Butterflies of North America. 2. Scientific names list for butterfly species of North America, North of Mexico

Tags: W.H. Edwards, Butterflies, Journal of the Lepidopterists' Society, species, the Lepidoptera, Fabricius, Boisduval, Hesperiidae, Nymphalidae, North America, pp, Journal of Research, Fabricius, 1775, Lepidopterists' Society, Entomological Society of America, Lepidoptera, Cramer, Linnaeus, 1758, Texas, Entomological News, Allyn Museum, Subfamily Morphinae Morpho Fabricius, subspecies, Cyllopsis R. Felder, Natural History, American Entomological Society, Biblidinae Biblis Fabricius, species problem, biological species concept, phylogenetic species concept, Bulletin of the American Museum of Natural History, Butterflies of North America, Family Hesperiidae, Limenitis Fabricius, Cynthia Fabricius, North American Butterfly Association, Subfamily Coliadinae Colias Fabricius, Lycaena Fabricius, Hypaurotis Scudder, Scudder, Xamia Clench, Satyrium Scudder, Habrodais Scudder, Incisalia Scudder, Mitoura Scudder, Kricogonia Reakirt, Vanessa Fabricius, American Museum of Natural History, Lepidoptera Research Foundation, Nathalis Boisduval, Canadian Journal of Zoology, Cyanophrys Clench, Universidade Estadual de Campinas, Entomological Society, Sandia Clench
Content: Butterflies of North America. 2. Scientific Names List for Butterfly Species of North America, north of Mexico by Paul A. Opler Department of Bioagricultural Sciences Colorado State University Fort Collins, CO 80523 [email protected] Research Associate, Department of Entomology Smithsonian Institution Washington, D.C. 20560 and Andrew D. Warren Department of Entomology Oregon State University Corvallis, OR 97331 [email protected] Research Associate, Museo de Zoologнa Facultad de Ciencias Universidad Nacional Autуnoma de Mйxico Apdo. Postal 70-399, Mйxico, D.F. 04510 Mйxico 1
January 20, 2004 This list generally follows the order found in the Miller and Brown (1981) Catalogue/Checklist of the Butterflies of North America and the supplement to that list by Ferris (1989c), both published as Memoirs of the Lepidopterists' Society. Generic name usage is conservative (when opinions vary on how to divide monophyletic groups) and generally follows that used in A Field Guide to Eastern Butterflies (Opler 1992, 1998), A Field Guide to Western Butterflies (Opler 1999) and the Stanford and Opler (1993) Atlas to Western USA Butterflies (updated as Opler et al. 2000). Some changes are made to conform with recent research results (e.g. Emmel 1998) and with Palaearctic and Neotropical publications (see References). The list that follows includes superfamily, family, and subfamily categories (with tribes for some Lycaenidae), generally in accord with the arrangement presented by de Jong et al. (1996) and Ackery et al. (1999). The Listing order of families, subfamilies, genera, and species does not necessarily imply relationships of taxa to each other. It is the intent of the authors to use the original spelling of species names as described. We would be pleased if anyone would point out where names herein do not conform with their original orthography. The main purpose of this list is to summarize our current knowledge of the nomenclature of North American butterflies at the species-level (and higher), and to generate interest in current nomenclatural problems. It is our hope that this list will serve as a point of reference for future studies and checklists; our goal will be met when arrangements adopted herein are proved incorrect through future research. Synonyms, homonyms, and subspecies are not listed. A complete catalog of North American butterflies including all synonyms and subspecies is in preparation by Jonathan P. Pelham (Seattle, Washington). Citations to original descriptions have been provided for some recently described taxa (see References); these are indicated in the list by an asterisk. We have tried to be as precise as possible in our listing of taxon authors and dates of publication, and thank Gerardo Lamas (Lima, Peru) and J. Pelham for a great amount of help with these issues. All changes (from other checklists) in authorship or dates of publication made herein will be fully explained in Pelham's upcoming catalog. We note here, however, that our spelling of Leconte's name follows Rehn (1954). This being a list of species, a few words on the species concepts held by the authors seem warranted. There are currently no fewer than 25 species concepts being used by taxonomists in various fields (Mayden 1997, Brower 1999, Sperling in press), and no fewer than five species concepts are currently applied in morphological and molecular studies of Lepidoptera taxonomy. Some of the more popular species concepts held by taxonomists today don't even recognize subspecies. While subspecies are frequently named to describe geographical variants, the criteria for recognizing such variants as "subspecies" are subjective and vary greatly from author to author (even when those authors 2
claim to apply the same species concept). The authors of this list apply two different species concepts when making taxonomic decisions. The senior author (PAO) holds views most consistent with Mayr's Biological Species Concept (Mayr 1957a,b). The most current version of this concept defines species as "groups of interbreeding natural populations that are reproductively isolated from other such groups" (see Wheeler & Meier 2000). Earlier versions of Mayr's concept required users to infer whether or not allopatric populations are capable of interbreeding, but later versions of the concept have abandoned this notion (since it is often impossible to make this inference anything other than a guess). The production of fertile offspring is an indication of a biological species. The views of the junior author (ADW) are most consistent with the Phylogenetic Species Concept (Nixon & Wheeler 1990) when making taxonomic decisions. A phylogenetic species is defined as the "smallest aggregation of (sexual) populations or (asexual) lineages diagnosable by a unique combination of character states in comparable individuals (semaphoronts)." Under this concept, species are identified by unique character states, which show no intermediacy or signs of introgression with related organisms. For a debate between proponents of these and other species concepts, see Wheeler & Meier (2000). Even with different species concepts, the authors of this list agreed on over 90% of the taxonomic decisions that have been incorporated herein, even before initial discussion. Subsequent discussion and re-evaluation of available evidence has resolved almost all the discrepancies in taxonomic opinions between the authors (only a few cases remain where the authors don't entirely agree). Therefore, the majority of the species listed herein can be considered biological and phylogenetic species, in cases where sufficient information is known (and most likely qualify as species under other species concepts). Annotations after species entries will hopefully explain differences between this listing and previous checklists, and enable users with alternate species concepts to extract necessary information from this list. 3
SUPERFAMILY HESPERIOIDEA, FAMILY HESPERIIDAE [Our arrangement of this family follows Evans (1951, 1952, 1953, 1955) and Voss (1952), as modified by Ackery et al. (1999) and Warren (2000, 2002). The higher-level classification of the family is currently being studied by ADW.] Subfamily Pyrrhopyginae Apyrrothrix Lindsey, 1921 1. araxes (Hewitson, 1867) [Burns & Janzen (2001) treat arizonae (Godman & Salvin, 1893) as a full species based on genitalic differences observed in eleven male specimens, compared from Arizona and the Mexico City area. However, populations from areas in between (Sinaloa and Nayarit) show intermediacy in all of the genitalic differences cited by Burns and Janzen, and several additional geographic variants are known (ADW, unpublished). See Mielke (2002) for generic change from Pyrrhopyge Hьbner, [1819].] Subfamily Pyrginae Phocides Hьbner, [1819] 2. pigmalion (Cramer, 1779) 2.1 belus Godman & Salvin, 1893 Observed and photographed at Bentsen-Rio Grande State Park, Hidalgo Co. Texas on April 13, 2003 by David J. Hanson. Photo identified by Andrew D. Warren (Hanson, Knudson, & Bordelon, 2003). 3. polybius (Fabricius, 1793) [Papilio palemon Cramer, 1777, being a primary homonym of Papilio palaemon Pallas, 1771, is not an available name (ICZN 1999).] Proteides Hьbner, [1819] 4. mercurius (Fabricius, 1787) Epargyreus Hьbner, [1819] 5. clarus (Cramer, 1775) 6. zestos (Geyer, 1832) 7. exadeus (Cramer, 1779) 8. species [Arizona records of "exadeus" almost certainly refer to E. windi H.A. Freeman, 1969* (a resident of central Sonora, Mexico), but specimens have not been examined; exadeus is unknown in western Mexico north of Nayarit (ADW, unpublished). Epargyreus aspina Evans, 1952* could possibly also stray into our area; dissection of all exotic Epargyreus specimens in our area is needed to confirm their specific identity.] Polygonus Hьbner, [1825] 9. leo (Gmelin, [1790]) 4
10. savigny (Latreille, [1824]) [Mielke & Casagrande (2002) treat manueli Bell & W.P. Comstock, 1948 as a junior synonym of savigny; also see Mielke (in press).] Chioides Lindsey, 1921 11. albofasciatus (Hewitson, 1867) [Raised to species level by Austin and Warren (2002); previously considered to be a subspecies of C. catillus (Cramer, 1779).] 12. zilpa (Butler, 1872) Aguna R.C. Williams, 1927 13. asander (Hewitson, 1867) 14. claxon Evans, 1952* 15. metophis (Latreille, [1824]) Typhedanus Butler, 1870 16. undulatus (Hewitson, 1867) Polythrix E.Y. Watson, 1893 17. octomaculata (Sepp, [1844]) 18. mexicanus H.A. Freeman, 1969* Zestusa Lindsey, 1925 19. dorus (W.H. Edwards, 1882) Codatractus Lindsey, 1921 20. alcaeus (Hewitson, 1867) 21. arizonensis (Skinner, 1905) 22. mysie (Dyar, 1904) [This species was placed in Codatractus by Burns (1996).] Urbanus Hьbner, [1807] 23. proteus (Linnaeus, 1758) 24. belli (Hayward, 1935) [Specimen reported from south Texas by Warren (1997).] 25. pronus Evans, 1952* 26. esmeraldus (Butler, 1877) 27. dorantes (Stoll, 1790) 28. teleus (Hьbner, 1821) 29. tanna Evans, 1952* 30. simplicius (Stoll, 1790) 31. procne (Plцtz, 1881) 32. doryssus (Swainson, 1831) Astraptes Hьbner, [1819] 33. fulgerator (Walch, 1775) 34. egregius (Butler, 1870) 35. alardus (Stoll, 1790) 36. alector (C. Felder & R. Felder, 1867) [Astraptes gilberti H.A. Freeman, 1969* is treated as a synonym of A. alector hopfferi 5
(Plцtz, 1881) by Scott (1986) and Warren (2000, 2002).] 37. anaphus (Cramer, 1777) Autochton Hьbner, 1823 38. cellus (Boisduval & Leconte, [1837]) 39. pseudocellus (Coolidge & Clemence, [1910]) [Apparently extirpated from the U.S portion of its range (Bailowitz & Brock 1991).] 40. cincta (Plцtz, 1882) Achalarus Scudder, 1872 41. lyciades (Geyer, 1832) 42. casica (Herrich-Schдffer, 1869) 43. albociliatus (Mabille, 1877) 44. toxeus (Plцtz, 1882) Thessia Steinhauser, 1989* 45. jalapus (Plцtz, 1881) Thorybes Scudder, 1871 46. drusius (W.H. Edwards, [1884]) 47. pylades (Scudder, 1870) 48. bathyllus (J.E. Smith, 1797) 49. confusis Bell, 1922 50. diversus Bell,1927 51. mexicana (Herrich-Schдffer, 1869) Cabares Godman & Salvin, 1894 52. potrillo (Lucas, 1857) Celaenorrhinus Hьbner, [1819] 53. fritzgaertneri (Bailey, 1880) 54. stallingsi H.A. Freeman, 1946* Spathilepia Butler, 1870 55. clonius (Cramer, 1775) Cogia Butler, 1870 56. hippalus (W.H. Edwards, 1882) 57. outis (Skinner, 1894) 58. caicus (Herrich-Schдffer, 1869) 59. calchas (Herrich-Schдffer, 1869) Arteurotia Butler & H. Druce, 1872 60. tractipennis Butler & H. Druce, 1872 Nisoniades Hьbner, [1819] 61. rubescens (Mцschler, 1877) Pellicia Herrich-Schдffer, 1870 62. arina Evans, 1953* 63. dimidiata Herrich-Schдffer, 1870 Noctuana Bell, 1937 6
