OF THE CHECKERED WHITE, PIERIS PROTODICE (PIERIDAE) RonALD L. Rutowski Department of Zoology, Arizona State University, Tempe, Arizona 85281 ABSTRACT. Courtship behavior leading to copulation is described for the check- ered white, Pieris protodice, from film records of courtships with perched virgin fe- males and written records of courtships elicited by releasing virgin females near free- flying males. Temporal and sequential patterns of successful courtships follow those seen in other pierid butterflies in that the interactions are simple and rapid, averaging about 3 seconds in duration. Unsuccessful courtships are also described and conform to patterns documented for other pierids. The checkered white differs from con- familials only in that there is no abdominal extension by females during successful courtship and after copulation a post-nuptial flight may occur. The possible func- tions of post-nuptial flights are discussed. Although ethologists have been interested in butterfly courtship for many years (Scott, 1974; Silberglied, 1975) few quantitative studies of the temporal and sequential structure of butterfly courtship can be found in the literature. Detailed information is essential for comparative studies of butterfly courtship which in the past have proven useful in deducing the ecological factors influencing the structure of both successful and unsuccessful courtships (Brower et al., 1965; Pliske, 1975; Rutowski, 1978a). In this paper the courtship of the checkered white, Pieris protodice Boisduval and Le Conte, is described in detail. Attention will be focused on the structure of successful courtships so that this account can be com- pared to previous studies of a similar nature, with special reference to the function of post-nuptial flights in butterflies. It should be noted that a brief description of P. protodice courtship was given in Abbott (1959). However, Shapiro (1970) has pointed out the interpretational problems surrounding that description, so it will not be dealt with here. METHODS All observations were made and all animals obtained from March through June in 1976 and 1977 at the Arizona State University Field Lab- oratory, Tempe, Arizona. To obtain virgin females, eggs were collected by placing field-caught females in tubular cheesecloth cages (1 m high, 0.25 m diameter) with cuttings of the local larval foodplant, Sisymbrium irio L.. The larvae from these eggs were reared to adulthood in the labora- tory on S. irio. The humidity and light regimen in the rearing area were not regulated and variable. VoLUME 33, NUMBER 1 43 All behavioral observations were made on clear days between 0900 and 1500 when the butterflies were most active. Naturally-occurring in- teractions between males and between males and females were observed and the form, outcome, and duration (as timed with a stopwatch) of each recorded. Similar records were made of interactions initiated by releasing virgins near free-flying males. During late May 1977, films of successful courtships were made at 24 and 70 frames per second using a Beaulieu 4008 ZM II super-8 movie camera. In all cases a virgin female was placed near the top of an ex- posed perch in a large patch of S. irio where there was a dense popula- tion of flying males. By activating the camera as a male approached a complete record of the courtship could be obtained. Temporal and se- quential data were gathered via frame-by-frame analysis of the film records. Where pertinent, summary statistics are given as mean + standard error of the mean. RESULTS I. Successful Courtship Using 38 virgin females, 27 successful courtships (= ending in copula- tion) were recorded on film and 58 were observed after releasing virgin females near free-flying males. Temporal data from the films will be summarized first to give a general impression of the structure of P. proto- dice courtship. A. Film records Successful courtship with a perched female began when the male's wings or legs made physical contact with the female. In 67 percent of the filmed courtships this contact was made with the legs as the male alit on the female and immediately walked toward her thorax. In all other film records the male broke and then renewed contact with the female at least once, and as many as four times, before positioning himself on the female’s thorax. Once on the female's thorax in a head-to-head orientation the male curled his abdomen out from between his hindwings and inserted the tip between the female’s hindwings, often after several unsuccessful attempts. It was not possible to determine when genital contact was made. The courtship ended when the male stopped moving his wings and assumed a quiescent posture. Males showed no preference for the side of the female from which they effected copulation (13 right vs. 14 lest — 0.057, p — 0:8). Fig. 1 shows the temporal pattern of the following major events in a successful courtship with a perched female: male contacts female, male 44 JOURNAL OF THE LEPIDOPTERISTS SOCIETY contacts ) oO begins probing(n=13) S| abdomen between ) wings (n=6) & stops moving (n= 27) a —— i SS aie O 0.5 1.0 1.5 20 22 3.0 3.5 40 125 SECONDS AFTER BEGINNING OF COURTSHIP (=c'CONTACTS 9) Fic. 1. Temporal relationships between the major behavioral events in the success- ful courtship of P. protodice. The mean time of occurrence, standard error (open bar), range line, and sample size (n) are shown for each behavior. See text for details. begins probing (abdomen appears from between hindwings), male ab- domen between female hindwings, male stops moving. There were no observed variations in the sequence of these events. In 45 percent of the courtships the female performed a low amplitude flutter response. Of these, 25 percent began before the male first contact and 75 percent after, and they ended most often before the male began probing but sometimes not until he had inserted his abdomen between the female’s hindwings. Detailed quantitative information on these points was difficult to gather because of 1) the viewing angle of many of the film records and 2) the low amplitude