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Home , Achalarus lyciades, Aglais milberti, Amblyscirtes hegon, Amblyscirtes vialis, Anaea andria, Ancyloxypha numitor

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University Microfilms International 300 N. Zeeb Road Ann Arbor, Ml 48106

8300197

Albrecht, Carl William, Jr.

THE , GEOGRAPHY, AND SEASONAL DISTRIBUTION OF RHOPALOCERA IN

The Ohio State University PH.D. 1982

University Microfilms International300 N. Zeeb Road, Ann Arbor, MI 48106

Copyright 1982

by Albrecht, Carl William, Jr. All Rights Reserved

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University Microfilms International

THE TAXONOMY, GEOGRAPHY, AND SEASONAL

DISTRIBUTION OF RHOPALOCERA IN OHIO

DISSERTATION

Presented in Partial Fulfillment of the Requirements for

the Degree Doctor of Philosophy in the Graduate

School of The Ohio State University

by

Carl William Albrecht, Jr., B.A., M.Sc.

sfr & Jc tfe *

The Ohio State University

1982

Reading Committee: Approved By

Charles A. Triplehorn, Professor

D. Lyle Goleman, Professor

David J. Horn, Professor

Department of Entomology ACKNOWLEDGMENTS

Numerous persons have helped me in acquiring the data on which

this study is based. It is my pleasure to acknowledge this

assistance, and to extend my sincere thanks to all of them. For

providing access to collections and making working space available,

I am indebted to the following curators and their institutions:

Frederick H. Rindge, American Museum of Natural History; the late

Rendell Rhoades, Ashland College; the late Harry K. Clench, Carnegie

Museum of Natural History; Richard A. Davis and Charles Oehler,

Cincinnati Museum of Natural History; Patricia Hellwig, Cleveland

Museum of Natural History; Robert^W. Alrutz, Denison University;

Arnnold W. Fritz, Malone College; Thomas E. Moore, Museum of Zoology,

University of ; Roland L. Fisher, Michigan State University

Insect Collection; William D. Field, National Museum of Natural

History; Roy W. Rings, Ohio Agriculture Research and Development

Center; William T. Schultz, Department of Natural History, The Ohio

Historical Center; Charles A. Triplehorn, The Ohio State University

Insect Collection; the late Edward S. Thomas, Museum of Zoology, The

Ohio State University; Glenn B. Wiggins, Royal Ontario Museum; Louis

J. Laux, Jr., Wittenberg University. Others who have been helpful in

providing records from their personal collections include: James W.

Amrine, Jr., Columbus, Ohio; Roger S. Boone, Springfield, Ohio; John

Calhoun, Westerville, Ohio; David C. Iftner, Worthington, Ohio; Joe and Calhoun, Westerville, Ohio; David C. Iftner, Worthington, Ohio; Joe

and Ella McMahon, Chillicothe, Ohio; David K. Parshall, Columbus,

Ohio.

My thanks are extended also to those who have been helpful and

welcome companions on many field collection trips, including:

Susan K. Ahearn, Stuart L. Alcott, Christine J. Chandler, Donald

Chandler, Glen Firebaugh, John Jenkinson, Eric H. Metzler, Frank

Moore, Beverley H. Moseley, David K. Parshall, Melanie Pratt, Robert

A. Restifo, Ann Skrei, John S. Thomas, Tom Wood.

This project has benefited from the many discussions I have had

with several zoologists, including John M. Condit, Roy W. Rings,

William T. Schultz, and David H. Stansbery. My adviser, Charles A.

Triplehorn, has been supportive throughout with his encouragement,

patience, and good humor. His suggestions and criticism of this manuscript are most appreciated, as are those of D. Lyle Goleman and

David J. Horn. All errors are, of course, my responsibility.

To my parents, Carl W. Albrecht, Sr., and Mabel M. Albrecht, I

owe special thanks for their support and encouragement throughout the

long period of my formal education. My wife, Joann, through her

infinite patience and tact, has given me an essential environment in which the completion of this project was possible.

Finally, the staff of The Ohio State University Instruction and

Research Computer Center and the University Libraries provided invaluable support in making their facilities available, and in giving assistance whenever required. Gratitude also is extended to the library staff of The Ohio Historical Center, in particular to

Mary Ellen West and Conrad Weitzek, both of whom were very helpful with interlibrary loans and the use of their facilities. To Charles

C. King, and the staff of the Ohio Biological Survey, my thanks are also extended. VITA

8 October 1932. . . . Born - Columbus, Ohio

1950-1952 ...... Ohio Academy of Science Scholar, Miami University, Oxford, Ohio

1954 . B .A. , Miami University, Oxford, Ohio

1954-1955 ...... University Scholar, Graduate School, The Ohio State University, Columbus, Ohio

1955-1958 ...... United State Army

1960-1963, 1965-1967. Teaching Assistant and Instructor, Departments of Zoology and Biology, The Ohio State Universityt Columbus, Ohio

1962 . M.Sc., The Ohio State University, Columbus, Ohio

1970-1978 ...... Curator, Natural History Department, The Ohio Historical Center, Columbus, Ohio

1975-1982 ...... Manager, Data Processing Department, The Ohio Historical Center, Columbus, Ohio

PUBLICATIONS

"The of Cedar Bog., Champaign Co., Ohio. I. An Annotated Check List of the Rhopalocera." Ohio J. Sci. 74(2):126-132, 1974.

"A Flexible Computer Program for the Production of Insect Labels." Great Lakes Entomol. 7(l):27-29, 1974, with Russell V. Skavaril.

FIELDS OF STUDY

Major Field: Entomology

Studies in Systematics. Professor Charles A. Triplehom

v TABLE OF CONTENTS

Page

ACKNOWLEDGMENTS...... ii

VITA...... v

LIST OF TABLES...... vii

LIST OF FIGURES...... viii

INTRODUCTION...... 1

METHODS...... 7

THE TAXA WITH KEYS FOR DETERMINATION...... 13

HESPERIIDAE...... 21 PAPILIONIDAE...... 150 ...... 167 ...... 204 ...... 263 LIBYTHEIDAE...... 268 HELICONIDAE...... 270 ...... 271 SATYRIDAE...... 327 DANAIDAE...... 345

DISCUSSION...... 349

SUMMARY...... 368

LIST OF REFERENCES...... 370

APPENDIX 378 LIST OF TABLES

Table Page

1. of Rhopalocera known from Ohio and neighboring states...... 14

2. Taxonomic composition of the rhopalocerous fauna of Ohio....349

3. Taxonomic composition of the rhopalocerous fauna reported from the area surrounding Ohio, but not from Ohio itself, with the number of species reported from each state/province...... 350

4. Summary of the Rhopalocera of Ohio by status...... 351

5. The and skippers of Ohio with notation relative to the position of Ohio within the overall range of the species...... 353

6. Resident Rhopalocera in Ohio which are more are less restricted to the Southern District of the state...... 366

7. Resident Rhopalocera in Ohio which are more or less restricted to the Northern District of the state...... 366

8. Species of Rhopalocera, not known from Ohio, but reported from neighboring states...... 508

9. The butterflies and skippers of Ohio with notation relative to the status of each species in the state...... 509

vii LIST OF FIGURES

Figure Page

1. Parts of Ohio included in regional or county reports of Rhopalocera...... 378

2. Wing vein terminology used in this work...... 379

3. Ohio's counties and their names...... 380

4. Geographical distribution of in Ohio 381

5. Seasonal distribution of adult Epargyreus clarus in Ohio....381

6. Geographical distribution of Autochton cellus in Ohio...... 382

7. Seasonal distribution of adult Autochton cellus in Ohio.....382

8. Geographical distribution of lyciades in Ohio 383

9. Seasonal distribution of adult Achalarus lyciades in Ohio...383

10. Geographical distribution of Thorybes bathyllus in Ohio 384

11. Seasonal distribution of adult Thorybes bathyllus in Ohio...384

12. Geographical distribution of Thorybes pylades in Ohio...... 385

13. Seasonal distribution of adult Thorybes pylades in Ohio 385

14. Geographical distribution of Staphylus hayhurstii in Ohio...386

15. Seasonal distribution of adult Staphylus hayhurstii in Ohio.386

16. Geographical distribution of Erynnis icelus in Ohio...... 387

17. Seasonal distribution of adult Erynnis icelus in Ohio...... 387

18. Geographical distribution of Erynnis brizo in Ohio...... 388

19. Seasonal distribution of adult Erynnis brizo in Ohio... 388 LIST OF FIGURES (continued)

Figure Page

20. Geographical distribution of Erynnis .juvenalis in Ohio 389

21. Seasonal distribution of adult Erynnis juvenalis in Ohio....389

22. Geographical distribution of Erynnis horatius in Ohio...... 390

23. Seasonal distribution of adult Erynnis horatius in Ohio 390

24. Geographical distribution of Erynnis martialis in Ohio 391

25. Seasonal distribution of adult Erynnis martialis in Ohio....391

26. Geographical distribution of in Ohio...... 392

27. Seasonal distribution of adult Erynnis lucilius in Ohio 392

28. Geographical distribution of Erynnis baptisiae in Ohio...... 393

29. Seasonal distribution of adult Erynnis baptisiae in Ohio....393

30. Geographical distribution of in Ohio...... 394

31. Seasonal distribution of adult Erynnis persius in Ohio 394

32. Geographical distribution of centaureae in Ohio .395

33. Seasonal distribution of adult Pyrgus centaureae in Ohio....395

34. Geographical distribution of Pyrgus communis in Ohio...... 396

35. Seasonal distribution of adult Pyrgus communis in Ohio...... 396

36. Geographical distribution of in Ohio 397

37. Seasonal distribution of adult Pholisora catullua in Ohio...397

38. Geographical distribution of numitor in Ohio....398

39. Seasonal distribution of adult in Ohio..398

40. Geographical distribution of Thymelicus lineola in Ohio 399

41. Seasonal distribution of adult Thymelicus lineola in Ohio...399

ix LIST OF FIGURES (continued)

Figure Page

42. Geographical distribution of Nastra lherminier in Ohio 400

43. Seasonal distribution of adult Nastra lherminier in Ohio....400

44. Geographical distribution of Hylephila phyleus in Ohio 401

45. Seasonal distribution of adult Hylephila phyleus in Ohio....401

46. Geographical distribution of leonardus in Ohio 402

47. Seasonal distribution of adult Hesperia leonardus in Ohio...402

48. Geographical distribution of Hesperia metea in Ohio...... 403

49. Seasonal distribution of adult Hesperia metea in Ohio...... 403

50. Geographical distribution of Hesperia sassacus in Ohio 404

51. Seasonal distribution of adult Hesperia sassacus in Ohio....404

52. Geographical distribution of coras in Ohio.... 405

53. Seasonal distribution of adult Polites coras in Ohio...... 405

54. Geographical distribution of Polites themistocles in Ohio...406

55. Seasonal distribution of adult Polites themistocles in Ohio.406

56. Geographical distribution of Polites origenes in Ohio...... 407

57. Seasonal distribution of adult Polites origenes in Ohio 407

58. Geographical distribution of Polites mystic in Ohio...... 408

59. Seasonal distribution of adult Polites mystic in Ohio...... 408

60. Geographical distribution of Polites vibex in Ohio...... 409

61. Seasonal distribution of adult Polites vibex in Ohio... 409

62. Geographical distribution of Wallengrenia egeremet in Ohio..410

63. Seasonal distribution of adult Wallengrenia egeremet in Ohio...... 410

x LIST OF FIGURES (continued)

Figure Page

64. Geographical distribution of Pompeius vema in Ohio...... 411

65. Seasonal distribution of adult Pompeius vema in Ohio...... 411

66. Geographical distribution of in Ohio..412

67. Seasonal distribution of adult Atalopedes campestris in Ohio...... 412

68. Geographical distribution of Atrytone logan in Ohio...... 413

69. Seasonal distribution of adult Atrytone logan in Ohio...... 413

70. Geographical distribution of massasoit in Ohio...... 414

71. Seasonal distribution of adult in Ohio.....414

72. Geographical distribution of Poanes hobomok in Ohio...... 415

73. Seasonal distribution of adult Poanes hobomok in Ohio...... 415

74. Geographical distribution of Poanes zabulon in Ohio...... 416

75. Seasonal distribution of adult Poanes zabulon in Ohio...... 416

76. Geographical distribution of in Ohio...... 417

77. Seasonal distribution of adult Poanes viator in Ohio...... 417

78. Geographical distribution of dion in Ohio...... 418

79. Seasonal distribution of adult Euphyes dion in Ohio...... 418

80. Geographical distribution of in Ohio...... 419

81. Seasonal distribution of adult Euphyes dukesi in Ohio...... 419

82. Geographical distribution of Euphyes conspicua in Ohio..... 420

83. Seasonal distribution of adult Euphyes conspicua in Ohio....420

84. Geographical distribution of Euphyes bimacula in Ohio...... 421

85. Seasonal distribution of adult Euphyes bimacula in Ohio.... 421 LIST OF FIGURES (continued)

Figure Page

86. Geographical distribution of Euphyes ruricola in Ohio...... 422

87. Seasonal distribution of adult Euphyes ruricola in Ohio 422

88. Geographical distribution of Atrytonopsis hianna in Ohio....423

89. Seasonal distribution of adult Atrytonopsis hianna in Ohio..423

90. Geographical distribution of hegon in Ohio.....424

91. Seasonal distribution of adult Amblyscirtes hegon in Ohio...424

92. Geographical distribution of Amblyscirtes vialis in Ohio....425

93. Seasonal distribution of adult Amblyscirtes vialis in Ohio..425

94. Geographical distribution of ocola in Ohio...... 426

95. Seasonal distribution of adult Panoquina ocola in Ohio 426

96. Geographical distribution of Battus philenor in Ohio...... 427

97. Seasonal distribution of adult Battus philenor in Ohio 427

98. Geographical distribution of Eurytides marcellus in Ohio....428

99. Seasonal distribution of adult Eurytides marcellus in Ohio..428

100. Geographical distribution of polyxenes asterius in Ohio...... 429

101. Seasonal distribution of adult asterius in Ohio...... 429

102. Geographical distribution of Papilio cresphontes in Ohio....430

103. Seasonal distribution of adult Papilio cresphontes in Ohio..430

104. Geographical distribution of Papilio in Ohio...... 431

105. Seasonal distribution of adult in Ohio 431

106. Geographical distribution of Papilio troilus in Ohio...... 432

107. Seasonal distribution of adult Papilio troilus in Ohio 432

xii LIST OF FIGURES (continued)

Figure Page

108. Geographical distribution of Pontia protodice in Ohio...... 433

109. Seasonal distribution of adult Pontia protodice in Ohio 433

110. Geographical distribution of Artogeia virginiensis in Ohio...434

111. Seasonal distribution of adult Artogeia virginiensis in Ohio.434

112. Geographical distribution of Artogeia rapae in Ohio...... 435

113. Seasonal distribution of adult Artogeia rapae in Ohio...... 435

114. Geographical distribution of Falcapica midea in Ohio...... 436

115. Seasonal distribution of adult Falcapica midea in Ohio...... 436

116. Geographical distribution of philodice in Ohio ..437

117. Seasonal distribution of adult in Ohio...... 437

118. Geographical distribution of in Ohio ..438

119. Seasonal distribution of adult Colias eurytheme in Ohio. 438

120. Geographical distribution of cesonia in Ohio ....439

121. Seasonal distribution of adult in Ohio... 439

122. Geographical distribution of sennae eubule in Ohio...440

123. Seasonal distribution of adult eubule in Ohio.440

124. Geographical distribution of Phoebis philea in Ohio .441

125. Seasonal distribution of adult Phoebis philea in Ohio...... 441

126. Geographical distribution of lisa in Ohio...... 442

127. Seasonal distribution of adult in Ohio...... 442

128. Geographical distribution of Eurema nicippe in Ohio ....443

129. Seasonal distribution of adult Eurema nicippe in Ohio...... 443 LIST OF FIGURES (continued)

Figure Page

130. Geographical distribution of iole in Ohio...... 444

131. Seasonal distribution of adult in Ohio...... 444

132. Geographical distribution of Feniseca tarquinius in Ohio 445

133. Seasonal distribution of adult Feniseca tarquinius in Ohio....445

134. Geographical distribution of phlaeas in Ohio...... 446

135. Seasonal distribution of adult Lycaena phlaeas in Ohio...... 446

136. Geographical distribution of Hyllolycaena hyllus in Ohio...... 447

137. Seasonal distribution of adult Hyllolycaena hyllus in Ohio....447

138. Geographical distribution of Epidemla helloides in Ohio...... 448

139. Seasonal distribution of adult Epidemia helloides in Ohio.....448

140. Geographical distribution of Harkenclenus titus in Ohio...... 449

141. Seasonal distribution of adult Harkenclenus titus in Ohio...,.449

142. Geographical distribution of acadica in Ohio...... 450

143. Seasonal distribution of adult Satyrium acadica in Ohio...... 450

144. Geographical distribution of Satyrium edwardsii in Ohio ..451

145. Seasonal distribution of adult Satyrium edwardsii in Ohio 451

146. Geographical distribution of Satyrium calanus falacer in Ohio.452

147. Seasonal distribution of adult Satyrium calanus falacer in Ohio...... 452

148. Geographical distribution of Satyrium caryaevorum in Ohio 453

149. Seasonal distribution of adult Satyrium caryaevorum in Ohio...453

150. Geographical distribution of Satyrium liparops in Ohio...... 454

151. Seasonal distribution of adult Satyrium liparops in Ohio 454

xiv LIST OF FIGURES (continued)

Figure Page

152. Geographical distribution of Calycopis cecrops in Ohio...... 455

153. Seasonal distribution of adult Calycopis cecrops in Ohio...... 455

154. Geographical distribution of Mitoura gryneus in Ohio...... 456

155. Seasonal distribution of adult Mitoura gryneus in Ohio...... 456

156. Geographical distribution of augustus in Ohio...... 457

157. Seasonal distribution of adult Incisalia augustus in Ohio...... 457

158. Geographical distribution of Incisalia irus in Ohio ...... 458

159. Seasonal distribution of adult Incisalia irus in Ohio...... 458

160. Geographical distribution of Incisalia henrici in Ohio...... 459

161. Seasonal distribution of adult Incisalia henrici in Ohio...... 459

162. Geographical distribution of Incisalia niphon in Ohio...... 460

163. Seasonal distribution of adult Incisalia niphon in Ohio...... 460

164. Geographical distribution of Euristrymon Ontario in Ohio...... 461

165. Seasonal distribution of adult Euristrymon Ontario in Ohio.... 461

166. Geographical distribution of Parrhasius m-album in Ohio...... 462

167. Seasonal distribution of adult Parrhasius m-album in Ohio...... 462

168. Geographical distribution of melinus in Ohio...... 463

169. Seasonal distribution of adult Strymon melinus in Ohio...... 463

170. Geographical distribution of Erora laeta in Ohio...... 464

171. Seasonal distribution of adult Erora laeta in Ohio ...... 464

172. Geographical distribution of isola in Ohio...... 465

173. Seasonal distribution of adult Hemiargus isola in Ohio...... 465

xv LIST OF FIGURES (continued)

Figure Page

174. Geographical distribution of Everes comyntas in Ohio...... 466

175. Seasonal distribution of adult Everes comyntas in Ohio...... 466

176. Geographical distribution of ladon in Ohio...... 467

177. Seasonal distribution of adult in Ohio...... 467

178. Geographical distribution of Celastrina ebenina in Ohio ....468

179. Seasonal distribution of adult Celastrina ebenina in Ohio...... 468

180. Geographical distribution of lygdamus in Ohio...... 469

181. Seasonal distribution of adult Glaucopsyche lygdamus in Ohio....469

182. Geographical distribution of Lycaeides melissa in Ohio...... 470

183. Seasonal distribution of adult Lycaeides melissa in Ohio...... 470

184. Geographical distribution of borealis in Ohio...... 471

185. Seasonal distribution of adult Calephelis borealis in Ohio...... 471

186. Geographical distribution of Calephelis muticum in Ohio...... 472

187. Seasonal distribution of adult Calephelis muticum in Ohio...... 472

188. Geographical distribution of Libytheana bachmanii in Ohio...... 473

189. Seasonal distribution of adult Libytheana bachmanii in Ohio.....473

190. Geographical distribution of Euptoieta claudia in Ohio...... 474

191. Seasonal distribution of adult Euptoieta claudia in Ohio...... 474

192. Geographical distribution of Speyeria cybele in Ohio...... 475

193. Seasonal distribution of adult Speyeria cybele in Ohio...... 475

194. Geographical distribution of Speyeria in Ohio...... 476

195. Seasonal distribution of adult Speyeria aphrodite in Ohio...... 476

xv i LIST OF FIGURES (continued)

Figure Page

196. Geographical distribution of Speyeria idalia in Ohio...... 477

197. Seasonal distribution of adult Speyeria idalia in Ohio...... 477

198. Geographical distribution of Speyeria in Ohio...... 478

199. Seasonal distribution of adult Speyeria atlantis in Ohio...... 478

200. Geographical distribution of Clossiana myrina in Ohio....479

201. Seasonal distribution of adult Clossiana selene myrina in Ohio..479

202. Geographical distribution of Clossiana bellona in Ohio...... 480

203. Seasonal distribution of adult Clossiana bellona in Ohio...... 480

204. Geographical distribution of Charidryas nycteis in Ohio...... 481

205. Seasonal distribution of adult Charidryas nycteis in Ohio...... 481

206. Geographical distribution of Charidryas harrisii in Ohio...... 482

207. Seasonal distribution of adult Charidryas harrisii in Ohio...... 482

208. Geographical distribution of tharos in Ohio...... 483

209. Seasonal distribution of adult Phyciodes tharos in Ohio...... 483

210. Geographical distribution of phaeton in Ohio...... 484

211. Seasonal distribution of adult Euphydryas phaeton in Ohio...... 484

212. Geographical distribution of interrogationis in Ohio..485

213. Seasonal distribution of adult Polygonia interrogationis in Ohio...... 485

214. Geographical distribution of in Ohio...... 486

215. Seasonal distribution of adult Polygonia comma in Ohio...... 486

216. Geographical distribution of in Ohio...... 487

217. Seasonal distribution of adult Polygonia progne in Ohio...... 487 LIST OF FIGURES (continued)

Figure Page

218. Geographical distribution of Nymphalis vau-album in Ohio...... 488

219. Seasonal distribution of adult Nymphalis vau-album in Ohio...... 488

220. Geographical distribution of Nymphalis antiopa in Ohio...... 489

221. Seasonal distribution of adult Nymphalis antiopa in Ohio ....489

222. Geographical distribution of milberti in Ohio...... 490

223. Seasonal distribution of adult Aglais milberti in Ohio...... 490

224. Geographical distribution of Vanessa virginiensis in Ohio...... 491

225. Seasonal distribution of adult Vanessa virginiensis in Ohio...... 491

226. Geographical distribution of Vanessa cardui in Ohio...... 492

227. Seasonal distribution of adult Vanessa cardui in Ohio...... 492

228. Geographical distribution of Vanessa in Ohio...... 493

229. Seasonal distribution of adult Vanessa atalanta in Ohio...... 493

230. Geographical distribution of Junonia coenia in Ohio...... 494

231. Seasonal distribution of adult Junonia coenia in Ohio...... 494

232. Geographical distribution of Basilarchia arthemis astyanax in Ohio...... 495

233. Seasonal distribution of adult Basilarchia arthemis astyanax in Ohio...... 495

234. Geographical distribution of Basilarchia archippus in Ohio...... 496

235. Seasonal distribution of adult Basilarchia archippus in Ohio...... 496

236. Geographical distribution of Anaea andria in Ohio...... 497

237. Seasonal distribution of adult Anaea andria in Ohio...... 497

238. Geographical distribution of in Ohio...... 498

239. Seasonal distribution of adult Asterocampa celtis in Ohio...... 498

xviii LIST OF FIGURES (continued)

Figure Page

240. Geographical distribution of in Ohio .....499

241. Seasonal distribution of adult Asterocampa clyton in Ohio...... 499

242. Geographical distribution of anthedon in Ohio...... 500

243. Seasonal distribution of adult in Ohio...... 500

244. Geographical distribution of + appalachia in Ohio...... 501

245. Seasonal distribution of adult Satyrodes eurydice + appalachia in Ohio...... 501

246. Geographical distribution of gemma in Ohio...... 502

247. Seasonal distribution of adult Cyllopsis gemma in Ohio...... 502

248. Geographical distribution of Megisto cymela in Ohio...... 503

249. Seasonal distribution of adult Megisto cymela in Ohio...... 503

250. Geographical distribution of Cercyonis pegala in Ohio...... 504

251. Seasonal distribution of adult Cercyonis pegala in Ohio...... 504

252. Geographical distribution of Danaus plexippus in Ohio...... 505

253. Seasonal distribution of adult Danaus plexippus in Ohio...... 505

254. Ohio's counties, with number of Rhopalocera species known from each of three classes of counties shown as a percentage of the total number of species known from the entire state...... 506

255. The state of Ohio showing tentative division into two districts based upon distribution of Rhopalocera species within the state: the Northern and Southern Districts...... 507

xvix INTRODUCTION

The present study of the Rhopalocera of Ohio includes all species

of butterflies and skippers known from the state since the first

records were published by Kirtland in 1851. The purpose of this study

is to record, in an acceptable taxonomic system, the taxa known from

the state and surrounding areas, along with the geographical and

seasonal distribution of these species within Ohio. The specific

objectives are derived from, and related to, the three parts of the

purpose, namely the taxonomic, geographical, and seasonal distribution

systems. The taxa of butterflies and skippers in Ohio and surrounding

areas are reported in an expanded and annotated catalogue with keys

for their determination. Geographical and seasonal distribution data

are discussed for each species to the extent made possible by the

available data.

The significance of this research project is twofold. This is

the first study to include the Rhopalocera of the entire state of Ohio

since Hine's list was published in 1898. In addition, this is the

first systematic reporting of distributional data of Ohio Rhopalocera

in terms of space and time.

The present study reports 50% more species from Ohio than did

Hine's first list (Hine, 1898a) and includes additional species from

surrounding areas which may eventually be collected in Ohio. Hine expanded his list in 1899, and there have been additions since then, on a regional or county basis. Price (1970) studied the northwestern corner of the state and included 13 counties in his paper. Janus

(1973) prepared a report on northeastern Ohio including seven counties.

In addition, individual reports have been written on the following counties: Pickaway (Bales, 1909), Summit (Claypole, 1897), Hamilton

(Dury, 1878; Wyss, 1930), Seneca (Henninger, 1910; Porter, 1965),

Cuyahoga (Kirkpatrick, 1864; Kirtland, 1851), and Montgomery (Pilate,

1879 and 1882) . Figure 1 shows the Ohio counties covered by the reports cited above. Other reports have included relatively small numbers of species, and have been limited in geographic area. Single

Ohio records of Rhopalocera appear in many of the Season Summaries published by The Lepidopterists' Society in their Newsletter.

This research project was begun in the late summer of 1969 as a limited survey of selected localities within the state, and was expanded to include the entire state in 1973. Three primary areas of research have been investigated during this study: (1) existing literature has been examined pertaining to the distribution of butterflies and skippers in Ohio and surrounding areas; (.2) collections of butterflies, both public and private., have been examined; (3) field collections of specimens have been made in portions of the state. Ohio was divided into five arbitrary divisions in order to distribute collecting efforts among the various parts of the state. These divisions and their use are discussed more thoroughly under Methods below. The information provided in regional faunal studies is useful

in many ways: in larger faunal surveys, taxonomic revisions, bio-

geographical studies, and other projects. This study includes the

basic information expected in such a survey along with keys for

determination of the taxa.

The system of taxa used for the butterflies and skippers included

in this study has its foundation in the tenth edition of Linnaeus'

Systema Naturae (1758), in which all species of butterflies and

skippers then known were placed in a single , Papilio, within the

order Lepidoptera. Expansions of Linnaeus' system of taxa for these

have been made by many authors during the past 220 years with

significant early contributions being made by Fabricius, Hllbner,

Latreille, Swainson and others. Linnaeus did not use family-group names

for any of the taxa in his publications. The categories within the

family-group, falling between the order-group and genus-group, were

introduced later and have subsequently been used extensively, especially

in the insects (Blackwelder, 1967). The first family-group taxa of

Rhopalocera to be named were the HESPERIIDAE and PAPILIONIDAE, families

introduced by Latreille in 1809. Swainson (1827) was the first author

to name subfamilies of butterflies. Other categories within the

family-group, including superfamilies and tribes, have been used since

then.

A universally acceptable system of classifying the Lepidoptera is

not presently available (Ehrlich, 1958; Brock, 1971; Common, 1975).

A much better understanding of relationships within this group is needed before inferences can be made which will lead to a better system; The names for the taxa chosen for this work are all available in the

literature, although they have not all been used together in a single publication exactly as they are presented here. The limits of the taxa

themselves vary in some cases from those of the most recent literature.

In such cases justifications are cited.

The butterflies and skippers are placed in the suborder Ditrysia, a taxon embracing all heteroneurous Lepidoptera with two genital openings in the female. Although there is some difference of opinion as to where this taxon should be placed within the hierarchy of categories, there is agreement among at least some authors today (Brock, 1971; Common,

1975; Clench, 1975) that it should exist. The name Rhopalocera, used by many to represent a suborder (as recently as 1975 by Higgins) is herein used as a division between suborder and superfamilies. • It includes those heteroneurous Lepidoptera in which the apex of the antenna is more or less expanded into a terminal club, and the fore- and hind-wings overlap without a frenulum in most cases; these are the

Lepidoptera covered by this study within the specified area of the study.

In the family-group taxa, two superfamilies are used. This agrees with most modern authors, although some (e.j;. Higgins, 1975 and Ferris

& Brown, 1981) use four superfamilies. Within the two superfamilies

(HESPERIOIDEA and ) 11 families are recognized in North

America, following many modern authors (Klots, 1951; dos Passos, 1964;

Howe, 1975; Miller & Brown, 1981). Subfamilies and tribes used in this work follow Miller & Brown (1981). 5

The genera used in this work are those generally used in the current

literature. Supergenera and subgenera are not used in this work.

Subspecific names are used only in those cases where geographical barriers clearly separate a group of populations, including those in the Ohio area, from other parts of the range of a particular species.

This satisfies the definitions of Mayr (1969) and Blackwelder (1967) which require that a subspecies must occupy a distinct geographical area. Also, following the same definitions, the recognized subspecies must differ taxonomically from other populations of the same species.

Keys have been prepared for determination of all taxa included in this study. The geographical limits for the keys have been set beyond the artificial limits of Ohio, to the greatest extent that is possible. This has been done in order to give them maximum utility without making them too cumbersome to use. In general, all keys relating to family-group taxa, except where otherwise indicated, include all of America north of . Most keys to the genus-group taxa include all states east of the Mississippi River (except ) as well as those portions of directly north of this area. The species-group taxa included in keys are those expected to be found in

Ohio, , eastern Kentucky, western West , western

Pennsylvania, southern Ontario, and southern Michigan.