64. stator (Godman, 1899) [Documented from Hidalgo Co., Texas, by Bob Stewart (Anonymous 2000).] Windia H.A. Freeman, 1969* 65. windi H.A. Freeman, 1969* [Specimen collected in Guadalupe Canyon, Cochise Co., Arizona, by Kilian Roever. Determined by J. Burns.] Bolla Mabille, 1903 66. brennus (Godman & Salvin, 1896) 67. clytius (Godman & Salvin, 1897) Staphylus Godman & Salvin, 1896 68. ceos (W.H. Edwards, 1882) 69. mazans (Reakirt, [1867]) 70. hayhurstii (W.H. Edwards, 1870) Gorgythion Godman & Salvin, 1896 71. begga (Prittwitz, 1868) [Listed as G. vox Evans, 1953* by Neck (1996), although the U.S. specimens have been confirmed as begga. Dissection is required for certain identification. Also see Stanford (2002).] Sostrata Godman & Salvin, 1895 72. nordica Evans, 1953* [Treatment as a full species follows Warren (2000, 2002); S. bifasciata (Mйnйtriйs, 1829) is a South American species. The relationship between these two taxa is currently being studied in detail by G.T. Austin and ADW.] Carrhenes Godman & Salvin, 1895 73. canescens (R. Felder, 1869) Xenophanes Godman & Salvin, 1895 74. tryxus (Stoll, 1780) [The species name is often misspelled (e.g. Neck 1996).] Antigonus Hьbner, [1819] 75. emorsa (R. Felder, 1869) Systasea W.H. Edwards, 1877 76. pulverulenta (R. Felder, 1869) 77. zampa (W.H. Edwards, 1876) Achlyodes 77.1 pallida (R. Felder, 1869) [Photographed in the lower Rio Grande Valley in October, 2003 and reported by Warren et al., 2003.] Eantis Boisduval, 1836 [Our treatment of Eantis is based on Warren (1996); these species were previously placed in Achlyodes Hьbner, [1819]. Also see Brown & Heineman (1972).] 78. tamenund (W.H. Edwards, 1871) [Eantis thraso (Hьbner, [1807]) is a Central and South American species (Warren 7
1996).] Grais Godman & Salvin, 1894 79. stigmaticus (Mabille, 1883) Timochares Godman & Salvin, 1896 80. ruptifasciata (Plцtz, 1884) Anastrus Hьbner, [1824] 80.1 sempiternus (Butler & H. Druce, 1872) [Anastrus sempiturnus (Butler and Druce) was first collected in Starr Co., Texas by Charles Bordelon (Knudson, 2002). Chiomara Godman & Salvin, 1899 81. georgina (Reakirt, 1868) [Chiomara asychis (Stoll, 1780) is a South American species that does not occur in Costa Rica (Janzen et al. 1998) or Mexico (Warren 2000, 2002); however, its relationship to georgina is currently being studied in detail by G. Austin and ADW.] 82. mithrax (Mцschler, 1879) Gesta Evans, 1953 83. invisus (Butler & H. Druce, 1872) [Austin and Warren (2002) treat G. invisus and G. gesta (Herrich-Schдffer, 1863), a West Indian and South American taxon, as separate species.] Ephyriades Hьbner, [1819] 84. brunnea (Herrich-Schдffer, 1865) Erynnis Schrank, 1801 85. icelus (Scudder & Burgess, 1870) 86. brizo (Boisduval & Leconte, [1837]) [Research is needed to determine if one or more subspecies should be elevated to the species-level.] 87. juvenalis (Fabricius, 1793) [Research is needed to determine if clitus (W.H. Edwards, 1882) should be elevated to the species-level.] 88. telemachus Burns, 1960 89. propertius (Scudder & Burgess, 1870) 90. meridianus Bell, 1927 91. scudderi (Skinner, 1914) 92. horatius (Scudder & Burgess, 1870) 93. tristis (Boisduval, 1852) 94. martialis (Scudder, [1870]) 95. pacuvius (Lintner, 1878) 96. zarucco (Lucas, 1857) 97. funeralis (Scudder & Burgess, 1870) 98. baptisiae (Forbes, 1936) 99. lucilius (Scudder & Burgess, 1870) 100. afranius (Lintner, 1878) 8
101. persius (Scudder, 1863) [Research is needed to better define relationships among and within the above three taxa; see Burns (1964).] Pyrgus Hьbner, [1819] 102. centaureae (Rambur, [1842]) [Tentatively includes wyandot (W.H. Edwards, 1863), which was treated as a specieslevel taxon by Shapiro (1974) and Gochfield & Burger (1997) (based on genitalic differences between it and freija (Warren, 1924) cited by Forbes (1960); but see Lindsey (1928)). Its relationship to other North American and Eurasian taxa associated with centaureae remains to be studied in detail. Also see Warren (1926, 1935, 1951), de Jong (1972), Devyatkin (1990) and Tuzov et al. (1997).] 103. ruralis (Boisduval, 1852) 104. xanthus W.H. Edwards, 1878 105. scriptura (Boisduval, 1852) 106. communis (Grote, 1872) 107. albescens Plцtz, 1884 [Treatment as a full species follows Burns (2001).] 108. philetas W.H. Edwards, 1881 109. oileus (Linnaeus, 1767) [The above four taxa were placed in Syrichtus Boisduval, [1834] by Durden (1982). This is incorrect because Syrichtus is a synonym of Pyrgus (see Cowan 1970). There is no published evidence indicating that these four species belong in a Palaearctic genus other than Pyrgus (see de Jong 1972, 1978); however, further study is needed.] Heliopyrgus Herrera, 1957 [Treatment of Heliopyrgus is based on Austin & Warren (2001); also see Mielke & Casagrande (2002).] 110. domicella (Erichson, [1849]) Heliopetes Billberg, 1820 111. ericetorum (Boisduval, 1852) 112. macaira (Reakirt, [1867]) 113. laviana (Hewitson, 1868) 114. arsalte (Linnaeus, 1758) Celotes Godman & Salvin, 1899 115. nessus (W.H. Edwards, 1877) 116. limpia Burns, 1974* Pholisora Scudder, 1872 117. catullus (Fabricius, 1793) 118. mejicanus (Reakirt, [1867]) Hesperopsis Dyar, 1905 119. libya (Scudder, 1878) 120. alpheus (W.H. Edwards, 1876) 121. gracielae (MacNeill, 1970)* 9
Subfamily Heteropterinae Carterocephalus Lederer, 1852 122. palaemon (Pallas, 1771) [The relationships between New and Old World taxa of palaemon remain to be studied in detail.] Piruna Evans, 1955* 123. pirus (W.H. Edwards, 1878) 124. haferniki H.A. Freeman, 1970* 125. polingii (Barnes, 1900) 126. aea (Dyar, 1912) [Previously known as P. cingo Evans, 1955*, which was treated as a synonym of aea by Warren (2000, 2002).] 127. penaea (Dyar, 1918) [Previously misidentifed as P. microsticta (Godman, 1900) (see McGuire & Rickard 1976), a southwestern Mexican species (Freeman & Warren, in preparation).] Subfamily Hesperiinae [Includes genera formerly placed in the Megathyminae, following Ackery et al. (1999).] Synapte Mabille, 1904 128. pecta Evans, 1955* [Warren (2000, 2002) treated S. pecta and S. malitiosa (Herrich-Schдffer, 1865) as separate species.] 129. species [We are uncertain if it is Synapte syraces (Godman, 1901), or the similar S. shiva Evans, 1955* (or both?) that is known from Arizona (Bailowitz & Brock 1991), since we have not examined specimens; both species occur in Sonora, Mexico. Synapte shiva was described as a subspecies of syraces, but was treated as a full species by Warren (2000, 2002).] 130. salenus (Mabille, 1883) Corticea Evans, 1955* 131. corticea (Plцtz, 1882) Vidius Evans, 1955* 132. perigenes (Godman, 1900) Monca Evans, 1955* 133. crispinus (Plцtz, 1882) [Mielke and Casagrande (2002) treat tyrtaeus (Plцtz, 1882) as a junior synonym of crispinus. Monca telata (Herrich-Schдffer, 1869) is a South and Central American species (Bell 1941) which flies sympatrically with crispinus (= tyrtaeus) in Guatemala (Austin et al. 1996).] Nastra Evans, 1955* 10
134. lherminier (Latreille, [1824]) 135. neamathla (Skinner & R.C. Williams, 1923) 136. julia (H.A. Freeman, 1945) Cymaenes Scudder, 1872 137. tripunctus (Herrich-Schдffer, 1865) 138. trebius (Mabille, 1891) [Cymaenes trebius was treated as a species distinct from C. odilia (Burmeister, 1878) by Warren (2000, 2002); however, relationships among taxa placed with odilia by Evans (1955) are under study by G. Austin and ADW.] Lerema Scudder, 1872 139. accius (J.E. Smith, 1797) 140. liris Evans, 1955* [The relationship between L. liris and L. ancillaris (Butler, 1877) where they meet in Central America needs further study, as noted by Warren (2000).] Vettius Godman, 1901 141. fantasos (Cramer, 1780) Perichares Scudder, 1872 142. philetes (Gmelin, [1790]) Rhinthon Godman, 1900 143. osca (Plцtz, 1882) [Treatment as a distinct species from cubana (Herrich-Schдffer, 1865) follows McGuire & Rickard (1976), Ferris (1989c) and Warren (2000, 2002), but see Mielke & Casagrande (2002).] Decinea Evans, 1955* 144. percosius (Godman, 1900) Conga Evans, 1955* 145. chydaea (Butler, 1877) Ancyloxypha C. Felder, 1862 146. numitor (Fabricius, 1793) 147. arene (W.H. Edwards, 1871) Oarisma Scudder, 1872 148. garita (Reakirt, 1866) 149. poweshiek (Parker, 1870) 150. edwardsii (Barnes, 1897) Copaeodes Speyer, 1877 151. aurantiaca (Hewitson, 1868) 152. minima (W.H. Edwards, 1870) Adopaeoides Godman, 1900 153. prittwitzi (Plцtz, 1884) Thymelicus Hьbner, [1819] 154. lineola (Ochsenheimer, 1808) Hylephila Billberg, 1820 11
155. phyleus (Drury, 1773) Pseudocopaeodes Skinner & R.C. Williams, 1923 156. eunus (W.H. Edwards, 1881) Stinga Evans, 1955* 157. morrisoni (W.H. Edwards, 1878) Hesperia Fabricius, 1793 158. uncas W.H. Edwards, 1863 159. juba (Scudder, 1874) 160. comma (Linnaeus, 1758) 161. assiniboia (Lyman, 1892) [Species-level status follows Layberry et al. (1998), but further elaboration on the relationship between this taxon and the next is needed.] 162. colorado (Scudder, 1874) [Hesperia colorado (Scudder, 1874) has priority over H. harpalus (W.H. Edwards, 1881) and H. idaho (W.H. Edwards, 1883). The Hesperia comma complex was discussed by MacNeill (1964, 1975) and by Scott (1975, 1998). Relationships of taxa placed with assiniboia, comma, and colorado are currently being studied by several researchers.] 163. woodgatei (R.C. Williams, 1914) 164. ottoe W.H. Edwards, 1866 165. leonardus Harris, 1862 166. pahaska (Leussler, 1938) 167. columbia (Scudder, 1872) 168. metea Scudder, 1864 169. viridis (W.H. Edwards, 1883) 170. attalus (W.H. Edwards, 1871) 171. meskei (W.H. Edwards, 1877) 172. dacotae (Skinner, 1911) 173. lindseyi (Holland, 1930) 174. sassacus Harris, 1862 175. miriamae MacNeill, 1959 176. nevada (Scudder, 1874) Atalopedes Scudder, 1872 177. campestris (Boisduval, 1852) Polites Scudder, 1872 [See Burns (1994a) for synonymy of Yvretta Hemming, 1935 under Polites.] 178. rhesus (W.H. Edwards, 1878) 179. carus (W.H. Edwards, 1883) 180. peckius (W. Kirby, 1837) 181. sabuleti (Boisduval, 1852) 182. draco (W.H. Edwards, 1871) 183. mardon (W.H. Edwards, 1881) 12
184. themistocles (Latreille, [1824]) 185. baracoa (Lucas, 1857) 186. origenes (Fabricius, 1793) 187. mystic (W.H. Edwards, 1863) 188. sonora (Scudder, 1872) 189. vibex (Geyer, 1832) Wallengrenia Berg, 1897 190. otho (J.E. Smith, 1797) [Tentatively includes clavus (Erichson, [1849]) and curassavica (Snellen, 1887); see Burns (1994b). The complex is being studied by J.Y. Miller.] 191. egeremet (Scudder, 1864) Pompeius Evans, 1955* 192. verna (W.H. Edwards, 1862) Atrytone Scudder, 1872 193. arogos (Boisduval & Leconte, [1837]) Anatrytone Dyar, 1905 [Generic combination follows Burns (1994b).] 194. logan (W.H. Edwards, 1863) 195. mazai (H.A. Freeman, 1969)* Problema Skinner & R.C. Williams, 1924 196. byssus (W.H. Edwards, 1880) 197. bulenta (Boisduval & Leconte, [1837]) Ochlodes Scudder, 1872 198. sylvanoides (Boisduval, 1852) 199. agricola (Boisduval, 1852) 200. yuma (W.H. Edwards, 1873) [Our treatment of Ochlodes species follows Chiba and Tsukiyama (1996).] Poanes Scudder, 1872 201. hobomok (Harris, 1862) 202. zabulon (Boisduval & Leconte, [1837]) 203. taxiles (W.H. Edwards, 1881) 204. melane (W.H. Edwards, 1869) 205. massasoit (Scudder, 1864) 206. viator (W.H. Edwards, 1865) 207. yehl (Skinner, 1893) 208. aaroni (Skinner, 1890) Paratrytone Godman, 1900 [See Burns (1992) for generic combination.] 209. snowi (W.H. Edwards, 1877) Quasimellana Burns, 1994* 13