of the flutter response. In any event, flutter responses had no significant effect on the duration of courtship (courtships with flutter response: 2.8 + 0.37 sec, n = 12; courtships without flutter response: 3.86 + 0.93 sec, n = 15; t = 0.968, p = 0.66). Shortly after coupling some males attempted to fly away from the perch carrying the female. In the 11 courtships where these attempts were filmed the males broke contact with the perch on the average 3.71 + 0.586 sec (range = 1.71-7.34 sec) after they had stopped moving their wings. In 8 of these attempted “post-nuptial flights” (Brower et al., 1965) the female did not release her grip on the perch and the male dangled from the female’s abdomen at the end of the attempt. In the other 3 cases the male flew off carrying the female to a distant and usually less exposed perch. Systematic data on these points or on the duration of successful post-nuptial flights was not collected. VoLuME 33, NUMBER 1 45 wl? i Ss Q Tying a 8 Mmedian=7 sec es a= SxS) =/6 O U4 5 ieee 2 Bib peere|n Y) lid] 10) O O Ea) VY) Le perched O median=6 sec aia n=9 A 3 ele 2 6 an on Mic. Sac... 2 eee DURATION (SEC) Fic. 2. A frequency histogram of the duration of successful courtships with perched and flying females. Courtships were elicited by releasing females near free- flying males and were timed with a stopwatch. n = number of courtships. B. Written records The successful courtships elicited by releasing virgin females near free- flying males were timed beginning when the two animals arrived within 1 to 2 cm of each other. The distance at which these observations were made did not allow the use of contact as a criterion for when courtship began. This coupled with the lack of precision in determining when courtship ended resulted in the apparent greater length of these court- ships relative to those timed from film records. These courtships were divided into two groups depending on whether the female was flying or perched when the male arrived within 1 to 2 cm of her (Fig. 2). The dif- ferences between these two types of courtships in the distribution of durations and median durations were slight suggesting that aerial com- ponents were of short duration. In fact, flying females typically landed immediately when approached by a male. The form and sequence of all 46 JoURNAL OF THE LEPIDOPTERISTS SOCIETY ground components of these courtships were exactly like those described from the film records. Post-nuptial flights were attempted in 27 of the 58 successful courtships with released females. This should not be taken as an absolute indicator of the frequency of attempted post-nuptial flights because no firm proto- col was established for how long a copulating pair should be watched after the courtship to wait for the onset of a post-nuptial flight attempt. Of the observed attempts 9 were unsuccessful, i.e., the female did not re- lease her grip on the perch where copulation occurred. The duration of copulation was timed for 15 pairs. Thirteen of these pairs copulated for less than 35 min (27.2 +1.17 min). The other two pairs copulated for 179 and 236 minutes, respectively. II. Unsuccessful Courtships Interactions between males and virgin females that did not result in copulation are here referred to as unsuccessful courtships. Film records of unsuccessful courtships were not made because of the great variation in their form and duration. Data on the following five general types were derived from written records of 34 unsuccessful courtships between males and released virgins. The number of observations (n) and the percent of all documented unsuccessful courtships is given for each type. All types terminated when the male departed. Type A. The female continued flying on a level course (n = 17, 50%). These interactions were of short duration (2 to 15 sec). The males were probably of low courtship persistence like those seen by Rutowski (1978b) in Colias butterflies. Type B. The female continued flying and initiated an ascending flight (n = 1, 3%). As well as the one interaction with a virgin female, seven naturally-occurring interactions were seen in which the females (presumably mated, see Shapiro, 1970) initiated ascending flights when courted by males. The interactions ranged in dura- tion from 5 to more than 35 sec. Since all observed ascending flights were between males and females and since none ended in copulation it is assumed that they were attempts by mated females to curtail the courtship of persistent males as has been documented for the ascending flights of Colias butterflies (Rutowski, 1978b). Type C. The female alit on vegetation or on the ground; the male may or may not have attempted copulation (n= 7, 20%). Since these unsuccessful courtships ranged in duration from 3 to 10 sec, the males were probably of low persistence as in type A. Type D. The female alit on vegetation or on the ground and performed a flutter response, a pierid mate refusal posture, or both (n =9, 26%). These interactions were from 10 to 68 sec in duration. When perched females spread their wings and elevated the abdomen they were said to have performed the mate refusal posture first described by Obara (1964) for P. rapae and reported for several other pierids (Scott, 1973). Abbott (1959) previously and incorrectly descibed the mate-refusal posture as an immediate invitation to copulation in P. protodice. This posture mechanically impedes the male’s copulatory attempts. Females achieved the same effect in other situations by performing the flutter response, a rapid opening and closing of the wings (Obara, 1964; Rutowski, 1978a). Type E. The male was displaced by another male (n= 1, 3%). This was only seen once with a virgin although I have also seen it occur with wild females. A VoLUME 33, NUMBER 1 47 courting male terminated his courtship attempt when another male approached the female and began courtship. The second male was also unsuccessful. The female in this case was perched during the displacement. In naturally occurring unsuccessful courtships two males courting the same perched or flying female