Lepidoptera Of North America 6 Butterflies Of Oregon Their .

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Lepidoptera of North America 6Butterflies of OregonTheir Taxonomy, Distribution, and BiologybyAndrew D. WarrenDepartment of ZoologyOregon State UniversityCorvallis, OR 97331warrena@science.oregonstate.eduResearch Associate:Museo de Zoología “Alfonso L. Herrera”Departamento de Biología EvolutivaFacultad de CienciasUniversidad Nacional Autónoma de MéxicoApdo. Postal 70-399, México, D. F. 04510 MéxicoC. P. Gillette Museum of Arthropod DiversityDepartment of Bioagricultural Sciences and Pest ManagementColorado State UniversityFort Collins, CO 80523Section of Invertebrate ZoologyCarnegie Museum of Natural History4400 Forbes AvenuePittsburgh, PA 15213March 15, 20051

Front cover: Extremes in variation seen among females of the Eriogonum marifoliumfeeding segregate of the Euphilotes battoides complex, ventral view. Specimens fromDeschutes Co., vic. north base of Mt. Batchelor, ca. 6400’, 17 July 2002, 5 July 2003,Andrew D. WarrenBack cover: Extremes in variation seen among females of Euphilotes glaucon (Mt. Hoodsegregate), ventral view. Specimens from Clackamas / Hood River Co., Mt. Hood, aboveTimberline Lodge, ca. 6000’, 20 July 2003, Andrew D. WarrenISBN 1084-8819This publication and others in the series may be ordered from the C. P. Gillette Museumof Arthropod Diversity, Department of Bioagricultural Sciences and Pest Management,Colorado State University, Fort Collins, Colorado 80523 USACopyright 2005 2

TABLE OF CONTENTSIntroduction 7Methods and Procedures 8Results 10Acknowledgments and Dedication 16List of Species 18Epargyreus clarus (Cramer, 1775) . 18Thorybes pylades (Scudder, 1870) 20Thorybes diversus Bell, 1927 .21Thorybes mexicana (Herrich-Schäffer, 1869) .22Erynnis icelus (Scudder & Burgess, 1870) . 23Erynnis propertius (Scudder & Burgess, 1870) 24Erynnis pacuvius (Lintner, 1878) . 25Erynnis persius (Scudder, 1863) .26Pyrgus ruralis (Boisduval, 1852) . 27Pyrgus communis (Grote, 1872) . 28Heliopetes ericetorum (Boisduval, 1852) . 30Pholisora catullus (Fabricius, 1793) .31Hesperopsis libya (Scudder, 1878) . 32Carterocephalus palaemon (Pallas, 1771) 32Oarisma garita (Reakirt, 1866) 34Hesperia uncas W. H. Edwards, 1863 .35Hesperia juba (Scudder, 1874) . 36Hesperia colorado (Scudder, 1874) .38Hesperia columbia (Scudder, 1872) . 41Hesperia lindseyi (Holland, 1930) 42Hesperia nevada (Scudder, 1874) .43Atalopedes campestris (Boisduval, 1852) .45Polites peckius (W. Kirby, 1837) .46Polites sabuleti (Boisduval, 1852) 47Polites mardon (W. H. Edwards, 1881) 49Polites themistocles (Latreille, [1824]) .50Polites sonora (Scudder, 1872) .51Ochlodes sylvanoides (Boisduval, 1852) .53Ochlodes agricola (Boisduval, 1852) . 55Ochlodes yuma (W. H. Edwards, 1873) . 55Euphyes vestris (Boisduval, 1852) 57Amblyscirtes vialis (W. H. Edwards, 1862) .58Parnassius clodius Ménétriés, 1857 . 59Parnassius smintheus Doubleday, [1847] .60Battus philenor (Linnaeus, 1771) . 61Papilio machaon Linnaeus, 1758 . 62Papilio zelicaon Lucas, 1852 .63Papilio indra Reakirt, 1866 . 65Papilio multicaudata W. F. Kirby, 1884 . 65Papilio rutulus Lucas, 1852 .663