210. eulogius (Plцtz, 1882) Euphyes Scudder, 1872 211. pilatka (W.H. Edwards, 1867) 212. conspicua (W.H. Edwards, 1863) 213. berryi (Bell, 1941) 214. dion (W.H. Edwards, 1879) [Includes E. alabamae (Lindsey, 1923) as a subspecies, and E. macguirei H.A. Freeman, 1975* as a synonym, following Shuey (1989); also see Shuey (1993).] 215. bayensis Shuey, 1989* 216. dukesi (Lindsey, 1923) 217. bimacula (Grote & Robinson, 1867) 218. arpa (Boisduval & Leconte, [1837]) 219. vestris (Boisduval, 1852) Asbolis Mabille, 1904 220. capucinus (Lucas, 1857) Atrytonopsis Godman, 1900 221. hianna (Scudder, 1868) 222. loammi (Whitney, 1876) 223. deva (W.H. Edwards, 1877) 224. lunus (W.H. Edwards, 1884) 225. vierecki (Skinner, 1902) 226. pittacus (W.H. Edwards, 1882) 227. python (W.H. Edwards, 1882) 228. cestus (W.H. Edwards, 1884) 229. edwardsi Barnes & McDunnough, 1916 Amblyscirtes Scudder, 1872 230. exoteria (Herrich-Schдffer, 1869) 231. cassus W.H. Edwards, 1883 232. aenus W.H. Edwards, 1878 [Includes erna H.A. Freeman, 1943, following Scott (1977).] 233. linda H.A. Freeman, 1943 234. oslari (Skinner, 1899) 235. elissa Godman, 1900 236. hegon (Scudder, 1864) 237. texanae Bell, 1927 238. carolina (Skinner, 1892) 239. reversa W.M. Jones, 1926 240. aesculapius (Fabricius, 1793) 241. nereus (W.H. Edwards, 1876) 242. nysa W.H. Edwards, 1877 243. eos (W.H. Edwards, 1871) 244. vialis (W.H. Edwards, 1862) 14
245. alternata (Grote & Robinson, 1867) 246. celia Skinner, 1895 247. belli H.A. Freeman, 1941 248. tolteca Scudder, 1872 [Includes prenda Evans, 1955*, following Freeman (1993).] 249. phylace (W.H. Edwards, 1878) 250. fimbriata (Plцtz, 1882) 251. simius W.H. Edwards, 1881 [While Burns (1990) removed this species from Amblyscirtes and left it "incertae sedis," we list simius here until a generic placement is proposed.] Lerodea Scudder, 1872 252. eufala (W.H. Edwards, 1869) 253. arabus (W.H. Edwards, 1882) [Includes dysaules Godman, 1900 as a synonym (Warren 2000, 2002).] Oligoria Scudder, 1872 254. maculata (W.H. Edwards, 1865) Calpodes Hьbner, [1819] 255. ethlius (Stoll, 1782) Panoquina Hemming, 1934* 256. panoquin (Scudder, 1864) 257. panoquinoides (Skinner, 1891) 258. errans (Skinner, 1892) 259. ocola (W.H. Edwards, 1863) 260. lucas (Fabricius, 1793) [Panoquina sylvicola (Herrich-Schдffer, 1865) was treated as a synonym of lucas by Robbins et al. (1996); also see Mielke & Casagrande (2002).] 261. hecebolus (Scudder, 1872) 262. evansi (H.A. Freeman, 1946)* Nyctelius Hayward, 1948 263. nyctelius (Latreille, [1824]) Thespieus Godman, 1900 264. macareus (Herrich-Schдffer, 1869) Agathymus H.A. Freeman, 1959* 265. neumoegeni (W.H. Edwards, 1882) [Includes chisosensis (H.A. Freeman, 1952), following Roever (1975).] 266. polingi (Skinner, 1905) 267. evansi (H.A. Freeman, 1950) 268. aryxna (Dyar, 1905) 269. baueri (D. Stallings & Turner, 1954) 270. gentryi Roever, 1998* [Treatment of this taxon and baueri as subspecies of aryxna is unsupported by any compelling data (Cassie et al. 2001), and does not consider problems associated with 15
the name aryxna; see dos Passos (1960) and Roever (1975).] 271. mariae (Barnes & Benjamin, 1924) [Tentatively includes gilberti H.A. Freeman, 1964*, although the status of this taxon is uncertain (see Roever 1975).] 272. estelleae (D. Stalllings & Turner, 1958) [Includes valverdiensis H.A. Freeman, 1966, following Roever (1975).] 273. stephensi (Skinner, 1912) 274. alliae (D. Stallings & Turner, 1957) Megathymus Scudder, 1872 275. yuccae (Boisduval & Leconte, [1837]) [Includes coloradensis C.V. Riley, 1877, following Roever (1975).] 276. streckeri (Skinner, 1895) 277. cofaqui (Strecker, 1876) 278. ursus Poling, 1902 Stallingsia H.A. Freeman, 1959* 279. maculosus (H.A. Freeman, 1955) 16
SUPERFAMILY PAPILIONOIDEA FAMILY PAPILIONIDAE [We have followed the generic arrangement proposed for this family by Miller (1987). We feel further study is needed on the generic relationships of swallowtails in order to subdivide them into smaller monophyletic groups with confidence.] Subfamily Parnassiinae Parnassius Latreille, 1804 280. eversmanni [Mйnйtriйs], [1850] 281. clodius Mйnйtriйs, 1855 282. phoebus (Fabricius, 1793) 283. behrii W.H. Edwards, 1870 284. smintheus Doubleday, [1847] [Split of P. phoebus complex is discussed by Shepard & Manley (1998), Bird et al. (1995) and Layberry et al. (1998).] Subfamily Papilioninae Battus Scopoli, 1777 285. philenor (Linnaeus, 1771) 286. polydamas (Linnaeus, 1758) Parides Hьbner, [1819] 287. eurimedes (Stoll, 1782) 288. alopius (Godman & Salvin, 1890) [Single specimen collected in Cochise Co., Arizona, by Neil Dankert; see Bailowitz & Brock (1991).] Eurytides Hьbner, [1821] 289. marcellus (Cramer, 1777) 290. philolaus (Boisduval, 1836) Papilio Linnaeus, 1758 291. machaon Linnaeus, 1758 [Includes bairdii W.H. Edwards, 1866, and oregonius W.H. Edwards, 1876, following Sperling (1987, 1993a) and Sperling and Harrison (1994); but see Eitschberger (1993) and Pyle (2002).] 292. brevicauda Saunders, 1869 293. joanae Heitzman, 1973* 294. polyxenes Fabricius, 1775 295. zelicaon Lucas, 1852 [Papilio nitra W.H. Edwards, 1883 is considered by Fisher (1977) to be an eastern subspecies of zelicaon containing rare dark individuals, which may represent introgression with polyxenes.] 296. indra Reakirt, 1866 17
297. glaucus Linnaeus, 1758 [Pterourus appalachiensis Pavulaan & D. Wright, 2002* was recently described, but convincing evidence for its treatment as a full species is lacking. It seems to be allopatric to glaucus itself, and it might be a high altitude, single-brooded population of glaucus; or it could be related to canadensis. Further information is needed to determine its status.] 298. canadensis Rothschild & Jordan, 1906 [Specific distinctness of P. canadensis is detailed by Hagen et al. (1991), and further supported by Sperling (1993b).] 299. rutulus Lucas, 1852 300. eurymedon Lucas, 1852 301. multicaudata W.F. Kirby, 1884 302. pilumnus Boisduval, 1836 303. troilus Linnaeus, 1758 304. palamedes Drury, 1773 305. garamas (Geyer, [1829]) [Papilio garamas abderus Hoppfer, 1856 was reported from south Texas by Bordelon & Knudson (2000).] 306. victorinus Doubleday, 1844 [Treated as a subspecies of P. menatius Hьbner [1819] by Tyler et al. (1994: but see page 26) and Lamas (in press); however, we feel that further study of the situation is needed (see DeVries 1987). First reported from the U.S. by Adams (1984).] 307. thoas Linnaeus, 1771 308. cresphontes Cramer, 1777 309. astyalus Godart, 1819 310. ornythion Boisduval, 1836 311. aristodemus Esper, 1794 312. andraemon (Hьbner, [1823]) 313. androgeus Cramer, 1775 314. anchisiades Esper, 1788 315. rogeri Boisduval, 1836 [Papilio pharnaces Doubleday, 1846 is treated as a subspecies of P. rogeri Boisduval, 1836 by Lamas (in press); the two taxa were also considered to be conspecific by Tyler et al. (1994).] *[Miller (1987) does not recognize Heraclides Hьbner [1819], Priamides Hьbner [1819] or Pterourus Scopoli, 1777 as valid genera. Tyler et al. (1994) present a considerably different generic arrangement for our swallowtail taxa; also see Lamas (in press).] FAMILY PIERIDAE Subfamily Dismorphiinae 18
Enantia Hьbner, [1819] 316. albania (H.W. Bates, 1864) [Texas specimen of this species illustrated by Kendall (1974a); see Llorente (1984) for notes on Enantia taxonomy.] Subfamily Pierinae Catasticta Butler, 1870 317. nimbice (Boisduval, 1836) Neophasia Behr, 1869 318. menapia (C. Felder & R. Felder, 1859) 319. terlooii Behr, 1869 [Spelling corrected to original orthography, following Llorente et al. (1997) and Lamas (in press).] Appias Hьbner, [1819] 320. drusilla (Cramer, 1777) [This species is often placed in the genus Glutophrissa Butler, 1887, e.g. Hemming (1967) and Lamas (1981, in press); but see Klots (1933), d'Almeida (1939a), Comstock (1943) and Brown & Heineman (1972).] Leptophobia Butler, 1870 321. aripa (Boisduval, 1836) [Collected in Santa Ana National Wildlife Refuge, Hidalgo Co., Texas, by T. Kral (20X-1988). Specimens deposited in C.P. Gillette Museum, Colorado State University.] Pontia Fabricius, 1807 322. beckerii (W.H. Edwards, 1871) 323. protodice (Boisduval & Leconte, [1830]) 324. occidentalis (Reakirt, 1866) 325. sisymbrii (Boisduval, 1852) Pieris Schrank, 1801 326. rapae (Linnaeus, 1758) 327. oleracea Harris, 1829 328. marginalis Scudder, 1861 [Relationship of meckyae Eitschberger, 1983* to angelika and marginalis needs clarification.] 329. angelika Eitschberger, 1983* [Study of type specimens is necessary; this may be a synonym of P. pseudobryoniae Barnes & McDunnough, 1917, which is an elevation of P. napi variety frigida form pseudobryoniae Verity, [1908], fide J. Pelham.] *[Pieris napi Linnaeus, 1758, a Palaearctic taxon, does not occur in North America as per several authors. North American species are treated as either three species by Eitschberger (1981, 1983) or four species by Geiger and Shapiro (1992). This entire complex needs much more study before relationships can be understood with confidence. Also see Bowden (1972), Shapiro (1984) and Kudrna & Geiger (1985).] 19
330. virginiensis (W.H. Edwards, 1870) *[See Robbins & Henson (1986) for use of Pieris over Artogeia Verity, 1947.] Ascia Scopoli, 1777 331. monuste (Linnaeus, 1764) Ganyra Billberg, 1820 332. josephina (Godart, 1819) 333. howarthi (Dixey, 1915) [Bailowitz (1988) describes the distribution and biology of this species in contrast with G. josephina; also see Beutelspacher (1986) and Llorente et al. (1997).] Euchloe Hьbner, [1819] 334. ausonides (Lucas, 1852) [Includes ogilvia Back, [1991]*, although this was described as a full species; see Layberry et al. (1998).] 335. naina Kozhanchikov, 1923 [Reported for North America by Kondla and Pelham (1995). Specific identity is described by Dubatolov and Kosterin (1994).] 336. guaymasensis Opler, 1987* [Specimen collected in Bisbee, Cochise Co., Arizona, by Sandy Upson. Identity verified by R. Bailowitz, PAO and ADW.] 337. olympia (W.H. Edwards, 1871) 338. creusa (Doubleday, [1847]) 339. hyantis (W.H. Edwards, 1871) 340. lotta Beutenmьller, 1898 [Above two species separated on basis of research by Opler (1965, 1967a,b, 1970, 1971, 1974) as well as unpublished data and communications. Detailed investigation has been initiated by PAO.] Anthocharis Boisduval, Rambur, [Dumйnil] & Graslin, [1833] 341. cethura C. Felder & R. Felder, 1865 [Includes pima W.H. Edwards, 1888. These two form a gradual cline between California, Nevada, and Arizona, as discussed by Emmel et al. (1998b: p. 132).] 342. sara Lucas, 1852 343. julia W.H. Edwards, 1872 344. stella W.H. Edwards, 1879 345. thoosa (Scudder, 1878) [The sara species group is split, in part, on basis of research by Geiger and Shapiro (1986); however, the overall distributions of these taxa remain somewhat unclear and the above arrangement is tentative. The complex is currently under study by PAO.] 346. lanceolata Lucas, 1852 347. midea (Hьbner, [1809]) *[Paramidea Kusnezov, 1929 was described as a monotypic genus for the Asian scolymus (Butler, 1866). It was applied to lanceolata and midea by Ferris (1989c) without justification. It has been treated by several authors as a subgenus of 20
Anthocharis, and is herein excluded as relating to any North American species.] Subfamily Coliadinae Colias Fabricius, 1807 348. philodice Godart, 1819 349. eurytheme Boisduval, 1852 350. occidentalis Scudder, 1862 351. christina W.H. Edwards, 1863 [This complex includes (among others) krauthii Klots, 1935, kluanensis Ferris, 1981*, pseudochristina Ferris, 1989*, astraea (W.H. Edwards, 1872), and several enigmatic eastern Oregon populations (see Ferris 1993). Layberry et al. (1998) treat occidentalis and christina (which includes all populations that are variably UV reflective and containing some orange individuals) as separate species, due to Ferris' statement that the variable Oregon populations were highly localized. However, these Oregon populations have proven to be very widespread over a large geographic area, and show a mosaic of variable segregates with different average character combinations. The complex is currently under study by Paul Hammond and David McCorkle.] 352. alexandra W.H. Edwards, 1863 353. harfordii Hy. Edwards, 1877 [Status of C. harfordii is detailed by Burns (1975) and Ferris (1988, 1993).] 354. meadii W.H. Edwards, 1871 355. johanseni Troubridge & Philip, 1990* 356. hecla Lefиbvre, 1836 357. canadensis Ferris, 1982 358. tyche Bцber, 1812 [Includes boothii Curtis, 1835 and thula Hovanitz, 1955, following Lafontaine & Wood (1997) and Layberry et al. (1998).] 359. nastes (Boisduval, 1832) 360. scudderii Reakirt, 1865 361. gigantea Strecker, 1900 362. pelidne Boisduval & Leconte, [1830] 363. interior Scudder, 1862 364. palaeno (Linnaeus, 1761) [North American populations of this species have been referred to as species chippewa W.H. Edwards, 1872 (see Tuzov et al. 1997, Verhulst 2000, and Guppy & Shepard 2001), restricting palaeno to the Palaearctic region. Opinions vary, however (see Gorbunov 2001) and until a revision of the group is presented we retain palaeno for our populations.] 365. behrii W.H. Edwards, 1866 Zerene Hьbner, [1819] 366. cesonia (Stoll, 1790) 367. eurydice (Boisduval, 1855) 21