may leave together while circling each other in rapid flight. Systematic observations of unsuccessful courtships with mated females were not made but casual observation suggests that they may only be of type A, B, D, or E. Shapiro (1970) noted that such courtships may be very lengthy in duration, up to 30 min or more. DISCUSSION Successful courtship in Pieris protodice is rapid and highly stereotyped. There are no prolonged aerial components and once the female alights on vegetation or on the ground the male does little more than land on her thorax and couple with her. This description closely fits that given for the temporal and sequential characteristics of the courtships of other pierids including Eurema lisa Boisduval (Rutowski, 1978a), Colias eury- theme Boisduval, C. philodice Latreille (Silberglied and Taylor, 1978), and four species of Pieris in Japan (Suzuki et al., 1977). Pieris protodice also shares components of its courtship with Leptidea synapsis Linnaeus although the latter's courtship is apparently longer and includes some striking male displays (Wiklund, 1977). However, the most obvious dif- ference between the courtship of species in the genus Pieris and that of other pierids is the lack of an abdominal extension response on the part of the female. In E. lisa, L. synapsis, and Colias species the male cannot couple with the female unless she extends her abdomen ventrally out from between the hindwings. Chemical and tactile cues delivered as the male courts the female elicit this response in E. lisa females (Rutowski, 1977). A male of P. protodice must insert his abdomen between the female’s hind- wings to reach her abdomen and couple. At present the proximate and ultimate causes of this variation are unknown. The unsuccessful courtships of P. protodice also follow patterns ob- served in other pierids. Males vary in persistence and females utilize mate refusal postures, flutter responses, and ascending flights to curtail or im- pede the copulatory attempts of persistent males. Interestingly, all three behavior patterns were displayed by virgins as well as mated females sug- gesting that males vary in their attractiveness to virgin females which may be selective in their choice of a mating partner. Similar responses to males by virgin E. lisa females have been hypothesized to serve the same func- tion (Rutowski, 1978a). The rejection responses of virgin Colias females definitely play a role in avoiding courtships with males of the wrong species (Taylor, 1973). Post-nuptial flights have been previously reported only for danaids by 48 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Brower et al. (1965) and Pliske (1975). It should be noted that these post-nuptial flights are spontaneous and as such are distinguished from the induced flights of copulating pairs summarized in Shields and Em- mel (1973). To date the only function attributed to post-nuptial flights is that of removing the copulating pair from the place where courtship ac- tivity has made them conspicuous to one where they are less visible (Brower et al., 1965). In P. protodice courtships, females often land on exposed perches. Presumably this facilitates copulation since males fre- quently appear to have trouble locating and mating with females that land in dense vegetation. Thus, post-nuptial flights in this species may also serve an anti-predator function. As an extension of this hypothesis, post-nuptial flights may also remove animals from exposed perches where thermal stress might become a problem, especially on warm days. The selective pressures of predation and thermal stress should act equally on copulating males and females. However, the data clearly in- dicate that males and females are not equally willing to participate in post-nuptial flights. I suggest that there may be conflict between males and females as a result of male—male competition. Most courtships occur in or near stands of larval foodplant where males look for receptive ovi- positing females or newly-eclosed virgin females. During copulation, pairs are often buffeted by single males in a way that makes them conspicuous to potential predators or may result in the separation of the pair. From the male’s perspective both would be detrimental to his fitness and select for copulating males who initiate post-nuptial flights and move to areas of low male density. However, from the female’s perspective it may be ad- vantageous to remain in the vicinity of a stand of larval foodplant for the following reason. Because males do not defend resources of interest to females, females may select males only on the basis of behavioral cues that are correlated with high genetic quality. One of these cues may be the male’s ability to defend the pair against interruptions by other males during copulation. If so, then it may be to the female’s advantage to stay in an area of high male density thereby forcing the male to fend off the onslaughts of other males. If he cannot, then the female might benefit by copulating with another male. In summary, I propose that selection does not act equally on males and females with respect to their participation in post-nuptial flights. In par- ticular, there appear to be ecological circumstances in which males bene- fit from post-nuptial flights but at least some females do not because the potential benefits of preventing post-nuptial flights and inciting male- male competition outweigh the potential costs of increased predation and VOLUME 33, NUMBER 1 49 thermal stress. The ecological conditions that give rise to this situation remain to be defined. ACKNOWLEDGMENTS I thank Dr. John Alcock for his critical reading of the manuscript and Mr. Larry Marshall for his assistance in the field. This study was sup- ported in part by an Arizona State University Faculty Grant-in-Aid and National Science Foundation Grant BNS 78-11211. LITERATURE CITED AspspotT, W. 1959. Local autecology and behavior in Pieris protodice Boisduval and Leconte with some comparisons to Colias eurytheme Boisduval. Wasmann J. Biology 17; 279-298. Brower, L. P., J. V. Z. BRowEer & F. P. Cranston. 