Comments on voltinism, as well as adult flight-times are included in the individual species accounts. Some remarks concerning both voltinism and life span are given here. The contemporary set of adults from a single generation usually has a flight period of from 30 to 60 6

days (Clench, 1975:28; Scott, 1981:30). Individuals from a single

generation may live from three or four days up to eight months to a

year. In the case of temperate zone species which have diapause in

some stage other than the adult, the average life span of the adult

has been determined by one worker to be eight days (nine days for

females, seven for males) (Scott, 1974:225; Scott, 1981:30). In

contrast, overwintering adults may live up to a year.

Multivoltine species usually exhibit total adult flight periods of

more than 60 days within a year, with a decrease or absence of adult

activity between two periods of increased activity. In some cases,

however, there is considerable overlap of the generations thereby more or less obliterating the intergenerational gaps, such that only the

total length of the flight period will give any indication of more than

a single generation. In some cases, the last generation of the season may show a much lower level of adult activity than the first. This is

often referred to as a partial generation, and usually indicates that

part of the penultimate generation goes into diapause without further

reproduction, and gives the adults of the first generation of the

following year. The remainder of the last generation develops into

adults which reproduce, giving another generation which overwinters.

Bivoltinism, often occuring in Ohio butterflies and skippers, is probably always marked by an increased period of adult activity, usually in excess of 60 days. Bivoltinism in these species also may be indicated by a partial or complete break between two periods of

relatively high activity, or between an early period of high and a later period of low activity. METHODS

The species included in this study are those known to exist in

the state of Ohio through specimens and/or literature records. Also

included are species known to exist in neighboring states which might

eventually be found in Ohio.

Collecting sites used during this study were selected on the basis

of an arbitrary division of Ohio into five divisions. Existing systems

of regions within the state (such as physiographic, soil , bedrock,

and Pleistocene glaciation regions) were rejected for this since no

correlations are known to exist between these systems and the distri­

bution of Rhopalocera in Ohio. These existing regional systems are

described by several authors in Lafferty (1979). The five divisions

which were used in this work are as follows: 1) CENTRAL - Franklin

County and its six contiguous counties, representing both the Glaciated

Appalachian Plateau Region and the Till Plains Region; 2) NORTHEASTERN -

made up of those counties that lie mostly to the north and east of 40°

N. Lat. and 83° W. Long., with parts of the Glaciated and Unglaciated

Appalachian Plateau, Lake Plains, and Glaciated Till Plains Regions;

3) SOUTHEASTERN - including those counties south and east of the

coordinates given above, and almost entirely in the Unglaciated

Appalachian Plateau Region; 4) NORTHWESTERN - to the north and west of

7 the coordinates above, including parts of the Till Plains and Lake Plains

Regions; 5) SOUTHWESTERN - to the south and west of the coordinates above, including parts of the Till Plains and Glaciated Appalachian

Plateau, as well as the Unglaciated Appalachian Plateau, and the unglaciated Bluegrass Region. Collecting sites were selected from all of these divisions such that all of the physiographic regions were represented, and that other collection data were augmented. These divisions are shown in Figure 3.

The discussion for each species given below includes the valid name, diagnosis (if not given in the key for that group), distribution

(both general and specific within Ohio), and some comments on the biology of the species. These discussions include information and data on nomenclature, synonymy, common names, foodplants, habitat preference, and geographical and seasonal distribution.

Available and valid names, as well as unavailable names and syno­ nyms, are included under the nomenclatorial and systematic status of each species. Original reference citations for these names will be found in Brock (1971), Brown & Heineman (1972), Higgins (1975), and

Miller & Brown (1981).

Common names are listed for each species. A commonly known reference is cited with each name, without regard for the first use of the name. The "Law of Priority", as a guiding principle, has no more place in the usage of common names than it does in the use of

the language which contains them. 9

The wing surface of insects is often divided into areas or regions

in order to facilitate the study of certain characters, or to add

precision to the use of these characters in the diagnosis of taxonomic

units. The wing margins and angles are also named by authors. The

terminology used to describe wing areas, margins, and angles in this

work follows Klots (1951) and Ehrlich & Ehrlich (1964).

The terminology used by various authors in describing the wing

veins and cells of butterflies and skippers has been reviewed by

Miller (1970). The wing venation nomenclature preferred by Miller is

essentially the same as that used by Borror, Delong and Triplehorn

(1976:470) and Clench (1975:14), based on the Comstock-Needham system

of insect wing venation. The cell terminology differs, however,

in that the cells other than the costal, discal, and anal cells are denoted by the veins both anterior and posterior to the cell. This practice was suggested by Miller (op.cit.:48) to reduce confusion between

the English system in which a cell is named for its posterior- limiting vein, and the system used by most American authors in which

the anterior-limiting vein is used to indicate the cell. Figure 2 illustrates the wing vein terminology used in this work.

The terminology used in describing other external morphological characters follows Clench (1975).

In this study wing span of an adult is sometimes reported. This is measured on a mounted specimen as the maximum distance between fore-wing 10 apices, with the wings spread so that the inner margin of the fore-wing is perpendicular to the longitudinal body axis. In measuring an individual fore-wing, measurement is made from the base of the wing to the apex at the outer angle.

The reported foodplant records come for the most part from three sources: Klots (1951), Howe (1975), and Pyle (1981). Additional data are included, especially when relating specifically to Ohio, and some discussion is included.

The citation of the name of a subsequent user of a scientific name, other than that of the author, is done here according to Article 51(b) of the International Code of Zoological Nomenclature (hereafter called the Code) (Stoll, et^ al., 1964), with a distinctive separation of the name of the taxon from the author's name. Three methods are used in this work to distinguish a subsequent user from the author of a name.

Two of these employ a semicolon. For example, all three of the following citations refer to Munroe's subsequent use of the tribal name , which was authored by Kirby in 1896: 1)

LEPTOCIRCINI Kirby, 1896:307; Munroe, 1961:15. 2) LEPTOCIRCINI Kirby,

1896:307 sensu Munroe, 1961:15. 3) LEPTOCIRCINI; Munroe, 1961:15.

References are sometimes found in the literature to the Life Zone in which a particular species of butterfly or is found. This system of zones as it applies to the Rhopalocera is described by Klots

(1951) and Clench (1975). is divided into seven zones, each consisting of a more or less continuous region (except in montane areas). These extend from the Artic-Alpine zone in the North to the 11

Subtropical zone in southern Florida and . Ohio is mainly within the Upper Austral zone, although a portion of the northeastern corner of the state is included in the Transition zone by both Klots and Clench

(op.cit.). As described by these authors, the Transition zone has a mean temperature of the warmest month of about 64-72° F., and is characterized in in the East by deciduous hardwood forest of beech, maple, and hemlock, and sometimes including eastern White Pine forest. The Upper Austral zone has a mean temperature of the warmest month of about 72-79° F., and is characterized by oak-hickory-maple forest in the East. Both authors cited above list Rhopalocera species which they consider characteristic of these zones. Since these relate to relatively broad, continental interpretation of the zones, the species will not be listed here. Rather, discussion of some of these will be included in the individual species accounts below.

Outline maps of Ohio illustrating the distribution of most species within Ohio are included in the appendix. Those species for which there are very few records, and those not known from Ohio but which might be expected to be found, are discussed in the text. Each map is marked with a hexagon for each county from which the species has been reported.

A solid hexagon represents one or more records based upon specimens personally examined by myself. A hollow hexagon represents a record based upon a literature reference, or some other source where it was impossible to examine the specimens. For purposes of this study a

"record" is meant to be a report of a single species collected at one 12

locality, at a particular time, by the same collector(s), without regard

for numbers of specimens collected.

Also, for most species there will be found in the appendix a graph

showing the seasonal distribution of adults. The months of March through

October, during which most adult butterflies are encountered in Ohio, are divided into thirds. A single asterisk on a chart represents a

single adult record (from both specimens and literature) for that

species during the time indicated. Again, the number of specimens

collected is not taken into account. In some cases, where adults of a species are frequently encountered during the period outside the

limits of these graphs (November through February), mention is made

in the discussion of that species.

Figure 3 shows the counties in Ohio with their names. THE TAXA WITH KEYS FOR DETERMINATION

The taxa included in this study generally are limited to the

following geographical areas: America north of Mexico for the family

group names, eastern United States, except Florida for the genera, and

Ohio and neighboring states for the species. These taxa are listed in

Table 1. Each taxon is discussed briefly. The information included for each species usually covers these areas of interest: nomen- clatorial and systematic status, diagnosis (unless necessary diag­ nostic characteristics have been included in the keys), geographical distribution, and biology including seasonal distribution.

Suborder Ditrysia BBorner, 1939

The higher classification of the Lepidoptera has been reviewed recently by Common (1975). He includes the families known commonly as the butterflies and skippers in the suborder Ditrysia. This sub­ order contains all heteroneurous Lepidoptera with two genital openings in the female, and an internal ductus seminalis which connects the two internal complexes which lead to the external openings. One of these openings, the ostium oviductus, vents the primary genitalia (ovaries, oviducts and vestibulum) which are involved in production and delivery. The second opening the ostium bursae, vents the bursa

13 Table 1. Species of Rhopalocera known from Ohio and neighboring states (an asterisk indicates a species not known from Ohio, but reported from neighboring states).

Species

HESPERIIDAE: PYRGINAE 1. Epargyreus clarus (Cramer, 1775) 2. *Urbanus ;.j (Linnaeus., ,1758):/ 3. Autochton cellus (Boisduval & Leconte, 1837) 4. Achalarus lyciades (Geyer, 1832) 5. Thorybes bathyllus (Smith, 1797) 6* Thorybes pylades (Scudder, 1870) 7. *Thorybes confusis Bell, 1922 8. Staphylus hayhurstii (Edwards, 1870) 9. Erynnis icelus (Scudder & Burgess, 1870) 10. Erynnis brizo (Boisduval & Leconte, 1834) 11. Erynnis juvenalis (Fabricius, 1793) 12. Erynnis horatius (Scudder & Burgess, 1870) 13. Erynnis martialis (Scudder, 1869) 14. *Erynnis zarucco (Lucas, 1857) 15. Erynnis lucilius (Scudder & Burgess, 1870) 16. Erynnis baptisiae (Forbes, 1936) 17. Erynnis persius (Scudder, 1864) 18. Pyrgus centaureae (Rambur, 1840) 19. Pyrgus communis (Grote, 1872) 20. Pholisora catullus (Fabricius, 1793) HESPERIIDAE: HESPERIINAE 21. Ancyloxypha numitor (Fabricius, 1793) 22. Thymelicus lineola (Ochsenheimer, 1808) 23. Nastra lherminier (Latreille, 1824) 24. * (Smith, 1797) 25. Hylephila phyleus (Drury, 1773) 26. *Hesperia ottoe Edwards, 1866 27. Hesperia leonardus Harris, 1862 28. Hesperia metea Scudder, 1864 29. Hesperia attalus (Edwards, 1871) 30. Hesperia sassacus Harris, 1862 31. Polites coras (Cramer, 1775) 32. Polites themistocles (Latreille, 1823) 33. Polites origenes (Fabricius, 1793) 34. Polites mystic (Edwards, 1864) 35. Polites vibex (Geyer, 1832) 36. Wallengrenia egeremet (Scudder, 1864) 37. (Edwards, 1862) ) 38. Atalopedes campestris (Boisduval, 1852) 39. Atrytone logan (Edwards, 1863) 40. Poanes massasoit (Scudder, 1864) 15

Table 1(continued).

Species

HESPERIIDAE: HESPERIINAE (continued) Poanes hobomok (Harris, 1862) 42. Poanes zabulon (Boisduval & Leconte, 1834) 43. Poanes viator (Edwards, 1865) 44. Euphyes dion (Edwards, 1879) 45. Euphyes dukesi (Lindsey, 1923) 46. Euphyes conspicua (Edwards, 1863) 47. Euphyes bimacula (Grote & Robinson, 1867) 48. Euphyes ruricola (Boisduval, 1852) 49. Atrytonopsis hianna (Scudder, 1868) 50. Amblyscirtes hegon (Scudder, 1864) 51. Amblyscirtes vialis (Edwards, 1862) 52. * eufala (Edwards, 1869) 53. * (Stoll, 1782) 54. Panoquina ocola (Edwards, 1863) PAPILIONIDAE: 55. Battus philenor (Linnaeus, 1771) 56. Eurytides marcellus (Cramer, 1779) 57. Papilio polyxenes asterius Stoll, 1782 58. Papilio cresphontes Cramer, 1777 59. Papilio glaucus Linnaeus, 1758 60. Papilio troilus Linnaeus, 1758 PIERIDAE: 61. Pontia protodice (Boisduval & Leconte, 1829) 62. *Artogeia napi (Linnaeus, 1758) 63. Artogeia virginiensis (Edwards, 1870) 64. Artogeia rapae (Linnaeus, 1758) PIERIDAE: ANTHOCHARINAE 65. * olympia (Edwards, 1871) 66. Falcapica midea (HUbner, 1809) PIERIDAE: 67. Colias philodice Godart, 1819 68. Colias eurytheme Boisduval, 1852 69. *Colias interior Scudder, 1862 70. Zerene cesonia (Stoll, 1790) 71. Phoebis sennae eubule (Linnaeus, 1767) 72. Phoebis philea (Johansson, 1763) 73. Eurema lisa (Boisduval & Leconte, 1829) 74. Eurema nicippe (Cramer, 1779) 75. Nathalis iole Boisduval, 1836 LYCAENIDAE: GERYDINAE 76. Feniseca tarquinius (Fabricius, 1793) Table 1(continued).

Species

LYCAENIDAE: 77. Lycaena phlaeas (Linnaeus, 1761) 78. Hyllolycaena hyllus (Cramer, 1775) 79. Epidemia epixanthe (Boisduval & Leconte, 1833) 80. Epidemia dorcas (Kirby, 1837) 81. Epidemia helloides (Boisduval, 1852) LYCAENIDAE: 82. Atlides halesus (Cramer, 1777) 83. Harkenclenus titus (Fabricius, 1793) 84. Satyrium acadica (Edwards, 1862) 85. Satyrium edwardsii (Grote & Robinson, 1867) 86. Satyrium calanus falacer (Godart, 1824) 87. Satyrium caryaevorum (McDunnough, 1942) 88. Satyrium liparops (Leconte, 1833) 89. Calycopis cecrops (Fabricius, 1793) 90. Mitoura gryneus (Hllbner, 1819) 91. Incisalia augustus (Kirby, 1837) 92. *Incisalia polios Cook & Watson, 1907 93. Incisalia irus (Godart, 1824) 94. Incisalia henrici (Grote & Robinson, 1867) 95. Incisalia niphon (Httbner, 1823) 96. Euristrymon Ontario (Edwards, 1868) 97. Parrhasius m-album (Boisduval & Leconte, 1833) 98. Strymon melinus HUbner, 1818 99. Erora laeta (Edwards, 1862) LYCAENIDAE: 100. Hemiargus isola (Reakirt, 1866) 101. Everes comynfas (Godart, 1824) 102. Celastrina ladon (Cramer, 1780) 103. Celastrina ebenina Clench, 1972 104. Glaucopsyche lygdamus (Doubleday, 1841) 105. Lyeaeides melissa (Edwards, 1873) RIODINIDAE 106. ^Calephelis virginiensis (Guerin-Meneville, 1831) 107. Calephelis borealis (Grote & Robinson, 1866) 108. Calephelis muticum McAlpine, 1937 LIBYTHEIDAE 109. Libytheana bachmanii (Kirtland, 1851) HELICONIIDAE 110. *Agraulis vanillae (Linnaeus, 1758) Table 1(continued).

Species

NYMPHALIDAE: ARGYNNINAE 111. Euptoieta claudia (Cramer, 1775) 112. Speyeria diana (Cramer, 1775) 113. Speyeria cybele (Fabricius, 1775) 114. Speyeria aphrodite (Fabricius, 1787) 115. Speyeria idalia (Drury, 1773) 116. Speyeria atlantis (Edwards, 1862) 117. Clossiana selene myrina (Cramer, 1777) 118. Clossiana bellona (Fabricius, 1775) NYMPHALIDAE: MELITAEINAE 119. *Charidryas gorgone (Hllbner, 1810) 120. Charidryas nycteis (Dobleday & Hewitson, 1847) 121. Charidryas harrisii (Scudder, 1864) 122. Phyciodes tharos (Drury, 1773) 123. (Reakirt, 1865) 124. Euphydryas phaeton (Drury, 1773) NYMPHALIDAE: 125. Polygonia interrogationis (Fabricius, 1798) 126. Polygonia comma (Harris, 1842) 127. *Polygonia (Edwards, 1862) 128. Polygonia progne (Cramer, 1776) 129. Nymphalis vau-album (Denis & Schiffermllller, 1775) 130. Nymphalis antiopa (Linnaeus, 1758) 131. Aglais milberti (Godart, 1819) 132. Vanessa virginiensis (Drury, 1773) 133. Vanessa cardui (Linnaeus, 1758) 134. Vanessa atalanta (Linnaeus, 1758) 135. Junonia coenia Hllbner, 1822 NYMPHALIDAE: 136. *Basilarchia arthemis arthemis (Drury, 1773) 137. Basilarchia arthemis astyanax (Fabricius, 1775) 138. Basilarchia archippus (Cramer, 1776) NYMPHALIDAE: 139. Anaea andria Scudder, 1875 NYMPHALIDAE: 140. Asterocampa celtis (Boisduval & Leconte, 1834) 141. Asterocampa clyton (Boisduval & Leconte, 1833) SATYRIDAE: ELYMNIINAE 142. *Enodia portlandia (Fabricius, 1781) 143. Enodia anthedon Clark, 1936 144. *Enodia creola (Skinner, 1897) 145. Satyrodes eurydice (Johansson, 1763) 146. (Chermock, 1947) 18

Table 1(concluded).

Species

SATYRIDAE: 147. Cyllopsis gemma (Hllbner, 1808) 148. sosybius (Fabricius, 1793) 149. mitchellii French, 1889 150. Megisto cymela (Cramer, 1777) 151. Cercyonis pegala (Fabricius, 1775) DANAIDAE 152. Danaus plexippus (Linnaeus, 1758) 153. Danaus gilippus (Cramer, 1775) copulatrix (corpus bursae, ductus bursae, and atrium) which

receives and stores sperm prior to fertilization (Clench, 1975)

Division Rhopalocera Boisduval, 1840.

The name Rhopalocera is used here as a grouping of

convenience. It is not intended to have any nomenclatorial

status. Rather, it is used to include those Lepidoptera which

have clubbed antennae, and usually lack either a frenulum or

jugum. These are commonly referred to as the butterflies and

skippers. In regard to these common names, some authors (Klots

1951 and Howe, 1975) include the skippers with the butterflies, while others (Ehrlich & Ehrlich, 1961) restrict the term butterfly so that the skippers are excluded.

Key to the superfamilies of the Rhopalocera in America north of Mexico.

1. Head broad, antennae separated at base by two to four times diameter of the flagellum, usually terminating with constricted, pointed apiculus beyond the thickened club; fore-wing veins extending from discal cell to margin unbranched ...... HESPERIOIDEA

1'. Head narrow, antennae separated at base by about one diameter of the flagellum always terminating in thickened club; fore-wing with some veins branched between discal cell and margin...... PAPILIONOIDEA 20

HESPERIOIDEA Latreille, 1809.

These are the Lepidoptera commonly called the skippers. As

with other Rhopalocera, the butterflies (PAPILIONOIDEA), there

is a thickened, club-like swelling at or near the end of the

antenna in the adult. As stated in the key above these may be

separated from the PAPILIONOIDEA by the wide spacing of the

antennae at their bases, the relatively broad head, and the

unbranched fore-wing veins between the discal cell and wing

margin. Also, the hind tibia usually possesses a spur about

midway along its length, not present in the PAPILIONOIDEA.

Key to the families of the HESPERIOIDEA in America north of Mexico.

1. Head stout, nearly as wide or wider than thorax; club partly or wholly recurved into an apiculus, or wing span of mounted specimen less than 30mm; hind tibia usually with two pairs of spurs HESPERIIDAE.

1'. Head narrower than thorax; antennal club straight, not recurved; wing span usually greater than 50mm; hind tibia with only one pair of spurs..MEGATHYMIDAE.

MEGATHYMIDAE Comstock & Comstock, 1895.

Adults of this family are stout-bodied, broad-headed and

are strong flyers. As such they resemble many of the

HESPERIIDAE. In America north of Mexico members of this

family are found in the western and southwestern United States,

in the Gulf states and in Florida, and the Carolinas

(Klots, 1951; Freeman, 1969). 21

HESPERIIDAE Latreille, 1809.

These skippers are distinguished from the MEGATHYMIDAE in the key and discussion above. They are widespread throughout

America north of Mexico, with over 200 species represented. In

Ohio and surrounding areas they account for about one-third of the known species of Rhopalocera.

Key to the Nearctic subfamilies of the HESPERIIDAE.

1. Antenna with distal end always recurved and with large, blunt, stout club in recurved portion, with widest part of club beyond recurve; all legs with 4th tarsal segment shorter than 5th (tropical, with a single U.S. species in the Southwest)...... PYRRHOPYGINAE,

1'. Antenna with distal end recurved or not, if recurved then with narrow apiculus always beyond recurve; middle and hind leg with 4th tarsal segment usually equal to or slightly longer than 5th...... 2.

2(1). Fore-wing vein M 2 arises from discal cell about midway between veins M^ and Mg, and mesotibia spineless; fore-wing vein CUg arises closer to wing base than Cu^; fore-wing discal cell usually 2/3 wing length or more; color usually dark brown, grayish, or black...... PYRGINAE.

2'. Fore-wing vein M 2 arises from cell closer to vein M^ than M^, or mesotibia spined; fore­

wing vein CU 2 arises midway between wing base and vein Cu^ or closer to Cu^; fore-wing discal cell less than 2/3 wing length; color usually medium brown and/or fulvous...... HESPERIINAE. 22

PYRGINAE Burmeister, 1878.

These skippers are separated from the other two Nearctic subfamilies in the key above (following the family discussion).

They are generally darker and larger than the HESPERIINAE, and the wings are usually held horizontally at rest. In some species the adult male possesses a fold of the costal edge of the fore-wing backward over the dorsal wing surface. This fold produces a pocket which contains androconial scales (specialized male scent scales) and functions as a scent-producing organ. The odors produced by these scales are presumably involved in attracting females. The male hind tibia may have tufts of specialized hairs.

The larvae feed mainly on dicotyledonous plants (MacNeill,

1975), often on legumes (Klots, 1951). Adults of this subfamily often bask in sunlight with wings spread either partly open or fully open (i. je. parallel to the substrate), with both fore- and hind- wings in the same plane.

As with the HESPERIINAE the PYRGINAE are widely distributed throughout America north of Mexico, and are well represented in

Ohio, although to a lesser extent in diversity of species than the

HESPERIINAE.

The PYRGINAE were divided into tribes by Clark (1948) who closely follwed the "groups" of Lindsey, Bell and Williams (1931).

Clark's classification has not been repeated in any major publication since his use, and since a revision of the group is 23 well beyond the scope of this work, the "groups" of Lindsey, Bell and Williams (1931), also used by Macy and Shepard (1941), and by

Klots (1951), will be followed here.

Key to the groups of the PYRGINAE in North America.

1. Fore-wing discal cell more than 2/3 length of wing, o£ antenna with slender, reflexed or recurved apiculus; labial palp erect, second segment appressed to face, with third segment not protruding in front of second segment...... Group A.

1'. Fore-wing discal cell 2/3 length of wing or less; antenna lacking apiculus; labial palp porrect, with third segment protruding in front of second segment Group B.

Group A.

Five genera are currently recognized in this group in the eastern United States, excluding Florida. The tailed-skippers of the Ohio area are included here, as are those with spectacular silver markings of the ventral hind-wing surface. This group is distinguished from Group B in the key above.

Key to the genera of Group A in the eastern United States (except Florida).

1. Hind-wing with large, ventral, white patch crossing more than two veins...... 2 .

1'. Hind-wing not as above; white spots, if present, cross no more than one vein...... 3. 24

2(1). Hind-wing with ventral* white patch centrad and with sharply defined edges...... Epargyreus.

2'. Hind-wing with ventral, white patch adj acent to outer margin, and with poorly defined, difuse edges...... Achalarus.

3(1'). Hind-wing with tail extending posteriorly, extending inner margin by at least 50%...... Urbanus.

3'. Hind-wing not as above; tail, if present, short...... 4.

4(3'). Fore-wing with dorsal, median gold band extending from anterior edge nearly to tomus ...... Autochton.

4'. Fore-wing not as above, lacking gold band...... Thorybes. 25

Epargyreus Hllbner, [1819]

NOMENCLATORIAL STATUS

Epargyreus Hllbner, [1819]:105. [type-species: Papilio tityrus Fabricius, 1775, selected by Scudder, 1872]

SYSTEMATIC STATUS

Epargyreus Hllbner, [1819]. [valid name (Evans, 1952:44; Dos Passos, 1964:29; Miller & Brown, 1981:3)]

Eridamus Burmeister, 1875. [subjective synonym (Evans, 1952:44; Dos Passos, 1964:29; Hemming, 1967:167]

DIAGNOSTIC CHARACTERISTICS

Fore-wing pointed apically, hind-wing produced at anal angle, most

species with a silver-colored band or group of spots on ventral surface.

Fore-wing with yellow or orange, hyaline spots. Fore-wing of male with

costal fold. Male lacking hair tufts on wings or legs. Labial palp

short, with stout third segment just visible beyond scaling of second

segment.

DISCUSSION

This is a New World genus with most of its 15 or 16 species

occuring in the tropics. Three of these species occur in America north of Mexico. One of these occurs in the Ohio area (MacNeill, 1975).

Larvae of this genus feed on a variety of Leguminaceae (Ferris and

Brown, 1981). 26

Epargyreus clarus (Cramer, 1775)

Silver-spotted Skipper

NOMENCLATORIAL STATUS

Papilio clarus Cramer, 1775. [type-locality: restricted to Dayton, Rockingham Co., Virginia by Dixon, 1955 (Irwin & Downey, 1973:14)]

Papilio tityrus Fabricius, 1775. [preoccupied by Papilio tityrus Poda, 1761 (Miller & Brown, 1981:3)]

Epargyreus tityrus; Holland, 1898. [preoccupied name]

SYSTEMATIC STATUS

Epargyreus clarus (Cramer, 1775). [valid name (MacNeill, 1975:571)]

COMMON NAME

Silver-spotted Skipper (Pyle, 1981:719; Klots, 1951:206)

DIAGNOSTIC CHARACTERISTICS

Dorsal fore-wing with prominent, translucent gold spots forming band across discal area. Ventral hind-wing with well-defined, silver patch in discal area, hence the common name. One of the larger skippers in Ohio.

DISTRIBUTION

MacNeill (1975) reports that this species ranges from Quebec to British Columbia and southward to Florida, Texas, northwestern

Mexico and Baja . Ohio records extend across most of the state (Fig. 4).

BIOLOGY

Larvae feed on woody legumes, including species of Robinia

(locusts), Wisteria and Gleditsia (Honey Locust) (MacNeill, 1975). 27

Adults common in old fields, grassy areas, and wood edges.

Klots (1951) reports one generation northward and two or three in the South, with adults appearing in May at 40° N. Latitude. Ohio records extend from April through August (Fig. 5). Peaks of adult activity in May and again in July/August, separated by a short decrease, suggest the possibility of two overlapping generations. 28

Urbanus Hllbner, 1807

NOMENCLATORIAL STATUS

Urbanus Hllbner, 1807. [type-species: Papilio proteus Linnaeus, 1758, selected by Hemming, 1933]

Goniurus Hllbner, 1819. [junior objective synonym of Urbanus Hllbner, 1807 (Hemming, 1967)]

Goniuris Westwood, 1852. [incorrect subsequent spelling]

Eudamus Swainson, 1831. [junior objective synonym of Urbanus Hllbner, 1807 (Hemming, 1967)]

SYSTEMATIC STATUS

Urbanus Hllbner, 1807. [valid name (Miller & Brown, 1981; Evans, 1952)]

DIAGNOSTIC CHARACTERISTICS

Hind-wing produced into tail at anal angle. Thorax and wing base often metallic green or blue. Medium to large in size for skippers.

DISCUSSION

This is a New World genus, mainly tropical, with species occuring from the United States to . A single species,

Urbanus proteus, is known from the eastern United States. Several members of this genus have been reported to feed on monocotyledonous plants. The supposed rule of dicotyledonous foodplants for the subfamily may not be valid. 29

Urbanus proteus (Linnaeus, 1758)

Long Tailed Skipper

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio proteus Linnaeus, 1758. [type-locality: "America"]

Urbanus proteus (Linnaeus, 1758). [valid name (Miller & Brown, 1981:7)]

Eudamus proteus; Holland, 1898:321.

Goniurus proteus; Lindsey, Bell & Williams, 1931:23.

COMMON NAMES

Long Tailed Skipper (Klots, 1951:207)

Bean Leaf Roller (Klots, 1951:207)

DIAGNOSIS

This is the only skipper found in the Ohio area with the hind-

wing produced into a tail which extends the length of the inner

margin of the wing by at least 50%. Head, thorax and wing base

conspicuously green dorsally.

DISTRIBUTION

This skipper is distributed from south to Florida,

west through Arkansas and Texas to and California, thence

southward to Argentina (MacNeill, 1975:559). There is a single, museum specimen with a Columbus, Ohio label (without date) in the

Ohio State University Insect Collection. This species has been

reported from Michigan, , and Kentucky, and could occur

in Ohio as a stray. Until confirmed by other specimens, this should not be considered part of Ohio's skipper fauna. 30

BIOLOGY

Urbanus proteus feeds frequently on agricultural legumes including several beans, turnips, and cabbage (MacNeill, 1975:559).

It becomes at times' a pest earning its common name of the Bean

Leaf Roller (Klots, 1951:207). Klots (1951:210) reported three generations, but it is doubtful that adults from each generation stray into the northern range area, which would include Ohio. 31

Autochton HUbner, 1823

NOMENCLATORIAL AND SYSTEMATIC STATUS

Cecrops Hllbner, 1818. [homonym of Cecrops Leach, 1816 (Hemming, 1967:103; Evans, 1952:120)]

Autochton Hllbner, 1823. [type-species: Autochton itylus Hllbner, 1823, by monotypy]

Autochton Hllbner, 1823. [valid name (Evans, 1952:120)]

Cecropterus Herrich-Schaeffer, 1869.

Rhabdoides Scudder, 1889.

DIAGNOSIS

In America north of Mexico, adults of this genus are easily

recognized by the wide, gold band crossing the discal area of the

dorsal fore-wing from the costal edge to near the toxmus. Antenna with well-developed, long apiculus. Males lack both costal fold of

fore-wing and metatibial hair tuft. Medium to large for skippers.