Papilio eurymedon Lucas, 1852 67Neophasia menapia (C. Felder & R. Felder, 1859) . 68Pontia beckerii (W. H. Edwards, 1871) 70Pontia protodice (Boisduval & Le Conte, [1830]) . 71Pontia occidentalis (Reakirt, 1866) . 72Pontia sisymbrii (Boisduval, 1852) . 73Pieris rapae (Linnaeus, 1758) . 75Pieris marginalis Scudder, 1861 76Euchloe ausonides (Lucas, 1852) . 78Euchloe hyantis (W. H. Edwards, 1871) .79Euchloe lotta Beutenmüller, 1898 . 81Anthocharis sara Lucas, 1852 . 82Anthocharis lanceolata Lucas, 1852 .84Colias philodice Godart, 1819 . 85Colias eurytheme Boisduval, 1852 86Colias occidentalis Scudder, 1862 .87Colias christina W. H. Edwards, 1863 . 89Colias alexandra W. H. Edwards, 1863 92Colias pelidne Boisduval & Le Conte, [1830] .93Colias interior Scudder, 1862 94Phoebis sennae (Linnaeus, 1758) . 94Lycaena arota (Boisduval, 1852) . 95Lycaena phlaeas (Linnaeus, 1761) . 97Lycaena cupreus (W. H. Edwards, 1870) . 98Lycaena xanthoides (Boisduval, 1852) .98Lycaena editha (Mead, 1878) . 100Lycaena gorgon (Boisduval, 1852) .102Lycaena rubidus (Behr, 1866) . 103Lycaena heteronea Boisduval, 1852 .104Lycaena helloides (Boisduval, 1852) 107Lycaena nivalis (Boisduval, 1869) . 108Lycaena mariposa (Reakirt, 1866) . 111Habrodais grunus (Boisduval, 1852) . 112Atlides halesus (Cramer, 1777) .113Introduction to Oregon’s green Callophrys species .114Callophrys affinis (W. H. Edwards, 1862) . 115Callophrys perplexa Barnes & Benjamin, 1923 116Callophrys sheridanii (W. H. Edwards, 1877) (1852) .119Callophrys gryneus (Hübner, [1819]) .125Callophrys spinetorum (Hewitson, 1867) .134Callophrys johnsoni (Skinner, 1904) 135Callophrys augustinus (Westwood, 1852) 136Callophrys mossii (Hy. Edwards, 1881) . 137Callophrys polios (Cook & Watson, 1907) . 140Callophrys eryphon (Boisduval, 1852) .141Satyrium fuliginosa (W. H. Edwards, 1861) .1424

Satyrium semiluna Klots, 1930 . 144Satyrium behrii (W. H. Edwards, 1870) . 146Satyrium titus (Fabricius, 1793) 147Satyrium californica (W. H. Edwards, 1862) . 149Satyrium sylvinus (Boisduval, 1852) 150Satyrium auretorum (Boisduval, 1852) 152Satyrium tetra (W. H. Edwards, 1870) 153Satyrium saepium (Boisduval, 1852) 154Strymon melinus Hübner, 1818 . 155Leptotes marina (Reakirt, 1868) . 157Brephidium exilis (Boisduval, 1852) 157Cupido comyntas (Godart, [1824]) . 158Cupido amyntula (Boisduval, 1852) . 160Celastrina echo (W. H. Edwards, 1864) .161Glaucopsyche piasus (Boisduval, 1852) .166Glaucopsyche lygdamus (Doubleday, 1841) 167Philotiella leona Hammond & McCorkle, 2000 169Introduction to Euphilotes .170Euphilotes [on Eriogonum marifolium] 171Euphilotes glaucon (W. H. Edwards, 1871) . 173Euphilotes [on Eriogonum heracleoides] . 176Euphilotes baueri (Shields, 1975) .178Euphilotes enoptes (Boisduval, 1852) . 179Euphilotes columbiae (Mattoni, 1954) . 182Euphilotes ancilla (Barnes & McDunnough, 1918) . 184Plebejus idas (Linnaeus, 1761) .185Plebejus anna (W. H. Edwards, 1861) . 187Plebejus melissa (W. H. Edwards, 1873) . 189Plebejus saepiolus (Boisduval, 1852) .190Plebejus icarioides (Boisduval, 1852) .192Plebejus shasta (W. H. Edwards, 1862) . 196Plebejus acmon (Westwood, [1851]) 197Plebejus lupini (Boisduval, 1869) .199Plebejus [on Eriogonum pyrolifolium var. coryphaeum] . 202Plebejus podarce (C. Felder & R. Felder, 1865) . 203Apodemia mormo (C. Felder & R. Felder, 1859) . 204Danaus plexippus (Linnaeus, 1758) . 205Danaus gilippus (Cramer, 1776) .206Euptoieta claudia (Cramer, 1776) 206Speyeria cybele (Fabricius, 1775) .207Speyeria coronis (Behr, 1864) . 208Speyeria zerene (Boisduval, 1852) . 209Speyeria callippe (Boisduval, 1852) .212Speyeria egleis (Behr, 1862) .215Speyeria hesperis (W. H. Edwards, 1864) 2165