1965. Courtship behavior of the queen butterfly. Zoologica (N.Y.) 50: 1-39. Oxpars, Y. 1964. Mating behavior of the cabbage white, Pieris rapae crucivora. II. The ‘mate-refusal posture’ of the female. Dobut. Zasshi 73: 175-178. Opara, Y. & T. Hipaxa. 1964. Mating behavior of the cabbage white, Pieris rapae crucivora. I. The ‘flutter response’ of the resting male to flying males. Dobut. Aassmi(o- V31—135. PutskE, T. 1975. Courtship behavior of the monarch butterfly, Danaus plexippus L. Ann. Entomol. Soc. Amer. 68: 143-151. Rurowsk1, R. L. 1977. Chemical communication in the courtship of the small sul- phur butterfly Eurema lisa (Lepidoptera, Pieridae). J. Comp. Physiol. 115: 75-85. 1978a. The courtship behavior of the small sulphur butterfly Eurema lisa (Lepidoptera, Pieridae). Anim. Behav. 26: 892-903. . 1978b. The form and function of ascending flights in Colias butterflies. Behav. Ecol. Sociobiol. 3: 163-172. Scott, J. A. 1973. Mating of butterflies. J. Res. Lepid. 11: 99-127. SHAPIRO, A. M. 1970. The role of sexual behavior in density-related dispersal of pierid butterflies. Am. Nat. 104: 367-372. SHIELDs, O. & J. F. EMMEL. 1973. A review of carrying pair behavior and mating times in butterflies. J. Res. Lepid. 12: 25-64. SILBERGLIED, R. E. 1977. Communication in the Lepidoptera. In: How Animals Communicate (ed. T. Sebeok). Indiana University Press, Bloomington. SILBERGLIED, R. E. & O. R. Taytor, Jr. 1978. Ultraviolet reflection and its be- havioral role in the courtship of the sulphur butterflies Colias eurytheme and C. philodice (Lepidoptera, Pieridae). Behav. Ecol. Sociobiol. 3: 203-243. Suzuki, Y., A. NAKxaAnisni, H. Sura, O. Yata & T. Satcusa. 1977. Mating be- haviour in four Japanese species of the genus Pieris (Lepidoptera, Pieridae). Kontyu 45: 300-313. Taytor, O. R., JR. 1973. Reproductive isolation in Colias eurytheme and C. philo- dice (Lepidoptera: Pieridae): Use of olfaction in mate selection. Ann. En- tomol. Soc. Amer. 66: 621-626. Wrixtunp, C. 1977. Courtship behaviour in relation to female monogamy in Lep- tidea synapsis (Lepidoptera). Oikos 29: 275-283. Journal of the Lepidopterists’ Society 33(1), 1979, 50-55 A NEW TECHNIQUE FOR THE PROSPECTIVE SURVEY OF SEX CHROMATIN USING THE LARVAE OF LEPIDOPTERA WINIFRED Cross AND ALISON GILL Department of Genetics, University of Liverpool, P.O. Box 147, Liverpool, L69 3BX, WRK ABSTRACT. A way of examining the heteropyknotic body is described, using cells from prolegs amputated from living larvae. Larval survival rate is high and the results are accurate. Prospective testing for the presence or absence of sex chromatin is par- ticularly valuable in studying intersexes, e.g., in Lymantria dispar, where the adult phenotype is not necessarily an indication of the chromosome constitution of the larva. It is well known that in Lepidoptera a heteropyknotic body may be found in the somatic cells of the female whereas it is lacking in the male (Smith, 1945). However, there are some exceptions to this rule (Traut and Mosbacher, 1968); for example, in Papilio machaon L. the male is polymorphic for the character (Clarke et al., 1977). There is also good evi- dence that in the female the body is derived from the W(=Y) chromo- some (Suomalainen, 1969; Traut and Rathjens, 1973) and that where it is present in the male it is associated with a particular autosome (Clarke et al., 1977). Testing for the “Smith” status has usually been carried out on freshly killed larvae or adults, but Daker (1977) showed that in Hypolimnas bo- lina L. it was possible to assess it from a spine taken from a living larva which thereafter usually developed normally. In the present paper we show that it is also possible to obtain good preparations using a proleg of last instar larvae. We, in fact, found that our preparations from prolegs were of better quality than those from spines. Moreover, this method is particularly useful in dealing with lar- vae which have no spines. MATERIALS AND METHODS The species investigated were Papilio glaucus L., Papilio dardanus Brown, Ewploea core amymone Cr., Hypolimnas bolina L. and Lymantria dispar L. The P. glaucus stock was bred at Caldy, Wirral, and derived from two females from Virginia, U.S.A., kindly supplied by Prof. J. J. Murray. Mrs. Jennifer Maddison of Ibadan sent the butterflies from which all the Nigerian P. dardanus stock was bred and Mrs. Gweneth Johnston posted to us living E. core butterflies from Hong Kong which produced the tested larvae. In H. bolina, race hybrid stock was used, the parent forms coming from Sarawak (from Mr. Stephen Kueh) and Sri Lanka (from Mr. VoLUME 33, NUMBER 1 51 Fic. 1. Proleg tip of H. bolina larva showing tissue scraped out. (As seen under dissecting microscope. ) Fic. 2. Nuclei of tissue cells containing heteropyknotic body from a larva which developed into a female butterfly. (Using x90—oil immersion—objective. ) 52 JOURNAL OF THE LEPIDOPTERISTS SOCIETY Fic. 3. Nuclei of tissue cells lacking heteropyknotic body from a larva which de- veloped into a male butterfly. (Using x90—oil immersion—objective.) Editor's note: photograph reduced to *4 size of original. P. B. Karunaratne. The pure Japanese broods of L. dispar originated from a wild Nagoya female supplied by Dr. Shigeru Ae, and the hybrid brood was from a mating between a German female (from Herr Willy Schultz) and a bred Nagoya male. In the early experiments, the larva to be tested was lightly anesthetized, and then the extreme tip of one of the abdominal prolegs (Fig. 1) was removed with a sharp pair of dissecting scissors. Later it was found that better survival was obtained without an anesthetic. Enough tissue can be scraped from the inside of the proleg to make one good preparation; the material is teased out and spread as thinly as possible. After the amputa- tion each larva was kept separately. The cells are not fixed before staining. Two drops of 2% orcein in 457% acetic acid are placed over the tissue and a coverslip added immediately. After 10-15 minutes the coverslip is firmly pressed to make a “squash” preparation. The “Smith” body when present can be clearly seen under the 40 objective as well as under the <90 (oil immersion) objective (Figs. 2&3). RESULTS Results are shown in Table 1. The accuracy of the method is assessed by noting the sex of the butterfly or moth when it emerges, and in the 53 VoLuME 33, NUMBER | Pep Arey, oy} Alo}eUNzAOFU_ -Isod ccGFTUG,, a19M G ‘pooiq siy} UlOIZ posiIOWla DALY so[eUloZ MOTIAA F ‘oQep OF, ‘s][99 NS UO 9ATIeROU ,YUIC,, se POULIIFUOD SeA }S9} 94} PUL PUNQIIOW UOYM Poe[L{ 19M syoosuT © VSO], ¢ *JS0} ALOJLUITFUOD V WIOFIOd 0} 9}e1OUNSOp 00} 919M Ie[onU [90 94} pue ‘yUsUIdO[aAep JO sasvj}S SNOTIVA }e Poa}se} “Pood sty UI Soxes YO JO SJOSUL OF JSvoT }V UL BSuUIpuIF dATzISOd A[UO 9Y} seM SIT, z ‘OATILSOU . YPIUIG,, 919M PUL DAT} ‘OVAIL] OATJESOU , .YYIWIS,, WOIZ podojsAsop ovdnd s[ewiez g 9S9y} JO OM], t — — soyeul Z dAT}eESOU FG al ossouedef P XK UPULION 6 — dAI}ISOd Z SO[VULIF OT dAT}ISOd GT SSOIO 9081 [Ty WDdsIp “"T G86FT —_— dAT}esOU ¢ sojeul ¢ 9AT}eSB9U Q al o0ei osouede[ = oATySOd T soyeulog G sartsod g wdsip "T P86F1I ® E86FT — OAT}VSOU Q sojeul C aAesou [TT GG eyuey Wg X years — aAT}IsOd G SoTeuloy g aAT}Isod [TT spriqdy puyog ‘]{ JO suorzye19Ues [W19A0S8 0} UBATS ‘OU JUTO[ ZPZG — dAT}eSOU T Q[eul T OAT}VBIU G (@ auowhwup aloo — — Q[euloFy T 9AT}Isod T “Yq JO SuOT}eI0UNS [eIOAOS 0} UAAIS ‘OU UTOL ZEOQG — ATeSOU € SoTBU Q ‘SoTBUOF ZF OAT}VBOU CT PI woosoddy seule; xo snunpiop _— dAT}IsOd T — ,9Ar}sod [| ‘qd JO suonesoues [eloAos 0} UAATS ‘OU JUIOL GT ZST soyeul ¢ dAesoU F — OAT}VBOU G 6I UWLIOF OTeWOT {SO[PULOF 9 aAryIsod Cc Q[VUIOF MOT[IA T 9AT}yIsOd QT MOTI9A XO SnonD)S “J FEOST — — eeu T OAT}VSOU T 6 UWILOF O[BUIOJ — aAr}Isod Z so[eUuloZ Yoryq 9g aArTyIsod g yoryq xo snonnjs' ‘J GOST A][eu1e}xe poxos sosvys [ednd » BAILY Po}So} OVAILT Apoq ,,q}1urg,, sotoeds » ‘ou poolg eednd pose} SuUII OJUIMIDAQGC [WAIL UL syyeOd WOIF SOUND SIOUN JO JUOUIssosse ul}eUOIyO xog IO} pose} OVAILT JO ‘ON ‘popoid [BAe] OY} WOJF S[[PO BUISN UTZeUIOIYO XOS IOF po}so} v19}dopidayT Jo soroeds [e1oAs UWOIF ( A][VUIO}xo9 poxos) ovdnd pur sooussi0uld YOpV ‘“T Alavy 54 JOURNAL OF THE LEPIDOPTERISTS SOCIETY case of overwintering insects, by scoring the sex of the pupa by its external appearance. Concordance between the larval score and the adult or pupal sex is high. DiIscussION Several points are of interest: In P. glaucus, it had been reported previously (Clarke et al., 1976) that the black females were “Smith” positive and the yellow ones (which are male-like) and the males were negative. It is now clear that frequently yellow females are positive, and there appears to be a polymorphism for the character in the yellow form. All black females have, however, so far been positive. In P. dardanus, in the present material, the hippocoon females are con- sistently negative, though previously a few insects of this and other fe- male forms have been positive, so that here again there is a polymorphism. This information in both these species could clearly have been obtained without sexing the larva, but prospective testing has the great advantage that it is possible to select and breed from a female of known “Smith” status. Moreover, it obviates the necessity of testing her immediately after death which is obligatory because rapid degeneration of the cell nuclei occurs post-mertem. The most valuable application of the method, however, will become evident when L. dispar is further studied. Here, in race crosses, inter- sexes may occur (Goldschmidt, 1933), and it will be most informative to relate the sex chromatin status to the phenotype and to the gonadal mor- phology. ACKNOWLEDGMENTS We are extremely grateful to Sir Cyril Clarke F.R.S. for his help in writing this paper and for the use of his facilities and livestock, to Mr. Maurice Gill for taking the photographs and to all those named in the text who kindly provided the living material. LITERATURE CITED Ciarke, C. A., P. M. SHepparp & U. Mirrwocu. 1976. Heterochromatin poly- morphism and colour pattern in the tiger swallowtail butterfly Papilio glaucus L. Nature, Lond. 263: 585-587. CLanke, C, A., U. Mirrwocu & W. Traur. 1977. Linkage and cytogenetic studies in the swallowtail butterflies Papilio polyxenes Fab. and Papilio machaon L. and their hybrids. Phil. Trans. R. Soc. Lond. B 198: 385-399. Daker, M. G. 1977. Report of R.E.S. Meeting. Antenna, Bull. R. Entomol. Soc. Lond. 1: 22 GoLpscumipr, R. 1933. Lymantria. Bibliographia Genetica 11: 1-186. VOLUME 33, NUMBER 1 Do SmirH, S. G. 1945. The diagnosis of sex by means of heteropycnosis. Sci. Agric. 25: 566-571. SUOMALAINEN, E. 1969. On the sex chromosome trivalent in some Lepidoptera females. Chromosoma ( Berl.) 28: 298-308. Traut, W. & G. C. Mospacuer. 1968. Geschlechtschromatin bei Lepidopteren. Chromosoma ( Berl.) 25: 343-356. Traut, W. & B. Ratyyens. 1973. Das W-Chromosom von Ephestia kuehniella (Lepidoptera) und die Ableitung des Geschlechtschromatins. Chromosoma (Berl. ) 41: 437-446. Journal of the Lepidopterists’ Society 33(1), 1979, 55 BOOK REVIEW BurrERFLIES. Text by Jo Brewer, photographs by Kjell B. Sandved. 1976. Harry N. Abrams, New York. 176 pp., ill. Price: hardcover, $18.95; softcover, $9.95, U.S. In the last few years a number of fine popular volumes with exquisitely colored plates have been published, but none has had such an innovative and refreshing ap- proach as this book. Since it is broad in scope and supplies the necessary basic in- formation for the study of Lepidoptera in the clear, concise manner for which Brewer is duly noted, the book should stimulate an interest in and appreciation for the insect group from a technical as well as an aesthetic point of view. It is well illustrated with 245 photographs (133 in color) and additional line drawings and scanning elec- tron micrographs. The organization of the book is quite a departure from traditional treatments. There is a section on the economic impact of butterflies on man (“Historical Notes on Butterflies, Moths and Men”). The section on “Butterflies in Art, Heraldry and Religion” which chronicles the symbolic impact of butterflies on man in