DISCUSSION

This New World genus is mainly tropical with a dozen or more

species ranging from the United States to Argentina. Two of these

occur in the United States, with only one of these occuring east of

the Mississippi River.

Autochton cellus (Boisduval & LeConte, 1837)

Golden-banded Skipper

NOMENCLATORIAL AND SYSTEMATIC STATUS

Eudamus cellus Boisduval & LeConte, 1837. [type-locality: North America] 32

Autochton cellus (Boisduval & Leconte, 1837). [valid name (Miller & Brown, 1981:10)]

Achalarus cellus; Holland, 1898:326.

Rhabdoides cellus; Holland, 1931:338.

Cecropterus cellus; Lindsey, Bell & Williams, 1931:31.

COMMON NAMES

Golden-banded Skipper (Holland, 1931:338; Pyle, 1981:730)

Gold-banded Skipper (Macy & Shepard, 1941:183)

DIAGNOSIS

Dorsal fore-wing surface with continuous, golden band extending across the discal area from costal margin to near the tornus. Ventral hind-wing lacking silver areas of Epargyreus and

Achalarus.

DISTRIBUTION

Disjunct, mainly local distribution from to Ohio and

Missouri, southward to Florida and westward to Arizona (MacNeill,

1975:556). Ohio records are limited to several southeastern, unglaciated areas (Fig. 6).

BIOLOGY

Authors report the Hog Peanut (Amphicarpa pitcheri) as the primary food source. Adults in wooded ravines, roadsides, and stream sides. Two to three generations, with adults first appearing in late May in Virginia (Klots, 1951:211). Ohio records extend from mid-June to late July (Fig. 7). Achalarus Scudder, 1872

NOMENCLATORIAL AND SYSTEMATIC STATUS

Achalarus Scudder, 1872. [type-species: Proteides lyciades Geyer 1832, by original designation; valid name (Miller & Brown, 1981:10)]

DIAGNOSIS

These are rather large for skippers, usually brown, with a short, slender costal fold present in males of most species. The single species found east of the Mississippi River in the United

States has gold-colored spots on the dorsal surface of the fore­ wing, similar to Autochton and Epargyreus. The broad, silver- colored, smear-like band along the outer edge of the ventral hind- wing surface distinguishes Achalarus from those other two genera.

DISCUSSION

This is another of the New World genera with five or more species ranging from the United States into northern

Three of these species are known from the United States.

Achalarus lyciades (Geyer, 1832)

Hoary Edge

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio lycidas Smith, 1797. [unavailable junior homonym, preoccupied by Papilio lycidas Cramer, 1777 (Miller & Brown, 1981:10)]

Proteides lyciades Geyer, 1832. [type-locality: Bahia] 34

Achalarus lyciades (Geyer, 1832). [valid name (Miller & Brown, 1981:10)]

Achalarus lycidas; Holland, 1898. [preoccupied]

COMMON NAMES

Hoary Edge (Pyle, 1981:731; Klots, 1951:210)

Frosted Skipper (Mitchell & Zim, 1964:74)

DIAGNOSIS

Ventral hind-wing with broad, silver-colored marking in limbal area. The position of this area on the wing, together with its diffuse edge, distinguish this skipper from Epargyreus clarus with which it is often confused.

DISTRIBUTION

MacNeill (1975:554) reports this species as being distributed from New Hampshire to Florida, and westward to Minnesota and northeastern Texas. Ohio records indicate it is fairly widespread

(Fig. 8).

BIOLOGY

Authors list for this species a number of foodplants, including tick trefoils (Desmodium spp.) and Lespedeza spp. A single annual generation with adults flying from May into July is reported by authors for the northern part of the range. A second generation is presumed to account for flight into the autumn in the southern part of its range. Ohio records show adult activity from early May into July, with one record in August (Fig. 9). Peak activity in Ohio occurs in late June, and there is no indication of more than a single generation. Adults of Achalarus lyciades are found in grassy areas, roadsides, and wooded edges. 36

Thorybes Scudder, 1872

NOMENCLATORIAL STATUS

Thorybes Scudder, 1872. [type-species: Papilio bathyllus Smith, 1797, by original designation]

Lintneria Butler, 1877. [preoccupied by Lintneria Butler, 1876 (Miller & Brown, 1981:11)]

SYSTEMATIC STATUS

Thorybes Scudder, 1872. [valid name (Miller & Brown, 1981:)]

Cocceius Godman & Salvin, 1894.

DIAGNOSIS

Dark brown with rounded hind-wing. Fore-wing with small,

hyaline spots. Males with fore-wing costal fold in some species; metatibial hair tuft lacking.

DISCUSSION

The seven species listed in this genus by MacNeill (1975:547)

range from the United States to . Three of these occur east

of the Mississippi River, and all are known from the Ohio area.

They feed on legumes (Ferris & Brown, 1981:70). Adults of all three

species are found in open, grassy areas, often near woods.

Key to the species of Thorybes in the Ohio area.

1. Male, with costal wing fold (fore-wing).... pylades.

1'. Male, lacking costal wing fold, or female.. 2 . 37

2(1'). Dorsal fore-wing surface with median, white spots large, extending from vein to vein, forming nearly continuous diagonal band across wing; labial palp with ventral surface white, strongly contrasting with darker head color...... bathyllus.

2'. Dorsal fore-wing surface with median, white spots small, not extending from vein to vein, widely separated by darker background and forming a discontinuous line across anterior % to 2/3 of wing rather than band (spots absent in some specimens); labial palp with ventral surface brown or dark gray, not strongly contrasting with head color...... 3.

3(2’). Ventral hind-wing surface with transverse bands lighter in color than background, but with clear, dark outlines giving tricolored contrast; ventral hind-wing surface uniformly flecked with single, pale, whitish scales...... confusis.

3'. Ventral hind-wing surface with transverse bands closely similar in color to background, and with weak outlines, poorly contrasting with other colors; ventral hind-wing surface dusted with bluish scales (contrasting little with ground color) near outer margins, lacking uniform flecking with whitish scales as above; female...... pylades.

Thorybes bathyllus (Smith, 1797)

Southern Cloudywing

NOMENCLATORIAL STATUS

Papilio bathyllus Smith, 1797. [type-locality: Georgia]

SYSTEMATIC STATUS

Thorybes bathyllus (Smith, 1797). [valid name (MacNeill, 1975:548)]

Eudamus bathyllus; Skinner, 1898:96.

COMMON NAMES

Southern Cloudywing (Pyle, 1981:733)

Southern Dusky Wing (Macy & Shepard, 1941:184) 38

DIAGNOSIS

This species, and the two species following, are characterized

in the key above.

DISTRIBUTION

This species is reported by authors to range from Massachusetts

southward to Florida, and westward to , and Texas.

Ohio records indicate distribution throughout the state (Fig. 10).

BIOLOGY

The reported foodplants include Lespedeza spp., Strophostyles

spp. and Cracca ambigua. One annual generation in the northern and

two in the southern parts of the range are reported. Ohio records

run from May into early September with peak adult activity indicated

for late June (Fig. 11). This corresponds with MacNeill (1975:548) who says that adults are most abundant in late June.

Thorybes pylades (Scudder, 1870)

Northern Cloudywing

NOMENCLATORIAL STATUS

Eudamus pylades Scudder, 1870. [type-locality: not stated, but probably Massachusetts (Miller & Brown, 1981:11)]

Eudamus bathyllus Harris, 1862:312 (nec Smith, 1797). [misiden- tification]

SYSTEMATIC STATUS

Thorybes pylades (Scudder, 1870). [valid name (Miller & Brown, 1981:11)] 39

Eudamus pylades ab. immaculata Skinner, 1911.

Thorybes pylades ab. integra Lanktree, 1968.

COMMON NAMES

Northern Cloudywing (Pyle, 1981:734)

Northern Dusky Wing (Macy & Shepard, 1941)

DISTRIBUTION

This species has a wider range than does Thorybes bathyllus, extending from Quebec to Flprida, and westward to British

Columbia, California, Baja California, and south-central Mexico.

Ohio records occur throughout the state (Fig. 12).

BIOLOGY

Foodplants include clovers (Trifolium), Lespedeza and Desmodiuro

(Klots, 1951:213). MacNeill (1975:549) reports a single generation

(May to early July) in the northern and three or more in the southern parts of this species' range. Klots (1951:213), on the other hand, says that there are two generations in the northern part of its range. Ohio records extend from May into September with greatest adult activity in late May and June (Fig. 13). More data are needed to determine whether there are two generations in

Ohio. 40

Thorybes confusis Bell, 1922

Confused Cloudywing

NOMENCLATORIAL AND SYSTEMATIC STATUS

Thorybes confusis Bell, 1922. [type-locality: Tampa, Florida; valid name (Miller & Brown, 1981:12)]

COMMON NAMES

Confused Cloudy Wing (Klots, 1951:231)

Eastern Cloudywing (Pyle, 1981:735)

DISTRIBUTION

Authors report distribution of this species as extending from

Maryland to Florida, and westward through Pennsylvania and Kentucky to , and Texas. Although no records are known from

Ohio, it may be expected to be found in the southern or southeastern part of the state. Confusion of this species with Thorybes pylades may have resulted in collectors overlooking this species.

BIOLOGY

MacNeill (1975:552) reports that the early stages of this species are still unknown. No food plants are listed by authors.

In the northern part of its range there is adult activity in July and August, probably with a single generation. 41

Group B.

Four genera of this group are found in the eastern United

States excluding Florida. These are generally the darkest and most dully colored of our skippers. The key to groups with the subfamily discussion of the PYRGINAE above distinguishes these from Group A.

Key to the genera of Group B in the eastern United States

(except Florida).

1. Hind-wing with conspicuous white markings on dorsal surface, contrasting strongly with darker background...... Pyrgus.

1'. Hind-wing lacking dorsal markings as above, or with single submarginal row of small, light colored spots...... 2.

2(1'). Antennal club bent at or near its middle...... Staphylus.

2'. Antennal club bent at or near its base...... 3.

3(2'). Fore- and hind-wing uniformly black dorsally in ground color; fore-wing usually with dorsal hyaline spots; antennal club bent proximally...... Pholisora.

3'. Wings above mottled with varying shades of brown and gray, especially on the fore-wing; fore-wing with or without hyaline spots; antennal club bent shortly distal to its base...... Erynnis. 42

Staphylus Godman & Salvin, 1896

NOMENCLATORIAL AND SYSTEMATIC STATUS

Staphylus Godman & Salvin, 1896. [type-species: Helias ascalaphus Staudinger, 1876, by original designation; valid name (Miller & Brown, 1981:15)]

DIAGNOSIS

Small (wing span 25-32mm), dark brown to black, with very few light colored wing markings. Male with costal fold on fore-wing, but lacking metatibial hair tufts. Hind-wing scalloped along outer margin opposit discal cell. Antenna long and slender with apiculus bent near its center. This genus is separated from the similar- appearing Pholisora in the key above.

DISCUSSION

There are more than a dozen species in this New World genus, most of which are tropical. Three of these occur in the United

States with only one found east of the Mississippi River.

According to Ferris and Brown (1981:89) foodplants are various members of the Chenopodiaceae and Amaranthaceae.

Staphylus hayhurstii (Edwards, 1870)

Scalloped Sootywing

NOMENCLATORIAL STATUS

Hesperia hayhurstii Edwards, 1870. [type-locality: Missouri]

Pholisora hayhursti Holland, 1898. [misspelling] 43

SYSTEMATIC STATUS

Staphylus hayhurstii (Edwards, 1870). [valid name (Miller & Brown, .1981:15]

Pholisora hayhurstii; Klots, 1951:218.

Staphylus mazans hayhurstii; dos Passos, 1964:23.

COMMON NAMES

Scalloped Sootywing (Pyle, 1981:741)

Southern Sooty Wing (Klots, 1951:218)

Hayhurst's Skipper (Holland, 1931:346)

DISTRIBUTION

Reported to be from Pennsylvania to Florida, and westward to

Colorado and Texas. It is reported from several central and southern Ohio counties, and three northern counties in Ohio (Fig.

14). Ohio is at the northern limit of this species.

BIOLOGY

Authors report Chenopodium and Alternanthera as foodplants.

NacNeill (1975:541) states that there are at least three generations with adults being collected from March to November. Klots (1951:

219) refers to "at least" three generations in the southern part of the range. Ohio records extend from May to the end of August, with little indication of multiple generations other than the long length of the flight period (Fig. 15). 44

Erynnis Schrank, 1801

NOMENCLATORIAL STATUS

Erynnis Schrank, 1801. ftype-species: Papilio tages Linnaeus, 1758, selected by Scudder, 1872:71]

Thymele Fabricius, 1807. [junior objective synonym of Erynnis Schrank, 1801 (Hemming, 1967:442)]

Thymele [Illiger], 1807. [suppressed by 1CZN in Opinion 232 and placed on Official Index of Rejected and Invalid Generic Names in Zoology (Burns, 1964:23)]

Thymale Oken, 1815. [junior objective synonym of Erynnis Schrank, 1801 (Hemming, 1967:442); also, a misspelling of Thymele Fabricius, 1807 (Cowan, 1970:32)]

Thymale Rafinesque, 1815. [misspelling of Thymele Fabricius, 1807 (Cowan, 1970:69)]

Astycus HUbner, 1822. [junior objective synonym of Erynnis Schrank, 1801 (Hemming, 1967:442)]

Thanaos Boisduval, [1834]. [junior objective synonym of Erynnis Schrank, 1801 (Hemming, 1967:442)]

Thanatos [Dunning & Packard], 1858. [misspelling of Thanaos Boisduval, [1834] (Hemming, 1967:437)]

Thanaus Kirby, 1897. [misspelling of Thanaos Boisduval, [1834] (Hemming, 1967:437)]

Hallia Tutt, 1906. [junior homonym of Hallia Edwards & Haime, 1850 (Hemming, 1967:205)]

SYSTEMATIC STATUS

Erynnis Schrank, 1801. [valid name (Evans, 1953:205)]

Nisoniades Hllbner, 1819. [subjective synonym: before being restricted to Neotropical species, various authors used this genus to include the Holarctic species (Scudder, 1889; Stevens, 1850 (Bums, 1964:23))]

Erynnides Burns, 1964. 45

DIAGNOSIS

These skippers are small to medium (wing span 25-44mm). Their wings are dark gray-brown and are marked with various combinations of brown and black markings, as well as small, hyaline spots in some

species. The antennae are relatively short, usually slightly less than the fore-wing length, and the apiculus is short. Males possess fore-wing costal folds, and in some species the metatibial hair tufts.

DISCUSSION

Burns (1964) has studied the New World species ("about thirty") of this Holarctic genus very thoroughly. Seventeen species occur in

America north of Mexico, with eight found in the Ohio area. Although some of these eight species are more common than others, distribution of the genus appears to be fairly general throughout the state.

Authors and students of skippers find this group very difficult to separate at the species level, and keys are often inadequate. It is hoped that the combination of key and discussion below will aid in the determination of species included in this work. In deference to the key and discussion, diagnostic characteristics have been omitted from the individual species accounts below. Burns (op. cit.) has pointed out and discussed several problems with the understanding of this genus, suggesting that species concepts may change with further study. Foodplant information reported in the species accounts below comes from Burns (1964), Klots (1951), and MacNeill (1975). 46

Problems are often encountered with determination of the species

of Erynnis. One pair of species which frequently gives concern is IS.

juvenalis and horatius. As pointed out by Klots (1951:223) the

presence or absence of two pale spots on the ventral hind-wing (see

couplet number 4 in the key to species below), while reliable for

females in the northern part of their ranges, is not always reliable

for males. Males of these two species may be confirmed by examining

the distal parts of the genitalia iri situ, under low magnification.

First, the terminal abdominal scales are removed with a fine-haired brush or jeweler's forceps. This exposes the distal parts of the male genitalia. Then, the distal part of each valva (also called the or clasper by some authors) is examined from the lateral aspect. The distal part of the valva is actually the ventral process of a three- part valva, and is the part of the male genitalia which extends farthest distad. In _E. juvenalis this ventral process of the left valva is fairly uniformly broad throughout its length, except at the posterior end where it tapers to a dull point. In IS. horatius, however, the ventral process of the left valva basally is relatively narrow, expanded in width distally, and broadly truncate at its terminus.

Erynnis zarucco has distinctive male genitalia, which will help in separating it from both IS. martialis and baptisiae. The ventral process of the left valva in IS. zarucco is long, thin, and tapering distally to a small hook at the end. It is gently curved dorsad throughout its length. The ventral process of the right valva, 47 however, is much shorter than the left, and is truncate distally. In contrast with 15. zarucco, the ventral processes of both valvae in 15. baptisiae are subequal in length, and both are gently curved dorsally throughout their lengths, and are long, thin, and tapering distally to a dully pointed end. However, they are without the small hook found on the left process in zarucco. In contrast with both JS. baptisiae and zarucco, the ventral processes of both valvae in j5. martialis are bent dorsad at a pronounced angle near the distal end, especially on the left process. This gives the appearance of far more of a bend than a small, terminal hook (in which the length of the hook is no greater than the width of the ventral process proximal to the hook).

In 15. martialis, the portion of the ventral process beyond the bend is much longer than the greatest width of the process.

In confirming separation of males of Erynnis icelus and brizo, the ventral processes of the valvae in icelus are subequal in length.

In _E. brizo, however, the right is much shorter than the left, and the dorsal edge of the left is dentate (smooth in icelus) .

The male genitalia are not reliable in separating _E. persius, lucillius, and baptisiae. Characters in the key below will have to be used. Authors sometimes use larval food preferences in separating them. A key to the species of Erynnis in the Ohio area. 48

1. Fore-wing with distinct, hyaline spots; antenna apex sharp...... 3.

1'. Fore-wing lacking distinct hyaline spots, sometimes with one or two clouded spots next to costal margin; antenna apex blunt...... 2.

2(1'). Labial palp with tip extending beyond end of palpal hairs for at least 1/3 eye diameter; male metatibia with dense brush of hairs, proximal hairs being longer than tibia; female sternum VII with scales covering proximal % or less, scales of terminal tergite (VIII) light gray-brown in color...... icelus.

2'. Labial palp with tip barely free of palpal hairs, extending far less than 1/3 eye diameter; male metatibia with hairs, but not a dense brush, none longer than tibia; female sternum VII entirely scales, scales of terminal tergite dark brown...... brizo.

3(1). Antennal nudum with 14-17 segments...... 5.

3'. Antennal nudum with 18-21 segments...... 4.

4(3'). Ventral hind-wing with two pale, subapical spots; male with metatibial hair tuft; dorsal fore-wing of male with whitish hairs elevated above wing...... juvenalis.

4'. Ventral hind-wing usually lacking spots as above; male dorsal fore-wing with brownish rather than whitish hairs elevated above wing surface...... horatius.

5(3). Dorsal hind-wing with distinct, blackish bar at distal end of discal cell, and with medial and submarginal row each of distinct, blackish spots, many of which have a lighter spot just distad; dorsal fore-wing often with purplish cast, especially in fresh specimens; male lacking metatibial hair tuft...... martialis.

5'. Dorsal hind-wing without above combination of characters; male with metatibial hair tuft •...... 6.

6(5'). Dorsal fore-wing very dark, almost black, usually with a pale, brownish patch at distal end of discal cell; dorsal fore-wing with hyaline white spots usually much reduced...... zarucco.

6'. Dorsal fore-wing brownish, with contrasting marks of varying shades of brown, lacking pale brown patch at distal end of discal cell; dorsal fore-wing with hyaline white spots usually distinct...... 7. 7(6'). Male dorsal fore-wing with many hairlike scales, ^9 rising from wing surface; female sternum VII unsealed; dorsal fore-wing with contrasting markings generally obscured toward base, hyaline white spots usually limited to small group near costal edge...... persius.

7'. Male dorsal fore-wing with few hairlike scales, lying flat to surface; female sternum VII heavily scaled, although scales may be partially or wholy lost in worn specimens; dorsal fore-wing usually with at least a few hyaline white spots in limbal area...... 8.

8(7'). Small, wing span generally 23-31mm; dorsal fore-wing with contrasting markings fairly well defined throughout; ventral hind-wing with distinct, clear cut submarginal pale spots...... lucilius.

8'. Medium sized, wingspread generally 29-40mm; dorsal fore-wing with contrasting markings more or less obscured, especially toward base; ventral hind-wing with pale, submarginal spots obscure...... baptisiae. 50

Erynnis icelus (Scudder & Burgess, 1870)

Dreamy

NOMENCLATORIAL AND SYSTEMATIC STATUS

Nisoniades icelus Scudder & Burgess, 1870. [type-locality: New England]

Erynnis icelus (Scudder & Burgess, 1870). [valid name (Miller & Brown, 1981:2)]

Nisoniades bautista PlBtz, 1884.

Nisoniades hamamaelidis Scudder, 1889.

Thanaos icelus; Holland, 1898.

Nisoniades icelus; Skinner, 1898.

COMMON NAME

Dreamy Duskywing (Pyle, 1981:748)

DISTRIBUTION

From Nova Scotia to British Columbia in the North, and south­ ward to Georgia, Ohio, Indiana, and Minnesota. West of the prairie areas it extends southward to New Mexico, Arizona, Nevada, and

California. Ohio records extend across the northern and southern counties, and, to a lesser extent, occur in the central part of the state (Fig. 16).

BIOLOGY

Foodplants include species of Salix, , and Betula

(MacNeill, 1975; Burns, 1964). MacNeill (op. cit.) reports that this species seems to fly from April to August, being ordinarily more common in occurrence during the early part of this period. 51

Ohio records are somewhat more restricted than this, extending from

April only through June (Fig. 17). Peak activity in Ohio appears to be in late May.

Erynnis brizo (Boisduval & LeConte, 1834)

Sleepy Duskywing

NOMENCLATORIAL AND SYSTEMATIC STATUS

Thanaos brizo Boisduval & LeConte, 1834. [type-locality: not stated]

Erynnis brizo (Boisduval & LeConte, 1834). [valid name (Miller & Brown, 1981)]

Nisoniades brizo; Skinner, 1898.

COMMON NAME

Sleepy Duskywing (Pyle, 1981:748)

DISTRIBUTION

From Massachusetts southward to Florida, westward to Manitoba and eastern Texas. Records from Ohio indicate a fairly broad range across the state (Fig. 18).

BIOLOGY

Authors report species of Quercus (oaks) and Castanea

(American chestnut) as foodplants for this species. Adult flight activity is reported to be in the early spring. Ohio records are from April and May, peaking in early May (Fig. 19). 52

Erynnis juvenalis (Fabricius, 1793)

Juvenal1s Duskywing

NOMEN CLATORIAL AND SYSTEMATIC STATUS

Hesperia juvenalis Fabricius, 1793. [type-locality: America]

Erynnis juvenalis (Fabricius, 1793). [valid name (Miller & Brown, 1981:20)]

Nisoniades costalis Westwood, 1852.

Nisoniades ennius Scudder & Burgess, 1870.

Nisoniades plautus Scudder & Burgess, 1870.

Nisoniades .juvenalis; Skinner, 1898.

Thanaos .juvenalis; Holland, 1898.

COMMON NAME

Juvenal's Duskywing (Pyle, 1981:749)

DISTRIBUTION

From Nova Scotia to Florida, thence westward to Manitoba and

Texas. It is probably the most common member of its genus to be found in Ohio, and is found throughout much of the state (Fig. 20).

BIOLOGY

As with the preceding species, Erynnis juvenalis is an oak

(Quercus) feeder. In most of its range it is reported to have a single generation, with adults flying in the spring. A second generation appears in Florida and Arizona according to MacNeill

(1975:522). Except for a single, September record, adult activity in Ohio is known to extend from April into June, peaking in the period of late April to early May (Fig. 21). 53

Erynnis horatius (Scudder & Burgess, 1870)

Horace's Duskywing

NOMENCLATORIAL AND SYSTEMATIC STATUS

Nisoniades horatius Scudder & Burgess, 1870. [type-locality: New England and Texas]

Erynnis horatius (Scudder & Burgess, 1870). [valid name (Miller & Brown, 1981:20)]

Nisoniades virgilius Scudder & Burgess, 1870.

Nisoniades petronius Lintner, 1881.

Thanaos horatius; Holland, 1898.

COMMON NAME

Horace's Duskywing (Pyle, 1981:752)

DISTRIBUTION

From Massachusetts to Florida, westward to Minnesota, Colorado,

New Mexico, and Texas. Records for Ohio include northern, central, and south-central parts of the state (Fig. 22).

BIOLOGY

Species of oak (Quercus) are reported as the foodplants. It is reported to have two generations in the northern and three in the southern parts of its range. This appears confirmed by records from Ohio where adults fly from April into September, with two minor peaks in May and July (Fig. 23). 54

Erynnis martialis (Scudder, 1869)

Mottled Duskywing

NOMENCLATORIAL AND SYSTEMATIC STATUS

Nisoniades martialis Scudder, 1869. [type-locality: New England]

Erynnis martialis (Scudder, 1869). [valid name (Miller & Brown, 1981:21)]

Thanaos quercus Butler, 1870.

Thanaos martialis; Holland, 1898.

COMMON NAMES

Mottled Duskywing (Pyle, 1981:753)

Martial's Dusky-wing (Holland, 1931)

DISTRIBUTION

From New England south to Georgia, and west to South Dakota and

Texas. The rather sparse Ohio records come from the northern and southern parts of the state, and are probably not representative of the whole distribution of this species, within the state (Fig. 24).

BIOLOGY

Although authors have reported many different foodplants for this skipper, probably is the primary source of larval food (MacNeill, 1975:526). B u m s (1964:147) reports this species to be multivoltine in eastern North America. This is confirmed for Ohio, to a certain extent, in that adult captures extend from May through August, too long a period for the univoltine species of this genus in Ohio (Fig. 25). Erynnis zarucco (Lucas, 1857)

Zarucco Duskywing

NOMENCLATORIAL AND SYSTEMATIC STATUS

Thanaos zarucco Lucas, 1857. ftype-locality: Cuba]

Erynnis zarucco (Lucas, 1857). [valid name (Miller & Brown, 1981: 21)]

Nisoniades terentius Scudder & Burgess, 1870.

Nisoniades ovidius Scudder & Burgess, 1870.

Nisoniades maevius Lintner, 1881.

Antigonous diogenes PlBtz, 1884.

Thanaos terentius; Holland, 1931.

COMMON NAME

Zarucco Duskywing (Pyle, 1981:755)

DISTRIBUTION

This is a southeastern species ranging from to

Florida and west to and Louisiana. Although not confirmed

for Ohio, records from Illinois (Irwin and Downey, 1973), Indiana

(Shull and Badger, 1972), Pennsylvania (Clench, 1971), and Kentucky

(Covell, 1974) strongly suggest the possibility of its occurence

in Ohio. Confusion between this species and other species of

Erynnis may have led to collectors overlooking Erynnis zarucco

in Ohio in the past.

BIOLOGY

MacNeill (1975:529) reports legumes as the primary foodplants for this species, although he cautions against the reliability of 56

published records because of the confusion between this species and

Erynnis baptisiae. Burns (1964:168) refers to this species as being

strongly multivoltine.

Erynnis lucilius (Scudder & Burgess, 1870)

Columbine Duskywing

NOMENCLATORIAL AND SYSTEMATIC STATUS

Nisoniades lucilius Scudder & Burgess, 1870. [type-locality: New England]

Erynnis lucilius (Scudder & Burgess, 1870). [valid name (Miller & Brown, 1981:21)]

Thanaos lucilius; Holland, >898:333.

COMMON NAMES

Columbine Duskywing (Pyle, 1981:756)

Lucilius' Dusky-wing (Hollands 1931:351)

DISTRIBUTION

This northeastern species ranges from Quebec and Ontario to

New Jersey and Pennsylvania, thence westward to Minnesota. The very few Ohio records come from the northern and southeastern parts of the state (Fig. 26). Ohio is at the southern limit of this species.

BIOLOGY

Columbine (Aquilegia sp.) provides larval foodplants for this species (MacNeill, 1975:530). Authors report this species to be multivoltine, but Ohio records are too few to be able to state more than that it flies here in late spring and early summer (Fig. 27). 57

Erynnis baptisiae (Forbes, 1936) '

Wild Indigo Duskywing

NOMENCLATORIAL AND SYSTEMATIC STATUS

Thanaos baptisiae Forbes, 1936. [type-locality: Woods Hole, Mass.]

Erynnis baptisiae (Forbes, 1936). [valid name (Miller & Brown, 1981:21)]

COMMON NAMES

Wild Indigo Duskywing (Pyle, 1981:757)

Indigo Dusky Wing (Macy & Shepard, 1941:195)

DISTRIBUTION

From Massachusetts to northern Florida, westward to Illinois,

Nebraska, Kansas and Texas. Ohio records are limited to rather isolated areas in northern, eastern, and central parts of the state.

BIOLOGY

Wild indigos (Baptisia spp.) are the only known foodplants for the larvae of this species (MacNeill, 1975:530). Erynnis baptisiae has been reported to be multivoltine, and in Ohio flies from May into September (Fig. 29). 58

Erynnis persius (Scudder, 1864)

Persius Duskywing

NOMENCLATORIAL AND SYSTEMATIC STATUS

Nisoniades persius Scudder, 1864. [type-locality: New England]

Erynnis persius (Scudder, 1864). [valid name (Miller & Brown, 1981:21)]

Thanaos persius; Holland, 1898.

COMMON NAME

Persius Duskywing (Pyle, 1981:758)

DISTRIBUTION

This is a northern species extending from New England south in the Appalachian Mountains to Tennessee, and westward to Alaska and, in the mountains, New Mexico and Arizona. Ohio records are mostly from northern counties, with a single, old literature (Bales,

1909) record for Pickaway County.

BIOLOGY

; Burns (1964) reports that eastern populations of this species feed on species of Salix () and Populus. It is univoltine throughout its range according to MacNeill (1975:532). Ohio rec'ords, limited to May and June, suggest a single generation

(Fig. 31). 59

Pyrgus HUbner, 1819

NOMENCLATORIAL STATUS

Pyrgus HUbner. 1819. [type-species: Papilio alveolus HUbner, 1803, selected by Westwood, 1841.

Syricthus Boisduval, 1840. [misspelling of Syrichtus Boisduval, 1834, a subjective synonym]

Scelothrix Scudder, 1875. [misspelling of Scelotrix Rambur, 1858, a subjective synonym]

Bremeria Moore, 1896. [junior homonym of Bremeria Alpheraky, 1892]

Bremeria Tutt, 1906. [junior homonym of Bremeria Alpheraky, 1892]

Teleomorpha Warren, 1926. [junior objective synonym of Scelotrix Rambur, 1858]

SYSTEMATIC STATUS

Pyrgus HUbner, 1819. [valid name (Miller & Brown, 1981:22)]

Syrichtus Boisduval, 1834.

Scelotrix Rambur, 1858.

Hemiteleomorpha Warren, 1926.

Ateleomorpha Warren, 1926.

Heliopyrgus Herrera, 1957.

DIAGNOSIS

The black and white checkered wings distinguish these from all other skippers in the area covered by this report.