Anteos Hьbner, [1819] 368. maerula (Fabricius, 1775) 369. clorinde (Godart, [1824]) Phoebis Hьbner, [1819] 370. sennae (Linnaeus, 1758) 371. argante (Fabricius, 1775) [Stray specimens are known from Kansas (Field 1940) and Texas (Neck 1996, Stanford 2002).] 372. agarithe (Boisduval, 1836) 373. philea (Linnaeus, 1763) 374. neocypris (Hьbner, [1823]) Aphrissa Butler, 1873 375. statira (Cramer, 1777) 376. orbis (Poey, 1832) *[Recognition of Aphrissa follows Brown (1931), d'Almeida (1939b), Smith et al. (1994), Llorente et al. (1997), and Lamas (in press); but see Klots (1933). Also see Brown (1929) and d'Almeida (1940). No recent information to support or refute the lumping of Aphrissa with Phoebis has been presented, but further research is warranted.] Kricogonia Reakirt, 1863 377. lyside (Godart, 1819) Eurema Hьbner, [1819] 378. daira (Godart, 1819) 379. boisduvaliana (C. Felder & R. Felder, 1865) [Treated as a subspecies of E. arbela Geyer, 1832, by Lamas (in press).] 380. mexicana (Boisduval, 1836) 381. salome (C. Felder & R. Felder, 1861) 382. albula (Cramer, 1775) [First U.S. record reported by Chuah & Cushing (1995).] Pyrisitia Butler, 1870 383. messalina (Fabricius, 1787) 384. proterpia (Fabricius, 1775) 385. lisa (Boisduval & Leconte, [1830]) 386. nise (Cramer, 1775) 387. dina (Poey, 1832) Abaeis Hьbner, [1819] 388. nicippe (Cramer, 1779) *[Although the above three genera are sometimes lumped under Eurema (e.g. Ferris 1989c), recent DNA analysis by Pollock et al. (1998) supports our arrangement.] Nathalis Boisduval, 1836 389. iole Boisduval, 1836 22
FAMILY LYCAENIDAE [Our arrangement of this family follows Ackery et al. (1999), which corroborates Eliot (1990), who unites the coppers, blues and hairstreaks into a single subfamily; also see Eliot (1973).] Subfamily Miletinae Feniseca Grote, 1869 390. tarquinius (Fabricius, 1793) Subfamily Lycaeninae Tribe Lycaenini (Coppers) Lycaena Fabricius, 1807 [We place all of our coppers in the genus Lycaena following the lead of Klots (1936); also see Pratt et al. (1993) and Lamas (in press). We feel further study is needed on the generic relationships of copper species worldwide (but see Miller & Brown 1979).] 391. arota (Boisduval, 1852) 392. phlaeas (Linnaeus, 1761) [Relationships between eastern and western North American, Eurasian, and African populations of phlaeas need further study (see Larsen 1991).] 393. cupreus (W.H. Edwards, 1870) [Tentatively includes snowi (W.H. Edwards, [1881]), although further elaboration on its status is needed.] 394. xanthoides (Boisduval, 1852) 395. dione (Scudder, 1868) 396. editha (Mead, 1878) 397. gorgon (Boisduval, 1852) 398. hyllus (Cramer, 1775) [The name hyllus appears to be a nomen dubium (see ICZN, 1999), and the use of thoe (Guйrin-Mйneville, [1831]) in its place may be warranted; see Koзak (1983).] 399. rubidus (Behr, 1866) 400. ferrisi K. Johnson & Balogh, 1977* 401. heteronea Boisduval, 1852 402. epixanthe (Boisduval & Leconte, [1835]) 403. helloides (Boisduval, 1852) 404. dorcas (W. Kirby, 1837) 405. dospassosi McDunnough, 1940 [Treated as a distinct species by Layberry et al. (1998) and Handfield (1999). Further study is needed to delineate this and related taxa (dorcas and helloides) with confidence; see Ferris (1977) and Scott (1978).] 406. nivalis (Boisduval, 1869) 407. mariposa (Reakirt, 1866) 408. hermes (W.H. Edwards, 1870) Tribe Theclini (Hairstreaks) 23
Hypaurotis Scudder, 1876 409. crysalus (W.H. Edwards, 1873) Habrodais Scudder, 1876 410. grunus (Boisduval, 1852) Tribe Eumaeini (Hairstreaks) [The order of genera within the Eumaeini follows Robbins (in press).] Eumaeus Hьbner, [1819] 411. toxea (Godart, [1824]) [The occurrence of this species in our area needs to be verified; see Kendall & McGuire (1984).] 412. atala (Poey, 1832) Atlides Hьbner, [1819] 413. halesus (Cramer, 1777) Callophrys Billberg, 1820 414. affinis (W.H. Edwards, 1862) [Tentatively includes apama (W.H. Edwards, 1882) and homoperplexa Barnes & Benjamin, 1923, in accord with the treatment of Scott (1986); however, further elaboration on the relationship betwe en affinis and homoperplexa is very badly needed.] 415. perplexa Barnes & Benjamin, 1923 [Callophrys perplexa and C. affinis occur together in the Pacific Northwest while maintaining separate hosts, habitats, and larval characteristics, according to Pyle (2002); also see Hinchliff (1996).] 416. dumetorum (Boisduval, 1852) [Previously known as C. viridis W.H. Edwards, 1862, a synonym with the same type locality (Emmel et al. (1998a).] 417. sheridanii (W.H. Edwards, 1877) [Includes comstocki Henne, 1940, and lemberti Tilden, 1963, partly on evidence presented by Austin (1998b). The complex is being studied by ADW, J. Pelham & R. Stanford.] 418. hesseli (Rawson & Ziegler, 1950) 419. nelsoni (Boisduval, 1869) [Includes C. rosneri K. Johnson, 1976*; The status of C. byrnei K. Johnson, 1976* is uncertain (see Ferris 1991). Putative areas of contact between nelsoni plicataria K. Johnson, 1976* and gryneus barryi K. Johnson, 1976* need further study (see Hinchliff 1994).] 420. thornei (J.W. Brown, 1983)* [Differences between this species and nearby juniper-feeding populations are detailed by Brown (1983, 1993).] 421. muiri (Hy. Edwards, 1881) [Identity described by Tilden (1952), where it occurs parapatrically with nelsoni. Species status was further supported by Nice and Shapiro (2001).] 24
422. gryneus (Hьbner, [1819]) [Includes all juniper-feeding taxa, including barryi, loki (Skinner, 1907), siva (W.H. Edwards, 1874), and sweadneri (Chermock, 1945), among others; see Ferris (1991).] 423. spinetorum (Hewitson, 1867) [Includes millerorum Clench, 1981* as a synonym following Robbins (1990).] 424. johnsoni (Skinner, 1904) 425. xami (Reakirt, [1867]) 426. mcfarlandi P. Ehrlich & Clench, 1960 427. augustinus (Westwood, 1852) [Guppy and Shepard (2001) treat iroides (Boisduval, 1852) as a distinct species but provide little evidence to support the split; also see Kondla (1999).] 428. mossii (Hy. Edwards, 1881) 429. fotis (Strecker, [1878]) 430. polios (Cook & Watson, 1907) 431. irus (Godart, [1824]) 432. henrici (Grote & Robinson, 1867) [Includes solatus (Cook & Watson, 1909) as a subspecies, although this is sometimes treated as a full species (e.g. Durden 1982).] 433. niphon Hьbner, [1819] 434. eryphon (Boisduval, 1852) 435. lanoraieensis (Sheppard, 1934) *[We have followed Clench (1961) and Robbins (in press) in our generic arrangement of Callophrys (s.l.), and treat Deciduphagus Johnson, 1992*, Loranthomitoura Ballmer & Pratt, 1992*, Xamia Clench, 1961*, Sandia Clench & P. Ehrlich, 1960, Mitoura Scudder, 1872 and Incisalia Scudder, 1872 as subgenera; but see Ballmer & Pratt (1992) and Johnson (1992).] Cyanophrys Clench, 1961* 436. goodsoni (Clench, 1946) 437. herodotus (Fabricius, 1793) 438. miserabilis (Clench, 1946) Rekoa Kaye, 1904 439. palegon (Cramer, 1780) 440. marius (Lucas, 1857) Arawacus Kaye, 1904 441. jada (Hewitson, 1867) Satyrium Scudder, 1876 442. favonius (J.E. Smith, 1797) 443. ilavia (Beutenmьller, 1899) 444. polingi (Barnes & Benjamin, 1926) 445. titus (Fabricius, 1793) 446. acadica (W.H. Edwards, 1862) 447. californica (W.H. Edwards, 1862) 25
448. sylvinus (Boisduval, 1852) [Includes dryope (W.H. Edwards, 1870).] 449. caryaevorus (McDunnough, 1942) 450. edwardsii (Grote and Robinson, 1867) 451. calanus (Hьbner, [1809]) 452. kingi (Klots & Clench, 1952) 453. liparops (Leconte, 1833) 454. auretorum (Boisduval, 1852) 455. tetra (W.H. Edwards, 1870) 456. saepium (Boisduval, 1852) 457. behrii (W.H. Edwards, 1870) 458. fuliginosa (W.H. Edwards, 1861) [Pyle (2002) noted that there may be two or more species in the Pacific Northwest. This possible complex is under investigation by PAO. Several apparent relatives are found in temperate Asia (Tuzov et al. 2000).] *[Species previously placed in Fixsenia Tutt, 1907 and Harkenclenus dos Passos, 1970, are now placed in Satyrium by Robbins (in press).] Phaeostrymon Clench, 1961* 459. alcestis (W.H. Edwards, 1871) Ocaria Clench, 1970 460. ocrisia (Hewitson, 1868) Chlorostrymon Clench, 1961* 461. simaethis (Drury, 1773) 462. maesites (Herrich-Schдffer, 1865) 463. telea (Hewitson, 1868) Allosmaitia Clench, [1964] 464. strophius (Godart, [1824]) [Allosmaitia pion (Godman & Salvin 1887) is a synonym, fide Robbins (in press); also see Ferris (1989c: p. 28).] Electrostrymon Clench, 1961* 465. sangala (Hewitson, 1868) [Previously known as E. endymion cyphara (Hewitson, 1874), a name placed in synonymy by Robbins (in press).] 466. joya (Dognin, 1895) [Previously referred to as E. canus (Druce, 1907), a name placed in synonymy by Robbins (in press).] 467. angelia (Hewitson, 1874) Calycopis Scudder, 1876 468. cecrops (Fabricius, 1793) 469. isobeon (Butler & H. Druce, 1872) Strymon Hьbner, 1818 470. melinus (Hьbner, [1813]) 26
471. avalona (W.G. Wright, 1905) 472. rufofusca (Hewitson, 1877) 473. albata (C. Felder & R. Felder, 1865) 474. alea (Godman & Salvin, 1887) 475. bebrycia (Hewitson, 1868) 476. yojoa (Reakirt, [1867]) 477. cestri (Reakirt, [1867]) 478. martialis (Herrich-Schдffer, 1865) 479. istapa (Reakirt, [1867]) [Formerly known as columella (Fabricius, 1793), which was determined to be a Caribbean species by Robbins & Nicolay (1999).] 480. bazochii (Godart, [1824]) 481. acis (Drury, 1773) 482. limenia (Hewitson, 1868) 483. serapio (Godman & Salvin, 1887) Tmolus Hьbner, [1819] 484. echion (Linnaeus, 1767) Ministrymon Clench, 1961* 485. leda (W.H. Edwards, 1882) 486. clytie (W.H. Edwards, 1877) 487. azia (Hewitson, 1873) Strephonota K. Johnson, Austin, Le Crom & Salazar, 1997* 488. tephraeus (Geyer, 1837) Oenomaus Hьbner, [1819] 489. ortygnus (Cramer, 1779) Parrhasius Hьbner, [1819] 490. m-album (Boisduval & Leconte, 1833) Hypostrymon Clench, 1961* 491. critola (Hewitson, 1874) Erora Scudder, 1872 492. laeta (W.H. Edwards, 1862) 493. quaderna (Hewitson, 1868) Tribe Polyommatini (Blues) [Our generic arrangement of this tribe follows Bбlint & Johnson (1995).] Leptotes Scudder, 1876 494. cassius (Cramer, 1775) 495. marina (Reakirt, 1868) Zizula Chapman, 1910 496. cyna (W.H. Edwards, 1881) Brephidium Scudder, 1876 497. exilis (Boisduval, 1852) 498. pseudofea (Morrison, 1873) 27