Speyeria hydaspe (Boisduval, 1869) .218Speyeria mormonia (Boisduval, 1869) . 219Boloria selene ([Denis & Schiffermüller], 1775) . 220Boloria bellona (Fabricius, 1775) .221Boloria epithore (W. H. Edwards, 1864) .222Chlosyne leanira (C. Felder & R. Felder, 1860) .224Chlosyne palla (Boisduval, 1852) .225Chlosyne acastus (W. H. Edwards, 1874) (1870) .229Chlosyne hoffmanni (Behr, 1863) . 233Phyciodes orseis W. H. Edwards, 1871 .234Phyciodes pallida (W. H. Edwards, 1864) . 235Phyciodes mylitta (W. H. Edwards, 1861) 236Phyciodes cocyta (Cramer, 1777) . 237Phyciodes pulchella (Boisduval, 1852) 239Euphydryas gillettii (Barnes, 1897) . 242Euphydryas anicia (Doubleday, [1847]) .243Euphydryas chalcedona (Doubleday, [1847]) . 246Euphydryas editha (Boisduval, 1852) .249Junonia coenia Hübner, [1822] 254Polygonia faunus (W. H. Edwards, 1862) 255Polygonia satyrus (W. H. Edwards, 1869) . 256Polygonia gracilis (Grote & Robinson, 1867) .257Polygonia oreas (W. H. Edwards, 1869) .258Aglais milberti (Godart, 1819) .260Nymphalis vaualbum (Denis & Schiffermüller, 1775) . 261Nymphalis californica (Boisduval, 1852) . 262Nymphalis antiopa (Linnaeus, 1758) 263Vanessa atalanta (Linnaeus, 1758) .264Vanessa cardui (Linnaeus, 1758) . 265Vanessa annabella (Field, 1971) . 266Vanessa virginiensis (Drury, 1773) . 267Limenitis archippus (Cramer, 1775) . 268Limenitis weidemeyerii W. H. Edwards, 1861 .269Limenitis lorquini (Boisduval, 1852) 270Adelpha californica (Butler, 1865) . 272Coenonympha tullia (Müller, 1764) . 273Cercyonis pegala (Fabricius, 1775) . 275Cercyonis sthenele (Boisduval, 1852) . 278Cercyonis oetus (Boisduval, 1869) . 280Erebia epipsodea Butler, 1868 . 281Neominois ridingsii (W. H. Edwards, 1865) .281Oeneis nevadensis (C. Felder & R. Felder, 1867) 282References .284Index of plant names .383Index of butterfly names 398Maps of Oregon . 4066

INTRODUCTIONThe purpose of this document is to provide an update on the taxonomy,distribution and biology of Oregon’s 171 naturally occurring butterfly species. Apreliminary attempt has been made to identify patterns of geographic and ecologicalvariation displayed by each of Oregon’s species. The distribution of each butterflyspecies is recorded at the county level. Available life history information for eachspecies is summarized, and foodplant records for Oregon’s butterfly species arereviewed. I have attempted to trace information related to Oregon’s butterflies to itsoriginal source, and all documents consulted during literature searches have been cited.Additionally, a considerable amount of new taxonomic, distributional and ecologicalinformation is presented herein, based on extensive fieldwork I have conductedthroughout the state between April 1999 and March 2005, together with study in variouspublic and private collections.This document is not a field guide to the identification of Oregon’s butterflyspecies. Nor is this document meant to serve as a single, stand-alone reference on thebutterflies of Oregon. Rather, this document is intended to serve as a companion to otherworks on, or related to, Oregon’s butterflies. Specifically, this volume is meant to be acompanion to the following western butterfly books. These, along with hundreds ofpapers published in technical journals (see References), have laid the basic foundationupon which this document is based: Pyle (1974, 1981, 1984a, 2002), Neill & Hepburn(1976), Dornfeld (1980), Christensen (1981), Ferris & Brown (1981), Garth & Tilden(1986), Tilden & Smith (1986), Stanford & Opler (1993), Hinchliff (1994), Emmel(1998), Opler (1999) and Neill (2001). Readers without all of these references should atleast consult, if possible, Ernst Dornfeld’s (1980), The Butterflies of Oregon, TomEmmel’s (1998) Systematics of Western North American Butterflies, Crispin Guppy &Jon Shepard’s (2001) Butterflies of British Columbia, Bob Pyle’s (2002) The Butterfliesof Cascadia, and most importantly, John Hinchliff’s (1994) An Atlas of OregonButterflies. Taxonomic changes proposed in Emmel (1998) are herein discussed andevaluated for all cases that potentially apply to Oregon.In many ways, this document remains very incomplete. A tremendous amount offield and laboratory research remains to be done before a thorough knowledge of thetaxonomy, distribution and biology of Oregon’s butterfly species can be attained. This isbut an initial attempt to synthesize the available information on Oregon’s butterflies,identify areas where further research is necessary, and stimulate interest in filling gaps inour knowledge. Hopefully, within just a few years, most information presented hereinwill be outdated. The structure of this document has very much been dictated by timeconstraints. I sincerely wish it would have been possible to fully illustrate this volume,together with distribution maps for each species, but constraints on time cannot allow forthat at present. Perhaps, in the future, all populations mentioned herein can be mappedand figured in a separate volume. I apologize upfront for the lack of photos anddistribution maps, as well as illustrations of genitalia. As a substitute, I have tried toreference sources where such may be obtained. I also apologize for all errors in thisdocument, and accept full responsibility for them.7