DISCUSSION

More than thirty species represent this genus in Eurasia and the New World, with about a dozen of these ranging from Alaska and

Labrador to Patagonia. Two species are found in the Ohio area. Key to the species of Pyrgus in the Ohio area.

1. Dorsal fore-wing with light-colored median spots elongate, at least twice as wide as high, in two complete rows (proximal row may lack some posterior spots) from anterior to posterior margins; outer margin of fore-wing with two or three complete rows of small, white spots, marginal and submarginal (visible at least ventrally)...... communis

1'. Dorsal fore-wing with median spots subquadrate, proximal row weak or incomplete toward posterior margin, distal row complete but sinuate toward anterior margin; submarginal rows of small spots absent or incomplete in some specimens...... 2

2(1'). Submarginal rows of small spots on fore-wing present, at least on ventral surface, but may lack some posterior spots (female)...... communis

2'. Submarginal spots as above lacking...... centaureae

Pyrgus centaureae (Rambur, 1840)

Grizzled Skipper

NOMENCLATORIAL AND SYSTEMATIC STATUS

Hesperia centaureae Rambur, 1840. [type-locality: "Andalusia"]

Pyrgus centaureae (Rambur, 1840). [valid name (Miller & Brown, 1981:22)]

Hesperia centaureae; Holland, 1898.

COMMON NAMES

Grizzled Skipper (Klots, 1951:215; Holland, 1931:340)

Alpine Checkered Skipper (Pyle, 1981:759)

DIAGNOSIS

This, and the following species, are characterized in the key above. 61

DISTRIBUTION

Pyrgus centaureae is a holarctic species with distribution

across Canada and northern United States. It is found as far south

as North Carolina in the East and Colorado in the West, at higher

elevations. In Ohio it has been reported only from ridge tops in

the unglaciated Appalachian plateau area of the southeastern part

of the state (Pig. 32). Ohio is near the southern limit of this

species.

BIOLOGY

The early stages of this species are unknown (MacNeill, 1975:

515; Ferris and Brown, 1981:81). Adult flight is reported to occur

in May in eastern North America (MacNeill, 1975:515). Ohio records

occur in the last half of April and first week of May (Fig. 33).

Pyrgus communis (Grote, 1872)

Checkered Skipper

NOMENCLATORIAL STATUS

Syrichthus communis Grote, 1872. [type-locality: central Alabama]

Hesperia tessellata Scudder, 1872. [preoccupied by Hesperia tessellata Hewitson, 1866]

SYSTEMATIC STATUS

Pyrgus communis (Grote, 1872). [valid name (Pyle, 1981:762)]

Hesperia albovittata Grote, 1873.

Pyrgus insolatrix PlBtz, 1884. 62

Pyrgus varus PlUtz, 1884.

Hesperia montivaga; Holland, 1898 (in part).

Hesperia communis; Holland, 1931.

COMMON NAMES

Checkered Skipper (Mitchell & Zim, 1964:77; Klots, 1951:215)

Common Checkered Skipper (Pyle, 1981:762)

Common Checker-spot (Holland, 1931:342)

DIAGNOSIS

The key above characterizes this species.

DISTRIBUTION

This species is widely distributed from Canada to Argentina.

It has been found across much of Ohio except for a band of eastern and southern counties extending from Ashtabula in the Northeast to Brown in the Southwest (Fig. 34), which are mostly, but not entirely, in unglaciated parts of the state.

BIOLOGY

Larval foodplants are reported to include a number of genera of the Malvaceae. Ohio records indicate adult activity from May into October, suggesting two, possibly three overlapping generations (Fig. 35). 63

Pholisora Scudder, 1872

NOMENCLATORIAL AND SYSTEMATIC STATUS

Pholisora Scudder, 1872. [type-species: Hesperia catullus Fabricius, 1793, by original designation; valid name (Miller & Brown, 1981:25)]

Hesperopsis Dyar, 1905.

DIAGNOSIS

The even, dark brown to black color of these skippers, leads to the common name, the sooty wings. They are generally small in size with delicate bodies and weak flight. The labial palps are usually held prominently porrect.

DISCUSSION

This genus is North American in its distribution with from two to five species depending upon the authority. The single species found east of the Mississippi River is discussed below.

Pholisora catullus (Fabricius, 1793)

Common Sootywing

NOMENCLATORIAL AND SYSTEMATIC STATUS

Hesperia catullus Fabricius, 1793. [type-locality: "In Indiis"; probably Georgia (Miller & Brown, 1981:25)]

Pholisora catullus (Fabricius, 1793). [valid name (Miller & Brown, 1981:25)] 64

COMMON NAMES

Common Sootywing (Pyle, 1981:769)

Pigweed Skipper (Macy & Shepard, 1941:189)

Sooty-wing (Holland, 1931:345)

DIAGNOSIS

Small in size (wing span 22-30mm) with sooty black wings.

Dorsal fore-wing surface with tiny, white subapical and subterminal spots.

DISTRIBUTION

This species ranges across most of the United States excepting

Florida. Ohio records extend across much of the state (Fig. 36).

The apparent gaps in eastern and southern counties may result from incomplete collecting.

BIOLOGY

Authors list various species of Chenopodiaceae and

Amaranthaceae as larval foodplants for this species. MacNeill

(1975:509) states that it may be bivoltine, with first adult activity in April in the northern part of its range. Ohio records extend from late April continuously to the end of September, suggesting the occurrence of more than one generation (Fig. 37). 65

HESPERIINAE Latreille, 1809.

These are the so-called branded or tawny skippers. An androconial

macule, made up of clusters of androconial scales, is on the dorsal

fore-wing surface of the males of many species. These macules, called

stigmata (singular stigma) by many authors, give the common name

"branded skippers" to this group. The stigma provides a diagnostic

character in some taxa. The macules usually appear as elongate, dark-

colored spots or patches centrad on the dorsal fore-wing surface.

The HESPERIINAE lack the costal fore-wing fold of some males in the

PYRGINAE. The tawny yellow or fulvous brown color found on the wings

of many species of this group leads to the other common name, the

"tawny skippers". The male hind tibia lacks the hair tufts foiind in

some PYRGINAE.

This taxon is separated from the other two nearctic subfamilies in

the key under the family discussion above. Additionally, it should be noted that the HESPERIINAE are generally smaller than the PYRGINAE as

adults. Wing span size ranges will be given with the species accounts below. The larvae of the HESPERIINAE usually feed on grasses, those

of the PYRGINAE feed mostly on dicotyledonous plants (MacNeill, 1975).

At rest adults of the HESPERIINAE often hold the fore-wings diagonally upward, and the hind-wings horizontally.

Species of this subfamily are widespread throughout America north of Mexico, and are well-represented in Ohio. 66

The HESPERIINAE were divided into by tribes by Clark (1948) who

closely followed the "groups" of Lindsey, Bell and Williams (1931).

Clark's classification has not been repeated in any major publication

since his paper, and since a revision of the group is well beyond the

scope of this work, the "groups" of Lindsey, Bell and Williams (1931),

also used by Macy and Shepard (1941), and by Klots (1951), will be

followed here.

Key to the groups of the HESPERIINAE in North America.

1. Antennal club with apiculus...... 2.

1'. Antennal club lacking apiculus...... 3.

2(1). Fore-wing discal cell less than 2/3 wing length; fore-wing lacking recurrent vein; male usually with discal stigma...... Group C.

2'. Fore-wing discal cell about 2/3 wing length; fore-wing with recurrent vein, at least as vestige; male lacking stigma...... Group D.

3(1'). Labial palp porrect; hind-wing discal cell much

longer than 1/2 wing length; fore-wing vein M 2 straight at base and arising midway between M^ and Group A.

3'. Labial palp not porrect, being upturned, oblique, or appressed to face; hind-wing discal cell 1/2 wing length

or shorter; fore-wing vein M 2 curved at base and arising closer to M^ than M^ ...... 4.

4(3'). Labial palp upturned, 3rd segment long and slender...... Group B.

4'. Labial palp not as above...... Group C.

1 67

Group A.

Carterocephalus Lederer, 1852 is the sole genus of this group

found in eastern North America (Klots, 1951). This Holarctic genus

is represented in the Ohio area by a single species, Carterocephalus

palaemon (, 1771), which occurs across the continent in the

northern United States and Canada, from Nova Scotia to Washington

(state) and Alaska, including northern Michigan and parts of

Pennsylvania (MacNeill, 1975). Although commonly known as the

Arctic Skipper, Klots (1951) states that it is characteristic of

Canadian Zone forested areas. Probably it is not to be expected in

Ohio.

Group B.

These skippers have been characterized in the key to the groups

of the HESPERIINAE above. To this may be added that they are all

relatively small (wing span 13-32mm) compared to other HESPERIINAE

(hence the common name for the group, the lesser skippers), and that

they exhibit relatively weak flight behavior. The long, slender

terminal segment of the labial palp is needlelike in appearance.

The antenna is short (approximately 0.4 the length of the fore-wing) with a flattened club, and either lacks arl apiculus or has only a

tiny, pointed terminal segment. 68

Key to the genera of Group B in the eastern United States

(except Florida).

1. Wings broadly rounded, especially outer margin of fore­ wing and tornus of hind-wing; mesotibia unspined...... Ancyloxypha.

1'. All angles of wings well-marked, wings not broadly rounded, rather somewhat triangular; mesotibia spined...... 2.

2(1'). Antennal club very long, equal to more than h length of shaft; wings with dorsal surfaces mostly dark, with limited fulvous color in costal area of fore-wing, near wing base, or along some veins...... Oarisma.

2'. Antennal club short, equal to less than % length of shaft; dorsal surfaces of wings mostly fulvous with limited areas of contrasting darker color...... 3.

3(2'). Hind-wing with dorsal margin darker than and contrasting with central fulvous area...... Thymelicus.

3'. Hind-wing with dorsal surface almost completely fulvous without contrasting, darker margins around entire wing...... . 69

Ancyloxypha Felder, 1862

NOMENCLATORIAL STATUS

Ancyloxypha Felder, 1862. [type-species: Hesperia numltor Fabricius, 1797, by original designation]

Ancyloxipha Auctorum; Dos Passos, 1964:14. [misspelling of Ancyloxypha Felder, 1862]

SYSTEMATIC STATUS

Ancyloxypha Felder, 1862. [valid name (Miller & Brown, 1981:30)]

DIAGNOSIS

These are small (wing span under 30mm), delicate skippers with

long and slender bodies. The protibia, mesotibia, and metatibia are

spineless. The terminal segment of the labial palp is about the same

length as the protibia. These skippers are mostly orange and the male

is without a stigma. The antenna has a greatly reduced apiculus,

consisting of a tiny, pointed terminal segment.

DISCUSSION

MacNeill (1975:497) reports that this genus contains seven species,

all in the Western Hemisphere, and ranging from Canada to Patagonia.

Only a single species, discussed below, is found east of the

Mississippi River. 70

Ancyloxypha numitor (Fabricius, 1793)

Least Skipper

NOMENCLATORIAL STATUS

Hesperia numitor Fabricius, 1793. [type-locality: "In Indiis"]

SYSTEMATIC STATUS

Ancyloxypha numitor (Fabricius, 1793). [valid name (Miller & Brown, 1981:30)]

Hesperia bion Fabricius, 1798.

Thymelicus puer Hllbner, [1823].

Heteropterus marginatus Harris, 1862.

Ancyloxypha longleyi French, 1897.

DIAGNOSIS

Wing span 19-26mm. Dorsal wing surface mostly dark brown to black,

except discal area of hind wing, which is fulvous. Ventral fore-wing mostly black. Ventral hind-wing uniformly bright yellow.

DISTRIBUTION

From Nova Scotia and Quebec to Florida, and westward to Saskatchewan

and Texas. Ohio records indicate probable distribution throughout the

state (Fig. 38).

BIOLOGY

Larvae have been reported to feed on various grasses including

Marsh Millet, cultivated rice, and (MacNeill, 1975:498). Adults fly 71 in moist, grassy areas, particularly in ditches and along ponds and streams. This species appears to be multivoltine throughout most of its range, with three generations reported for New England and four in

Texas (Klots, 1951:231; MacNeill, 1975:498). Ohio records extend from

May into September, with peaks in early June, late July, and late

August, indicating possibly three generations (Fig. 39). 72

Oarisma Scudder, 1872

NOMENCLATORIAL STATUS

Oarisma Scudder, 1872:75. [type-species: Hesperia powesheik Parker, 1870:271, by original designation]

SYSTEMATIC STATUS

Oarisma Scudder, 1872:75. [valid name (Evans, 1955:305)]

Paradopaea Godman, [1900].

DIAGNOSIS

Wing span 19-32mm. Protibia and mesotibia spined. Antennal club long, nearly as long as the pre-club shaft. Antenna lacking apiculus.

Males of North American (north of Mexico) species without stigma.

DISCUSSION

Nine species in this New World genus range from Canada south­ ward to northern South America, and into the . A single species, Oarisma powesheik (Parker, 1870) (characterized in the key to genera bove), has been reported east of the Mississippi River.

It is not known from Ohio or any of its immediate neighbors. 73

Copaeodes Speyer, 1877

NOMENCLATORIAL STATUS

Copaeodes Speyer, 1877. [type-species: Heteropterus procris Edwards, 1871:215, selected by Kirby [1879]:139]

SYSTEMATIC STATUS

Copaeodes Speyer, 1877. [valid name (Hemming, 1967:125)]

DIAGNOSIS

Wing span 13-22mm. Mesotibia spined, protibia and metatibia without spines. Antennal club lacking apiculus. Male with long, slender stigma.

Wing yellow-orange to orange.

DISCUSSION

This mainly Neotropical genus embraces three species, one of which is found east of the Mississippi River in America north of

Mexico ( (Edwards, 1870)), but not in the Ohio area.

Copaeodes minima is characterized in the key to genera above. 74

Thymelicus Hllbner, [1819]

NOMENCLATORIAL STATUS

Thymelicus HUbner, [1819]. [type-species: Papilio acteon Rottemburg, 1775:30, selected by Butler, 1870:94]

Pelion Kirby, 1858. [unavailable because of private printing, and junior objective synonym of Adopoea Billberg, 1820 (Hemming, 1967:349)]

Thymelinus Stephens, 1875:405. [misspelling]

Adopaea Scudder, 1875 (not McDunnough, 1938). [misspelling of Adopoea Billberg, 1820]

SYSTEMATIC STATUS

Thymelicus HUbner, [1819]. [valid name (Hemming, 1967:442)]

Adopoea Billberg, 1820. [subjective synonym (Dos Passos, 1964:13)]

DIAGNOSIS

Mesotibia spined, protibia and metatibia spineless. Fore-wing of male with small, slender stigma. Antenna lacking apiculus.

DISCUSSION

This genus was formerly palearctic, and is made up of eight species.

One of these species, Themelicus lineola (Ochsenheimer, 1808), was introduced into North America within the past 100 years, and is now well established in Ohio.

Thymelicus lineola (Ochsenheimer, 1808)

European Skipper

NOMENCLATORIAL STATUS

Papilio lineola Ochsenheimer, 1808. [type-locality: Europe] 75

SYSTEMATIC STATUS

Thymelicus lineola (Ochsenheimer, 1808). [valid name (Miller & Brown, 1981:32)]

Papilio virgula HUbner, [1813].

Pamphila ludoviciae Mabille, 1883.

Adopaea lineola; Holland, 1931:366.

COMMON NAMES

European Skipper (Klots, 1951:232; Pyle, 1981:783)

New English Skipper (Holland, 1931:366)

DIAGNOSIS

Wing span 19-26mm. Dorsal fore-wing with distal ends of veins darkened. Dorsal fore-wing in male with stigma approximately central in position, and parallel to costal edge of wing, thereby unlike any other butterfly in the Ohio area.

DISTRIBUTION

This species is found over much of the Palearctic Region, and in the New World from New Brunswick south to , and west through Ontario to Michigan, Illinois and Kentucky. It is also established in British Columbia (Burns, 1966). It was found first in Ohio in 1927 (Findlay, Hancock Co.) by George W. Rawson (Rawson,

1931). The next report for Ohio was for a colony found in Columbus in 1952 (Thomas, 1952). Ohio records now indicate distribution over much of the northern part of the state (Fig. 40). Its distribution in North America has been reviewed by Bums (1966). BIOLOGY

Various authors report timothy (Phleum. pratense) as the preferred larval foodplant, with other grasses mentioned as used occasionally. Adults are found flying in grassy areas including roadsides and meadows. MacNeill (1975:493) reports North American adult flight in June, July, and possibly into August. Ohio records confirm this with two records in August, and expand the time period a bit with two records in the last third of May (Fig. 41). Higgins and Riley (1970) report adult flight in Great Britain and Europe to be from May to August in a-single generation. 77

Group C.

Male of most species with stigma. Female of many species noticeably darker than corresponding male, with dark-brown replacing much of the light, tawny or orange-brown. This is the largest group of hesperiine skippers in America north of Mexico. Most of the hesperiine genera and species known from Ohio and surrounding areas are included in this group.

Key to the genera of Group C in eastern North America.

1. Antenna with portion of nudum on apiculus equal to or less than portion on club; antenna with club constricted before apiculus, or apiculus absent...... '...... 3.

11. Antenna with portion of nudum on apiculus greater than portion on club; club not constricted before apiculus...... 2.

2(1'). Wings with ventral surface mostly yellow-brown or dull yellow-brown, especially hind-wing; apiculus consisting of eight segments...... Nastra.

2'. Wings with ventral surface dark brown, hind-wing often with violet or light brown bands, but not with yellow- brown; apiculus consisting of nine segments...... Lerema.

3(1'). Antenna distinctly longer than width of thorax...... 4.

3'. Antenna relatively short, scarcely longer than width of thorax; ventral hind-wing bright yellow with many small, dark brown spots...... Hylephila.

4(3). Mesotibia spined...... 7.

4'. Mesotibia without spines, or if present short, covered with scales or scarcely protruding...... 5. 78

5(4'). Labial palp with distal (third) segment long, extending clearly beyond scaling of second segment...... .

5'. Labial palp with distal (third) segment extending only slightly beyond scaling of second segment...... 6.

6(5')’. Antenna with nudum less than 14 segments; metatibia with short spines between two pairs of spurs...... Atrytone.

6'. Antenna with nudum greater than or equal to 14 segments...... Euphyes.

7(4) . Antenna with apiculus at least as long as thickness of club, slender, and abruptly constricted, or if not abruptly constricted then longer than thickness of club...... 8.

7’. Antenna with apiculus shorter than thickness of club, or if about same length as thickness then not sharply constricted..12.

8(7). Apiculus slender, about twice as long as thickness of club; ventral hind-wing brownish-black with three, distinct light-colored median spots...... Oligoria.

8'. Apiculus shorter than above, or thick; if ventral hind- wing brownish-black then with more than three, or lacking median spots...... 9.

9(8’). Mesotibia with prominent spines...... 10.

9'. Mesotibia smooth, or with short, inconspicuous spines (Florida only)...... Choranthus.

10(9). Fore-wing with vein much nearer to Cu^ than to wing base...... 11.

10' Fore-wing with vein almost intermediate between Cu^ and wing base; fore-wing with dorsal surface dark brown, with three small subapical white spots, one large spot posterior to these, and another smaller one at end of discal cell; fore-wing with ventral surface brown suffused with gray...... Atrytonopsis. 79

11(10). Antenna with apiculus long and slender, length exceeding diameter of club; wing with ventral surface red or blackish, sometimes with small white spots; male stigma with two oval spots, one on either

11'. Apiculus length about equal to diameter of club, tapered, not abruptly constricted; wing with ventral surface with bright yellow areas, or brownish black with large white markings behind inner angle of fore­ wing; male without stigma, or with stigma slender and hardly visible...... Poanes.

12(7'). Fore-wing apically produced, hind-wing lobed; apiculus very short, length about equal to 1/3club width...... 13.

12'. Fore- and hind-wing moderate, not as above; apiculus usually moderately long, length greater than or equal to % width of club...... 14.

13(12). Apiculus of four segments; male stigma slender with light-colored central streak; hind-wing of female less strongly lobed than that of male...... Hesperia.

13'. Apiculus of seven segments; male stigma a large blotch, not colored as above; hind-wing of female much like that of male...... Atalopedes.

14(12'). Hind-wing dorsal surface with little fulvous color, at most a transverse band; apiculus of four segments or less; male with stigma...... 15.

14'. Dorsal hind-wing with large amount of fulvous color; apiculus of six segments; male with or without stigma...... 16.

15(14). Antenna with apiculus of 6-7 segments...... Pompeius.

15'. Antenna with apiculus of 4-5 segments...... Polites.

16(14'). Fore-wing vein M 2 well curved toward base, arising much nearer M^ than M^; color tawny or blackish...... 17.

16'. Vein M 2 as above only slightly curved, arising only slightly nearer M^ than M^; color blackish or grayish- brown , never tawny...... 18. 80

17(16). Antennal club length equals about 1/3 shaft length; labial palp with terminal segment hidden by scales of second (subapical); mesotibia spined...... Poanes.

17'. Antennal club length equals about 1/4 shaft length; labial palp with terminal segment extending free of scales of second; mesotibia smooth or with short, inconspicuous spines (Florida only)...... Choranthus.

18(16'). Antenna with apiculus fine, shorter than thickness of club; fore-wing produced apically; male without stigma..Lerodea.

18'. Apiculus longer than thickness of club; male with stigma...... Amblyscirtes. Nastra Evans, 1955

NOMENCLATORIAL STATUS

Nastra Evans, 1955:125. [type-species: Hesperia lherminier Latreille, [1824], by original designation]

SYSTEMATIC STATUS

Nastra Evans, 1955:125. [valid name (Dos Passos, 1964:16)]

DIAGNOSIS

In addition to the characters used in the key to genera of this group (above) the apiculus is longer than the width of the antennal club, and the male is without a stigma.

DISCUSSION

Most of the approximately 12 species in this genus are neotropical

(MacNeill, 1975:502). Three of these are found east of the Mississippi

River, with one, Nastra lherminier (Latreille, [1824]), in Ohio.

Nastra lherminier (Latreille, [1824])

Swarthy Skipper

NOMENCLATORIAL STATUS

Hesperia lherminier Latreille, [1824]. [type-locality: Carolina]

SYSTEMATIC STATUS

Nastra lherminier (Latreille, [1824]). [valid name (Miller & Brown, 1981:28)]

Hesperia fusca Grote & Robinson, 1867.

Megistias lherminieri; Holland, 1931:395.

Lerodea lherminieri; Klots, 1951:266.

Pamphila lherminieri; Forbes, 1960:89. 82

COMMON NAMES

Swarthy Skipper (Klots, 1951:266; Pyle, 1981:774)

Fuscous Skipper (Holland, 1931:395)

DIAGNOSIS

Small, wing span 22-26mm. Wings brown to brownish gray above,

yellowish brown below, frequently with veins of ventral hind-wing a

paler yellow than the adjacent cell color. Wings without spots both

dorsally and ventrally, although the dorsal fore-wing surface sometimes

has poorly defined pale areas which may appear to be spots.

DISTRIBUTION

From New York to Florida, west to Missouri and Texas. MacNeill

(1975:502) questions the validity of records for Mexico and South

America reported by Macy and Shepard (1941). Ohio records (Fig. 42 ),

although sparse, suggest that this species may be found throughout much

of the state, which is near the northern limit of this species.

BIOLOGY

The larval foodplant for this species is scoparius

(MacNeill, 1975:502). Adults may be found in old fields and other open areas. This species is reported to have two generations in the northern part of its range, with adult flight occurring from May to September.

Ohio records (Fig. 43) extend from early June into September, with a single record in mid-October. The length of this flight period in Ohio, alone, suggests that two generations may occur. 83

Lerema Scudder, 1872

NOMENCLATORIAL STATUS

Lerema Scudder, 1872. [type-species: Papilio accius Smith, 1797, by original designation]

SYSTEMATIC STATUS

Lerema Scudder, 1872. [valid name (Miller & Brown, 1981:29)]

Sarega Mabille, 1904.

DIAGNOSIS

Antenna relatively long, about equal to % fore-wing, and with a relatively long apiculus. Upper wing surface mainly dark brown, with limited white spots on the dorsal fore-wing surface. Mesotibia spined, protibia and metatibia spineless or weakly spined. Labial palp slender with short, conical third segment. Male with curved, broad stigma.

DISCUSSION

Eight species are included in this mainly neotropical genus (Evans,

1955:162), with a single species, Lerema accius (Smith, 1797), in the area covered by this study.

Lerema accius (Smith, 1797)

Clouded Skipper

NOMENCLATORIAL STATUS

Papilio accius Smith, 1797. [type-locality: Georgia]

SYSTEMATIC STATUS

Lerema accius (Smith, 1797). [valid name (Miller & Brown, 1981:29)]

Hesperia monoco Scudder, 1864. 84

COMMON NAMES

Clouded Skipper (Klots, 1951:260; Pyle, 1981:777)

Accius Skipper (Holland, 1931:396)

DIAGNOSIS

Dorsal fore-wing dark brown with a short series of three, white,

subapical spots running at right angle away from costal edge. Male with stigma, also usually with very small, white spot in discal cell,

and an additional spot distal to end of discal cell. Female dorsal fore­ wing with similar maculation, but with spots larger. Ventral hind-wing both sexes with three bands of light brown or violet-gray parallel to

outer margin. Wing span 26-36mm.

DISTRIBUTION

From New England south to Florida, west to Illinois and Texas,

thence southward through to northern South America.

This species has not been recorded from Ohio, but has been found in

Illinois (Irwin & Downey, 1973), Kentucky (Covell, 1974), Pennsylvania

(Clench, 1971), and Indiana (Shull & Badger, 1972). Although it appears

to be rather scarce in the northern part of its range (Klots, 1951:261;

MacNeill, 1975:500) it is a distinct possibility for capture in Ohio. 85

BIOLOGY

Several grasses have been reported as food for this species including

Wooly Beard Grass (Erianthus alopecuroides) • Authors report that adults frequent open, grassy glades and meadows, flying close to the ground.

Klots (1951:261) reports two generations northward. Shull & Badger

(1972) report a July 3rd date in Indiana, and Irwin & Downey (1973) report an August 26th date in Illinois. 86

Hylephila Billberg, 1820

NOMENCLATORIAL STATUS

Hylephila Billberg, 1820. [type-species: Papilio phyleus Drury, [1773], selected by Scudder, 1875:193]

Euthymus Scudder, 1872. [junior homonym of Euthymus Gray, 1867 (Evans, 1955:310)]

SYSTEMATIC STATUS

Hylephila Billberg, 1820. [valid name (MacNeill, 1975:489)]

Andinus Hayward, 1940.

Cordillana Hayward, 1941.

DIAGNOSIS

Antenna very short, about 1/3 fore-wing length. Apiculus rudimentary, with length equal to or less than width of club. All tibiae spined. Male dorsal fore-wing usually tawny, and usually with a prominent stigma.

DISCUSSION

Eleven species comprise this mainly neotropical genus, with but a single species in North America, Hylephila phyleus (Drury, 1773).

Hylephila phyleus (Drury, 1773)

Fiery Skipper

NOMENCLATORIAL STATUS

Papilio phyleus Drury, 1773. [type-locality: "Antigua, Nevis, etc."]

Hylephila phlaeus; Holland, 1898:354; Holland, 1931:377. [misspelling] SYSTEMATIC STATUS

Hylephila phyleus (Drury, 1773). [valid name (Miller & Brown, 1981:33)]

Papilio phareus Panzer, 1785.

Hesperia carin Hilbner, 1823.

Pamphila bucephalus Stephens, 1828.

Pamphila hala Butler, 1870.

COMMON NAME

Fiery Skipper (Pyle, 1981:784; Holland, 1931:377)

DIAGNOSIS

The very short antennae are distinctive for this species, in

America north of Mexico (MacNeill, 1975:489).

DISTRIBUTION

Widely distributed, extending from Connecticut southward and westward to California, Central America, and South America to Chile.

Authors report it to be common southward, rarer in the northern part of its range. Ohio records are limited to the northern 2] 3 of the state

(Fig. 44).

BIOLOGY

Larvae are reported to feed on a wide variety of grasses including sugarcane, Bermuda Grass () and bent grass ( sp.).

Adults are found flying in grassy areas, in clearings and along wooded edges. Authors report this species to have two or more generations.

Ohio records begin in late July, skip August, and then extend from

September through October (Fig. 45). This may indicate two generations, but more data are needed. Hesperia Fabricius, 1793

NOMENCLATORIAL STATUS

Hesperia Fabricius, 1793. [type-species: Papilio comma Linnaeus, 1758:484, selected by Dalman, 1816:200]

Pamphila Fabricius, 1807. [objective synonym (Hemming, 1964:332)]

Hesperia Htlbner, 1819. [junior homonym (Hemming, 1967:216)]

Symmachia Sodovskii, 1837. [objective synonym (Hemming, 1964:421)]

Hespena Edwards, 1872. [misspelling]

Urbicola Tutt, 1905. [junior objective synonym (Hemming, 1967:454)]

SYSTEMATIC STATUS

Hesperia Fabricius, 1793. [valid name (Dos Passos, 1964:11)]

Ocytes Scudder, 1872.

Anthomaster Scudder, 1872.

DIAGNOSIS

All tibiae spined, metatibia with two pairs of spurs. Antenna

length about % fore-wing length. Apiculus short, length about 1/3 club

width. Labial palp with third segment conspicuously pointed. Male

stigma with a silver colored line in the middle, distinctive for this

genus. In some species the tufted, brush-like "eyelash" at the outer

antennal base is used in diagnosis.

DISCUSSION

This holarctic genus has 16 to 18 species, depending on the author who lists them. All are found in the New World ranging from Labrador

and Alaska southward to central Mexico. Five of these are found in the

Ohio area, and are discussed below. This group has been studied

extensively by Lindsey (1942) and MacNeill (1964). 89

Key to species of Hesperia in the Ohio area.

1. Dorsal wing surface mostly dark brown, or olive- brown; ventral hind-wing with light yellow or whitish V-shaped (pointing distad) limbal band contrasting strongly with dark brown discal area...... metea.

I1. Dorsal wing surface mostly yellow-orange; or ventral hind-wing not as above, but if yellow spots in limbal area appear as band, then the contrasting discal area is dark, reddish-brown...... 2.

2(1'). Ventral hind-wing with light colored spots, sometimes combined into band and clearly contrasting with darker surrounding area...... 3.

2'. Ventral hind-wing without distinct spots as above; if present they are combined into band and contrast only weakly with surrounding area, and often extend distad along veins...... 4.

3(2). Ventral hind-wing dark reddish-brown with contrasting, pale yellow spots...... :.... leonardus.

3'. Ventral hind-wing yellow-orange or olive-brown, spots sometimes very small [some male specimens without spots key out below]...... attalus.

4(2'). Ventral hind-wing with yellow band in limbal area, only weakly contrasting with surrounding area, and with yellow color often extending distad along veins...... sassacus.