[Brown & Heineman (1972), Riley (1975) and Smith et al. (1994) treat pseudofea as a separate species from exilis (based on genitalic differences between them reported by Comstock & Huntington (1943)), and treat isophthalma (Herrich-Schдffer, 1862) as a subspecies of exilis. Note that Bordelon & Knudson (2000) mention possible intermediates between exilis and pseudofea from Texas, and Pavulaan & Gatrelle (1999) suggest exilis and pseudofea may be conspecific (as treated by Scott (1986)), but provide little evidence to support their claim; further study is badly needed.] Cupido Schrank, 1801 499. comyntas (Godart, [1824]) 500. amyntula (Boisduval, 1852) [We follow Karsholt & Razowski (1996), Hesselbarth et al. (1995: 547), and Gorbunov (2001) in treating Everes Hьbner, [1819] as a subgenus of Cupido.] Celastrina Tutt, 1906 501. ladon (Cramer, 1780) 502. lucia (W. Kirby, 1837) [Evidence of this taxon being a full species is provided by Pavulaan (1995), Kondla (1999), Nielsen (1999), and Oehlenschlager & Huber (2002).] 503. neglecta (W.H. Edwards, 1862) 504. echo (W.H. Edwards, 1864) [Treated as a full species by Guppy & Shepard (2001).] *[Further elaboration on the relationships between echo, nigrescens (Fletcher, 1903), sidara (Clench, 1944), cinerea (W.H. Edwards, 1883), gozora (Boisduval, 1870), and Baja California populations (see Brown et al. 1992) is needed to determine their status; some or all of which may represent species-level taxa.] 505. nigra (Forbes, 1960)* [Celastrina ebenina Clench, 1972* is a synonym; see Scott & Wright (1993).] 506. neglectamajor Opler & Krizek, 1984* 507. humulus Scott & D. Wright, 1998* 508. idella Wright & Pavulaan, 1999* *[Additional species-level entities, including at least one undescribed species, have been recognized by some (see Pratt et al. (1994), Wright (1995) and Layberry et al. (1998)), including violacea (W.H. Edwards, 1866).] *[It is uncertain which, if any, of our North American Celastrina taxa are conspecific with argiolus (Linnaeus, 1758), where they have traditionally been placed (see Eliot & Kawazoй 1983).] Glaucopsyche Scudder, 1872 509. piasus (Boisduval, 1852) 510. lygdamus (Doubleday, 1841) [The extinct G. xerces (Boisduval, 1852) was suggested to be a subspecies of lygdamus by Scott (1986).] Philotes Scudder, 1876 511. sonorensis (C. Felder & R. Felder, 1865) 28
Philotiella Mattoni, [1978] 512. speciosa (Hy. Edwards, 1877) 513. leona Hammond & McCorkle, 2000* Euphilotes Mattoni, [1978] [Most recent treatments of this genus are by Mattoni (1989), Pratt (1988, 1994, 1999), Pratt & Ballmer (1993), and Pratt & Emmel (1998). It is likely that additional specieslevel taxa will be delineated in the future.] 514. battoides (Behr, 1867) [Tentatively includes, among others, centralis (Barnes & McDunnough, 1917).] 515. bernardino (Barnes & McDunnough, 1916) 516. intermedia (Barnes & McDunnough, 1917) [Includes oregonensis (Barnes & McDunnough, 1917).] 517. ellisi (Shields, 1975)* 518. baueri (Shields, 1975)* 519. enoptes (Boisduval, 1852) 520. ancilla (Barnes & McDunnough, 1918) 521. mojave (Watson & W.P. Comstock, 1920) 522. rita (Barnes & McDunnough, 1916) 523. pallescens (Tilden & Downey, 1955) 524. spaldingi (Barnes & McDunnough, 1917) Hemiargus Hьbner, 1818 525. ceraunus (Fabricius, 1793) [Note that Brown & Heineman (1972), Riley (1975), Smith et al. (1994) and Lamas (in press) treat ceraunus as a subspecies of hanno (Stoll, 1790); however, Clench (1977) and Johnson & Bбlint (1995) consider the two as separate species, and Schwartz (1989) reported hanno and ceraunus as widely sympatric on Hispaniola. The situation needs further study.] Echinargus Nabokov, 1945* 526. isola (Reakirt, [1867]) Cyclargus Nabokov, 1945* 527. thomasi (Clench, 1941) 528. ammon (Lucas, 1857) [This species was first reported in our region by Beck (1985), from Big Pine Key, Monroe Co., Florida; it was subsequently photographed there by Krizek (1998).] *[Generic treatment of Hemiargus, Echinargus, and Cyclargus follows Nabokov (1945), Smith et al. (1994), Bбlint & Johnson (1995), Johnson & Bбlint (1995), and Lamas (in press); also see Brown & Heineman (1972).] Plebejus Kluk, 1780 529. idas (Linnaeus, 1761) 530. anna (W.H. Edwards, 1861) [This taxon was treated as a full species by Bбlint & Johnson (1997), Guppy & Shepard (2001), and Pyle (2002); also see Nice & Shapiro (1999). More fieldwork and 29
laboratory investigations are needed to clarify the situation in North America and Eurasia (see Tuzov et al. 2000).] 531. melissa (W.H. Edwards, 1873) [See Lane & Weller (1994) for details on the taxonomic status of P. melissa populations, including samuelis (Nabokov, 1944).] 532. saepiolus (Boisduval, 1852) 533. emigdionis (F. Grinnell, 1905) 534. icarioides (Boisduval, 1852) 535. shasta (W.H. Edwards, 1862) 536. acmon (Westwood, [1851]) 537. lupini (Boisduval, 1869) [The previous two species appear to represent a species complex containing, at present, an unknown number of species. The group appears to be much more diverse than was suggested by Goodpasture (1973, 1974). Recently, Scott (1998) transferred several subspecies from acmon to lupini, but presented almost no data to support the changes. The complex is currently under study by PAO (Opler, 2003).] 538. neurona (Skinner, 1902) 539. optilete (Knoch, 1781) 540. glandon (de Prunner, 1798) 541. podarce (C. Felder & R. Felder, 1865) 542. cassiope (J. Emmel & T. Emmel, 1998)* [Relationships of the above three taxa were discussed by Emmel & Emmel (1998), but are still under debate. Some authors recognize additional species, including rusticus (W.H. Edwards, 1865) and franklinii (Curtis, 1835); see Ferris (1989c).] *[Our treatment of Plebejus follows that presented by Gorbunov (2001). He refers to it as a "supergenus" with many subgenera, which in our area include Lycaeides Hьbner, [1819], Plebulina Nabokov, [1945], Icaricia Nabokov, [1945], Vacciniina Tutt, 1909, and Agriades Hьbner, [1819]. Bбlint & Johnson (1997) present a considerably different arrangement of the group, placing our members in Aricia [Reichenbach], 1817, Albulina Tutt, 1909 and Plebejus. None of these authors retain the use of Lycaeides, Icaricia, Vacciniina or Plebulina as valid genera.] FAMILY RIODINIDAE [Our treatment of the Riodinidae as a family follows Robbins (1988a,b) and Callaghan and Lamas (in press), but should be considered a tentative arrangement only. The relationship of this group to the Lycaenidae and Danainae requires further study, as noted by Campbell et al. (2000).] Subfamily Riodininae Calephelis Grote & Robinson, 1869 543. virginiensis (Guйrin-Mйneville, [1832]) 544. borealis (Grote & Robinson, 1866) 30
545. muticum McAlpine, 1937 546. nemesis (W.H. Edwards, 1871) 547. perditalis Barnes & McDunnough, 1918 548. wrighti Holland, 1930 549. rawsoni McAlpine, 1939 [Scott (1986) lumped freemani McAlpine, 1971* and arizonensis McAlpine, 1971* as subspecies of rawsoni, yet presented no data in support of these changes (but did describe genitalic differences between these taxa). We do not accept his combinations, and stress that further study of this genus is badly needed; Neck (1996) treats freemani as a full species. The status of C. dreisbachi McAlpine, 1971* (one specimen from Arizona in the type series) needs clarification, and is tentatively not included in our list of species.] 550. freemani McAlpine, 1971* 551. arizonensis McAlpine, 1971* Caria Hьbner, 1823 552. ino Godman & Salvin, 1886 Lasaia H.W. Bates, 1868 553. sula Staudinger, 1888 554. maria Clench, 1972* [Specimen collected in Guadalupe Canyon, Cochise Co., Arizona, by Kilian Roever; determination confirmed by R. Bailowitz.] Melanis Hьbner, [1819] 555. pixe (Boisduval, 1836) Emesis Fabricius, 1807 556. zela Butler, 1870 557. ares (W.H. Edwards, 1882) [Callaghan and Lamas (in press) treat ares as a synonym of zela cleis (W.H. Edwards, 1882), but we feel that further study of the types is needed, since two species indeed appear to be involved in our region (e.g. Bailowitz & Brock 1991).] 558. emesia (Hewitson, 1867) 559. tenedia C. Felder & R. Felder, 1861 Apodemia C. Felder & R. Felder, 1865 560. mormo (C. Felder & R. Felder, 1859) 561. duryi (W.H. Edwards, 1882) [See Toliver et al. (1994) for notes on the specific status of duryi.] 562. virgulti (Behr, 1865) 563. mejicanus (Behr, 1865) [Species-level relationships of the mormo complex (above four taxa) remain poorly understood, and our arrangement is tentative. Our treatment is based, in part, on information in Pratt & Ballmer (1991), Cary & Holland (1992) and Toliver et al. (1994). The group has not been considered as a whole since the analysis of Opler & Powell (1962).] 31
564. palmerii (W.H. Edwards, 1870) 565. hepburni Godman & Salvin, 1886 566. walkeri Godman & Salvin, 1886 567. multiplaga Schaus, 1902 568. phyciodoides Barnes & Benjamin, 1924 [Possibly extirpated in U.S. portion of range (Bailowitz & Brock 1991); also see Holland & Forbes (1981).] 569. nais (W.H. Edwards, 1877) 570. chisosensis H.A. Freeman, 1964* FAMILY NYMPHALIDAE [Our subfamily and generic arrangement generally follows Harvey (1991), as modified by Ackery et al. (1999), Freitas (1999), Brower (2000), Wahlberg (2002), Freitas & Brown (in preparation) and Lamas (in press), although studies are under way to further resolve relationships. It is possible that the family, as presented here, is not a monophyletic group, since the exact placement of the Libytheinae and Danainae remains uncertain (see Campbell et al. 2000). ] Subfamily Libytheinae Libytheana Michener, 1943 571. carinenta (Cramer, 1777) [Many authors (e.g. Field 1940, Friedlander 1984, Shields 1984, 1985, Okano 1987, Ferris 1989c, Motono 1993, and Austin & Emmel 1998) treat bachmanii (Kirtland, 1851) and carinenta as separate species. However, little information on populations across North and Central America is available, and reported genitalic differences between the two do not hold up in series (Kawahara 2001, pers. comm. 2002).] 572. motya (Hьbner, [1823]) [Heitzman & Heitzman (1973) reported this species from Texas as rare strays or temporary colonists from Cuba; see Alayo and Hernбndez (1981).] Subfamily Danainae [In our area this subfamily includes the tribes Danaini and Ithomiini; see Ackery & Vane Wright (1984), Ackery (1987), and Ackery et al. (1999).] Danaus Kluk, 1780 573. plexippus (Linnaeus, 1758) 574. gilippus (Cramer, 1775) 575. eresimus (Cramer, 1777) Lycorea Doubleday, [1847] 576. halia (Hьbner, 1816) [Central American populations are sometimes treated as species cleobaea (Godart, 1819), e.g. Miller & Brown (1981), Schwartz (1989), Brown (1992), and Smith et al. 32
(1994); but see Lamas (in press).] Dircenna Doubleday, 1847 577. klugii (Geyer, 1837) Subfamily Heliconiinae Dione Hьbner, [1819] 578. moneta Hьbner, [1825] Agraulis Boisduval & Leconte, [1835] 579. vanillae (Linnaeus, 1758) Dryas Hьbner, [1807] 580. iulia (Fabricius, 1775) Dryadula Michener, 1942 581. phaetusa (Linnaeus, 1758) Eueides Hьbner, 1816 582. isabella (Stoll, 1781) Heliconius Kluk, 1780 583. charithonia (Linnaeus, 1767) [Spelling corrected to original orthography (Brower 1994).] 584. erato (Linnaeus, 1758) Euptoieta Doubleday, 1848 585. claudia (Cramer, 1775) 586. hegesia (Cramer, 1779) Speyeria Scudder, 1872 587. diana (Cramer, 1777) 588. cybele (Fabricius, 1775) [Includes leto (Behr, 1862), although this is treated as a full species by some (e.g. Kondla 1999).] 589. aphrodite (Fabricius, 1787) 590. idalia (Drury, 1773) 591. nokomis (W.H. Edwards, 1862) 592. edwardsii (Reakirt, 1866) 593. coronis (Behr, 1864) 594. zerene (Boisduval, 1852) 595. carolae (dos Passos & Grey, 1942) [Although somewhat intermediate between coronis and zerene, this isolated, disjunct taxon was considered a full species by Emmel & Austin (1998).] 596. callippe (Boisduval, 1852) 597. egleis (Behr, 1862) 598. adiaste (W.H. Edwards, 1864) 599. atlantis (W.H. Edwards, 1862) 600. hesperis (W.H. Edwards, 1864) [Pyle (2002) treats the above two taxa as a single species based on unpublished 33