everyday life and in legend is especially noteworthy. The remaining sections delve into those areas which man finds so curiously fascinating: metamorphosis, ornamentation of the wings, the compound eye and protective devices. The section on the wings not only examines the physical aspects in terms of wing scales, pigmentation and wing forma- tion, but also the mechanics involved in temperature regulation and flight, all through the enchanted photographic eye of Kjell Sandved. In “Protective Devices,” decep- tion, warning coloration and camouflage are discussed. There is also a brief explana- tion of Batesian and Miillerian mimicry, along with a discussion of larval specificity on certain toxic hostplants and the important role which these plants play in mimetic associations. In such a volume which includes an array of photographs, there are some or- ganizational problems in fitting the plates with the appropriate text. The last 25 pages illustrate further intricate designs and structural iridescence, so intriguing to the natural observer. While these are interesting, they seem somewhat superfluous. In a few cases the identifications are incorrect or not in keeping with current litera- ture such as Thecla syncellus (=Panthiades bitias ). The above points by no means diminish the utility and significance of this book for its intended audience. Its true value will be realized indeed by the enthusiasm and appreciation generated for this diverse biological group in both aspiring and pro- fessional lepidopterists alike. JACQUELINE Y. MILLER, Allyn Museum of Entomology, 3710 Bay Shore Road, Sarasota, Florida 33580. Journal of the Lepidopterists’ Society 33(1), 1979, 56-57 GENERAL NOTES OVIPOSITION OF THE BUTTERFLY BATTUS BELUS VARUS (PAPILIONIDAE ) Members of the swallowtail genus Battus use as larval foodplants woody vines in the family Aristolochiaceae (Brower & Brower 1964, Zoologica 49: 137-159; Ehrlich & Raven 1965, Evolution 18: 586-608; Young 1971, Rev. Biol. Trop. 19: 210-240; Tyler 1975, The Swallowtail Butterflies, Naturegraph, California). The Aristolochia- ceae, like many vines, reach their greatest diversity in the New World tropics (Pfeifer 1966, Ann. Missouri Bot. Garden 53: 1-114), where both Battus and the closely related Parides exploit them. Although the neotropical B. polydamas carefully de- posits small clusters of eggs on various species of Aristolochia in Costa Rica (Young, op. cit.; pers. obs. ), the temperate zone B. philenor is known to oviposit on plants other than Aristolochiaceae, which are not acceptable to larvae (Tyler, op. cit.). Parides always deposits eggs precisely on Aristolochiaceae (Young 1973, Psyche 80: 1-21; 1976, J. Lep. Soc. 31: 100-108). Battus belus varus Kollar ranges from Vera Cruz, Mexico to northeastern Ecuador and northern Venezuela (Rothschild & Jordan, 1906, Fic. 1; _ Battus belus varus ovipositing on Melothria guadalupensis (Curcubita- ceae ) at Minca La Tigra, near La Virgen de Sarapiqui, Heredia Prov., Costa Rica, 19 February 1977, 1300 hrs. VoLUME 33, NuMBER 1 57 Noyitates Zool. 13: 27-753; Tyler, op. cit.); larval foodplants are various species of Aristolochia (e.g., Tyler, op. cit.). This note reports an observation of oviposition of B. belus varus on a plant other than a larval foodplant. On 19 February 1977, a female B. belus varus was observed flying among several clumps of woody and herbaceous vines in a secondary forest at Finca La Tigra, near La Virgen de Sarapiqui, Heredia Province, Costa Rica. She finally began ovipositing on a vine, Melothria guadelupensis (Spreng.) Cogn. (Cucurbitaceae), intertwined with another vine, Aristolochia constricta Griseb.; the leaves of the two plants were similar in size and general shape. Oviposition lasted several minutes (Fig. 1). Forty- three eggs were deposited in a tight cluster on a single leaf of M. guadelupensis and no eggs were found on the A. constricta. In the laboratory, the freshly hatched larvae did not accept leaves of M. guadelupensis, but fed briefly on A. ringens Vahl (ob- tained from H. W. Pfeifer in 1971; locality not specified) before dying. Aristolochia constricta was not available for testing. Tyler (op. cit.) mentions that B. philenor accepts only certain species of Aristolo- chiaceae as foodplants. Foodplant specificity is apparent where different species of Aristolochia occur in the same region (Scriber & Feeny 1976, J. Lep. Soc. 30: 70-71). An Australian Aristolochia-feeding swallowtail, Ornithoptera priamus, has been ob- served to deposit eggs on an introduced species of Aristolochia, and the larvae perished (Straatman 1962, J. Lep. Soc. 16: 99-103). The refusal of A. ringens, a plant spe- cies native to Costa Rica, by B. belus larvae, supports the possibility that Battus specializes on restricted larval foodplants within the Aristolochiaceae. Eggs of B. vhilenor have been found on Convolvulaceae and Polygonaceae, vines which generally look like Aristolochia. Larvae of another Aristolochia-feeding swallowtail, Polydorus aristolochiae (Fabricius ), have been seen on various Cucurbitaceae in India, but their larvae refused to accept these plants in captivity (Ghosh 1914, Mem. Dept. Agr. India, Entomol. Sec. V(1): 53-587). Only certain species of Aristolochia are food- plants of P. aristolochiae (Munshi & Moiz 1967, J. Lep. Soc. 21: 127-128). It is pos- sible that B. belus varus mistook the cucurbit vine for an Aristolochia. Perhaps the very close proximity of the A. constricta vine contributed to this confusion, by provid- ing odoriferous and visual properties of a correct foodplant. Alternatively, the ovi- position on the intertwined cucurbit might have been deliberate, possibly representing an adaptation to avoid waiting egg parasites and predators. Under this explanation, the newly hatched larvae would have rapidly found the correct foodplant. Further observations are needed to distinguish between these two hypotheses. If an adaptation for avoiding egg parasites and predators, such behavior might be more prevalent among vine-feeding butterflies in the tropics, where the intertwining of unrelated