4'. Ventral hind-wing not as above...... 5.

5(4'). Male dorsal fore-wing largely dark brown, with dark brown coloration extending along at least part of costal edge proximal to outer angle, and covering at least half of area between stigma and outer edge; male [females and some males key out above]...... attalus.

5'. Male dorsal fore-wing largely yellow-orange, with edge proximal to outer angle yellowish, and dark brown coloration covering much less than half of area between stigma and outer edge; female ventral hind- wing without light, contrasting spots, although very pale spots may be present...... ottoe. 90

Hesperia ottoe Edwards, 1866

Ottoe Skipper

NOMENCLATORIAL STATUS

Hesperia ottoe Edwards, 1866. [type-locality: Kansas]

SYSTEMATIC STATUS

Hesperia ottoe Edwards, 1866. [valid name (MacNeill, 1975:471)]

Erynnis ottoe; Holland, 1931:374.

Pamphila ottoe; Forbes, 1960:93.

COMMON NAME

Ottoe Skipper (Klots, 1951:239; Pyle, 1981:790)

DISTRIBUTION

This prairie species is found from South Dakota to Iowa, and westward to Colorado and Texas. An isolated, relict population has been well documented in southern Michigan (MacNeill, 1975:471). This species has not been reported in 0,hio, but it should be looked for in prairie remnants in western and northwestern parts of the state.

BIOLOGY

Several grasses including Fall Witchgrass (Leptoloma cognatum) have been reported by authors as larval food plants. Pyle (1981:791) reports the habitat for this species to be undisturbed prairies, rangeland oak scrub and swamps, and nearby roads. Adult flight is reported to be from June into August. 91

Hesperia leonardus Harris, 1862

Leonardus Skipper

NOMENCLATORIAL STATUS

Hesperia leonardus Harris, 1862. [type-locality: Massachusetts]

SYSTEMATIC STATUS

Hesperia leonardus Harris, 1862. [valid name (Miller & Brown, 1981:36)]

Erynnis leonardus; Holland, 1931:376.

Pamphila leonardus; Forbes, 1960:94.

COMMON NAMES

Leonardus Skipper (Klots, 1951:237; Pyle, 1981:791)

Leonard's Skipper (Holland, 1931:376; Macy & Shepard, 1941:211)

DIAGNOSIS

Wing span 22-36mm. This is the only skipper in the area covered by this study in which the ventral hind-wing is brick red (some authors

call this red, or rust in males and chestnut in females) with prominent,

limbal whitish or yellow spots.

DISTRIBUTION

From Ontario and Nova Scotia south to South Carolina, thence westward to Minnesota, Missouri and Louisiana. Ohio records, while not extensive, come from all of the major parts of the state (Fig. 46).

BIOLOGY

Larvae feed on grasses with preference for bent grass (Agrostis), panic grass (), and tumble grass (Eragrostis) (MacNeill, 1975:

472; Pyle, 1981:791). Found in open fields, wet meadows and oak 92 openings. Price (1970) found adults to be often abundant at flowers of blazing star (Liatris sp.) in the Oak Openings Park in Lucas County, and on Purple Boneset () at Mud Lake in Williams

County. Adult flight is reported by authors to occur in a single generation during August and September. Ohio records (Fig. 47) confirm this.

Hesperia metea Scudder, 1864

Cobweb Skipper

NOMENCALTORIAL STATUS

Hesperia metea Scudder, 1864. [type-locality: Connecticut]

SYSTEMATIC STATUS

Hesperia metea Scudder, 1864. [valid name (MacNeill, 1975:474)]

Erynnis metea; Holland, 1931:375.

COMMON NAME

Cobweb Skipper (Klots, 1951:238; Pyle, 1981:793)

DIAGNOSIS

Wing span 24-36mm. As characterized in the key to species of

Hesperia above, this is the only skipper in the area of this study with a combination of mostly dark brown or olive-brown dorsal fore-wing, and a light yellow or whitish V-shaped limbal band on the ventral hind- wing. Veins of ventral hind-wing at least partly white. "Eyelash" long, usually more than half the greatest eye diameter. 93

DISTRIBUTION

From to Florida, westward to Minnesota and Texas. In

Ohio it has been found on an open ridge top in eastern Vinton County

in the Lake Hope area. Also, it has been collected in Athens County

(Fig. 48). Hesperia metea has been reported for Kentucky (Covell,

1974:254), Michigan (Moore, 1960:29), Pennsylvania (Clench, 1971),

and Indiana (Shull & Badger, 1972).

BIOLOGY

Grasses provide the larval food plants for this species. MacNeill

(1975:475) and Pyle (1981:794) give Bluestem Beard Grass (Andropogon

scoparius), also known as Little Bluestem, as the preferred food.

Weishaupt (1967:169) describes this as a dominant of some large areas of original grasslands in the United States, often in drier sites

than the Big Bluestem () with distribution throughout most of Ohio. Pyle (1981:794) describes the habitat for Hesperia metea as fields and clearings, hillsides, rocky sites, limestone outcrops, burns and barrens. MacNeill (1975:475) describes this skipper as a transient species in areas where the larval foodplant, a fire pioneer following a burning, is replaced as succession proceeds. In barren areas where soil conditions limit some competitive plants, Andropogon scoparius may form a more stable part of the flora and Hesperia metea may be a permanent resident. This skipper, then, may be expected to be found with more thorough collecting in other areas within Ohio, in at least some of the areas where the foodplant exists. However, 94

since it has been found only recently (1979) in Ohio for the first

time, and in only two limited areas, there may very well be other

limiting factors, presently unknown, which affect this species'

geographical distribution in Ohio. Authors report adult flight in

May in areas around Ohio. Both Ohio records are from May (Fig. 49).

Hesperia attalus (Edwards, 1871)

Dotted Skipper

NOMENCLATORIAL STATUS

Pamphila attalus Edwards, 1871. [type-locality: Waco, Texas]

SYSTEMATIC STATUS

Hesperia attalus (Edwards, 1871). [valid name (Miller & Brown, 1981:36)]

Ocytes seminole Scudder, 1871:76.

Erynnis quaiapen Scudder, 1889:1655.

Erynnis attalus; Holland, 1931:374.

COMMON NAMES

Dotted Skipper (Klots, 1951:239; Pyle, 1981:794)

Attalus Skipper (Holland, 1931:374; Macy & Shepard, 1941:212)

DIAGNOSIS

Intraspecific variation makes this species difficult to

characterize. The key to species of the genus Hesperia above should

suffice for specimens from the Ohio area. Additionally, the veins

of the ventral hind-wing are dark. The "eyelash" is short, less

than half the greatest eye diameter. 95

DISTRIBUTION

Hesperia attalus is found from Massachusetts south to Florida, west to Wisconsin, Nebraska and Texas. A single record for Ohio is from Cuyahoga County (Janus, 1973). The origin of Klots' (1951:239) reference to Ohio is not known. It may have come from Macy & Shepard

(1941:213). This species probably is a stray in Ohio, which is near the northern edge of the range.

BIOLOGY

Larval foodplant unknown. Pyle (1981:794) reports this to be

an "apparent sand barrens dweller." Klots (1951:239) reports that

this species is not well known. Adults in June in Nebraska (Klots,

1951:239). Hesperia sassacus Harris, 1862

Indian Skipper

NOMENCLATORIAL STATUS

Hesperia sassacus Harris, 1862. [type-locality: Massachusetts]

SYSTEMATIC STATUS

Hesperia sassacus Harris, 1862. [valid name (MacNeill, 1975:478)]

Erynnis sassacus; Holland, 1931:373.

Pamphila sassacus; Forbes, 1960:93.

COMMON NAMES

Indian Skipper (Comstock & Comstock, 1904:276; Klots, 1951:238; Pyle 1981:797)

Sassacus Skipper (Holland, 1931:373)

DIAGNOSIS

In addition to the characters used in the key above, the

"eyelash" is long, usually at least two-thirds the greatest eye diameter.

DISTRIBUTION

From Maine and southern Ontario south to Virginia and Tennessee west to Wisconsin and Iowa. In Ohio it has been found in the north­ eastern and central parts of the state (Fig. 50). It is known to occur in Pennsylvania (Clench, 1971), West Virginia (Drees & Butler,

1978), Michigan (Moore, 1960:29), and Indiana (Shull & Badger, 1972)

It has not been reported from Kentucky (Covell, 1974) and is known from but a single record in Illinois (Irwin & Downey, 1973:10).

Further collecting in Ohio may expand the known range. 97

However, Pyle's (1981:797) remark that acid soil scrub is a habitat

type for this species suggests a factor, the acidity of the soil, which may limit the distribution of this species within Ohio.

Generally, the soils of eastern Ohio, including those of eastern Lake

Plains, Glaciated Appalachian Plateau, and the Unglaciated

Appalachian Plateau regions are acidic with occasional neutral or mildly alkaline areas. The soils of western Ohio, on the other hand, are usually neutral to alkaline, with some acid areas (Holowaychuclc,

1979). If one compares the Ohio distribution map for Hesperia sassacus (Fig. 50) with Holowaychuck's (op. cit.) map for Ohio soil regions, a very close correspondence can be seen between this skipper's distribution and the largely acid soils of eastern Ohio.

Further study is required to determine the significance of this relationship, and other states should be included.

BIOLOGY

Authors report several grasses as larval foodplants for this species.

Included are prairie grass (Panicum), fescue (Festuca), and Crabgrass

(Digitaria sanguinalis). Pyle (1981: 797) states that preferred habitats for this species include dry, old fields, acid soil scrub, and damp meadows and roadsides. As suggested in the distribution discussion above, it may be possible that these habitats are limited to acid soil areas.

Adult flight is reported to occur in a single generation from May into

July in the northern part of the range of this skipper. Ohio records

(Fig. 51) extend from mid-May into the first week of July. 98

Polites Scudder, 1872

NOMENCLATORIAL STATUS

Polites Scudder, 1872. [type-species: Hesperia peckius Kirby, 1837, by original designation]

Pyrrhosydia Evans, 1955. [misspelling of Phrrhosidia Scudder, 1874, a subjective synonym of Polites Scudder, 1872]

SYSTEMATIC STATUS

Polites Scudder, 1872. [valid name (Miller & Brown, 1981:37)]

Hedone Scudder, 1872.

Limochores Scudder, 1872.

Pyrrhosidia Scudder, 1874.

Talides of authors (not Htlbner, 1819.) ---

DIAGNOSIS

These are small to medium skippers (wing span 20-32mm). Antenna slightly less than half the fore-wing length, with club between 1./3 and % the shaft length. Apiculus short, four or fewer segments. All tibiae spined. Labial palp with third segment short, barely protruding beyond scales of second segment. Male stigma prominent, often sigmoid.

DISCUSSION

Ten species are included in this New World genus which ranges from

Canada to Argentina. Five of these, described below, are known from the

Ohio area. Grasses provide the larval food. Records for sedges and dicotyledonous plants are oviposition, rather than feeding, records according to Stanford (1981:117). 99

Key to the species of Polites in the Ohio area.

1. Ventral hind-wing with yellow or yellow-orange contributing at least 50% of the wing color...... 2.

1'. Ventral hind-wing without large areas of yellow or yellow-orange, generally uniformly dark, similar to dark coloration of ventral fore-wing; or with median row of faint yellow spots contributing far less than 50% of the wing coloration...... 4.

2(1). Ventral hind-wing with central yellow area contrasting strongly with a continuous darker area of costal, outer, and inner margins...... coras.

2'. Ventral hind-wing nearly entirely yellow to yellow- orange with at most a few, irregularly placed contrasting darker spots...... 3.

3(2'). Ventral hind-wing with postmedian yellow area contrasting somewhat with slightly darker yellow- orange of rest of wing...... mystic.

3'. Ventral hind-wing with lighter coloration uniformly yellow-orange...... vibex.

4(1'). Male with stigma consisting of three parts; female with ventral hind-wing uniform in color, and with costal area of dorsal fore-wing usually orange or yellow-orange...... themistocles.

4'. Male stigma consisting of two parts; female with coastal area of dorsal fore-wing same color as rest of wing, and with the ventral hind-wing usually showing a short, median row of lighter colored, faintly contrasting spots...... origenes. 100

Polites coras (Cramer, 1775)

Peck's Skipper

NOMENCLATORIAL STATUS

Papilio coras Cramer, 1775. [type-locality: "Surinam"]

Polites packius Tietz, 1972:509. [misspelling]

SYSTEMATIC STATUS

Polites coras (Cramer, 1771). [valid name (Miller & Brown, 1981:37)]

Hesperia peckius Kirby, 1837:300.

Hesperia wamsutta Harris, 1862:318.

Pamphila peckius; Forbes, 1960:^5

COMMON NAMES

Peck's Skipper (Holland, 1931:382: Klots, 1951:246)

Yellow Spot (Comstock & Comstock, 1904:282)

Yellowpatch Skipper (Pyle, 1981:800)

DIAGNOSIS

Wing span 19-26mm. Ventral hind-wing surface with basal and post-median yellow bands, each consisting of large, rectangular spots.

DISTRIBUTION

Eastern Canada south to Georgia, west to British Columbia, Oregon and Arizona. Ohio records (Fig. 52) indicate statewide distribution.

BIOLOGY

Larvae feed on grasses () and have been reared on a variety of grasses in the laboratory, but the preferred species in nature are not known (MacNeill, 1975:483; Pyle, 1981:800). These skippers are 101

usually seen in open, grassy areas along roadsides and in marshes and

pastures. Authors report multiple, overlapping generations from May

to September, with one or two generations in the northern part of the

range. Ohio records (Fig. 53) extend from mid-May to the last third

of September, with peaks in late May/early June, and again in August.

Polites themistocles (Latreille, 1823)

Tawny-edged Skipper

NOMENCLATORIAL STATUS

Papilio taumas Fabricius, 1787:84. [nomen oblitum; Miller & Brown, 1981:213 favor suppression of this senior synonym. Such action is possible under Article 23(b) of the Code if, in fact, the name has remained unused in the primary zoological literature for more than 50 years. Action of the ICZN would be required]

Papilio Fabricius, 1793:327. [preoccupied by Papilio thaumas Hufnagel, 1766:62]

Hesperia phocion Fabricius, 1798:431. [preoccupied by Hesperia phocion Fabricius, 1793:335]

Hesperia themistocles Latreille, 1823. [type-locality: "Amerique meridionale"]

SYSTEMATIC STATUS

Polites themistocles (Latreille, 1823). [valid name (Miller & Brown, 1981:38)]

Hesperia c e m e s Boisduval & LeConte, 1834:pl. 76.

Hesperia ahaton Harris, 1862:316.

Limochores taumas; Holland, 1898:357.

Polites taumas; Holland, 1931:381.

Pamphila themistocles; Forbes, 1960:95. 102

COMMON NAME

Tawny-edged Skipper (Holland, 1931:381; Pyle, 1981:804)

DIAGNOSIS

.Wing span 19-26mm. See diagnosis under Polites origenes below

for further discussion.

DISTRIBUTION

From Nova Scotia to Florida, west to British Columbia and

California. As with the preceding species, Polites coras; this has

been collected widely in Ohio and probably is to be found throughout

the state (Fig. 54).

BIOLOGY

Authors report grasses, including Panicum, as larval foodplants.

Adults are usually found flying in grassy areas. This is reported to be univoltine in the northern part of its range, and bivoltine in the

southern, with flight between May and August (Klots, 1951:246; MacNeill,

1975:486; Pyle, 1981:805). Ohio records (Fig. 55) extend from the middle third of May through most of September, with one record in late

October. A break occurs in late June/early July, and two peaks occur, one in early June and a second in August. Considering as well the overall length of the flight period in-Ohio, spanning over five months, two generations are indicated. 103

Polites origenes (Fabricius, 1793)

Crossline Skipper

NOMENCLATORIAL STATUS

Hesperia origenes Fabricius, 1793. [type-locality: probably New York (Miller & Brown, 1981:38)]

SYSTEMATIC STATUS

Polites origenes (Fabricius, 1793). [valid name (Miller & Brown, 1981: 38)]

Hesperia manataaqua Scudder, 1864.

Limochores manataaqua; Holland, 1898:357.

Polites manataaqua; Klots, 1951:245.

COMMON NAME

Crossline Skipper (Pyle, 1981:805)

DIAGNOSIS

Somewhat larger (wing span 25-28mm) than Polites themistocles

(wing span 19-26mm) which it resembles. Pyle (1981:806) states that

these two species, P_. themistocles and JP. origenes, cannot be separated without dissection of the male genitalia. However, in all of the

specimens I have examined from the Ohio area, the males are clearly

distinguishable by the structure of the fore-wing stigma. In Polites

themistocles the stigma consists of three parts, arranged in such a way as to appear sinuate or S-shaped. The latter appearance can be seen with

the unaided eye, while the actual three-part structure can be seen only with magnification. Polites origenes, on the other hand, has a stigma consisting of only two parts, lying end-to-end, and appearing to be straight rather than S-shaped. Several authors, including Klots (1951) 104 and MacNeill (1975) mention the gross appearance only of these stigmata.

I have examined over 50 specimens each of both species from the Ohio area, and have found these character states to be consistent. It is possible, of course, that populations from other parts of their ranges may show variations in the stigma.

DISTRIBUTION

Polites origenes ranges from New England to Georgia, and westward to Montana and New Mexico. Based on Ohio records, this species may be expected to be found throughout most of the state (Fig. 56).

BIOLOGY

Larvae of this species feed on grasses. Tridens flavus is mentioned

specifically by MacNeill (1975:487) and Pyle (1981:805) for Polites origenes. This grass is listed for Ohio (under the synonym Triodia

flava) by Weishaupt (1967:92) as common in old fields and along roadsides. Pyle (1981:805) states that this skipper is univoltine in the northern part of its range with adult flight from May into August, and bivoltine with a longer flight period in the South. Ohio records

(Fig. 57) extend from early June through the last third of August.

This is longer than might be expected for a single generation, but the data available are insufficient to determine whether there is a second generation in Ohio. 105

Polites mystic (Edwards, 1864)

Long Dash

NOMENCLATORIAL STATUS

Hesperia mystic Edwards, 1864. [type-locality: White Mountains, N.H.]

SYSTEMATIC STATUS

Polites mystic (Edwards, 1964). [valid name (Miller & Brown, 1981:38)]

Thymelicus mystic; Holland, 1898:351.

COMMON NAME

Long Dash (Klots, 1951:247; Pyle, 1981:806)

DIAGNOSIS

In addition to characters used in the key to species above, the male dorsal fore-wing has the stigma connecting with the narrow, dark brown marking which extends toward the outer angle. The effect of this is to give the appearance of a very long stigma, much longer in proportion to the wing than in other hesperiine skippers. Wing span 25-32mm.

DISTRIBUTION

From New England south to Virginia, west to Colorado, Washington, and British Columbia. Present in northern Indiana (Shull & Badger,

1972:15), northern Illinois (Irvin & Downey, 1973:9), Michigan (Moore,

1960:31), and Pennsylvania (Clench, 1971). Ohio records (Fig. 58) are limited to the northern part of the state, as are those from Indiana and Illinois. Since this species is not known from West Virginia

(Drees & Butler, 1978) or Kentucky (Covell, 1974), it may be inferred that Ohio is at or very near the southern extent of this species in the area covered by this study. 106

BIOLOGY

Various grasses (Poaceae), including Poa (bluegrass) serve as

larval foodplants (MacNeill, 1975:487; Pyle, 1981:806).

Adults fly in grassy areas associated with water, such as stream banks,

marshes and wet meadows. Authors report that eastern populations are

univoltine in the North (May-July) and bivoltine in the South (May to

September). Ohio records (Fig. 59) extend from May to early July, with

a peak in early/mid-June, in what appears to be a single generation.

This, then, seems to place Ohio in the northern part of the range.

Polites vibex (Geyer, 1832)

Whirlabout

NOMENCLATORIAL STATUS

Thymelicus vibex Geyer, 1832. [type-locality: West Indies]

SYSTEMATIC STATUS

Polites vibex (Geyer, 1831). [valid name (Miller & Brown, 1981:39)]

Hesperia brettus Boisduval & Leconte, 1834.

Hesperia wingina Scudder, 1864.

Hesperia osyka Edwards, 1867.

Hesperia morganta PlBtz, 1883.

Hesperia unna PlHtz, 1883.

Thymelicus brettus; Holland, 1898.

COMMON NAMES

Whirlabout (Klots, 1951:247; Pyle, 1981:807)

Geyer's Skipper (Holland, 1931:384) 107

DIAGNOSIS

Wing span 25-32mm. Otherwise characterized in key above.

DISTRIBUTION

From Virgnia (occasionally in Connecticut) west to Arkansas and

Arizona, south to the West Indies and Argentina. In Ohio this species has been reported from Hamilton County in the southwestern part of the state, and from three counties in the northeastern part of the state

(Fig. 60). Of the neighboring states, only Pennsylvania has been reported to have this species (Clench, 1971). Ohio appears to be at the northern limit of this species.

BIOLOGY

Various grasses, including Paspalum, have been reported as larval foodplants. Adults fly in open, grassy areas, both wet and dry. In

Ohio adult flight has been reported from August and September (Fig. 61).

Since Pyle (1981:808) reports peak flight to occur in mid-summer, it appears that Ohio does not have a resident population, only occasional strays. 108 Wallengrenia Berg, 1897

NOMENCLATORIAL AND SYSTEMATIC STATUS

Wallengrenia Berg, 1897. [type-species: Hesperia premnas Wallengren, 1860, by original designation; valid name (Dos Passos, 1964:9)]

Catia Godman, 1901. [subjective synonym]

DIAGNOSIS

In North America the males of this genus may be separated from other hesperiine genera by the stigma which exists as two distinct, black spots separated by large gray scales, on either side of vein near its base. Both male and female can be separated from Polites by the long, slender apiculus which exceeds in length the diameter of the antennal club.

DISCUSSION

From Canada to Argentina, and on many of the Antillean islands

Brown & Heineman, 1972). A single species, Wallengrenia egeremet, is found in Ohio.

Wallengrenia egeremet (Scudder, 1864)

Northern Broken Dash

NOMENCLATORIAL STATUS

Hesperia egeremet Scudder, 1864. [type-locality: Massachusetts]

SYSTEMATIC STATUS

Wallengrenia egeremet (Scudder, 1864). [valid name (Miller & Brown, 1981:40)]

Hedone aetna Scudder, 1872 (nec Boisduval, 1840).

Pamphila ursa Worthington, 1880.

Hesperia cinna PlBtz, 1882. 109

Catia otho var. egeremet; Holland, 1931.

Wallengrenia otho egeremet; Klots, 1951.

COMMON NAMES

Northern Broken Dash (Pyle, 1981:809)

Broken Dash (in part, Klots, 1951:248)

Volcanic Skipper (Macy & Shepard, 1941:221)

Otho's Skipper (in part, Holland, 1931:386)

DIAGNOSIS

Wing span 24-32mm.

DISTRIBUTION

From Quebec and Ontario south to Florida, west to Minnesota and

Texas. This species is known from all states neighboring Ohio, and is

found throughout much of Ohio (Fig. 62).

BIOLOGY

Larval foodplants are grasses, including panic grass (Panicum).

Found in open, grassy areas, including shrub edges and old field.

Univoltine in the North, bivoltine in the South (Pyle, 1981:809).

Ohio records extend from May into September, with most in June and

July (Fig. 63), probably with a single generation. 110 Pompeius Evans, 1955

NOMENCLATORIAL STATUS

Pompeius Evans, 1955:335. [type-species: Hesperia pompeius Latreille, 1824:765, by original designation]

SYSTEMATIC STATUS

Pompeius Evans, 1955:335. [valid name (Dos Passos, 1964:9)]

DIAGNOSIS

Antenna with club length equal to 1/4 entire shaft length, apiculus

length equal to club width (in closely related Polites equal to % club

width or less); nudum extends to six or seven club segments and an equal

number on the apiculus (thereby differing from Polites in which nudum

is found on six or eight club segments, but is limited to four or five

segments of the apiculus). Male with well-marked, elongate, three-

part stigma.

DISCUSSION

Evans (1955:301) lists seven species in this genus, ranging from the

United States to Argentina. Only one species, Pompeius v e m a , is found

in the United States. It occurs in the area covered by this study, and

is discussed below.

Pompeius verna (Edwards, 1862)

Little Glassywing

NOMENCLATORIAL STATUS

Pamphila verna Edwards, 1862. [type-locality: Illinois] Ill

SYSTEMATIC STATUS

Pompeius verna (Edwards, 1862). [valid name: Miller & Brown, 1981:40)]

Pamphila pottawattomie Worthington, 1880:50.

Hesperia vetulina PlBtz, 1883:58.

Telesto sigida Mabille, 1891:177.

Euphyes verna; Holland, 1898:360.

Polites verna; Holland, 1931:380.

Polites sequoyah Freeman, 1942:104.

COMMON NAME

Little Glassywing (Klots, 1951:245; Pyle, 1981: 809)

DIAGNOSIS

Wing span 24-32mm. Fore-wing with relatively large, rectangular, whitish spot posterior to distal end of discal cell; nearly square in females, more elongate in males and lying parallel to stigma. This spot is visible on both dorsal and ventral surfaces. Hind-wing dark brown to blackish on both surfaces, usually marked with weakly defined spots which are slightly lighter in color than the background.

DISTRIBUTION

From New England southward to Georgia, westward to Nebraska and

Texas. Evans (1955:336) includes Mexico in its distribution. Ohio records indicate distribution throught the state (Fig. 64). Ohio is near the northern limit of this species. 112

BIOLOGY

Larvae feed on grasses, specifically Tridens flavus (MacNeill,

1975:480). Authors report this species to have one or more annual generations, depending on latitude, with adult flight between April and August. Ohio records (Fig. 65) are from June to late-August, with a break in late-July/early-August, suggesting the possibility of a second generation. 113 Atalopedes Scudder, 1872

NOMENCLATORIAL STATUS

Atalopedes Scudder, 1872. [type-species: Hesperia huron Edwards, 1863:16, by original designation]

SYSTEMATIC STATUS

Atalopedes Scudder, 1872:78. [valid name (Miller & Brown, 1981:40)]

Pansidia Scudder, 1872.

DIAGNOSIS

Antenna with apiculus of seven segments, nudum on seven club segments and seven apicular segments. Fore-wing of male with greatly enlarged stigma, whose width is greater than length. Females similar to Hesperia in general appearance, although the apiculus is longer, exceeding in length % width of club. All tibiae spined.

DISCUSSION

Three species are listed for this genus by Evans (1955), only one of which is known from the United States. The other two are found in the

Bahamas and the Greater Antilles (MacNeill, 1975:462).

Atalopedes campestris (Boisduval, 1852)

Sachem

NOMENCLATORIAL STATUS

Hesperia campestris Boisduval, 1852. [type-locality: California]

SYSTEMATIC STATUS

Atalopedes campestris (Boisduval, 1852). [valid name (Miller & Brown, 1981:41)]

Hesperia sylvanoides Boisduval, 1852 (female only).

Hesperia huron Edwards, 1863. 114 Pamphila kedema Butler, 1870.

Hesperia amphissa PlHtz, 1883.

Atalopedes huron; Scudder, 1889.

Pamphila campestris; Hine, 1898.

Atalopedes campestris; Godman, 1900.

Hylephila campestris; Comstock & Comstock, 1904.

COMMON NAMES

Sachem (Holland, 1931:385; Pyle, 1981:799)

Field Skipper (Mitchell & Zim, 1964:79)

DIAGNOSIS

Fore-wing of male with stigma very large, length subequal to length

of discal cell, stigmal width at distal end exceeding % stigmal length

(all other male skippers in area covered by this report with stigma much smaller, not as above). Fore-wing of female with distinctive, hyaline spot just posterior to distal end of discal cell. Wing span

25-35mm.

DISTRIBUTION

From east coast to west coast across southern half of United States where it overwinters, thence southward through Central and South America to and Ecuador. Records from northern states, from New York to

Colorado are from non-overwintering adults which stray northward during a given season (MacNeill, 1975:463). In Ohio, this species has been found across the state from southern to northern counties (Fig. 66)-

BIOLOGY

Larvae feed on Cynodon dactylon (Bermuda Grass) and Stenotaphrum seeundatum (St. Agustine Grass) according to Klots (1951:243) and 115

MacNeill (1975:463). Klots (1959:243) reported that overwintering probably occurs in the adult stage. Ohio records are from May, and from July through October (Fig. 67). The May records, which are from

Adams and Hocking counties in the southern part of the state, may represent an overwintering generation, or may be individuals which have flown into Ohio from overwintering sites further southward. Both

MacNeill (1975:463) and Pyle (1981:800) describe this skipper as a resident of the southern third to half of the United States, which moves into more northern areas including New York. New York records are from immigrants from July to October (Shapiro, 1974:25). Records from Pennsylvania are of the same period, July to October (Tietz,

1952:20). Indiana records are from July to September (Shull & Badger,

1972:15). Records from Illinois, however, include May (as does Ohio) as well as late summer and autumn according to Irwin and Downey

(J973:9), who believe that the May record is "probably overwintered individuals." Therefore, it seems likely that the May records in Ohio represent overwintering adults, while the later represent immigrants from the South or a second local generation. More data are needed concerning the regularity of May activity in Ohio. 116 Atrytone Scudder, 1872

NOMENCLATORIAL STATUS

Atrytone Scudder, 1872. [type species: Hesperia iowa Scudder, 1869:336, by original designation]

SYSTEMATIC STATUS

Atrytone Scudder, 1872. [valid name (Miller & Brown, 1981:41)]

Anatrytone Dyar, 1905.

DIAGNOSIS

Pro- and metatibia always spined, although often weakly. Mesotibia

either smooth or with short, scale-covered spines. Terminal segment of

labial palp stout, conical, protruding only slightly beyond scales of

second segment. Male without stigma. Antennal club relatively long,

equal in length to about % remaining flagellum.

DISCUSSION

Evans (1955) reports two species for this New World genus, ranging

from eastern United States to . Only one of these is known

from Ohio and surrounding states, and is discussed below.

Atrytone logan (Edwards, 1863)

Delaware Skipper

NOMENCLATORIAL STATUS

Hesperia logan Edwards, 1863. [type-locality: Lansing, Michigan]

SYSTEMATIC STATUS

Atrytone logan (Edwards, 1863). [valid name (Miller & Brown, 1981:41)]

Hesperia delaware Edwards, 1863:19.

Atrytone logan; Scudder, 1889.

Atrytone delaware; Holland, 1898. 117 Pamphila delaware; Hine, 1898.

Phycanassa vitellius; Comstock & Comstock, 1904.

COMMON NAMES

Delaware Skipper (Pyle, 1981:811)

Vitellius Skipper (Comstock & Comstock, 1904:288)

DIAGNOSIS

Dorsal fore-wing with veins conspicuously darker than yellow-orange central area, dark line marking distal end of discal cell. Outer margin of dorsal fore-wing in male dark brown, female with wider dark outer margin and with basal area mostly dark brown. Ventral hind-wing immaculate, orange to yellow-orange.