research by Paul Hammond. Until this evidence is presented, we follow the arrangement proposed by Scott et al. (1998); also see Ferris (1983).] 601. hydaspe (Boisduval, 1869) 602. mormonia (Boisduval, 1869) Boloria Moore, [1900] [We follow a broad usage of Boloria; see dos Passos & Grey (1945), dos Passos (1964), Grey (1957, 1989) and Gorbunov (2001). Clossiana Reuss, 1920 and Proclossiana Reuss, 1926 are considered to be subgenera or synonyms (but see Warren (1944), Miller & Brown (1981) and Aubert et al. (1996)).] 603. alaskensis (Holland, 1900) [Boloria napaea (Hoffmansegg, 1804) is a separate Eurasian species with differing genitalia that reportedly occurs sympatrically with alaskensis at some locations. See discussions by Crosson du Cormier (1977), Dubatolov (1992), Kosterin (2000), Tuzov et al. (2000), and Gorbunov (2001).] 604. eunomia (Esper, 1800) 605. selene ([Denis & Schiffermьller], 1775) 606. bellona (Fabricius, 1775) 607. frigga (Thunberg, 1791) 608. improba (Butler, 1877) [Includes acrocnema Gall & Sperling, 1980*.] 609. kriemhild (Strecker, 1879) 610. epithore (W.H. Edwards, 1864) 611. polaris (Boisduval, [1828]) 612. freija (Thunberg, 1791) 613. natazhati (Gibson, 1920) [Taxonomic status as a full species detailed by Troubridge & Wood (1995), Shepard et al. (1998), and Guppy & Shepard (2001).] 614. alberta (W.H. Edwards, 1890) 615. astarte (Doubleday, [1847]) [Tentatively includes distincta (Gibson, 1920), following dos Passos (1961); but see Wyatt (1958). The relationship of astarte to distincta and tschukotkensis (Wyatt, 1961) needs further elaboration. Guppy & Shepard (2001) very briefly summarize reasons for using tritonia (Bцber, 1812) for our North American populations, a combination also implied by Gorbunov (2000). However, no comprehensive review of the complex has been presented, and Tuzov et al. (2000) maintain tritonia and distincta as separate species.] 616. chariclea (Schneider, 1794) [Includes our former concept of titania (Esper, [1793]), a wholly Palaearctic taxon (see Shepard 1998); also includes montinus (Scudder, 1863). However, the possibility exists that more than one species is being represented in our area by this name (see Layberry et al. 1998).] 34
Subfamily Nymphalinae Poladryas Bauer, 1961 617. minuta (W.H. Edwards, 1861) 618. arachne (W.H. Edwards, 1869) [We do not feel that Scott (1974, 1986) presents sufficient data to lump the above two taxa. Several authors also maintain arachne and minuta as full species, e.g. Ferris (1989c) and Austin (1998a); however, further research in the U.S. and Mexico is needed to resolve the situation.] Chlosyne Butler, 1870 619. theona (Mйnйtriйs, 1855) [Includes chinatiensis (Tinkham, 1944), according to Austin & Smith (1998).] 620. cyneas (Godman & Salvin, 1878) 621. fulvia (W.H. Edwards, 1879) 622. leanira (C. Felder & R. Felder, 1860) 623. californica (W.G. Wright, 1905) 624. lacinia (Geyer, 1837) 625. definita (E.M. Aaron, [1885]) 626. endeis (Godman & Salvin, 1894) 627. janais (Drury, 1782) 628. rosita A. Hall, 1924 629. melitaeoides (C. Felder & R. Felder, 1867) [See Kendall & McGuire (1984) for reports from Texas.] 630. eumeda (Godman & Salvin, 1894) [Kons (2000) and Luis et al. (2003) consider eumeda, melitaeoides and marina (Geyer, 1837) to be separate species (these were lumped by Scott (1986) without justification). The report from Pima Co., Arizona, appears to be of the semi-desert species, eumeda; however, C. marina also occurs in Sonora, Mexico, in montane habitats.] 631. gorgone (Hьbner, [1810]) 632. nycteis (Doubleday, [1847]) [A petition to the International Commission on Zoological Nomenclature to suppress ismeria (Boisduval & Leconte, [1835]), a possible senior synonym of nycteis (see Gatrelle 1998, 2000), is in preparation; see ICZN (1999).] 633. harrisii (Scudder, 1864) 634. palla (Boisduval, 1852) 635. gabbii (Behr, 1863) 636. acastus (W.H. Edwards, 1874) [Includes neumoegeni (Skinner, 1895), among others.] 637. whitneyi (Behr, 1863) [Tentatively includes damoetas (Skinner, 1902), following Scott (1998) and Kons (2000), but further elaboration on the relationships of taxa in this group is needed.] 638. hoffmanni (Behr, 1863) *[Charidryas Scudder, 1872 (see Higgins 1960) and Thessalia Scudder, 1875 (see 35
Wahlberg & Zimmerman 2000) are considered synonyms of Chlosyne.] Microtia H.W Bates, 1864 639. elva H.W. Bates, 1864 Dymasia Higgins, 1960* 640. dymas (W.H. Edwards, 1877) Texola Higgins, 1959 641. elada (Hewitson, 1868) Phyciodes Hьbner, [1819] 642. graphica (R. Felder, 1869) [Phyciodes graphica, described in April, 1869, supersedes P. vesta (W.H. Edwards, 1869), which was described in September-October, 1869; see Lamas (in press).] 643. picta (W.H. Edwards, 1865) 644. orseis W.H. Edwards, 1871 645. pallida (W.H. Edwards, 1864) 646. mylitta (W.H. Edwards, 1861) 647. phaon (W.H. Edwards, 1864) 648. tharos (Drury, 1773) 649. cocyta (Cramer, 1777) [Supersedes selenis (W. Kirby, 1837), an apparent junior synonym; see Scott (1994).] 650. batesii (Reakirt, [1866]) 651. pulchella (Boisduval, 1852) [Lectotype designation of pulchella by Emmel et al. (1998a) displaces pratensis (Behr, 1863). See Miller & Brown (1981) for synonymy of campestris (Behr, 1863).] *[Our treatment of Phyciodes follows Wahlberg et al. (2003).] Anthanassa Scudder, 1875 [Recognition of this genus follows Wahlberg & Zimmerman (2000).] 652. frisia (Poey, 1832) 653. tulcis (H.W. Bates, 1864) 654. texana (W.H. Edwards, 1863) [Anthanassa seminole (Skinner, 1911) was suggested to be a species distinct from texana by Watts & Habeck (1991), and treated as such by Neck (1996); further study is needed to determine its status.] 655. ptolyca (H.W. Bates, 1864) 656. argentea (Godman & Salvin, 1882) [First U.S. record reported by Chuah & Cushing (1995); also see Bordelon & Knudson (2000). Anthanassa ardys (Hewitson, 1864) may have also strayed into Texas (see Excluded Species).] Tegosa Higgins, 1981* 657. anieta (Hewitson, 1864) Euphydryas Scudder, 1872 658. gillettii (Barnes, 1897) 659. phaeton (Drury, 1773) 36
660. editha (Boisduval, 1852) 661. chalcedona (Doubleday, [1847]) 662. anicia (Doubleday, [1847]) [We tentatively consider anicia to be a separate species from chalcedona following Ferris (1989b), Guppy & Shepard (2001) and Pyle (2002), however further elaboration on the relationship between the two taxa is needed. While the taxa clearly behave as two species in the Pacific Northwest, their relationship appears to be less well defined in other areas, e.g. Brussard et al. (1989), Austin & Murphy (1998). We tentatively consider colon (W.H. Edwards, 1881) to be conspecific with chalcedona, but again, much additional research is needed. A recent study by Zimmerman et al. (2000) did not resolve relationships among these taxa.] *[Following Wahlberg & Zimmerman (2000) and Zimmerman et al. (2000), Occidryas Higgins, 1978* and Hypodryas Higgins, 1978* are considered synonyms of Euphydryas.] Hypolimnas Hьbner, [1819] 663. misippus (Linnaeus, 1764) Junonia Hьbner, [1819] [Our use of Junonia instead of Precis Hьbner, [1819] follows de Lesse (1952), Turner & Parnell (1985), and Lamas (in press); but see Hemming (1934) and Comstock (1944).] 664. coenia Hьbner, [1822] 665. evarete (Cramer, 1779) [Includes nigrosuffusa Barnes & McDunnough, 1916 (contra Turner & Parnell 1985), following Lamas (in press).] 666. genoveva (Stoll, 1780) *[The relationships between these Junonia taxa need further elaboration; see Hafernik (1982), Turner & Parnell (1985), and Lamas (in press).] Anartia Hьbner, [1819] 667. jatrophae (Linnaeus, 1763) 668. fatima (Fabricius, 1793) [Sometimes treated as a subspecies of amathea (Linnaeus, 1758), e.g. Lamas (in press); but see Silberglied et al. (1979), Dasmahapatra et al. (2002), and Blum et al. (2003).] 669. chrysopelea (Hьbner, [1831]) [Reported as a subspecies of lytrea (Godart, 1819) by Anderson (1974), Bennett & Knudson (1976), Scott (1986), and Lamas (in press); but see Silberglied et al. (1979), Alayo & Hernбndez (1981), and Blum et al.(2003).] Siproeta Hьbner, [1823] 670. stelenes (Linnaeus, 1758) 671. epaphus (Latreille, [1813]) Polygonia Hьbner, [1819] 672. interrogationis (Fabricius, 1798) 37
673. comma (Harris, 1842) 674. satyrus (W.H. Edwards, 1869) 675. faunus (W.H. Edwards, 1862) 676. gracilis (Grote & Robinson, 1867) [Includes zephyrus (W.H. Edwards, 1870); see Scott (1984) and Layberry et al. (1998).] 677. progne (Cramer, 1775) 678. oreas (W.H. Edwards, 1869) [This species was treated as distinct from progne (contra Scott 1984) by Bird et al. (1995), Layberry et al. (1998), and Guppy & Shepard (2001).] Roddia Korshunov, 1995 Aglais Dalman, 1816 679. milberti (Godart, 1819) 680. urticae (Linnaeus, 1758) [Several individuals of this common Palaearctic butterfly have been captured or photographed (e.g. Glassberg 1992, Zirlin & Ingraham 1997, Zirlin 2002), and this species is likely either an occasional accidental introduction or, less likely, a periodic vagrant.] Nymphalis Kluk, 1780 681. vaualbum (Denis and Schiffermьller) [Guppy & Shepard (2001: pp. 256-257) indicate that vaualbum may be a nomen nudum (see ICZN 1999), and use l-album (Esper, 1781) in its place. This treatment is repeated by Wahlberg & Nylin (2003). A petition to the ICZN is being prepared to resolve the issue, and until the commission rules, we maintain the name in widest use, vaualbum, as recommended by the ICZN (1999). See Sattler & Tremewan (1984) for further discussion.] [The inclusion of l-album (formerly as vau-album) in Roddia Korshunov, 1995 by Guppy and Shepard (2001) and others is reversed by Wahlberg & Nylin (2003).] 682. antiopa (Linnaeus, 1758) 683. californica (Boisduval, 1852) [Miller & Miller (1990) treat californica as a subspecies of the Eurasian xanthomelas Esper [1781], but see Nylin et al. (2001).] *[Our arrangement of Nymphalis and Aglais follows Nylin et al. (2001) as modified by Wahlberg and Nylin (2003).] Vanessa Fabricius, 1807 684. atalanta (Linnaeus, 1758) 685. cardui (Linnaeus, 1758) 686. annabella (Field, 1971)* 687. virginiensis (Drury, 1773) [Some authors place the above three species in Cynthia Fabricius, 1807, following Field (1971), but several natural hybrids between atalanta and annabella are known (Comstock 1927, Dimock 1973, Tilden & Smith 1986).] 38
Hypanartia Hьbner, [1821] 688. lethe (Fabricius, 1793) Historis Hьbner, [1819] 689. odius (Fabricius, 1775) 690. acheronta (Fabricius, 1775) Smyrna Hьbner, [1823] 691. blomfildia (Fabricius, 1781) Subfamily Limenitidinae [Our treatment of the Limenitidinae and Biblidinae follows Freitas (1999), Wahlberg (2002) and Freitas & Brown (in preparation), although Lamas (in press) places this subfamily within the Biblidinae, and Marpesia may belong elsewhere; further study is needed.] Limenitis Fabricius, 1807 [Basilarchia Scudder, 1872 is considere d a synonym; see Chermock (1950), Niculescu (1986), Tuzov et al. (2000), and Gorbunov (2001).] 692. arthemis (Drury, 1773) 693. archippus (Cramer, 1775) 694. lorquini (Boisduval, 1852) 695. weidemeyerii W.H. Edwards, 1861 [Suggested to be conspecific with lorquini by Porter (1990), but see Boyd et al. (1999).] Adelpha Hьbner, [1819] 696. bredowii Geyer, 1837 697. fessonia (Hewitson, 1847) 698. basiloides (H.W. Bates, 1865) [Opler (1992) and Neck (1996) reported this species from Texas.] Marpesia Hьbner, 1818 699. chiron (Fabricius, 1775) 700. petreus (Cramer, 1776) 701. eleuchea Hьbner, 1818 702. zerynthia Hьbner, [1823] [This species is occasionally listed as M. coresia (Godart, [1824]), a junior synonym of zerynthia (see Neild 1996).] Subfamily Biblidinae Biblis Fabricius, 1807 703. hyperia (Cramer, 1779) Mestra Hьbner, [1825] 704. amymone (Mйnйtriйs, 1857) [This is sometimes treated as a subspecies of the Jamaican M. dorcas (Fabricius, 1775), e.g. Lamas (in press).] 39
Eunica Hьbner, [1819] 705. monima (Stoll, 1782) 706. tatila (Herrich-Schдffer, [1855]) Myscelia Doubleday, [1845] 707. ethusa (Doyиre, [1840]) 708. cyananthe C. Felder & R. Felder, 1867 Dynamine Hьbner, [1819] 709. dyonis Geyer, 1837 [Sometimes considered to be a subspecies of the Caribbean D. egaea (Fabricius, 1775), e.g. de la Maza & Turrent (1985) and Smith et al. (1994), but serina (Fabricius, 1775) has precedence over egaea (see Lamas in press).] Diaethria Billberg, 1820 710. species [The west Texas Diaethria record (see Kendall & McGuire 1984, Neck 1996) is apparently of D. anna (Guйrin-Mйneville, [1844]) or D. astala (Guйrin-Mйneville, [1844]), since these are the only Diaethria species resident in northeastern Mexico (de la Maza & Turrent 1985); however, the west Texas specimen has not been positively determined and is reportedly no longer extant. There are also unverified records for both anna and astala from south Texas (Stanford 2002). See Excluded Species.] Epiphile Doubleday, [1845] 711. adrasta Hewitson, 1861 Hamadryas Hьbner, [1806] 712. februa (Hьbner, [1823]) 713. amphichloe (Boisduval, 1870) 714. glauconome (H.W. Bates, 1864) 715. atlantis (H.W. Bates, 1864) 716. feronia (Linnaeus, 1758) 717. guatemalena (H.W. Bates, 1864) 718. iphthime (H.W. Bates, 1864) 719. amphinome (Linnaeus, 1767) Subfamily Charaxinae Anaea Hьbner, [1819] 720. troglodyta (Fabricius, 1775) [Includes floridalis F. Johnson & W.P. Comstock, 1941, in accordance with Lamas (in press), but see discussion by Smith et al. (1994: p. 65).] 721. aidea (Guйrin-Mйneville, [1844]) [Often considered a species-level taxon (e.g. DeVries 1987), but sometimes treated as a subspecies of troglodyta (e.g. Lamas in press).] 722. andria Scudder, 1875 Memphis Hьbner, [1819] 40