vines is common. ALLEN M. Younc, Invertebrate Division, Milwaukee Public Museum, Milwaukee, Wisconsin 53233. Journal of the Lepidopterists’ Society 33(1), 1979, 57-58 MALFUNCTION OF ECDYSIS ALLOWING IMAGINAL EMERGENCE BUT CAUSING DEATH OF ADULT HACKBERRY BUTTERFLY (NYMPHALIDAE) Insects must periodically shed their skins—a process known as ecdysis which al- lows growth or transformation of the individual. Each molt period is a dangerous time during which the insect is susceptible to predation or physiological malfunc- tioning, both of which may cause death. Natural selection has, therefore, perfected the process of ecdysis to such a degree that physiological failures are rare. I describe 58 JouRNAL OF THE LEPIDOPTERISTS SOCIETY below the partial failure of an act of ecdysis—a developmental malfunction which did not prevent transformation, but one that was eventually fatal to the butterfly involved. Such malfunctions are not uncommon in laboratory cultures; however, this observation is of interest because it involved a wild-caught adult. On 17 July 1977 at 1100 hours CDT, I was handed an adult female Asterocampa celtis antonia (Edwards) which had been hand-collected in a residential backyard in Austin, Travis County, Texas. Lack of worn spots on the fully expanded wings, occasional release of untransformed fluids and an egg-packed abdomen indicated recent emergence, probably that same morning. I soon discovered that the head was covered with partial exuviae which had not been properly shed. Practically the en- tire chrysalid head capsule was still present covering the greater part of the imaginal head. Additionally, the left side of the larval head capsule was still attached to the outside of the chrysalid head capsule. Upon metamorphosis of the prepupa to pupa, the left half of the final-instar larval head capsule failed to separate from the newly formed pupal epidermis. When ima- ginal emergence occurred, both sides of the pupal “head” failed to separate as a result of mechanical restriction caused by the still-present larval head capsule section. Although the adult was able to emerge and properly expand and dry its wings, damage to selected parts of the head effectively negated any chance that this in- dividual would reproduce. Both eyes appeared completely normal and allowed reac- tion to approaching objects. The left antenna was not visible, having been trans- formed into an unrecognizable mass associated with the remnant exuviae. The right antenna was free and fully developed; however, it was flexed laterally and horizontal- ly (about 75° from perpendicular) about 2 mm from its base. This antenna could be moved at its base, but such movements occurred only when the antenna was touched; no spontaneous movements were observed. The most significant damage affecting the fitness of this individual involved its proboscis. The proboscis was en- tirely nonfunctional because of failure of the two maxillae to properly fuse. The two halves adhered to each other in a haphazard manner and to the remaining exuviae; re- moval of the exuviae was accompanied by removal of the proboscis halves. Lack of a functional proboscis caused early death of this individual, because the butterfly was unable to feed or obtain moisture. This damaged individual grew pro- gressively weaker until it died approximately 75 hours following capture (in a cage at an ambient diurnal temperature range of about 25-35°C). I thank Patrick K. Neck for supplying the specimen. RayMonp W. Neck, Pesquezo Museum of Natural History, 6803 Esther, Austin, Texas 78752. Journal of the Lepidopterists’ Society 33(1), 1979, 58-60 AGGREGATIVE BEHAVIOR OF ANARTIA FATIMA (NYMPHALIDAE) IN- GUANACASTE PROVINCE, COSTA RICA DURING THE DRY SEASON The neotropical butterfly Anartia fatima Fabricius (Nymphalidae: Nymphalinae ) is widespread throughout the coastal wet and dry regions of Central America and northern South America (Godman & Salvin 1879, Biologia Centraliamericana, Insecta, Lepidoptera-Rhopalocera, vol. 1, 487 p.). Several larval foodplants, in the Acanthaceae, are shared with other nymphalines such as Siproeta (Young & Muyshondt 1974, Stud. Neotrop. Fauna 9: 155-176), but during the severe dry season of dry regions these plants exhibit leaf drop and become unsuitable for oviposition (Young & Stein 1975, Contr, Biol. & Geol., Milwaukee Pub. Museum, No. 8, 29 p.). Several years of observ- ing A, fatima populations in the lowland tropical dry forest region (Tosi 1969, Mapa VOLUME 33, NUMBER 1 59 ecologico de Costa Rica, Trop. Sci. Centr., San José, Costa Rica), Guanacaste Province, indicate that adults are abundant and active throughout the dry season, especially in fields with fully leaved shrubs and trees. This note discusses aggregative behavior of adult A. fatima in Guanacaste. During the afternoon (1530 h) of 26 January 1977, 12 individuals of A. fatima were seen fluttering about a low shrub in a field near Playas del Coco. The general area is about 4 km from the Pacific coast and highly exposed to strong gusty winds characteristic during the dry season. Most of these butterflies were worn. Eventually they settled inside the bush for the rest of the day and night. They were scattered in the leeward side of the bush. Most rested, with wings folded, by hanging from shaded branches. This behavior was observed again on 20 February 1977 at Playa Naranjo; six butterflies settled into a shrub at 1615 h remaining there until the fol- lowing morning. This bush was 400 m from the beach (Gulf of Nicoya) and ex- posed to strong gusty winds. During the following day, adults fluttered inside the bush, with occasional settling for short periods (1-6 min). At no time were more than one or two butterflies resting at once. Both days were sunny, with air tempera- ture near these bushes being 40-42°C. The air temperature inside them at the same time of day was 32—34°C. Anartia fatima was active, along with Phoebis ( Pieridae), in the surrounding habitat, visiting various flowers. These observations took place during the severe, 4—6 month dry season of this region. The bushes