METHODS AND PROCEDURESFIELDWORKBetween April 1999 and March 2005, I spent about 240 days studying butterfliesat various localities in 33 of Oregon’s 36 counties. Some of these trips were purelyexploratory, but many were conducted in order to study particular species. At eachlocality, effort was made to sample at least a few individuals of each butterfly speciesencountered, and long series of certain taxa have been obtained for morphological andgenetic studies. In addition, larval foodplant vouchers were collected for many species.Information on sampling and how to curate and store voucher specimens is not repeatedherein, since many references on butterflies published before about 1995 include thisinformation (e.g., Dornfeld 1980: 31-36). In particular, those interested in information oncollection and curation techniques should consult Winter (2000).The Oregon Atlas & Gazetteer (DeLorme 2001) proved to be very useful duringexploratory fieldwork. However, not all roads indicated in that volume are open to thepublic. Additionally, some localities in the Columbia River Gorge discussed by Jolley(1988) proved to be excellent sites for butterflies. The herbarium at Oregon StateUniversity was frequently consulted in order to determine sites where larval foodplants ofcertain butterfly species occur. More often than not, once relocated, those sites hostedpopulations of the butterflies being sought. This methodology was particularly successfulin locating populations of lycaenids, including Callophrys, Euphilotes and Plebejus.A detailed discussion of the various physiographic regions in Oregon wasprovided by Dornfeld (1980: 5-16), who summarized the butterfly species that occur ineach area. No attempt is made herein to duplicate his efforts. Two butterfly taxa inOregon are protected by the Endangered Species Act, and cannot be sampled withoutspecial permits. These are Plebejus icarioides fenderi and Speyeria zerene hippolyta.Neither of these taxa were personally studied during field research through 2004.REVIEW OF COLLECTIONSDornfeld (1980: 1-2) provided a good historical summary of the study ofbutterflies in Oregon and western North America (also see Garth & Tilden 1986: 41-49),with information on early collectors and collections. Between 1999 and 2004, I studiedparts of various private and institutional butterfly collections, with a special focus onspecies from Oregon. Institutional collections consulted include those at the OregonState Arthropod Collection (Oregon State University, Corvallis), the Burke Museum(University of Washington, Seattle) and the McGuire Center for Lepidoptera Research(Gainesville, Florida; including Nevadan material from the George Austin collection).Unfortunately, institutional collections in California were not examined, although theyare likely to contain many specimens from Oregon. Private collections consulted, in part,include those of Vern Covlin, Steve Kohler, David McCorkle, Harold Rice, Dana Ross,Erik Runquist, Donald and Paul Severns, Ray Stanford, Terry Stoddard and John Vernon.8

Statements made in the text about adult butterfly phenotypes are generated fromside-by-side visual comparison of pinned specimens. All references to “average”phenotypes herein are anecdotal, and were not derived from any statistical procedures.For some taxa, genitalic examination proved necessary for identification, especially forspecies of Erynnis, Pyrgus (communis), Cupido, Euphilotes (see p. 171 herein), Plebejusand Euphydryas. However, to date I have performed only a limited number of genitalicdissections on butterfly species from Oregon, only enough to ensure accurate specieslevel determinations. A great deal of laboratory work remains to be conducted onsamples obtained between 1999 and 2004.A few comments on potentially mislabeled museum specimens are needed.Occasionally, specimens may be accidentally mislabeled at some point during thecuration process. A few specimens (discussed in the text) found in the Oregon StateArthropod Collection are apparently mislabeled. Most of these are from the ElmerGriepentrog collection, although a few are from the Vick McHenry collection. It isunknown how these apparent mix-ups happened, but they certainly were unintentional.Mention in the text of potentially mislabeled specimens is made in order to clarifydistributional patterns of various species, and should in no way be interpreted as criticismof those researchers.John Hinchliff kept notebooks of data pertaining to butterfly species known fromeach county in Oregon. These notebooks were used to produce Hinchliff‘s (1994) AnAtlas of Oregon Butterflies,

taxonomy, distribution and biology of Oregon’s butterfly species can be attained. This is but an initial attempt to synthesize the available information on Oregon’s butterflies, identify areas where further research is necessary, and sti

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