723. glycerium (Doubleday, [1849]) [Some authors (e.g. Lamas in press) place glycerium in the genus Fountainea Rydon, 1971*, but the monophyly of Fountainea has not been demonstrated and is questionable.] 724. pithyusa (R. Felder, 1869) 725. echemus (Doubleday, [1849]) Subfamily Apaturinae Asterocampa Rцber, 1915 726. celtis (Boisduval & Leconte, [1835]) 727. leilia (W.H. Edwards, 1874) 728. clyton (Boisduval & Leconte, [1835]) 729. idyja (Geyer, [1828]) [It is possible that Asterocampa argus (H.W. Bates, 1864), currently treated as the continental subspecies of idyja, may comprise a separate species (see Smith et al. 1994).] *[Our treatment of Asterocampa species follows Friedlander (1987), although further study of the genus is needed.] Doxocopa Hьbner, [1819] 730. laure (Drury, 1773) 731. pavon (Latreille, [1809]) Subfamily Morphinae Morpho Fabricius, 1807 732. polyphemus Westwood, [1850] Subfamily Satyrinae [Our arrangement of this subfamily generally follows the classification presented by Miller (1968), as modified by Harvey (1991).] Enodia Hьbner, [1819] 733. portlandia (Fabricius, 1781) 734. anthedon A.H. Clark, 1936 735. creola (Skinner, 1897) Satyrodes Scudder, 1875 736. eurydice (Linnaeus, 1763) 737. appalachia (R.L. Chermock, 1947) Cyllopsis R. Felder, 1869 738. pyracmon (Butler, 1867) [Cyllopsis henshawi (W.H. Edwards, 1876) is treated as a seasonal form of pyracmon, as suggested by Bailowitz & Brock (1991) and Brock (1998); but see Miller (1974).] 739. pertepida (Dyar, 1912) 740. gemma (Hьbner, 1808) 41
Hermeuptychia Forster, 1964* 741. sosybius (Fabricius, 1793) [The status of sosybius versus hermes (Fabricius, 1775) is uncertain and requires detailed study; see Forster (1964), Miller & Brown (1981: p. 241, note 624), and Smith et al. (1994). The use of hermes prevails in the Neotropical literature, e.g. Lamas (in press), for what may be (at least in part) the same species that occurs in our region.] Neonympha Hьbner, [1818] 742. areolatus (J.E. Smith, 1797) 743. helicta (Hьbner, 1808) [Species status recently proposed by Gatrelle (1999); however, further research on this group is needed to clarify overall distributions and confirm the status of helicta.] 744. mitchellii French, 1889 Megisto Hьbner, [1819] 745. cymela (Cramer, 1777) [Includes viola (Maynard, 1891), following Catling & Calhoun (1997); however, it is still possible that more than one species is included under the name cymela, and further study is needed.] 746. rubricata (W.H. Edwards, 1871) Paramacera Butler, 1868 747. xicaque (Reakirt, [1867]) [Tentatively includes allyni L. Miller, 1972*, following Scott (1986), but see Miller (1972). Further study is needed.] Coenonympha Hьbner, [1819] 748. haydenii (W.H. Edwards, 1872) 749. tullia (Mьller, 1764) [Treated as a species complex by Ferris (1989c); also see Davenport (1941) and Brown (1955, 1961). Included here are several taxa sometimes considered full species (e.g. dos Passos 1958); these include (among others) ampelos W.H. Edwards, 1871, california Westwood, 1851, inornata W.H. Edwards, 1861, kodiak W.H. Edwards, 1869, ochracea W.H. Edwards, 1861, and nipisiquit McDunnough, 1939. Layberry et al. (1998) treated nipisiquit as a full species based on (then) unpublished evidence by R. Webster; Webster (1999) subsequently treated nipisiquit as a subspecies of tullia but stressed that additional research was needed to determine its status. Handfield (1999) gave nipisiquit species-level status. Although gene flow between purported species was demonstrated by Porter & Geiger (1988) (also see Porter & Mattoon 1989), it is unclear which, if any, of our taxa are conspecific with the Eurasian C. tullia; see Kondla (1999) and Guppy & Shepard (2001). It is likely that more than one species-level taxon is included in this complex. Further research is badly needed since there is currently no satisfactory taxonomic arrangement for this group.] Cercyonis Scudder, 1875 42
750. pegala (Fabricius, 1775) 751. sthenele (Boisduval, 1852) 752. meadii (W.H. Edwards, 1872) 753. oetus (Boisduval, 1869) Erebia Dalman, 1816 754. vidleri Elwes, 1898 755. rossii (Curtis, 1835) 756. disa (Thunberg, 1791) 757. mancinus Doubleday, [1849] [See Layberry et al. (1998) for separation of mancinus from disa.] 758. magdalena Strecker, 1880 759. mackinleyensis Gunder, 1932 [See Hilchie (1990) and Layberry et al. (1998) for separation of mackinleyensis from magdalena; however, Gorbunov (2001) treats the two as conspecific.] 760. fasciata Butler, 1868 761. discoidalis (W. Kirby, 1837) 762. pawloskii Mйnйtriйs, 1859 [North American populations of this species have been referred to as Erebia theano (Tauscher, 1809), but Guppy & Shepard (2001) note that Erebia theano is restricted to eastern Siberia and Mongolia. See Tuzov et al. (1997).] 763. youngi Holland, 1900 764. occulta Roos & Kimmich, 1983* [Our use of occulta over anyuica Kurentzov, 1966 follows Roos & Arnscheid (1984), Tuzov et al. (1997) and Belik & Zamolodchikov (2002); but see Dubatolov (1992) and Layberry et al. (1998).] 765. lafontainei Troubridge & Philip, 1983* 766. callias W.H. Edwards, 1871 767. epipsodea Butler, 1868 Gyrocheilus Butler, 1867 768. patrobas (Hewitson, 1862) Neominois Scudder, 1875 769. ridingsii (W.H. Edwards, 1865) [Scott (1998) described "Hipparchia (Neominois) ridingsii wyomingo" as a "new subspecies (or species?)", and suggested it is genetically isolated from ridingsii; Opler (1999) subsequently treated this as a full species. Additional field observations have shed some light on the situation but many questions remain and the status of wyomingo remains uncertain. The situation is currently under study by Matthew Garhart.] Oeneis Hьbner, [1819] 43
770. philipi Troubridge, 1988* [The name rosovi Kurentzov, 1970 (a potential senior synonym of philipi) cannot be used in any meaningful way until a lectotype is designated, since two syntypes exist (but see Layberry et al. 1998). Oeneis rosovi was treated as a subspecies of O. norna (Thunberg, 1791) by Lukhtanov (1989), and as a synonym of that species by Gorbunov (2001). See also Lukhtanov & Eitschberger (2000).] 771. polixenes (Fabricius, 1775) 772. jutta (Hьbner, [1806]) 773. melissa (Fabricius, 1775) 774. alpina Kurentzov, 1970 [Includes excubitor Troubridge, Philip, Scott & Shepard, 1982* as a synonym, following Layberry et al. (1998).] 775. bore (Esper, 1798) [Tentatively includes taygete Geyer, [1830], following Layberry et al. (1998), but further study is needed to clarify the relationship of these taxa; Tuzov et al. (1997) consider the taxa separate species.] 776. chryxus (Doubleday, [1849]) [Includes ivallda (Mead, 1878), following Porter & Shapiro (1991).] 777. alberta Elwes, 1893 778. nevadensis (C. Felder & R. Felder, 1867) 779. macounii (W.H. Edwards, 1885) 780. uhleri (Reakirt, 1866) [Hassler & Feil (2002) recently reported the occurrence of Oeneis nanna (Mйnйtriйs, 1859) in North America, with the description of a new subspecies (kluanensis). The authors did not examine sufficient material of uhleri, and the proper taxonomic placement of this taxon is currently being studied.] 44
Hawaiian Butterflies Although not part of the North American continent, Hawaii is one of the 50 United States. For sake of completeness we present a list of the butterflies recorded from the Hawaiian Islands, based on Riotte & Uchida (1979). Additional information is provided by Zimmerman (1958). Family Hesperiidae Erionota torus Evans, 1941 [Originally misdetermined as E. thrax (Linnaeus, 1767) by Riotte & Uchida (1979).] Hylephila phyleus (Drury, 1773) Family Papilionidae Papilio xuthus Linnaeus, 1767 Family Pieridae Pieris rapae (Linnaeus, 1758) Family Lycaenidae Tmolus echion (Linnaeus, 1767) Strymon bazochii (Godart, [1824]) Lampides boeticus (Linnaeus, 1767) Brephidium exilis (Boisduval, 1852) *Udara blackburni (Tuely, 1878) [See Eliot & Kawazoй (1983) for generic combination.] Family Nymphalidae Agraulis vanillae (Linnaeus, 1758) Vanessa virginiensis (Drury, 1773) Vanessa cardui (Linnaeus, 1758) Vanessa atalanta (Linnaeus, 1758) *Vanessa tameamea Eschscholtz, 1821 Danaus plexippus (Linnaeus, 1758) ___________________________________________________ *Endemic to Hawaiian Islands. All others are introduced exotics. 45
Excluded Species The species listed below have been excluded from the North American fauna due to erroneous or inadequate documentation, or recent change in status.\1 This is not an exhaustive list, but treats most discrepancies in the post-1980 literature that are not discussed in the text. See Calhoun (1997) for a complete list of erroneous Florida records. Hesperiidae Phocides urania (Westwood, [1852]) [Dubious Texas records reported by Aaron (1890); Arizona records are unsubstantiated (see Bailowitz & Brock 1991).] Polythrix asine (Hewitson, 1867) [Erroneously reported from the U.S. based on an unjustifiable assumption of conspecificity with mexicanus by Scott (1986); see Freeman (1969) and Austin & Warren (2002).] Polythrix procerus (Plцtz, 1880) [Misidentified but later corrected to P. octomaculata by Freeman (1967); procerus is now placed in Cephise Evans, 1952* (see Austin & Mielke 2000).] Codatractus melon (Godman & Salvin, 1893) [Unsubstantiated records; see Bailowitz & Brock (1991: p. 25).] Urbanus pronta Evans, 1952* [Record refers to U. pronus; see Kendall & McGuire (1984).] Urbanus albimargo (Mabille, 1876) [Unsubstantiated records from Texas apparently refer to doryssus; see Ferris (1989c: p. 6).] Astraptes galesus (Mabille, 1888) [Unsubstantiated records; see Bailowitz & Brock (1991: pp. 30-31). The species is unknown from Mexico (Warren 2000, 2002).] Thorybes valeriana (Plotz, 1881) [Our records refer to Codatractus mysie (see Burns 1996); however the identity of valeriana remains a mystery.] Dyscophellus euribates (Stoll, 1782) [The report of this species from Texas by Aaron (1890), as Eudamus hesus (Westwood, [1852]), is unsubstantiated (see Lindsey et al. 1931); euribates remains unknown from Mexico (Warren 2000, 2002).] Cogia cajeta (Herrich-Schдffer, 1869) [One unsubstantiated record from Texas (Stanford 2002).] Pellicia costimacula Herrich-Schдffer, 1870 [This South American species was shown to be separate from arina by Steinhauser (1989).] Pellicia angra Evans, 1953* [All reports from our area (e.g. Tilden 1974) apparently refer to Pellicia arina.] Staphylus azteca (Scudder, 1872) [Single individual reported from Texas by Freeman (1977) later identified as Staphylus ceos by 46