DISTRIBUTION

From Massachusetts to Florida, west to the Dakotas, Texas and northern Mexico. Most Ohio records are from the northern-most counties, a few from the central and southern part of the state (Fig. 68).

BIOLOGY

Larvae feed on various grasses. Klots (1951) reports adult flight in March in Florida, and late June in Massachusetts, with one generation in the North and up to three in the South. Ohio records extend from mid-June into early August without any indication of more than a single. generation (Fig. 69). 118 Problema Skinner & Williams, 1924

NOMENCLATORIAL STATUS

Problema Skinner & Williams, 1924. [type-species: Pamphila byssus Edwards, 1880:224, by original designation]

SYSTEMATIC STATUS

Problema Skinner & Williams, 1924. [valid name: Miller & Brown, 1981]

DIAGNOSIS

Mesotibia lacking spines, protibia usually spineless. Terminal

segment of labial palp relatively long, protruding beyond scales of

second segment. Male without stigma. Relatively large for hesperiine

skippers (wing span 32-38mm).

DISCUSSION

Two species are known from the United States east of the Mississippi

River, (Edwards, 1880) and (Boisduval

& Leconte, 1833). Both of these are southern; neither has been reported from Ohio or its immediate neighbors, although the former is known from

Illinois (Irwin & Downey, 1973). 119

Poanes Scudder, 1872

NOMENCLATORIAL STATUS

Poanes Scudder, 1872. [type-species: Hesperia massasolt Scudder, 1863: 171, by original designation]

Paratrytone Dyar, 1905. [homonym of Paratrytone Godman, 1900; subjective synonym of Poanes Scudder, 1872 (Dos Passos, 1964:7)]

SYSTEMATIC STATUS

Poanes Scudder, 1872. [valid name (Miller & Brown, 1981:42)]

DIAGNOSIS

Labial palp shaggy, semi-porrect, third segment mostly concealed by hairs of second segment. Tibiae of all legs spined. Antenna long, equal in length to about ^ costal margin of fore-wing. Antennal club long, equal in length to about 1/3 entire flagellum. Apiculus about equal in length to width of club. Male stigma absent in most species.

DISCUSSION

Eleven species ranging from the United States to Argentina in this New World genus (Evans, 1955). Four species in this genus are in the area covered by this study.

Key to the species of Poanes in the Ohio area.

1. Ventral hind-wing with clear, yellow areas contrasting with rest of wing...... 2.

1'. Ventral hind-wing not as above, at most with small yellow spots...... 4.

2(1). Ventral fore-wing with more than 75% of area in yellow or yellow-orange...... 3.

2'. Ventral fore-wing mostly dark, little or no yellow to yellow-orange massasoit ■ 120

3(2). Ventral hind-wing with solid, continous dark margin, contrasting with lighter, central yellow area, extending from outer angle around outer and inner margins to base of wing...... hobomok.

3'. Ventral hind-wing not as above, margin from outer angle to base with irregularly contrasting color, part of which is only slightly darker than central yellow area (male)...... zabulon.

4(1). Dorsal hind-wing with distinctly contrasting yellow-orange area...... viator.

4'. Dorsal hind-wing entirely dark, with small, light spots, or with indistinct or faint markings, not as above (female)...... 5.

5(4'). Dorsal fore-wing with light-colored spot in discal cell, at least faintly contrasting with ground color (female)...... hobomok form pocahontas.

5'. Dorsal fore-wing with discal cell immaculate; rest of wing may have distinct light-colored spots...... 6.

6(5'). Dorsal wing surface immaculate (female)..massasoit form suffusa.

6'. Dorsal wing surface with light-colored spots (female)..... zabulon.

Poanes massasoit (Scudder, 1864)

Mulberry Wing

NOMENCLATORIAL STATUS

Hesperia massasoit Scudder, 1864. [type-locality: "Carver, Massachusetts; Connecticut; New England"]

SYSTEMATIC STATUS

Poanes massasoit (Scudder, 1864). [valid name (Miller & Brown, 1981:43)]

Pamphila massasoit; Worthington, 1880.

Pamphila suffusa Laurent, 1892.

DIAGNOSIS

Wing span 24-28mm. Characterized in key above. 121

DISTRIBUTION

From middle New England south to Maryland, questionably to Georgia

(MacNeill, 1975), westward through Ontario to Minnesota and Nebraska.

Ohio records limited to a few counties in western half of state (Fig. 70).

DISCUSSION

Occurs in bogs and marshes, adults often found at flowers of

Swamp Milkweed () (Klots, 1951). Larvae feed on

Carex sp. (MacNeill, 1975). In Ohio adults have been captured in late

June and July (Fig. 71) , in Michigan from July 3 to August 19 (Moore,

1960), in Illinois from late June to mid August (Irwin & Downey, 1973),

and in Indiana in July (Shull & Badger, 1972). Considering these dates

together it appears that there is a single, annual generation in the area

including Ohio and neighboring states.

Poanes hobomok (Harris, 1862)

Hobomok Skipper

NOMENCLATORIAL STATUS

Hesperia hobomok Harris, 1862. [type-locality: Massachusetts]

Pamphila quadraquina Scudder, 1868. [misspelling of Hesperia.quadaquina Scudder, 1868, a subjective synonym of Poanes hobomok (Harris, 1862)]

Atrytone zabulon Holland, 1898:364. [misidentification (Holland, 1931: 390)] 122

SYSTEMATIC STATUS

Poanes hobomok (Harris, 1862). [valid name (Miller & Brown, 1981:43)]

Hesperia pocahontas Scudder, 1864. [female form]

Hesperia quadaquina Scudder, 1868.

Pamphila hobomok; Hine, 1898.

Poanes ridingsii Chermock & Chermock, 1940.

COMMON NAMES

Hobomok Skipper (Pyle, 1981:817)

Mormon (Comstock & Comstock, 1904:275)

DIAGNOSIS

Ventral hind-wing with large, median, yellow patch surrounded on three sides (outer margin, inner margin and basal area) by darker, brownish areas; yellow area not marked with darker spots. Dorsal hind-wing with large, yellow-orange median color patch, usually with thin, dark line crossing discal cell. Dark form of female resembles female of Poanes zabulon, but always has at least a trace of yellow in dorsal fore-wing discal cell (not in zabulon), and dorsal fore-wing yellow markings more diffuse (sharply limited in zabulon). Ventral wing surface with continuous purplish-brown border (discontinuous in zabulon). Wing span 26-35mm.

DISTRIBUTION

From southeastern Canada to Georgia and westward to Saskatchewan,

Kansas, Arkansas and Alabama. Ohio records from most of state (Fig. 72). 123

BIOLOGY

MacNeill (1975) refers to this as one of the more common skippers in the northeastern United States. Larvae feed on grasses. Klots (1951), and MacNeill (1975) refer to one generation annually, with adults flying from May to September. Pyle (1981:818) calls this an "extended brood."

Ohio records extend from April through most of September, with a strong peak in the last third of May and first third of June (Fig. 73). This is an unusually long flight period for a single generation of skippers, and may indicate a partial second generation. Poanes zabulon (Boisduval & LeConte, 1834)

Zabulon Skipper

NOMENCLATORIAL STATUS

Hesperia zabulon Boisduval & LeConte, 1834. ftype-locality: Georgia]

SYSTEMATIC STATUS

Poanes zabulon (Boisduval & LeConte, 1834). [valid name (Miller & Brown, 1981:43)]

Hesperia eratica PlHtz, 1883.

Pamphila zabulon; Hine, 1898.

Atrytone zabulon; Comstock & Comstock, 1904.

COMMON NAME

Zabulon Skipper (Holland, 1931:390; Pyle, 1981:818)

DIAGNOSIS

Compared with Poanes hobomok the ventral hind-wing of this species

in the male has more extensive yellow which interrupts the marginal,

darker border in most individuals. Unlike hobomok this yellow area

in zabulon is usually broken by a row of small, dark, red-brown spots

extending from the outer, submarginal area medially toward the inner margin. These spots are also usually visible on the female which, in

zabulon, has darker wings than the male. The female of Poanes zabulon has been compared with the dark-form female of Poanes hobomok in the discussion under hobomok above. Wing span 25-36mm.

DISTRIBUTION

More southern than Poanes hobomok, ranging from New England to

Georgia, westward to Wisconsin and Texas, and southward through Mexico 125 to Panama. Ohio records indicate a general distribution across the state (Fig. 74) .

BIOLOGY

Larvae of this species feed on grasses, with Tridens sp. and

Eragrostis specifically mentioned by MacNeill (1975). Klots (1951) reports two annual generations, possibly three in the southern part of the range, with adults flying in May and August. This appears to be confirmed by Ohio records which include May and June, and then August and September (Fig. 75). Like Poanes hobomok, zabulon is a common species in this area, found frequently in old fields and along wooded roads.

Poanes taxiles (Edwards, 1881)

Golden Skipper

Evans (1955) reports this species from Ohio, but this not confirmed in the literature or by museum specimens. As pointed out by

MacNeill (1975), this is a western species, and its occurence in Ohio is at the very least highly questionable.

Poanes viator (Edwards, 1865)

Broad-winged Skipper

NOMENCLATORIAL STATUS

Hesperia viator Edwards, 1865. [type-locality: N. Illinois and New Orleans] 126 SYSTEMATIC STATUS

Poanes viator (Edwards, 1865). [valid name (Miller & Brown, 1981:43)]

Pamphila viator; Worthington, 1880.

Phycanassa viator; Scudder, 1889.

Poanes viator; Lindsey, Bell & Williams, 1931.

COMMON NAME

Broad-winged Skipper (Pyle, 1981:821)

DIAGNOSIS

This is the largest member of the genus in the area covered by

this study (wing span 30-44mm). Fore-wing of both sexes largely brown dorsally, with orange spots in males and similar whitish spots in females.

Large spot in fore-wing discal cell of both sexes. Male without stigma.

Dorsal surface of hind-wing in both sexes with dark line usually present across distal end of discal cell.

DISTRIBUTION

Described by both Klots (1951:249) and MacNeill (1975:456) as being local, in scattered and isolated colonies, but ranging widely from Massachusetts and Ontario southward to Florida, and westward to

Minnesota, Nebraska and Texas. Ohio records are limited to the northern fifth of the state (Fig. 76).

BIOLOGY

Larval foodplants include Marsh Millet () and wild rice (Zizania sp.). Habitat in coastal areas seems to be limited to brackish marshes, and in inland areas may be mostly associated with marshes (MacNeill, 1975). Ohio records are from July and early August

(Feg. 77) indicating a single generation. 127 Euphyes Scudder, 1872

NOMENCLATORIAL STATUS

Euphyes Scudder, 1872. [type-species: Hesperia metacomet Harris, 1862:317, by original designation]

SYSTEMATIC STATUS

Euphyes Scudder, 1872. [valid name (Miller & Brown, 1981:45]

Arotis Mabille, 1904.

Pereneia Lindsey, 1925.

DIAGNOSIS

Members of this genus are usually tawny to brown with lighter areas

of color on the dorsal wing surface. The ventral wing surface is often

more extensively colored with light yellow or orange-brown, and the

veins are distinctively lined with even lighter color in some species.

The male stigma in some species consists of two oval spots forming a

broken band. In others it is elongate and slender. The mesotibia is without spines.

DISTRIBUTION

According to Evans (1955) there are 21 species, all in the Western

Hemisphere, ranging from the United States to , and in the

West Indies. Five species are known from Ohio and neighboring states.

Most of the species are associated with marsh or swamp habitats

(MacNeill, 1975:447).

Key to the species of Euphyes in Ohio and surrounding states.

1. Ventral hind-wing surface immaculate...... 2.

1'. Ventral hind-wing surface with areas of contrasting light color, much broader than vein width...... 3. 128 2(1). Ventral hind-wing dark brown; male stigma about five times as long as wide...... vestris.

2'. Ventral hind-wing orange or yellow-orange, with underlying darker scales; stigma slender, length much greater than five times width...... bimacula.

3(1'). Ventral hind-wing with two, broad, light streaks from base toward outer margin, central and posterior; veins on ventral surface lighter in color than cells...... dion.

3'. Ventral hind-wing not with above combination of characters...... 4.

4(3'). Ventral hind-wing with large, medial, yellow- orange spot contrasting with darker remaining area; veins darker than cells...... conspicua.

4'. Ventral hind-wing not as above, uniformly tan to brown, with light-colored veins...... dukesi.

Euphyes dion (Edwards, 1879)

Dion Skipper

NOMENCLATORIAL STATUS

Pamphila dion Edwards, 1879. [type-locality: Indiana]

SYSTEMATIC STATUS

Euphyes dion (Edwards, 1879). [valid name (Miller & Brown, 1981:45)]

Pamphila dion; Hine, 1898.

Atrytone dion; Lindsey, Bell & Williams, 1931:117.

COMMON NAMES

Dion Skipper (Holland, 1931:388; Klots, 1951:255)

Sedge Skipper (Pyle, 1981:824)

DIAGNOSIS

Ventral wing surface of both sexes is distinctly marked with two light, yellowish-orange streaks extending from the base of the wing toward the outer margin. The anterior one runs across the center of 129 the wing, the posterior runs between the discal cell and the second

anal vein. These streaks contrast with darker, orange-brown color

of the rest of the wing. Veins appear as yellowish lines against the darker, orange-brown. Wing span 32-41mm.

DISTRIBUTION

Occurs throughout much of the eastern United States from Ontario and New York southward to Florida, westward to Wisconsin and Texas.

Ohio records are limited to the northern half of the state, with

Franklin County at the southern extent (Fig. 78).

BIOLOGY

Associated with marshy areas. Larval foodplants include Scirpus spp. and spp. (Klots, 1951; MacNeill, 1975). Univoltine, July-

August in New Jersey (Pyle, 1981:824), June-July in Ohio (Fig. 79).

Euphyes dukesi (Lindsey, 1923)

Duke's Skipper

NOMENCLATORIAL STATUS

Atrytone (Euphyes) dukesi Lindsey, 1923. [type-locality: Mobile Co., Alabama]

SYSTEMATIC STATUS

Euphyes dukesi (Lindsey, 1923). [valid name (Miller & Brown, 1981:45] '

COMMON NAMES

Duke's Skipper (Klots, 1951:255)

Scarce Swamp Skipper (Pyle, 1981:825) 130 DIAGNOSIS

This species is distinguished from Euphyes dion by having relatively shorter and more rounded fore-wings, and by the relatively less extensive light color on the dorsal surface of the fore-wing.

The subapical light spots present dorsally on the fore-wing of Euphyes dion are lacking in this species. Euphyes dukesi has contrasting light- colored streaks on the ventral surface of the hind-wing, similar to those on dion, but more yellowish than in the former. Wing span 32-

38mm.

DISTRIBUTION

Euphyes dukesi is very local in its distribution, and is known from a relatively few, widely distributed colonies in the eastern United

State (MacNeill, 1975). In the region around Ohio it has been reported from Illinois (Irwin & Downey, 1973), Indiana (Shull & Badger, 1972), and Michigan (Moore, 1960). Ohio records occur in north-central and north-western counties, and in Union County in central Ohio (Fig. 80).

BIOLOGY

Larval foodplants for this species are unknown. MacNeill (1975) reports adults flying from August to October in Alabama and during both spring and fall in Virginia. In states near Ohio, as cited above, flight has been reported for July in Michigan and Indiana, and August-

September in Illinois. Ohio records include June-August with majority in July (Fig. 81). Euphyes conspicua (Edwards, 1863)

Black Dash

NOMENCLATORIAL STATUS

Hesperia conspicua Edwards, 1863:17. [type-locality: Lansing, Mich.]

SYSTEMATIC STATUS

Euphyes conspicua (Edwards, 1863:17). [valid name (Miller & Brown, 1981 45)]

Hesperia pontiac Edwards, 1863.

Hedone orono Scudder, 1872.

Pamphila pontiac; Worthington, 1880.

Limochores pontiac; Scudder, 1889.

Atrytone pontiac; Holland, 1931.

Atrytone conspicua; Macy & Shepard, 1941.

Atrytone buchholzi Ehrlich & Gillham, 1951.

COMMON NAMES

Black Dash (Pyle, 1981:825)

Pontiac Skipper (Macy & Shepard, 1941:230)

DIAGNOSIS

Ventral hind-wing distinctly marked with large, medial spot whose light, yellow-orange color contrasts with the somewhat darker orange- brown of the rest of the wing. This spot is also visible on the dorsal hind-wing surface. Male stigma about four times as long as broad.

Wing span 25-36mm.

DISTRIBUTION

From New England to Virginia, west through Ontario and Michigan to

Minnesota and Nebraska. Ohio records are from northern and west-central 132 counties (Fig. 82) . This species is also known from Indiana (Shull &

Badger, 1972), Illinois (Irwin and Downey, 1973), and Pennsylvania

(Clench, 1971). It has not been reported from West Virginia (Drees &

Butler, 1978) or Kentucky (Covell, 1974). Ohio, therefore, appears to be at the southern limit of distribution.

BIOLOGY

Klots (1951) reports that there appears to be a single, annual generation with adults in July and August. Ohio records extend from

June to August (Fig. 83). Restricted to marshes and boggy meadows, adults favoring flowers of Buttonbush (Cephalanthus) and Swamp Milkweed

(Asclepias incarnata) (Klots, 1951; MacNeill, 1975). MacNeill (1975) reports Carex sp. as a larval foodplant.

Euphyes bimacula (Grote & Robinson, 1867)

Two-spotted Skipper

NOMENCLATORIAL STATUS

Hesperia bimacula Grote & Robinson, 1867. [type-locality: Atlantic District, Philadelphia]

SYSTEMATIC STATUS

Euphyes bimacula (Grote & Robinson, 1867). [valid name (MacNeill, 1975: 449)]

Hesperia acanootus Scudder, 1868.

Hesperia illinois Dodge, 1872.

Limochores bimacula; Scudder, 1889.

Pamphila bimacula; Hine, 1898.

Atrytone bimacula; Lindsey, Bell & Williams, 1931.

Atrytone contradicta Leussler, 1933. [aberrant form] 133 COMMON NAME

Two-spotted Skipper (Pyle, 1981:827)

DIAGNOSIS

Dorsal hind-wing without contrasting pale colored regions, similar

to Euphyes vestris, but unlike Euphyes dion, dukesi or conspicua. Veins

of ventral hind-wing light yellow, as in Euphyes dion, but unlike dion

there are no light colored, broad streaks contrasting with the rather

consistant, olive-orange to yellow color. Male stigma quite narrow

when compared with conspicua, having a width much less than one quarter

the length. Wing span 28-32mm.

DISTRIBUTION

Similar to Euphyes conspicua (see above). Ohio records include

northern, central, and southwestern counties (Fig. 84). Like conspicua

this is described as a "northern species" by both Klots (1951) and

MacNeill (1975). Both authors point out its local distribution and

preference for bogs and swamps.

BIOLOGY

Larvae feed on sedges (Carex) (Pyle, 1981:827). Univoltine, adult

flight in Ohio in June and July (Fig. 85).

Euphyes ruricola (Boisduval, 1852)

Dun Skipper

NOMENCLATORIAL STATUS

Hesperia ruricola Boisduval, 1852:315. [type-locality: California] 134 SYSTEMATIC STATUS

Euphyes ruricola (Boisduval, 1852:315). [valid name (MacNeill, 1975:450)]

Hesperia vestris Boisduval, 1852:317. [junior page priority to Hesperia ruricola Boisduval, 1852:315]

Hesperia metacomet Harris, 1862:317.

Pamphila rurea Edwards, 1862.

Hesperia kiowah Reakirt, 1866.

Hesperia osyka Edwards, 1867.

Pamphila osceola Lintner, 1878.

Pamphila califomica Mabille, 1883.

Euphyes metacomet; Scudder, 1889.

Pamphila ruricola; Smith, 1891.

Pamphila vestris; Smith, 1891.

Pamphila metacomet; Hine, 1898.

Pamphila vestris; Wright, 1905.

Atrytone vestris; Comstock, 1927.

Atrytone ruricola; Holland, 1931.

Euphyes vestris; Dos Passos, 1964.

COMMON NAME

Dun Skipper (Holland, 1931:388; Pyle, 1981:827)

DIAGNOSIS

Male with characeristically dark wings, lacking dorsal and ventral light markings. Stigma about one-fifth its length in width. Female wing often shows a few postmedian, light spots on dorsal surface.

Females of this species may be confused with those of some other species, but the lack of mesotibial spines in this species will help to separate them. Wing span 24-32mm. 135 DISTRIBUTION

Found through much of North America from Atlantic coast to

California, and from southern Canada to northern Mexico. Records exist for much of Ohio, except for eastern and southern counties (Fig. 86).

BIOLOGY

Common, roadside species, with grasses and sedges as larval foodplants. Univoltine (midsummer) in the North, bivoltine (spring and late summer) in the South (Pyle, 1981:828). Ohio records extend from

June to September, with most in July, The length of the flight period, with diminished activity toward the end, suggests a possible, partial second generation (Fig. 87). 136 Atrytonopsis Godman, 1900

NOMENCLATORIAL STATUS

Atrytonopsis Godman, 1900:497. [type-species: Hesperia deva Edwards, 1876:592]

SYSTEMATIC STATUS

Atrytonopsis Godman, 1900:497. [valid name (Miller & Brown, 1981:46)]

DIAGNOSIS

Dorsal wing surface dark brown, gray scaling on much of ventral wing surface. Male fore-wing usually produced apically, stigma when present long, slender, and gray. Unlike many hesperiine genera, wing spots often whitish.

DISCUSSION

Evans (1955) lists eight species in this genus, limited to North

America between the United States and . One, Atrytonopsis hianna (Scudder, 1868) is known from Ohio. Members of this genus are often encountered on rocky hillsides and boulder-strewn stream beds, although they may also be found at flowers (MacNeill, 1975).

Atrytonopsis hianna (Scudder, 1868)

Dusted Skipper

NOMENCLATORIAL STATUS

Hesperia hianna Scudder, 1868:382. [type locality: Quincy and Dorchester, Mass.]

SYSTEMATIC STATUS

Atrytonopsis hianna (Scudder, 1868:382). [valid name (Miller & Brown, 1981:46)]

Hesperia grotei Plfltz, 1883:54. 137 Lerema hianna; Scudder, 1889.

Atrytonopsis hianna; Godman, 1900:497.

Atrytomopsis tumeri Freeman, 1948.

COMMON NAME

Dusted Skipper (Pyle, 1981:829)

DIAGNOSIS

Variable species with cellular white spots present on fore-wing

of some females, but absent from most males and many females. Ventral

hind-wing marked with gray scales mentioned under generic discussion

above. Some specimens have a single, whitish spot anterior to the

discal cell of the hind-wing, most easily seen on the ventral surface.

Males without stigma. Wing span 32-36mm.

DISTRIBUTION

Most of the eastern United States from New England to Georgia, westward to Manitoba, Nebraska and Arkansas. In Ohio it is known only

from the "Oak Openings" area of Lucas County (Fig. 88).

BIOLOGY

Univoltine northward with adult flight in May and June. Ohio records from June (Fig. 89). MacNeill (1975) describes an interesting larval foodplant relationship presumed for this species: the apparent foodplant is Beard Grass (Andropogon scoparius) which is associated as a pioneer species with areas recently disturbed by fire, and is replaced after a few years with other plants. Price (1970) describes this skipper as rare; he says that Andropogon is common in the Oak

Openings. 138

Amblyscirtes Scudder, 1872

NOMENCLATORIAL STATUS

Amblyscirtes Scudder, 1872:75. [type-species: Hesperia vialis Edwards, 1862:58, by original designation]

Ephiphyes Evans, 1955:386. [misspelling of Epiphyes Dyar, 1905, a subjective synonym of Amblyscirtes]

SYSTEMATIC STATUS

Amblyscirtes Scudder, 1872:75. [valid name (Miller & Brown, 1981:47)]

Stomyles Scudder, 1872.

Mastor Godman, 1900.

Epiphyes Dyar, 1905.

DISCUSSION

This North American genus has 23 species found in the United States and Mexico (Evans, 1955), two of which are found in Ohio.

DIAGNOSIS

Generally small (wing span 19-32mm), dark brown or gray. Fore­ wing, and in some species the hind-wing, marked with small, white or pale orange spots. Ventral hind-wing in many species dusted with grayish scales. An unusual character of this genus, compared to other skippers, is found in the male genitalia: the saccus and aedoeagus are very long and extend cephalad nearly to the thorax (Evans, 1955). 139

Key to the species of Amblyscirtes in the Ohio area.

1. Dorsal fore-wing with pale markings limited to subapical group near costal martin; ventral hind- wing dusted with violet-gray scales distally, and free of light spots...... vialis.

1'. Dorsal fore-wing with pale markings extending from costal margin in irregular, subapical row more than 2/3 way to posterior margin; ventral hind-wing dusted with greenish or greenish-gray scales, and with postmedian row of light spots...... hegon.

Amblyscirtes hegon (Scudder, 1864)

Pepper-and-salt Skipper

NOMENCLATORIAL STATUS

Hesperia hegon Scudder, 1864:176. [type-locality: White Mts., New Hampshire]

SYSTEMATIC STATUS

Amblyscirtes hegon (Scudder, 1864:176). [valid name (Miller & Brown, 1981:48)]

Hesperia samoset Scudder, 1864:176. [Hesperia hegon is the valid name for this taxon by reason of line priority over H. samoset, and by selection of the first reviser, Kirby (1871:613)]

Hesperia nemoris Edwards, 1864. [this name postdates both of Scudder's names by several months (Brown, 1964:221) and is therefore a junior synonym of Hesperia hegon Scudder, 1864]

Pyrgus argina PlBtz, 1884:22.

Amblyscirtes samoset; Scudder, 1889.

Stomyles samoset; Draudt, 1924.

Amblyscirtes hegon; Lindsey, Bell & Williams, 1931:129.

COMMON NAMES

Pepper-and-salt Skipper (Klots, 1951:263; Pyle, 1981:834)

Samoset Skipper (Macy & Shepard, 1941:235) 140

DIAGNOSIS

Small (wing span 22-26mm), dorsal fore-wing black, with irregular row of submarginal, white spots. Basally, dorsal surfaces of both wings greenish-gray. Ventral wing surface dusted with greenish-gray scales. Male stigma well-developed, but inconspicuous because of surrounding dark coloration.

DISTRIBUTION

A northern species, from southeastern Canada south to Georgia in the mountains, west to Manitoba, Wisconsin, and Iowa. Ohio records limited to southcentral and southwestern counties (Fig. 90). MacNeill

(1975) calls this a forest species, usually found in glades and other openings. This may partially account for the southern Ohio records since the most extensively forested portions of the state are in the southeastern part at the present time. Prior to European settlement of the Ohio area, and the clearing of the native forests, this species may have been more widespread in Ohio. Amblyscirtes hegon was not reported from Ohio in the literature prior to 1900, so it is not possible now to know what the distribution was prior to man-made changes in the forests.

BIOLOGY

Larvae feed on grasses. Univoltine northward, bivoltine south­ ward. Univoltine in Ohio with adults in May and June (Fig. 91). 141

Amblyscirtes vialis (Edwards, 1862)

Roadside Skipper

NOMENCLATORIAL STATUS

Hesperia vialis Edwards, 1862:58. [type locality: Rock Island, 111.]

SYSTEMATIC STATUS

Amblyscirtes vialis (Edwards, 1862:58). [valid name (Miller & Brown, 1981:49)]

Cobalus asella Herrich-SchHffer, 1869.

COMMON NAME

Roadside Skipper (Klots, 1951:262; Pyle, 1981:836)

DIAGNOSIS

Distinguished by its combination of nearly unmarked dorsal wing

surface, dark in color, with a few small, white subapical spots, and

the violet-gray scaling of the apical part of the fore-wing and distal

half of the hind-wing. Outer margins of both wings, dorsal and ventral,

distinctly marked with alternating dark and light brown rectangles.

Male stigma small and inconspicuous. Wing span 22-26mm.

DISTRIBUTION

Transcontinental, known from every state in the United States

(except Alaska and Hawaii), as well as being widely distributed in

southern Canada. Ohio records from southern and northeastern counties

(Fig. 92). Generally more common than A. hegon (Klots, 1951).

BIOLOGY

Larval foodplants are grasses. Two or more generations in the

South, one in the North. Ohio records from late-April into August with an early-summer break, indicating two generations (Fig. 93). 142 Lerodea Scudder, 1872

NOMENCLATORIAL AND SYSTEMATIC STATUS

Lerodea Scudder, 1872:80. [type species: Hesperia eufala Edwards, 1869:311, by original designation; valid name (Millier & Brown, 1981:49)]

DIAGNOSIS

Male without stigma. Fore-tibia is spineless. Apiculus about equal in length to width of club. Labial palp appressed, third segment

short, stout, not extending beyond distal ends of scales of second.

DISCUSSION

Twenty species in this genus from the United States to Patagonia

(Evans, 1955). MacNeill (1975), however, says there may be as few as

15 species. Three species in the United States, one of which (Lerodea eufala) is found in the Ohio area.

Lerodea eufala (Edwards, 1869)

Eufala Skipper

NOMENCLATORIAL STATUS tiesperia eufala Edwards, 1869:311. [type-locality: Apalachicola, Florida]

SYSTEMATIC STATUS

Lerodea eufala (Edwards, 1869:311). [valid name (Miller & Brown, 1981:49)]

Goniloba dispersa Herrich-SchMffer, 1869:197.

Pamphila floridae Mabille, 1876.

Carystus micylla Burmeister, 1878:272.

Lerodea obscura Mabille, 1904.

Pamphila Wright, 1905. 143 COMMON NAME

Eufala Skipper (Pyle, 1981:839)

DIAGNOSIS

Distinguished from other skippers in this area by the 3-5 white, hyaline spots on the fore-wing, and the relatively even dusting of gray color on the hind-wing ventral surface. Antenna with constriction between club and apiculus.

DISTRIBUTION

Most of the southern United States from mid-Atlantic states to central California. In the Ohio area this skipper has been reported from Indiana (Shull & Badger, 1972) and Illinois (Irwin & Downey, 1973), but not in Ohio.

BIOLOGY

Multi-generations in the South, two in the North. Various grasses serve as larval food plants, and it has been reared on , sugar­ cane, and Sorghum species. 144 Oligoria Scudder, 1872

NOMENCLATORIAL AND SYSTEMATIC- STATUS

Oligoria Scudder, 1872:82. [type species: Hesperia maculata Edwards, 1865:202, by original designation; valid name (Miller & Brown, 1981:50)]

DIAGNOSIS

Tibiae of all legs spined, although the fore-tibia to a lesser extent than mid- and hind-tibia. Males lack stigma. Apiculus length greater than width of club.

DISCUSSION

Monotypic genus, known from Texas to Florida in the South, with strays northward to Massachusetts and New York. Oligoria maculata

(Edwards, 1865) from Georgia and Florida westward to Louisiana and

Texas. Klots (1951), MacNeill (1975), and Pyle (1981:840) all mention the northward strays in Massachusetts and New York, but made no mention of Ohio and surrounding states. The single record for Ohio (Janus, 1973) must be considered in doubt without substantiation, since there is no extant specimen. 145 Group D.