used by A. fatima were low and dense with branches and leaves. Strong winds coming from the nearby coast were blocked, as indicated by holding a handkerchief on the lee side of the bushes. While it is generally well documented that unpalatable: butterflies exhibit highly structured communal nocturnal roosting (e.g., Crane 1957, Zoologica 42: 135-146; Brower & Brower 1964, Zoologica 49: 137-159; Urquart 1960, The Monarch Butter- fly, Univ. Toronto Press, 361 p.; Owen & Chanter 1969, J. Zool. (London) 157: 345- 374; Young & Thomason 1975, J. Lep. Soc. 29: 243-255), less is known about the nocturnal behaviors of supposedly palatable species. Other nymphalines, such as Marpesia berania (Hewitson), Hypolimnas bolina Linnaeus, and Smyrna karwinski (Geyer), exhibit highly structured nocturnal communal roosting (Barrett & Burns 1951, Butterflies of Australia and New Guinea, Seward, Melbourne, 187 p.; Benson & Emmel 1973, Ecology 54: 326-335; Muyshondt & Muyshondt 1974, J. Lep. Soc. 28: 924-229) although the adaptive role of this behavior has not been determined. The unstructured condition of A. fatima aggregates and their low numbers suggests that such behavior is a sheltering and thermoregulatory response to a dry, windy environ- ment. Prevailing high temperatures throughout the day, low air humidity, and strong evening winds induce flocking behavior in A. fatima, causing adults to aggregate in some bushes. Low, thick bushes offer protection from the sun and evening gusty winds. On the Pacific slopes of the Cordillera Central in Costa Rica, evening ag- gregates of another nymphaline, Siproeta stelenes Fabricius, are found in coffee bushes during the dry season (pers. ob.); these localities are also exposed to strong, gusty winds and dry conditions. The presence of worn adults suggests that sites of aggregation are not necessarily located near eclosion sites; both A. fatima and S. stelenes oviposit singly, and eggs are distributed over large areas, resulting in low densities of adults eclosing at one spot (Young & Stein 1975, op. cit., Young & Muyshondt 1974, op. cit.). On Grand Cayman Island, British West Indies, adults of Anartia jatrophe Linnaeus cling to the leeward sides of low clumps of the creeping vine Clitoria sp. (Leguminosae) on sunny after- noons in February (pers. obs. ). The aggregative behavior of nymphaline butterflies in exposed secondary habitats near the wind-blown coasts of Caribbean islands and mainland Central America during the dry season could be related to thermoregulation and physical protection from strong winds. The dark brown wing and body color of A. fatima and other species such as S. selenes undoubtedly result in considerable heat gain during afternoon 60 JOURNAL OF THE LEPIDOPTERISTS SOCIETY hours; the thermoregulatory problems of A. fatima have been discussed (Emmel 1972, Evolution 26: 96-107). Gradual heat gain on sunny days leads to shade-seeking be- havior by late afternoon and shady perches also provide nocturnal shelter from winds. It is known that for some insects living in hot climates, such as desert cicadas, gains in body heat result in definite periods of movement into shade and subsequent quiet periods (e.g., Heat & Wilkin 1970, Physiol. Zool. 43: 145-154). Dark butterflies in lowland tropical dry climates may have similar temperature-response problems. The above observations and comments suggest that aggregative behavior of A. fatima is an adaptive response to highly localized climatic or abiotic factors, having little or nothing to do with biotic factors such as vertebrate predators. A biotic result of such behavior, however, may be the maintenance of a cohesive adult population that survives until the following rainy season to oviposit on foodplants as they leaf out. This idea has been discussed with respect to S. karwinski (Muyshondt & Muyshondt 1974. op. cit.) and it may be generally true for other secondary habitat or pasture- dwelling tropical nymphalines which pass the dry season in the adult stage. This fieldwork was funded by N.S.F. Grant GB-33060 and friends of the Museum, Inc. of the Milwaukee Public Museum. ALLEN M. Younc, Invertebrate Division, Milwaukee Public Museum, Milwaukee, Wisconsin 53233. Journal of the Lepidopterists’ Society 33(1), 1979, 60-64 BISTON BETULARIA, OBLIGATE F. INSULARIA INDISTINGUISHABLE FROM F. CARBONARIA (GEOMETRIDAE) It is well known that there is sometimes difficulty in phenotypically recognizing and scoring f. insularia of Biston betularia. On the one hand it may be confused with carbonaria (Kettlewell 1973, The Evolution of Melanism, Oxford, London) and on the other with “typical,” particularly in the Isle of Man (Bailey et al. 1973, En- tomologist 106: 210-214). To help clarify the matter, various scoring methods have been devised for insularia, e.g., that of Lees and Creed (1977, Heredity 39: 66-73, and used by us), where I’ is the lightest and I® the darkest. Neveretheless, difficulties remain, and Lees and Creed (1977) report a brood, B/574, in which a mating be- tween two insularia I’/“typical” heterozygotes produced 66 “typical,” 149 T° in- sularia and 77 carbonaria which were thought to be I*/I®? homozygotes. They also quote Bowater (1914, J. Genet. 3: 299-315) who crossed a wild carbonaria male with a “typical” female, and all the resulting progeny were dark insularia, which is con- sistent with the I*/I* hypothesis. The present note shows that this cannot always be the explanation since the brood to be described could not have produced I*/I* homozygotes. On 13 June 1976 a a female “typical” (Fig. la) was caught in a mercury vapor trap on Hilbre Island near West Kirby, Wirral, England. She was placed in a tin for the following five nights and laid a few eggs, but these were infertile. On the night of 19 June 1976, she was put in a hanging cage with a male insularia (Fig. 1b) (score I’, confirmed by Lees and Creed, pers. comm.), which had been caught in the mercury vapor trap at Caldy, Wirral. Mating took place the same evening and the female then laid freely (brood 14672). 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