ADW, not mazans as reported by Cassie et al. (2001).] Pyrgus adepta Plцtz, 1884 [Two highly unlikely and unverified records exist from Texas (Stanford 2002).] Heliopyrgus sublinea (Schaus, 1902) [Unsubstantiated record for this species from Texas reported by Stanford (2002); see Austin & Warren (2001) for generic combination.] Piruna cyclosticta (Dyar, 1920) [The recent report of Piruna pirus from Jeff Davis Co., Texas (Bordelon 2000), may represent this species, as listed (with a "?") by Stanford (2002). We have not yet examined these specimens.] Callimormus saturnus (Herrich-Schдffer, 1869) [A single male of this species labeled from Texas (Evans 1955) is most likely mislabeled, although the species is common in tropical northern Mexico.] Repens florus (Godman, 1900) [A single unverified record exists from south Texas (Stanford 2002),although this species is resident in Sonora and Tamaulipas, Mexico; see Vargas et al. (1996), Warren et al. (1998) and Warren (2000, 2002) for generic combination.] Remella remus (Fabricius, 1798) [A single unsubstantiated record exists from Texas (Stanford 2002).] Decinea huasteca (H.A. Freeman, 1969)* [Although cited by Ferris (1989c), no U.S. voucher is known.] Polites subreticulata (Plцtz, 1883) [Reports by Freeman (1951) of this species apparently refer to P. carus; see Burns (1994a).] Anatrytone potosiensis (H.A. Freeman, 1969)* [Although cited by Ferris (1989c), no U.S. specimens are known; see Burns (1994b) for generic combination.] Choranthus radians (Lucas, 1857) [Unsubstantiated records from Florida; see Smith et al. (1994) and Calhoun (1997).] Choranthus haitensis Skinner, 1920 [Unsubstantiated records from Florida; see Smith et al. (1994).] Choranthus vitellius (Fabricius, 1793) [Single female of this species reportedly from Florida (Evans 1955) is most likely mislabeled; see Smith et al. (1994).] Quasimellana mexicana (Bell, 1942) [Cited by Miller & Brown (1981) and Ferris (1989c), but apparently no known U.S. specimen; see Bailowitz & Brock (1991) and Burns (1994b).] Atrytonopsis ovinia (Hewitson, 1866) [Burns (1983) treats A. ovinia and A. edwardsi as separate species.] Amblyscirtes fluonia Godman, 1900 [Reports of this species from Texas (Stanford 2002) are unsubstantiated.] Panoquina fusina (Hewitson, 1868) [Erroneously reported from North America based on an assumption of conspecificity with 47
evansi by Evans (1955) and Scott (1986).] Agathymus remingtoni (D. Stallings & Turner, 1958) [Confusion with estelleae based on Scott (1986) has led to erroneous reports of this species in the U.S.] Stallingsia smithi (H. Druce, 1896) [This was erroneously reported from the U.S. based on an assumption of conspecificity with maculosus by Scott (1986).] Papilionidae Battus devilliers (Godart, [1824]) [Unsubstantiated records from Florida; see Smith et al. (1994).] Eurytides celadon (Lucas, 1852) [Considered hypothetical for Florida; see dos Passos (1961) and Smith et al. (1994).] Papilio kahli F. Chermock & R. Chermock, 1937 [Excluded as hybrid on basis of Layberry et al.(1998); also see Klassen et al. (1989).] Papilio alexiares (Hoppfer, 1865) [A possible record from south Texas was reported by Stanford (2002); although this taxon is considered conspecific with glaucus by Tyler et al. (1994) and Llorente et al. (1997). Also see Scriber et al. (1989).] Pieridae Enantia mazai Llorente, 1984 [Report of this species by Stanford (2002) from Texas apparently refers to E. albania; see Kendall (1974a), Llorente (1984) and Llorente et al. (1997).] Pontia callidice (Hьbner, [1800]) [Reports of callidice from our area refer to P. occidentalis; the two were considered conspecific by Higgins & Riley (1970) and Scott (1986). Also see Shapiro (1976).] Pontia chloridice (Hьbner, [1813]) [Erroneously reported from our area by Scott (1986), based on an unsupported assumption of conspecificity with beckerii.] Pieris napi (Linnaeus, 1758) [A Palaearctic species; see text.] Pieris brassicae (Linnaeus, 1758) [Accidental introductions, but no known persistent North American colonies (see Mello 1999 [these are sight records], Cassie et al. 2001, and Zirlin 2002).] Anthocharis dammersi J.A. Comstock, 1929 [Considered to be a hybrid on basis of Emmel & Emmel (1973: p. 24) and Shields & Mori (1979).] Phoebis intermedia (Butler, 1872) [Reports of this taxon (e.g. Scott 1986) and of P. rurina (C. Felder & R. Felder, 1861) from our area refer to P. neocypris; intermedia is a synonym of neocypris virgo (Butler, 1870) (see Lamas in press).] 48
Pyrisitia chamberlaini (Butler, 1898) [Unsubstantiated report from Florida; see Scott (1986) and Smith et al. (1994).] Lycaenidae Eumaeus minyas (Hьbner, [1809]) [Misidentification of E. toxeus; see Kendall & McGuire (1984).] Pseudolycaena marsyas (Linnaeus, 1758) [Dubiously reported from our region by Pyle (1981); Pseudolycaena damo (H. Druce, 1875) is the continental species which remains a hypothetical stray for Texas.] Cyanophrys amyntor (Cramer, 1775) [Specimen from Brewster County, Texas, reported as this species, was misidentified. Its true identity as C. herodotus confirmed by Johnson & Le Crom (1997: p. 26).] Cyanophrys longula (Hewitson, 1868) [Reported from southeastern Arizona by unsubstantiated sight records (Bailowitz & Brock 1991: p. 182), and one unverified record exists for Texas (Stanford 2002); no specimen or photographic voucher has been located.] Rekoa zebina (Hewitson, 1869) [Excluded from the North American fauna by Robbins (1991); records refer to misdetermined R. marius.] Riodinidae Euselasia abreas (W.H. Edwards, 1881) [Described from Arizona, but likely based on mislabeled material (Powell 1975); it is unknown from Mexico (de la Maza et al. 1989) or Costa Rica (DeVries 1997), and its identity was unknown to D'Abrera (1994). However, this taxon may be a senior synonym of E. sergia (Godman & Salvin, 1885); see Lamas (in press).] Calephelis nilus (C. Felder & R. Felder, 1861) [Erroneously reported from our area by Scott (1986); records refer to perditalis (see McAlpine 1971).] Calephelis sinaloensis McAlpine, 1971* [Reported occurrence in Texas by Durden (1982) requires confirmation.] Caria domitianus (Fabricius, 1793) [Casually reported from Texas by DeVries (1997), but no details on this record have been presented.] Nymphalidae Greta polissena (Hewitson, 1863) [Highly questionable records from Texas summarized by Kendall & McGuire (1984: pp. 4141); the species is unknown from Mexico (de la Maza et al. 1989).] Philaethria diatonica (Fruhstorfer, 1912) [Sight record only, from Texas (as dido); see Bordelon (1991). Philaethria diatonica is considered to be a separate species from dido (Linnaeus, 1763) by Lamas (in press).] 49
Boloria titania (Esper, [1793]) [According to Shepard (1998), this is a strictly Palaearctic species.] Chlosyne ehrenbergii (Geyer, [1833]) [Highly questionable record from Texas summarized by Kendall & McGuire (1984: p. 26).] Chlosyne marina (Geyer, 1837) [This was erroneously reported from the U.S. based on an assumption of conspecificity with melitaeoides and eumeda by Scott (1986). See text.] Chlosyne erodyle (H.W. Bates, 1864) [Casually reported from Texas by Higgins (1960), but no authentic voucher specimen is known; see Neck (1996).] Anthanassa ardys (Hewitson, 1864) [While the single record of this species from Texas may be valid (see Stanford & Opler 1993), details have not been presented in the literature and we have not examined the specimen.] Anthanassa drusilla (C. Felder & R. Felder, 1861) [A single unsubstantiated record for this species exists from south Texas (Stanford 2002).] Inachis io (Linnaeus, 1758) [Three or four almost certainly human-related, accidental introductions are known; see Hinchliff (1994: p. 176), Anonymous (1995), and Zirlin (2002).] Vanessa carye (Hьbner, [1812]) [This South American species is occasionally reported from our area instead of annabella (e.g. Scott 1986); but see Field (1971) and Shapiro & Geiger (1989).] Smyrna karwinskii Geyer, [1833] [All records from North America apparently represent misidentifications of S. blomfildia (Kendall & McGuire 1984: p. 36-37).] Mestra cana (Erichson, [1849]) [Reported as cana floridana Strecker, 1900, but it is likely that the types were mislabeled specimens; see Kimball (1965) and Masters (1970). Lamas (in press) treats cana and floridana as synonyms of dorcas hersilia (Fabricius, 1776).] Dynamine tithia (Hьbner, 1823) [Texas specimen correctly identified and illustrated by Kendall & McGuire (1984: pp. 31-32); however it is a very unlikely long-distance stray and it may have been accidentally transported. The species is unknown from Mexico (de la Maza & Turrent 1985, de la Maza et al. 1989). Specimen is in Illinois Natural History Survey.] Dynamine postverta (Cramer, 1779) [Dubiously reported from Texas by Pyle (1986) as D. mylitta (Cramer, 1779) and by Stanford (2002); no vouchers are known.] Diaethria asteria (Godman & Salvin, 1894) [Summary of history surrounding the single specimen reported from Texas provided by Kendall & McGuire (1984: p. 33). This record was seriously doubted by de la Maza & Turrent (1985: p. 32), and shown to be virtually impossible by Luis et al. (1996).] Diaethria clymena (Cramer, 1775) [Some records of D. clymena from our area, e.g. Klots (1951), apparently refer to this taxon; 50
however, clymena is unknown from Mexico or the Caribbean (Kendall & McGuire 1984, de la Maza & Turrent 1985, Smith et al. 1994), and its occurrence in our area is assumed to be accidental.] Hamadryas fornax (Hьbner, [1823]) [Records of this species from the U.S. are based on misdetermined specimens (Jenkins 1983); but see Stanford (2002).] Morpho peleides Kollar, 1850 [Texas sighting reported by Stallings & Turner (1946), Freeman (1960), and Neck (1996: p. 162.); see Penz & DeVries (2002) for systematics of the genus Morpho.] Opsiphanes boisduvallii Doubleday, [1849] [Circumstances surrounding a specimen of uncertain origin are discussed by Cassie et al. (2001).] Erebia dabanensis Ershov, 1871 [Erroneously reported from North America based on an unsupported assumption of conspecificity with youngi by Scott (1986); see Troubridge & Philip (1983), Tuzov et al. (1997) and Belik & Zamolodchikov (2002).] Erebia kozhantshikovi Sheljuzhko, 1925 [Erroneously reported from North America based on an unsupported assumption of conspecificity with lafontainei by Scott (1986); see Troubridge & Philip (1983) and Tuzov et al. (1997).] Erebia inuitica Wyatt, 1966* [Known from only a single specimen whose authenticity is dubious; see Warren (1968) and Ferris (1989c: p. 59).] \1 Adequate documentation should consist of a specimen or photograph (when genitalic dissection is not necessary), identified by a specialist in the appropriate taxonomic group. Details should be reported in the literature (website or e-mail postings are not adequate), along with complete data, when possible. Preferably, it should be deposited in a major entomological institution [National Museum of Natural History, Washington, D.C.; American Museum of Natural History, New York; Natural History Museum of Los Angeles County; California Academy of Sciences, San Francisco; Allyn Museum of Entomology, Sarasota; Carnegie Museum of Natural History, Pittsburgh; Canadian National Collection, Ottawa, Ontario] or university. To our knowledge no major collections have established photographic documentation files, but they should. Sight records do not constitute adequate documentation for national records. 51
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Acknowledgments We thank the many people who have contributed information on specific records, scientific names, publications, species concepts, and who have provided us with important literature. Notable among them are A. Atkins, G.T. Austin, R. Bailowitz, C.W. Bordelon, Jr., J.P. Brock, E. Brockmann, A.V.Z. Brower, F.M. Brown, J.M. Burns, S.J. Cary, H. Chiba, R. de Jong, A.L. Devyatkin, R. Dirig, U. Eitschberger, J.F. Emmel, T.C. Emmel, C.D. Ferris, H.A. Freeman, A.V.L. Freitas, J. Glassberg, P. Hall, P.C. Hammond, R. Hirzel, R. Holland, R. Huber, K. Johnson, A.Y. Kawahara, R.O. Kendall, E. Knudson, N.G. Kondla, J.D. Lafontaine, I. Leeuw, A. Luis-Martнnez, J.E. Llorente-Bousquets, C.D. MacNeill, B. Mather, D.V. McCorkle, O.H.H. Mielke, J.S. Miller, J.Y. Miller, L.D. Miller, D.D. Mullins, C.C. Nice, B.A. O'Hara, T.W. Ortenburger, R.S. Peigler, J.A. Powell, G.F. Pratt, F.W & J.D. Preston, R.M. Pyle, M. Quinn, R.K. Robbins, K. Roever, R.L. Romeyn, D.F. Schweitzer, J.A. Scott, A.M. Shapiro, J.H. Shepard, J.A. Shuey, J. Slotten, F.A.H. Sperling, R.E. Stanford, S.R. Steinhauser, S.A. Upson, I. Vargas-Fernбndez, E.G. Voss, N. Wahlberg, D.J. & S.J. Warren, Q.D. Wheeler, D.M. Wright, and especially G. Lamas and J.P. Pelham who assisted in many ways. We are appreciative of their input, but they do not bear responsibility for the final result. 83

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