This small group contains only two genera and a single species in

the area covered by this study. The key to the groups of HESPERIINAE

above separates this from the other three groups found in North America.

Key to the genera of Group D in the eastern United States.

1. Mesotibia with spines on inner surface; wing span 44-56mm...... Calpodes.

1'. Mesotibia without spines; wing span 24-36mm...... Panoquina.

Calpodes Htlbner, 1819

NOMENCLATORIAL AND SYSTEMATIC STATUS

Calpodes Htlbner, 1819:107. [type-species: Papilio ethlius Stoll, 1782:212, selected by Scudder, 1872:82; valid name (Lindsey, Bell and Williams, 1931:136)]

Colpodes Auctorum. [misspelling]

DIAGNOSIS

Distinguished from Panoquina in the key above.

DISCUSSION

New World, single species (Calpodes ethlius (Stoll, 1782)) from southern United States to Argentina, with wide-spread distribution in between (Evans, 1955; MacNeill, 1975). Calpodes ethlius has been reported from Illinois (Irwin & Downey, 1973) and Pennsylvania (Clench,

1971). Klots (1951) refers to this as a species of the deep south.

MacNeill (1975), however, states that it has strong dispersal powers.

It may be found at some time in Ohio. Larval food plants include various species of the ornamental plant Canna (MacNeill, 1975). 146 Panoquina Hemming, 1934

NOMENCLATORIAL AND SYSTEMATIC STATUS

Panoquina Hemming, 1934:38. ftype-species: Hesperia panoquin Scudder, 1864:178, by original designation; valid name (MacNeill, 1975:427)]

Prenes Scudder, 1872:81. [invalid as a junior homonym of Prenes Gistler, 1848]

DIAGNOSIS

See key to genera above.

DISCUSSION

New World genus with 15 species recognized by Evans (1955), most

of which are tropical. Seven of these species are known from the United

States, none from Canada (MacNeill, 1975). Members of this genus feed

on various grasses including marsh grasses and at least two agricultural

crops, sugarcane and rice.

Panoquina ocola (Edwards, 1863)

Long-winged Skipper

NOMENCLATORIAL STATUS

Hesperia ocola Edwards, 1863:20. [type-locality: Georgia, Florida, and Texas]

SYSTEMATIC STATUS

Panoquina ocola (Edwards, 1863:20. [valid name (Lindsey, Bell & Williams, 1931:138)]

Prenes hecebolus Scudder, 1872.

Prenes Cornelius Draudt, 1924.

Prenes ocola; Lindsey, Bell & Williams, 1931. 147 DIAGNOSIS

Fore-wing usually without light spot in discal cell. Hind-

wing usually immaculate below, as well as above, unlike some other

members of the genus. Appearance of the adult is generally that of

uniformly brown wings and body, with several postmedian,

light-colored spots on the fore-wing.

DISTRIBUTION

Throughout most of the southeastern United States, from New Jersey

south, and westward through Kentucky and Ohio to Arkansas and Texas.

Scattered Ohio records extend from Lake Erie to south-central portions

of the state (Fig. 94).

BIOLOGY

Authors report this species commonly associated with disturbed

areas, with various grasses serving as larval food plants. Three

generations southward (Klots, 1951), fewer to the North. Ohio records

(Fig. 95) occur in June and August, with a break between, suggesting the possibility of two generations. 148 PAPILIONOIDEA Latreille, 1809.

As used here, this superfamily embraces all of the non-skipper

butterflies. This agrees with traditional and, in many cases, current

usage. This name was first used by Dyar (1902:1) for all of the

butterflies, including the skippers. Comstock and Comstock (1904:256)

segregated the skippers into the superfamily HESPERIOIDEA, and the

remaining butterflies into the PAPILIONOIDEA. More recently, many

authors have continued this system of two superfamilies (e.jg,.

McDunnough, 1938; Klots, 1964; Miller & Brown, 1981). However, some

authors now separate the PAPILIONOIDEA (sensu Comstock & Comstock, 1904)

into three superfamilies (PAPILIONOIDEA, LYCAENOIDEA, NYMPHALOIDEA)

based on characters including the adult thorax and its appendages, and

the (Higgins, 1975; Ferris & Brown, 1981).

Key to the families of PAPILIONOIDEA in America north of Mexico.

1. Labial palp as long as thorax, about four times as long as head...... LIBYTHEIDAE.

, 1'. Labial palp shorter than thorax...... 2.

2(1’). Eye emarginate at antenna; width of face between eyes less than length; hind-wing lacks humeral vein...... 3.

2'. Eye not emarginate; width of face between eyes about same as length ; hind-wing with humeral vein...... 4.

3(2). Hind-wing with humeral vein existing as a small spur off the base of Sc + R^, C thickened from wing base distad to humeral angle, without tails; antenna longer than fore-wing length...... RIODINIDAE.

3'. Hind-wing without humeral vein, C not thickened as above, with or without tails; antenna equal to or slightly less than \ fore-wing length...... LYCAENIDAE. 149

4(2'). Prothoracic legs about same size as metathoracic...... 5.

4'. Prothoracic legs reduced, less than % size of metathoracic legs...... 6.

5(4). Fore-wing discal cell with 4 veins arising from posterior margin; hind-wing with one anal vein, often with tail-like extension at vein M^; protibia with lateral epiphysis; tarsal claw simple, not bifid...... PAPILIONIDAE.

5 1. Fore-wing discal cell with three veins arising from posterior margin; hind-wing with two anal veins; protibia without epiphysis; tarsal claw bifid..... PIERIDAE.

6(4'). Fore-wing with some veins (especially Sc) swollen basally; antenna swollen apically, not distinctly clubbed; wings often with relatively large, round, eye-like spots...... SATYRIDAE.

6'. Fore-wing with veins not swollen as above; antenna more or less distinctly clubbed; few with eyespots on wings...... '...... 7.

7(6'). Hind-wing with humeral vein recurved, bent basad; fore-wing relatively long and narrow, wing span 64-92mm; (not in Ohio area)...... HELICONIIDAE.

7'. Hind-wing with humeral vein curving distad; fore-wing variable...... 8.

8(7') . Antenna scaled...... NYMPHALIDAE.

8'. Antenna without scales DANAIDAE. 150 PAPILIONIDAE Latreille, 1809.

In North America this family includes three subfamilies:

BARONIINAE, PARNASIINAE AND PAPILIONINAE. Ehrlich (1958) found that

the latter two are relatively more closely related than either is to

the more primitive BARONIINAE (Mexico only), which is close to deserving family rank. Munroe (1961) also retains BARONIINAE as a subfamily of the PAPILIONIDAE to clarify rather than obscure relationships. The family may be distinguished from others by the combination of: presence of protibial epiphyses, one anal vein in the hind-wing (two in the BARONIINAE), eversible osmaterium in the (immature stages unknown in Baronia, the single genus of the

BARONIINAE). A detailed discussion of the higher classification and evolution of this family is provided by Munroe (1961).

Key to the North American Subfamilies of the PAPILIONIDAE.

1. Hind-wing with one anal vein...... 2.

1'. Hind-wing with two anal veins (Mexico only)...... BARONIINAE.

2(1). Fore-wing with radius 5-branched, basal cubital- anal cross vein present; hind-wing usually tailed, if not then wing color mostly dark with lighter markings...... PAPILIONINAE.

2'. Fore-wing with radius 4-branched, lacking cubital- anal cross vein; hind-wing tailless, ground color white or gray with dark markings...... PARNASSINAE. 151

PARNASSIINAE

This subfamily is limited in North America to the single genus

Parnassius Latreille, 1804 (with three species), which ranges from

Alaska and western Canada southward through the mountains to central

California and New Mexico. It is absent from eastern North America.

PAPILIONINAE Latreille, 1809.

This subfamily is represented in North America north of Mexico by

four genera: Parides, Battus, Eurytides, and Papilio. The latter three

are found east of the Mississippi River and will be considered here.

The use of Papilio here is in the more traditional, broader sense of

many authors. Recently, some authors (including Ferris & Brown, 1981,

and Miller & Brown, 1981) have elevated subgenera to the level of full

genera without agreement as to how it should be done. The result is

somewhat confusing, especially since the common and wide-spread tiger

swallowtails have been placed in two different genera in the two

different works (Ferris & Brown, Miller & Brown, o£. cit.). Un.til

greater stability of nomenclature is achieved, the more conservative

use of Papilio is retained here.

Key to the New World tribes of PAPILIONINAE (after Munroe, 1961).

1. Tibia and tarsus scaled...... LEPTOCIRCINI.

1'. Tibia and tarsus scaleless...... 2.

2(1'). Tibia and tarsus with dorsal rows of spines separated from ventral rows by a spineless area...... PAPILIONINI.

2'. Tibial and tarsal spines not separated as above...... TROIDINI. 152

TROIDINI Ford, 1944

This tribe is represented in America north of Mexico by the genus Battus and occasionally in the southwestern United States by stragglers of the Central American genus Parides. Some species of the tribe have body fluids which are unpleasant to the taste of predators, resulting from the larvae having fed on plants of the family Aristolochiaceae (Klots, 1951). These "protected" species of butterflies serve as the models in systems of Batesian mimicry in which other, palatable species are the mimics. Some of these mimics will be mentioned below in species discussions.

Battus Scopoli, 1777

NOMEN CLATORIAL AND SYSTEMATIC STATUS

Battus Scopoli, 1777:433. [type-species: Papilio polydamus Linnaeus, 1758:460, designated by Lindsey, 1925:80]

Laertias Hllbner, 1819:84.

Ithobalus Hllbner, 1819:88.

DIAGNOSIS

This genus may be distinguished from other genera in the tribe,

on a worldwide basis, by several characters. The ventral sensory

pits of the antennae (called sensory grooves by some authors), usually

found in members of this tribe, are absent in Battus. The anal region

of the male hind-wing has a shiny, scaleless streak instead of the 153 conspicuous, wooly scent organ often found in this tribe. Also, some of the larval, thoracic tubercles are elongated instead of all being about the same length. (Munroe, 1961).

DISCUSSION

Battus is limited to the New World, with 14 species in North,

Central, and South America and in the West Indies (Tyler, 1975:154).

Battus philenor (Linnaeus, 1771)

Pipevine Swallowtail

NOMENCLATORIAL STATUS

Papilio philenor Linnaeus, 1771. [type-locality: "America"]

SYSTEMATIC STATUS

Battus philenor (Linnaeus, 1771). [valid name (Miller & Brown, 1981:60)]

Papilio astinous Drury, 1773.

Laertias philenor; Scudder, 1889.

COMMON NAMES

Pipevine Swallowtail (Pyle, 1981:325)

Blue Swallowtail (Mitchells Zim, 1964:21; Pyle, 1981:325)

Green Swallowtail (Mitchell& Zim, 1964:21)

DIAGNOSIS

This species can be distinguished from the other species of

PAPILIONIDAE in the Ohio area by the combination of the following

characters: hind-wing with irredescent blue or greenish color dorsally

and ventrally; no median spots on either wing; hind-wing above lacking

orange spots. 154

DISTRIBUTION

Battus philenor is found throughout much of the eastern United

States and southern Ontario, and in the Midwest, Southwest and parts of

California. Ohio records indicate distribution throughout the state

(Fig. 96).

BIOLOGY

Larval foodplants are mainly pipevines such as Aristolochia macrophylla (Dutchman's Pipe) and Aristolochia serpentaria (Snakeroot).

Collection records in Ohio indicate two generations (Fig. 97). This

species is reported by authors to be unpalatable to some predators and

in the Ohio area is presumed to be the model for several mimicry

complexes involving other butterflies of similar size and coloration: dark females of Papilio glaucus; both sexes of Limenitis astyanax and

Papilio polyxenes; females of Speyeria diana. 155 LEPTOCIRCINI Kirby, 1896 (=GRAPHINI Ford, 1944).

Members of this tribe are commonly known as the "kite swallowtails" after the long, slender tails found on the hind-wings of many species.

This is distinguished from the other two New World tribes in the key above. A single genus and species are found in eastern North America

(Eurytides marcellus (Cramer, 1779)).

Eurytides Hllbner, 1821

NOMENCLATORIAL AND SYSTEMATIC STATUS

Graphium Scopoli, 1777. [now restricted to Old World species]

Eurytides Hllbner, 1821: [92], [type species: Eurytides iphitas Hllbner, 1821, by monotypy; valid name (Miller & Brown, 1981:60)]

Protesilaus Swainson, 1822:pi. 93.

Cosmodesmus Haase, 1891:15.

DIAGNOSIS

In eastern North America this genus may be distinguished from the other two genera of PAPILIONIDAE by the very long hind-wing tails (tail length in excess of three times the widest tail width) as well as the characters of the tribe.

DISCUSSION

The single leptocircine species found in eastern North America, formerly included in the holarctic genus Scopoli, 1777

(Dos Passos, 1964; Howe, 1975), is now placed, along with other primitive neotropical LEPTOCIRCINI, in the genus Eurytides Hllbner,

1821 by Munroe (1961). 156

Eurytides marcellus (Cramer, 1779)

Zebra Swallowtail

NOMENCLATORIAL STATUS

Papilio marcellus Cramer, 1779. [type-locality: not stated]

Papilio a j a x Linnaeus, 1758 (in part). [name suppressed by I.C.Z.N., Opinion 286, for reasons of ambiguity in the original description]

Iphiclides ajax; Scudder, 1889. [suppressed name]

SYSTEMATIC STATUS

Erytides marcellus (Cramer, 1779). [valid name (Miller & Brown, 1981:60] Papilio marcellus; Jordan, 1907.

Graphium marcellus; Dos Passos, 1964.

COMMON NAMES

Zebra Swallowtail (Klots, 1951:197; Pyle, 1981:347)

Pawpaw Butterfly (Holland, 1931:321)

DIAGNOSIS

This is the only species of in the Ohio area with wing coloration made up of white (sometimes greenish white) ground color crossed longitudinally by blackish bands. Also, the bright, red spot distally located on the inner margin of the hind-wing is distinctive.

DISTRIBUTION

Throughout most of the United States east of the from

Minnesota, Michigan and southern Ontario in the North to the Gulf states and Central Florida. Ohio records indicate distribution throughout the state (Fig. 98). 157

BIOLOGY

Larvae feed on the Pawpaw (Asimina triloba). Price (1970) reports that in northwestern Ohio the adults are usually found around Pawpaw thickets. Eurytides marcellus also has been collected in a dense Pawpaw thicket in Westerville, Franklin County in central

Ohio (John Calhoun, 1982, personal communication). Additionally, adults are found in open deciduous woods, and near wood edges and along roadways and streams. Multivoltine, with two to four annual generations. In Ohio adults first appear in early April and can be found as late as mid-September (Fig. 99). The length of the Ohio flight period suggest two generations. 158 PAPILIONINI Latreille, 1809

These swallowtail butterflies are distinguished from the other two North American tribes in the key to tribes above. Many of these

(including all of the Ohio species) are the so-called "fluted swallowtails." They are so called because in both sexes the inner margin of the hind-wing, next to the abdomen, curves downward (ventrad) forming a depressed groove which parallels the abdomen.

Papilio Linnaeus, 1758

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio Linnaeus, 1758:458. [type-species: Papilio machaon Linnaeus, 1758:462, designated by Latreille, 1810:440, 350; valid name (Miller & Brown, 1981:61)]

Aeronauta Berge, 1842:106. [junior objective synonym]

Achivus Kirby, 1896:286. [junior objective synonym]

DIAGNOSIS

The characters which distinguish these butterflies from other genera are those of the tribe. Munroe (1961) places all species in this tribe in the single genus Papilio. This system is followed in the present study, although some more recent authors have elevated subgenera such that more than one genus is used (see discussion under the subfamily

PAPILIONINAE above).

DISCUSSION

This genus is found throughout most of North America except for the far northern part of the continent and some of the more extreme desert and alpine habitats. In the eastern United States and Canada there are nine species, with four of these found in the Ohio area. 159 Key to the species of Papilio in the Ohio area.

1. Hind-wing with black line crossing discal cell from anterior to posterior veins, approximately 2/3 distance distad from base of cell (best seen ventrally in dark females...... glaucus.

1'. Hind-wing not marked as above...... 2.

2(1'). Hind-wing tail with central yellow spot; spread wings showing dorsal band of yellow running from base of anal margin of hind-wing diagonally across hind-wing, continuing diagonally across fore-wing to apex...... cresphontes.

2'. Wing, markings not as above...... 3.

3(2'). Dorsal fore-wing with marginal and submarginal rows of yellow spots; dorsal hind-wing with median row of yellow spots...... polyxenes.

3'. Dorsal fore-wing with only marginal row of light- colored spots, lacking submarginal row; dorsal hind- wing lacking median row of yellow spots...... troilus. 160

Papilio polyxenes asterlus Stoll, 1782

Black Swallowtail

NOMENCLATORIAL STATUS

Papilio ajax Linnaeus, 1758 (in part). [name suppressed by I.C.Z.N., Opinion 286, for reasons of ambiguity in the original description]

Papilio asterius Stoll, 1782. [type-locality: New York, Carolina, Virgina]

Papilio asterias; Holland, 1898. [misspelled name]

Papilio polyxenes asterias; Clark, 1932. [misspelled name]

SYSTEMATIC STATUS

Papilio polyxenes Fabricius, 1775.

Papilio polyxenes asterius Stoll, 1782. [valid name (Miller & Brown, 1981:61]

Papilio ampliata Menetries, 1857.

Papilio asterioides Reakirt, 1866.

Papilio curvifascia Skinner, 1902.

COMMON NAMES

Black Swallowtail (Klots, 1951:172)

Parsleyworm (larva) (Sutherland, 1978:115)

Caraway-worm (larva) (Comstock & Comstock, 1904:62)

Common American Swallowtail (Holland, 1931:314)

Parsnip Swallowtail (Macy & Shepard, 1941:43)

Common Eastern Swallowtail (Mitchell & Zim, 1964:22)

Eastern Black Swallowtail (Pyle, 1981:327) 161

DIAGNOSIS

The presence of both marginal and median rows of yellow or yellowish-

white spots on fore- and hind-wing distinguish this species from other members of its family in the Ohio area.

DISTRIBUTION

This is one of the more common butterflies in Ohio, and is

probably found in all counties (Fig.100)• It ranges from southern

Canada to Florida, and from the east coast to the Rocky Mountains,

then southward into Mexico. The eastern populations are placed in

the subspecies asterius, while there is some difference of opinion

among authors about the taxonomic status of the forms found in the

southwestern United States. The nominate subspecies is in Cuba.

BIOLOGY

Larvae feed mainly on various species of the Umbelliferae, making

the species an occasional pest of home gardens and truck farms.

Adults are first seen in the early spring, and reproduction is almost continuous throughout the summer. Klots (1951) reports two or more

generations for the northern part of the range. Ohio records extend

from March into October suggesting two, or possibly three generations

(Fig. 101). 162

Papilio cresphontes Cramer, 1777

Giant Swallowtail

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio cresphontes Cramer, 1777. [type-locality: New York, South Carolina, ; valid name (Miller & Brown, 1981:64)]

Heraclides cresphontes; Scudder, 1889.

Papilio thoas var. cresphontes; Hine, 1898.

Papilio thoas; Comstock & Comstock, 1904.

COMMON NAMES

Giant Swallowtail (Klots, 1951:173; Pyle, 1981:337)

Orange Dog (larva) (Klots, 1951:173; Pyle, 1981:337)

DIAGNOSIS

This is the largest of the Ohio area butterflies in both adult and

larval stages. Adults are distinguished from other Ohio swallowtails in

the keys above.

DISTRIBUTION

In eastern North America this species is found from southern Canada, where it is rare, to Florida. It ranges across southern United States

to Arizona and California. Ohio records indicate its presence in most

of the state with the possible exception of the southeastern counties

(Fig. 102). 163

BIOLOGY

In the southern part of its range Papilio cresphontes feeds on various citrus fruits (Klots, 1951; Emmel, 1975). In Ohio the Prickly

Ash (Xanthoxylum americanum) is certainly one of the foodplants (Bales,

1909; Price, 1970) and other trees may be used as well. As reported by authors for the northern part of the range, there are two generations annually in Ohio (Fig. 103). 164

Papilio glaucus Linnaeus, 1758

Eastern Tiger Swallowtail

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio glaucus Linnaeus, 1758. [type-locality: "America"; valid name (Dos Passos, 1964)]

Papilio turnus Linnaeus, 1771.

Jasoniades glaucus; Scudder, 1889.

Pterourus glaucus; Miller & Brown, 1981:66.

Euphoeades glaucus; Ferris & Brown, 1981:187.

COMMON NAMES

Eastern Tiger Swallowtail (Macy & Shepard, 1941:45; Mitchell & Zim, 1964:27)

Tiger Swallowtail (Klots, 1951:175; Pyle, 1981:340)

DIAGNOSIS

This species, including its dark female form, is distinguished

from other Ohio swallowtails in the keys above.

DISTRIBUTION

Papilio glaucus is found from Alaska and British Columbia eastward

across Canada to the eastern coast of North America, and southward east

of the Rocky Mountains to Florida and the Gulf Coast. Dark form

females are more common in the southern part of the range than in the

North. However, these dark females are scarce in the southernmost

part of the range (Emmel, 1975; Tyler, 1975). Papilio glaucus (in­

cluding dark females) has state-wide distribution in Ohio (Fig. 104). 165

BIOLOGY

This species flies continuously in Ohio from early April through early September (Fig. 105). Higher adult capture numbers in April/

May and August, with somewhat lower numbers at other times, indicate

the occurrence of two generations in Ohio. This is consistent with published reports of two generations in the northern and three in the southern part of the range. The larval stage feeds on a variety of trees. Adults are found in deciduous woodland where they may be found flying along the edges of openings or along streams, trails and roadways. At times Papilio glaucus has been found to be common in urban areas (in New England, David J. Horn, 1982, personal com­ munication), including parks and gardens (Pyle, 1981:341). 166 Papilio troilus Linnaeus, 1758

Spicebush Swallowtail

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio troilus Linnaeus, 1758. [type-locality: "Indiis" (error; restriction to lower Delaware River suggested by Brown, 1968:81)]

Euphoeades troilus; Scudder, 1889.

Pterourus troilus; Miller & Brown, 1981:67; Ferris & Brown, 1981:190.

COMMON NAMES

Spicebush Swallowtail (Klots, 1951:178; Pyle, 1981:345)

Green-clouded Swallowtail (Comstock & Comstock, 1904:59; Pyle, 1981:345)

Mellow Bug (larva) (Macy & Shepard, 1941:48)

DIAGNOSIS

This species is distinguished in the keys above.

DISTRIBUTION

This species is found throughout most of the eastern United States from Canada to Florida and the Gulf states. It is less common west of the Mississippi River, and is absent west of the eastern slope of the

Rocky Mountains. In Ohio it is probably found throughout the state

(Fig. 106; .

BIOLOGY

Klots (1951) reports two generations in the North and three in the

South. Ohio records extend from March through September (Fig.107) suggesting two generations. Larvae are reported to feed on several species of plants. One of these, the spicebush (Benzoin sp.), gives the species its common name. 167 PIERIDAE Duponchel, 1832.

There are four nearctic subfamilies of this family as distinguished in the key below. One of these, the DISMORPHIINAE, is known in the

United States only from Mexican strays in southern Texas (Howe, 1975).

The other three subfamilies are found in both eastern and western United

States, and all three are represented in the Ohio area. The PIERIDAE are separated from other families in the key to the families of the

PAPILIONOIDEA above.

Another character, not mentioned before, which distinguishes pierids from all other butterflies is the presence of pterines as a major source of wing coloration in the family. These nitrogen-containing compounds, absent in other butterflies (Howe, 1975), include leucopterin (white), xanthopterin and chrysopterin (yellow) and erythropterin, (red)

(Chapman, 1971).

A key to the nearctic subfamilies of PIERIDAE.

1. Fore-wing with vein M 2 arising from discal cell nearer M^ than M^; fore-wing radius 3- to 5- branched, R. always arising from discal cell, other branches stalked; fore-wing cubitus appears trifid; length of antenna equal to or less than % fore-wing length...... 2.

1'. Fore-wing with vein M 2 arising from discal cell nearer M^ than M^; fore-wing radius always 5-branched, R 1 stalked with other R branches; fore-wing cubitus appears quadrifid; antenna length equal to or greater than h fore-wing length...... DISMORPHIINAE. 168 2(1). Wing color usually orange or yellow, sometimes white; labial palp usually close-scaled with few hair-like scales, 3rd segment less than 1/6 total length of labial palp; hind-wing humeral vein absent or greatly reduced...... COLIADINAE.

2'. Wing color usually white; labial palp usually hirsute, bristly with hair-like scales, 3rd segment at least 1/4 total length of labial palp; hind-wing humeral vein present, usually long and well-developed...... 3.

3(2'). Fore-wing radius 4- or 5- branched, if 4- then fore-wing apex falcate; hind-wing ventral surface marbled with yellow-greenish markings...... ANTHOCHARINAE.

3'. Fore-wing radius 3- or 4-branched, fore-wing apex rounded; hind-wing ventral surface white, if greenish markings present they follow veins...... PIERINAE. 169 PIERINAE Duponchel, 1832.

Adult members of this subfamily are commonly referred to as "whites" because of the extensive white areas on the wings. The apex of the fore­ wing is rounded, never falcate as in some members of the ANTHOCHARINAE.

In some, greenish markings occur on the ventral surface of the hind- wing, following the veins. On the ventral hind-wing surface of the

ANTHOCHARINAE, however, the greenish markings are mottled and cross both veins and cells. The PIERINAE are separated from the COLIADINAE and

DISMORPHIINAE in the key to subfamilies above.

Some authors treat this subfamily in a broader sense, including the

ANTHOCHARINAE and COLIADINAE as tribes (Klots, 1951, Ehrlich, 1958,and

Howe, 1975 for the former; Klots, 1951 for the latter). The arrangement of Miller & Brown (1981) is followed here.

The genus Pieris Schrank, 1801 traditionally has been used by authors

(Klots, 1951; Howe, 1975) as a worldwide (except Antarctica) taxon, consisting of the smaller PIERINAE (wing span about 30-45mm), with wing markings different on the two surfaces. The genus is now divided into three genera, mainly on the basis of secondary sexual characters of the male (Higgins, 1975; Miller & Brown, 1981; Ferris & Brown, 1981). Pieris

(now wholly Palearctic) is distinguished by the very long, narrow androconial scales of the male wing (about six times as long as the greatest width) as well as by male genitalic characters. In the other two genera (Artogeia Verity, 1947 and Pontia Fabricius, 1807) the androconial scales are short (less than four times as long as the greatest width). These scales are pyriform in Artogeia and straight­ sided in Pontia. 170 Key to the genera of PIERINAE in eastern North America (except Florida).

1. Fore-wing with length of beyond branching of R^+^ subequal to length of R^+^+ ^ stem; large, wing span 41-60mm...... Appias.

1'. Fore-wing R^ as above equal to about % length of R^+^+^ stem; wing span variable...... 2.

2(1'). Ventral hind-wing with yellow-green or green markings contrasting in mottled pattern with lighter background...... Pontia.

2'. Ventral hind-wing of uniform color, either light or dark, at most with contrasting coloration following veins...... 3.

3(2'). Small, fore-wing less than 30mm long...... Artogeia.

3'. Large, fore-wing greater than 30mm long...... Ascia.

Appias Hllbner, 1819

NOMENCLATORIAL AND SYSTEMATIC STATUS

Appias Hllbner, 1819. [type-species: Papilio zelmira Stoll, 1780, designated by Butler, 1870:49; valid name (Miller & Brown, 1981:69)]

Catophaga Hllbner, 1832.

Hiposcirta Geyer, 1832.

Trigonia Geyer, 1837. [preoccupied by Trigonia Grugier, 1789]

Tachyris Wallace, 1867.

Glutophrissa Butler, 1887.

Lade de Niceville, 1898.

DISCUSSION

This is a tropical genus with a single New World species, (Cramer, 1777), which is migratory, and is found throughout the neotropical region including southern Florida and Texas. Strays have been reported as far north as New York and Nebraska, but not in the

Ohio area, though it could be collected here. 171 Pontia Fabricius, 1807

NOMENCLATORIAL STATUS

Pontia Fabricius, 1807. [type-species: Papilio daplidice Linnaeus, 1758:468, designated by Curtis, 1824:pl. 48]

Mancipium Htlbner, 1807. [invalid name No. 214, I.C.Z.N., 1954]

Leucochloe RHber, 1907. [junior objective synonym]

SYSTEMATIC STATUS

Pontia Fabricius, 1807. [valid name (Miller & Brown, 1981:70)]

Pieris Schrank, 1801 (in part).

Synchloe Hllbner, 1818.

Parapieris de Niceville, 1897.

DISCUSSION

Four North American species are placed in this genus, one of which

(Pontia protodice) has been reported from Ohio. Pontia is distinguished from other members of its subfamily in the key and discussion above.

Pontia protodice (Boisduval & Leconte, 1829)

Checkered White

NOMENCLATORIAL AND SYSTEMATIC STATUS

Pieris protodice Boisduval & Leconte, 1829. [type-locality: probably Screven Co., Georgia (Miller & Brown, 1981:70); valid name (Miller & Brown, 1981:70)]

Pieris vernalis Edwards, 1864.

Pieris protodice; Holland, 1898:278.

COMMON NAMES

Checkered White (Klots, 1951:200; Pyle, 1981:355)

Common White (Holland, 1931:280; Pyle, 1981:355)

Southern Cabbageworm (Sutherland, 1978:116) 172

DIAGNOSIS

In the Ohio area the generic characters distinguish this species.

The immaculate summer generation differs from Artogeia rapae, which it resembles, in the absence of the dark, longitudinal marking on the ventral surface of the hind-wing, and from Artogeia virginiensis in the presence of yellow tinting on the ventral surface of the hind-wing (lacking in the latter).

DISTRIBUTION

This species ranges across North America from east to west coasts, and from the southern states north to southern New England, New York,

Michigan, , Montana, and Oregon, with rare occurence in southern Alberta, Manitoba, and Quebec (Klots, 1951; Forbes, 1960; Howe,

1975). It is now more common in the West than in the East (Pyle,

1981:356). Distribution records for Ohio (Fig. 108) indicate that it was common and widespread during the first half of this century. The lack of any records of capture in recent years, however, seems to indicate that it has been extirpated from the state. The last date of capture known for this species in Ohio was 1966. Since the local habitat preference is old fields, roadsides, and other disturbed areas

(Klots, 1951; Howe, 1975), the occurence of this species in Ohio probably increased from the time of early European settlement of the area as the landscape was altered. Reasons for population decline and probable extirpation from some parts of its range (including

Ohio) in recent years are not known. Klots (1951) speculates that competition from Artogeia rapae is an important factor in the 173

distribution of this species. Pyle (1981:356) offers an alternate

hypothesis: that changes in distribution patterns of this species may

have been caused by expansion and contraction of habitats through use

and disuse by humans. However, experimental studies are needed

concerning this species.

BIOLOGY

Pontia protodice is multivoltine with three generations occuring

in the northern part of its range. Adults first appear in late March

or early April. Ohio records extend from April into October

indicating at least two, possibly three generations (Fig. 109).

Larvae feed on many members of the family Cruciferae, and in this

respect are in potential competition with Artogeia rapae (Forbes,

1960; Howe, 1975). i 174 Artogeia Verity, 1947

NOMENCLATORIAL AND SYSTEMATIC STATUS

Artogeia Verity, 1947. [type-species: Papilio napi Linnaeus, 1758, by original designation; valid name (Miller & Brown, 1981:71)]

Pieris Schrank, 1801 (in part).

DISCUSSION

All three North American species of this holarctic genus are found in the Ohio area, and are discussed below.

Key to the species of Artogeia in the Ohio area.

1. Ventral hind-wing surface white, veins outlined with diffuse, grayish-brown or light gray shading....virginiensis.

1'. Ventral hind-wing surface shaded with yellow, veins outlined or not, but if outlined then with distinct dark gray shading...... 3.

2(1'). Ventral hind-wing with anterior and posterior ray­ shaped areas of dark shading, both narrow at wing base and spreading out in fan distally; the two areas of shading are often separated by a narrow, central light band running from base of wing through anterior portion of discal cell to outer margin of wing...... rapae.

2'. Ventral hind-wing without darkly shaded areas and central light band as above...... napi.

Artogeia napi (Linnaeus, 1758)

Mustard White

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio napi Linnaeus, 1758. [nec Linnaeus, 1761; type-locality: Sweden (designated by Verity, 1947)]

Artogeia napi (Linnaeus, 1758). [valid name (Pyle, 1981:358)]

Pieris napae; Edwards, 1881. [misspelling] 175 COMMON NAMES

Mustard White (Klots, 1951:201; Pyle, 1981:358)

Veined White (Pyle, 1981:358)

DIAGNOSIS

See key to species of Artogeia.

DISTRIBUTION

This holarctic species is found throughout northern Europe, Asia, and North America. It is the only species of butterfly known presently to inhabit the Aleutian Islands or the islands of the Bering Sea. In the United States it is found in the Great Lakes region and New England in the East, and in the Rocky Mountains as far south as New Mexico and

Arizona, and in the Pacific Northwest south to central California. In the Ohio area its distribution is limited to Michigan and southern

Ontario (Hovanitz, 1962). It is not known from Ohio. Several old literature records for Ohio are probably in error, caused by confusion with Artogeia virginiensis or the immaculate form of Artogeia rapae.

BIOLOGY

In the original description of this species Linnaeus (1758) reports it as being found in cabbage. Klots (1951) suggests that competition with A. rapae in eastern North America has restricted the habitat to the extent that its food is now limited to woodland species such as toothworts (Dentaria sp.), rock cresses (Arabis sp.), and winter and water cresses ( sp.). However, Chew (1981) has found that cympatric populations of Artogeia rapae and A. napi oleracea in northern Vermont exist with broad overlap without competition. 176

"Each species occupies a habitat or utilizes a foodplant not success­ fully exploited by the other." Chew (op. cit.) suggests that in areas where Artogeia napi oleracea has been excluded, this results from shifts in land use by humans and concomitant changes in crucifer flora, rather than from competition with Artogeia rapae. There are two to three annual generations in eastern North America and adults appear in late April in the latitude of the Ohio area. 177 Artogeia virginiensis (Edwards, 1870)

West Virginia White

NOMENCLATORIAL AND SYSTEMATIC STATUS

Pieris napi Linnaeus, 1758 (in part).

Pieris virginiensis Edwards, 1870. [type-locality: restricted to Kanawha district, West Virginia (Brown, 1973:97)]

Artogeia virginiensis (Edwards, 1870). [valid name (Pyle, 1981:359)]

Pontia napi virginiensis; Dyar, 1902.

DISTRIBUTION

This Nearctic species is found from northern Michigan eastward to Quebec and New England, and southward through eastern Ohio and

Pennsylvania, and in the Appalachian Mountains to western North Carolina

(Hovanitz, 1963). The range is primarily limited to the Transition

Life Zone, although the Upper Austral Zone is represented in Ohio and other limited areas. Distribution in Ohio is limited to the northcentral and northeastern parts of the state (Fig.110)*

BIOLOGY

This species is univoltine (although Forbes, 1960, records a rare, second summer generation near its southern limit). Ohio records extend from the middle of April to the end of May (Fig.111). Although the food­ plant is usually listed as the toothwort Dentaria diphylla, the food in at least one area in Ohio (Highbanks Metropolitan Park, Delaware County) is very likely Dentaria lacinatum, the only species of Dentaria known in that area (Edward S. Thomas, 1973, personal communication). As noted for other parts of its range (Klots, 1951) it is very local and is probably limited to wooded areas where the foodplant is found. 178

Artogeia rapae (Linnaeus, 1758)

Cabbage Butterfly

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio rapae Linnaeus, 1758. [type-locality: Sweden (designated by Verity, 1947)]

Pieris rapae; Klots, 1951:200.

Artogeia rapae (Linnaeus, 1758). [valid name (Pyle, 1981:360)]

COMMON NAMES

Cabbage Butterfly (Holland, 1931:282; Mitchell & Zim, 1964:37)

European Cabbage Butterfly (Klots, 1951:200)

Cabbage White (Pyle, 1981:360)

Small White (Pyle, 1981:360)

European Cabbage White (Pyle, 1981:360)

Imported Cabbageworm (larva) (Sutherland, 1978:116)

DISTRIBUTION

This Holarctic species is found in most of temperate and sub­ tropical North America and Eurasia. Also, it has been introduced onto many islands in the Pacific Ocean, including Hawaii (Howe, 1975).

It was introduced into North America about 1860 by man (after having been limited to the Palearctic Region), and subsequently at several different locations in North America. By the mid-1880's its range covered most of the continent between 30 and 60 degrees of north latitude (Scudder, 1889). In Ohio it is probably ubiquitous

(Fig. 112). 179

BIOLOGY

The larva of this species is a wide feeder including nearly all of the Cruciferae (Klots, 1951; Forbes, 1960). Because of its feeding habits as a larva it has become a garden and field pest of some importance in North America. Ohio records extend from March through October, suggesting almost continous reproduction (Fig. 113). Ascia Scopoli, 1777

NOMENCLATORIAL AND SYSTEMATIC STATUS

Ascia Scopoli, 1777. [type-species: Papilio monuste Linnaeus, 1764, designated by Scudder, 1871; valid name (Miller & Brown, 1981:73)]

DISCUSSION

This is a New World, mainly tropical genus, with a single species,

Ascia monuste (Linnaeus, 1764), found east of the Mississippi River, in

the South. The female of this species is dimorphic, with the dark form

showing migratory behavior, moving at times up the Mississippi River valley to Kansas and up the East coast at least to Virginia. It is not known from Ohio. 181 ANTHOCHARINAE Tutt, 1894.

The ANTHOCHARINAE (EUCHLOEINAE Auctorum) are small to medium-sized butterflies (wing span 30-45mm in the Ohio area) with distinctive greenish markings on the ventral hind-wing surface. They are distinguished from the other three Nearctic subfamilies in the key above. Species in

Ohio and surrounding states have wings predominantly white with some dark markings.

Key to the Nearctic genera of the EUCHLOEINAE.

1. Fore-wing R£ vein arises connate from discal cell with R ^ , or stalked with R^, radius 5-branched...... Euchloe.

1'. Fore-wing R 2 arises from discal cell free of R^, radius 4- or 5-branched...... Falcapica.

Euchloe Htlbner, 1819

NOMENCLATORIAL AND SYSTEMATIC STATUS

Euchloe Htlbner, 1819. [type-species: Euchloe ansonia var. esperi Kirby, 1871:506, designated by I.C.Z.N., Opinion 177; valid name (Miller & Brown, 1981:74)]

Phyllocharis Schatz, 1885. [preoccupied by Phyllocharis Dalman, 1824]

DISCUSSION

Four species of this holarctic genus occur in America north of

Mexico. They are univoltine, with adults flying for a short time in the spring. Typically, the larvae feed on buds, flowers, or fresh seed pods of the foodplant. 182

Euchloe olympia (Edwards, 1871)

Olympia

NOMENCLATORIAL STATUS

Anthocaris [sic.] olympia Edwards, 1871. [type locality: West Virginia]

SYSTEMATIC STATUS

Euchloe olympia (Edwards, 1871). [valid name (Pyle, 1981:364)]

Synchloe olympia; Scudder, 1889.

Zegris olympia; Barnes & McDunnough, 1917.

COMMON NAMES

Olympia (Klots, 1951:182)

Olympia Marblewing (Pyle, 1981:364)

Marbled White (Wagner, 1977:107)

DIAGNOSIS

This species may be distinguished from other pierids in the eastern

United States by the combination of white ground color of the wings, black bar at the distal end of the discal cell in the fore-wing, and irregular olive-green bands on the ventral surface of the hind-wing which do not reach the wing margin in the area of the cubital veins.

Neither sex shows the orange patches on the fore-wing which distinguish the males of Falcapica midea. 183

DISTRIBUTION

This species ranges from southern Ontario southward through

Pennsylvania to Maryland, and westward through West Virginia, Kentucky,

Indiana, and Illinois to central Montana, eastern Wyoming, eastern

Colorado, and northern Texas. Although no known records exist for the state of Ohio, it probably has been extant here in the past and may be present today.

Wagner (1977) described a Great Lakes "dune form" which differs from the "inland form" in foodplant preference, as well as some morphological characters. The "dune form" seems to prefer Arabis lyrata, while the "inland form" prefers Arabis drummondii. Further,

Wagner found that in Michigan A. lyrata is mainly found in sand dunes and shoreline sand strips, and A. drummondii is a plant of inland prairie regions. If Euchloe olympia is to be found in Ohio, it should be sought near Lake Erie, if the foodplant (probably A. lyrata) can be found in sandy areas, or in inland areas of prairie remnants in western and southern parts of the state.

BIOLOGY

Adults of this single-generation species appear in April. The larvae feed on cresses, as stated above, and Hedge Mustard

(Sisymbrium officinale), and probably other Cruciferae as well

(Klots, 1951). 184 Falcapica Klots, 1930

NOMBNCLATORIAL AND SYSTEMATIC STATUS

Falcapica Klots, 1930. [type-species: Mancipium midea Hllbner, 1809, by original designation]

Anthocharis Boisduval, Rambur & Graslin, 1833 (in part).

Midea Herrich-SchHffer, 1867. [preoccupied by Midea Bruzelius, 1855]

DISCUSSION

This Holarctic genus contains five species, three of which are

New World (Howe, 1975:385). One of these, Falcapica midea, is discussed below.

Falcapica midea (Htlbner, 1809)

Falcate Orangetip

NOMENCLATORIAL STATUS nec Papilio genutia Cramer, 1779. [available name, but applies to a species of oriental danaids; improperly cited in connection with species of EUCHLOEINAE (Hemming, 1967:192)]

Papilio genutia Fabricius, 1793. [invalid junior homonym of Papilio genutia Cramer, 1779.]

Mancipium midea Htlbner, 1809. [type-locality: probably vicinity of Savannah, Georgia (Miller & Brown, 1981:76)]

Anthocaris genutia Morris, 1860. [misspelling]

SYSTEMATIC STATUS

Falcapica midea (Htlbner, 1809). [valid name (Miller & Brown, 1981:76)]

Euchloe midea; Scudder, 1889.

Pieris lherminieri Godart, 1816.

Anthocharis genutia; Weidemeyer, 1864.

Midea genutia; Kirby, 1871. 185 Euchloe genutia; Holland, 1898.

Synchloe genutia; Dyar, 1902.

Anthocharis (Falcapica) genutia; Klots, 1930.

Anthocharis midea; McDunnough, 1938.

DIAGNOSIS

The produced (falcate) apex of the fore-wing in both sexes distinguishes this species from all other pierids in this area. The male is also characterized by the presence of an orange patch of color just median to the dark apex of the fore-wing.

DISTRIBUTION

This nearctic species occurs in the eastern United States from central Massachusetts southward on the east coast to Georgia, and westward to Illinois and Kansas, northeastern Texas and Arkansas. Except for a single, old literature record (Henninger, 1910), distribution in Ohio is mainly limited to the southern half of the state (Fig.114).

BIOLOGY

This species is univoltine in the northern part of its range and bivoltine in the southern part. Ohio records are limited to April and

May, indicating a single generation (Fig.115). The larvae feed on various members of the Cruciferae including Rock Cress (Arabis perfoliatum), Bitter Cress (Cardamine spp.), Winter Cress (Barbarea vulgaris), Mouse Ear Cress (Sisymbrium thaliana), and Shepherd's Purse

(Capsella bursapastoris). 186 COLIADINAE Swainson, 1827.

This subfamily includes the butterflies commonly called sulphurs

and yellows. In many, yellow and/or orange are the predominant wing

colors; white females are found in some species. Black wing borders

are found in many species, especially in females. Next to Artogeia

rapae some of these are the most common butterflies in Ohio.

Key to the genera of the COLIADINAE in eastern North America

except Forida.

1. Fore--wing radius 3-branched...... Nathalis.

1'. Fore-wing radius 4-branched...... 2.

2(1'). Tarsus with well-developed arolium and pulvilli...... 4.

2'. Arolium and pulvilli lacking...... 3.

3(2'). Fore-wing apex acutely pointed...... Zerene.

3'. Fore-wing apex rounded to bluntly pointed...... Colias.

4(2). Fore-wing length less than 29mm...... Eurema.

4'. Fore-wing length greater than 30mm...... Phoebis.

Colias Fabricius, 1807

NOMENCLATORIAL STATUS

Colias Fabricius, 1807. [type-species: Papilio hyale Linnaeus, 1758, designated by I.C.Z.N., 1943, Opinion 146]

Colias Htlbner, 1819. [junior homonym of Colias Fabricius, 1807]

Eurymus Horsfield, 1829. [junior homonym of Eurymus Rafinesque, 1815]

Megonostoma Reakirt, 1863. [junior objective synonym of Zerene Htlbner, 1819]

Meganostoma Kirby, 1871. [misspelling] 187 SYSTEMATIC STATUS

Colias Fabricius, 1807. [valid name (Miller & Brown, 1981:77]

Scalidoneura Butler, 1871.

Ericolias Watson, 1895.

DISCUSSION

Colias is a Holarctic genus with some representation in Africa,

southern Asia, and tropical America. It is found throughout North

America from the Arctic southward (Klots, 1975), including 16 species

(Miller & Brown, 1981). Species in this genus may be placed into three groups based on foodplant preferences: legume feeders (Leguminosae), heath feeders (), and willow feeders (Salicaceae). The number of generations each year is highly variable between species

(Klots, 1975).

Key to the species of Colias in the Ohio area.

1. Dorsal wing color usually predominantly orange (white in some), with black color of fore-wing apex extending about % distance from apex to discocellular spot in male, about 2/3 same distance in female...... eurytheme.

1'. Dorsal wing color usually predominantly yellow (white in some), with black fore-wing color as above extending about 1/3 distance in male, ^ in female...... 2.

2(1'). Ventral hind-wing with discocellular spot double; ventral wing surface with dark submarginal spots...... philodice.

2'. Ventral hind-wing with discocellular spot single; ventral wing surface without dark submarginal spots...... interior. 188 Colias philodice Godart, 1819

Common Sulphur

NOMENCLATORIAL STATUS

Colias philodice Godart, 1819. [type-locality: Virginia]

Colias philodin Edwards, 1872. [misspelling]

Eurymus philodice; Scudder, 1881. [Eurymus in this usage is a junior homonym of Eurymus Rafinesque, 1815]

SYSTEMATIC STATUS

Colias philodice Godart, 1819. [valid name (Miller & Brown, 1981:78]

Colias chrysotheme (Esper, 1781).

Colias doripe Godart, 1819.

Colias maria Edwards, 1884.

Colias notatus Clark & Clark, 1941.

The distinct separation of this species from its close relative,

Colias eurytheme Boisduval, 1852, has been questioned by authors, some of whom place the two as subspecies of a single species or as subspecies of the palearctic species Colias chrysotheme (Esper, 1781). At present,

Klots (1975) regards the two as separate species that hybridize widely, and that are distinct from the palearctic species. Klots (1975) believes that the two nearctic forms probably evolved nearly complete specific separation with almost complete geographical separation, with £. philodice ranging from Virginia northward into eastern Canada and with a westward extension in Canada, while eurytheme ranged over most of central and western North America. With European immigration and the accompanying clearing of forests and plowing of prairies, and the massive introduction of cultivated legumes, the two species were able to expand their ranges and become sympatric in most areas. 189 COMMON NAMES

Common Sulphur (Klots, 1951: 185; Pyle, 1981:371)

Clouded Sulphur (Klots, 1951:185)

Roadside Butterfly (Comstock & Comstock, 1904:92)

DIAGNOSIS

See key to the species of Colias above.

DISTRIBUTION

As mentioned above this species was probably limited in its distribution to the eastern coast of North America from Virginia north

to Canada, prior to European settlement. Today, its distribution includes most of North America northward into the Hudsonian Zone except for western California, peninsular Florida (where rarely found), and Alaska and adjacent Canada (Klots, 1975). Although not recorded from every county in Ohio, it probably is resident throughout the state

(Fig.116).

BIOLOGY

Clovers (Trifolium sp.) are the preferred foodplants. Others which have been reported include lupins (Lupinus sp.), alfalfa (Medicago sp.), and other Leguminosae. The species is bivoltine in the northern parts of its range and multivoltine with continuous, overlapping generations in the southern portions. In Ohio there are probably at least three generations annually, and the adults may be found from April through

October (Fig.117). Colias eurytheme Boisduval, 1852

Alfalfa Butterfly

NOMENCLATORIAL STATUS

Colias eurytheme Boisduval, 1852. [type-locality: California]

Colias enegthenu Edwards, 1872. [misspelling]

Eurymus eurytheme; Scudder, 1881. [Eurymus is not available; see discussion under Colias above]

SYSTEMATIC STATUS

Colias chrysotheme (Esper, 1781) (in part).

Colias eurytheme Boisduval, 1852. [valid name (Miller & Brown, 1981:79)]

COMMON NAMES

Alfalfa Butterfly (Klots, 1951:183; Pyle, 1981:372)

Alfalfa Caterpillar (larva)(Sutherland, 1978:116)

Orange Sulphur (Klots, 1951:183; Pyle, 1981:372)

Eurytheme Sulphur (Holland, 1931:297)

DIAGNOSIS

See key to species of Colias above.

DISTRIBUTION

Contemporary distribution of the nearctic species extends from the

Atlantic to Pacific Oceans, and from the Hudsonian Zone in Canada to

Florida, and into southern Mexico (Klots, 1951; Klots, 1975). Prior to

European settlement of North America this species probably was limited

to western North America. In his original description, Boisduval (1852) used specimens from California and said that the species was also known from Mexico and some parts of the United States. Other authors have written of the absence or rarity of this species in the eastern United

States until the early part of the twentieth century (Klots, 1951, 191 stated that as late as 1918 this species was considered a "rare prize" in New York). Although not recorded from every county in Ohio, this species probably is distributed throughout the state (Fig. 118).

BIOLOGY

Alfalfa (Medicago sativa) is the preferred foodplant of this species.

Adults fly from early spring into late fall, with variations from a single generation per year in the far North and at high altitudes, to many, continuous generations in the most southern parts of the range (Klots,

1951; Klots, 1975). In Ohio there are probably three to four overlapping generations annually, with records extending from April through October

(Fig. 119).

Colias interior Scudder, 1862

Pink-edged Sulphur

NOMENCLATORIAL AND SYSTEMATIC STATUS

Colias chrysotheme (Esper, 1781) (in part).

Colias interior Scudder, 1862. [type locality: north shore of Lake Superior, mouth of Saskatchewan River (Klots, 1951); valid name (Miller & Brown, 1981:83)]

Colias interier; Kirby, 1877. [misspelling]

Eurymus interior; Scudder, 1889. [Eurymus is not available; see discussion under Colias above]

COMMON NAMES

Pink-edged Sulphur (Macy & Shepard, 1941:57; Pyle, 1981:380)

DIAGNOSIS

See key to species of Colias above. 192

DISTRIBUTION

This is a Canadian Zone species distributed over most of Canada from the eastern coast westward to the Rocky Mountains, and southward into the United States in New England, New York, Virginia and West

Virginia and Pennsylvania (in the mountains), Michigan and Minnesota

(Klots, 1951; Klots, 1975). A single Ohio record for Seneca County in the literature (Henninger, 1910) has not been confirmed by specimens and is doubtful. Although this has been reported from the southern part of Michigan (Moore, 1960), it has not been found in either

Indiana or Illinois. Since the records from areas east of Ohio are montane, this species probably will not be found in Ohio. 193

Zerene HUbner, 1819

NOMENCLATORIAL AND SYSTEMATIC STATUS

Zerene HUbner, 1819. [type-species: Papilio cesonia Stoll, 1790, designated by Scudder, 1871; valid name (Miller & Brown, 1981: 84)]

Colias Fabricius, 1807 (in part).

Megonostoma Reakirt, 1863. [junior objective synonym]

Meganostoma Kirby, 1871. [misspelling]

DISCUSSION

Two species of this New World genus are found in America north

of Mexico. One of these, Zerene cesonia, occurs in the Ohio area and is discussed below.

Zerene cesonia (Stoll, 1790)

Dogface Butterfly

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio cesonia Stoll, 1790. [type-locality: Georgia]

Zerene cesonia (Stoll, 1790). [valid name (Miller & Brown, 1981:85)]

Colias coesnonia Kirtland, 1854. [misspelling]

Colias caesonia Morris, 1860. [misspelling]

Meganostoma cesonia Kirby, 1871. [misspelling]

Zerene caesonia Comstock & Comstock, 1904. [misspelling]

Colias cesonia; Klots, 1951.

COMMON NAMES

Dogface Butterfly (Pyle, 1981:384)

Dog's Head (Comstock & Comstock, 1904:90) 194

Southern Dog-face (Holland, 1931:292)

Eastern Dog's Head (Macy & Shepard, 1941:58)

Dog Face (Klots, 1951:189)

DIAGNOSIS

The pointed apex of the fore-wing along with the "dog's head

profile" (in both sexes in this species) marking of the yellow area

of the fore-wing distinguish this species from all other pierids in

the eastern United States.

DISTRIBUTION

This New World species is distributed across the entire southern

part of the United States and southward as far as Argentina. Although

uncommon to rare northward it has been reported from New York, Ontario,

Michigan, Minnesota, Wisconsin, Manitoba, North Dakota, and Wyoming

(Klots, 1951; Howe, 1975). Sporadic Ohio records are limited to the western half of the state from Cincinnati northward to the Toledo area

and as far eastward as Columbus (Fig. 120).

BIOLOGY

This species is trivoltine in the southern part of the United

States and univoltine in the North. Larvae feed chiefly on False

Indigo (), but other legumes such as clovers

(Trifolium), soy beans (Glycine), and alfalfa (Medicago) have been reported. Adults first appear in the early spring. The two known dates for Ohio adults are both from June (Fig. 121). 195 Phoebis HUbner, 1819

NOMENCLATORIAL AND SYSTEMATIC STATUS

Phoebis Htlbner, 1819. [type-species: Phoebis cypris Htlbner, 1819, nec Papilio cipris Cramer, 1777, designated by Butler, 1873; valid name (Miller & Brown, 1981:85)]

Metura Butler, 1873. [junior homonym of Metura Walker, 1855]

Catopsilia Htlbner, 1819.

Callidryas Boisduval & Leconte, 183.0.

Parura Kirby, 1896.

Prestonia Schaus, 1920.

DISCUSSION

This genus is found in the tropical and subtropical regions of the

New World, although some species range into the temperate regions. The large sulphurs of the Old World are presently placed in the genus

Catopsilia Htlbner, 1819, and are more closely related to Colias

Fabricius, 1807 than to Phoebis (Brown & Heineman, 1972:301). Some of the species of Phoebis are migratory and in the tropics are well known for their swarming and mass migrations. This behavior accounts for the presence of the genus in the Ohio area as autumnal strays.

Several species of Cassia (sennas) provide larval food for this genus

(Klots, 1951).

Key to the species of Phoebis in the Ohio area.

1. Dorsal hind-wing with marginal orange coloration...... philea.

1'. Dorsal hind-wing without orange...... sennae eubule. Phoebis sennae eubule(Linnaeus, 1767)

Cloudless Sulphur

NOMENCLATORIAL STATUS

Papilio eubule Linnaeus, 1767. [type-locality: probably vicinity of Charleston, South Carolina (Miller & Brown, 1981:86)]

SYSTEMATIC STATUS

Phoebis sennae eubule (Linnaeus, 1767). [valid name (Miller & Brown, 1981:86)]

Papilio drya Fabricius, 1793.

Catopsilia eubule; Holland, 1898.

Callidryas eubule; Holland, 1931.

COMMON NAMES

Cloudless Sulphur (Holland, 1931:289; Klots, 1951:190)

Giant Sulphur (Mitchell & Zim, 1964:33)

Cloudless Giant Sulphur (Pyle, 1981:387)

DIAGNOSIS

See key to species of this genus.

DISTRIBUTION

General distribution is discussed under the genus above. In Ohio

it is known from western and central parts of the state (Fig. 122).

BIOLOGY

Larvae of this species feed on various legumes including Partridge

Pea (Chamaecrista cinerea), sennas (Cassia), and clovers (Trifolium).

It is multivoltine in the South. Migrant adults are found in midsummer and fall in northern areas. Ohio records are mainly in August-October, with a single June record (Fig. 123). 197 Phoebis philea (Johansson, 1763)

Orange-barred Sulphur

NOMENCLATORIAL STATUS

Papilio philea Johansson, 1763. [type-locality: probably Surinam (Miller & Brown, 1981:86)]

Mancipium argante Htlbner, 1819. [preoccupied by Papilio argante Fabricius, 1775]

SYSTEMATIC STATUS

Phoebis philea (Johansson, 1763). [valid name (Miller & Brown, 1981:86)]

Papilio aricye Cramer, 1779.

Papilio melanippe Stoll, 1782.

Colias corday Htlbner, 1819.

Colias lollia Godart, 1819.

Catopsilia philea; Holland, 1898.

Callidryas philea; Holland, 1931.

COMMON NAMES

Orange-barred Sulphur (Mitchell & Zim, 1964:33)

Red-barred Sulphur (Comstock & Comstock, 1904:87)

Orange-barred Giant Sulphur (Pyle, 1981:388)

DIAGNOSIS

See key to species of this genus.

DISTRIBUTION

See discussion under genus above. A single record in Ohio is from

Franklin County (Fig. 124). 198 BIOLOGY

The larval foodplant in the United States is unknown. A legume has been reported as a foodplant in Costa Rica (Pyle, 1981:389). Adults fly year-round in the Tropics. The single Ohio record is from the second week in September (Fig. 125). 199 Eurema Hilbner, 1819

NOMENCLATORIAL AND SYSTEMATIC STATUS

Eurema Htlbner, 1819. [type-species: Papilio delia Cramer, 1780, designated by Butler, 1870:35; valid name (Miller & Brown, 1981:88)]

DISCUSSION

This genus is divided into three genera (Pyrisitia Butler, 1870

and Abaeis Htlbner, 1819 in addition to Eurema) by Miller & Brown, 1981 and Ferris & Brown, 1981, on the basis of "rather subtle structural differences" (Ferris & Brown, 1981:172). Howe (1971:371) considered

these to be subgenera of Eurema. Pyle (1981), on the other hand, retains

the more conservative and traditional use of the single genus (Eurema), an action which is followed here. Seven species of this genus are known from America north of Mexico, two of which are discussed below.

Key to the species of Eurema in the Ohio area.

1. Dorsal wing orange in discal area nicippe.

1'. Dorsal wing yellow in discal area ...lisa.

Eurema lisa (Boisduval & Leconte, 1829)

Little Sulphur

NOMENCLATORIAL STATUS

Xanthidia lisa Boisduval & Leconte, 1829. [type-locality: United States]

SYSTEMATIC STATUS

Eurema lisa (Boisduval & Leconte, 1829). [valid name: Pyle, 1981:392]

Terias clappi Maynard, 1891.

Terias lisa; Holland, 1898.

Pyrisitia lisa; Miller & Brown, 1981. 200

COMMON NAMES

Little Sulphur (Holland, 1931:302; Klots, 1951:198)

Little Yellow (Pyle, 1981:392)

DIAGNOSIS

See key to species of Eurema.

DISTRIBUTION

This species ranges from New England south to the Gulf states, and thence westward to the Great Lakes area, Mississippi River valley, and the West Indies and South America. Ohio records (Fig. 126) indicate statewide distribution.

BIOLOGY

Larval foodplants include various legumes. This species does not hibernate in the northern part of its range (including Ohio); adults migrate northward annually.^ Adult activity in Ohio occurs from June through October (Fig. 127).

Eurema nicippe (Cramer, 1779)

Sleepy Orange

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio nicippe Cramer, 1779. [type-locality: Virginia]

Eurema nicippe (Cramer, 1779). [valid name (Pyle, 1981:394)]

Terias nicippe; Holland, 1898.

Abaeis nicippe; Miller & Brown, 1981. COMMON NAMES

Sleepy Orange (Klots, 1951:197; Pyle, 1981:394)

Sleepy Yellow (Comstock & Comstock, 1904:97)

Sleepy Sulphur (Mitchell & Zim, 1964:35)

Nicippe Sulphur (Holland, 1931:301)

DIAGNOSIS

See key to species of the genus Eurema above.

DISTRIBUTION

This species is distributed from New York southward to Florida westward to Nebraska, Colorado, and southern California. Ohio distribution is probably statewide (Fig. 128).

BIOLOGY

Larvae feed on several species of legumes. Adults occur year- round in the South. This species cannot hibernate in the North and annually moves into northern areas (Howe, 1975:373; Pyle, 1981:395)

Ohio records occur in the spring, and again from July to October

(Fig. 129), indicating two generations. 202 Nathalis Boisduval, 1836

NOMENCLATORIAL AND SYSTEMATIC STATUS

Nathalis Boisduval, 1836. [type-species: Nathalis iole Boisduval, 1836, by monotypy]

Natalis Doubleday, 1836. [misspelling]

DISCUSSION

The distribution of this genus in America north of Mexico is that

of the only species found in the area, Nathalis iole Boisduval, 1836

(see discussion below).

Nathalis iole Boisduval, 1836

Dainty Sulphur

NOMENCLATORIAL AND SYSTEMATIC STATUS

Nathalis iole Boisduval, 1836. [type-locality: Mexico (Klots, 1951: 199); valid name (Miller & Brown, 1981:91)]

Nathalis .iole RHber, 1910. [misspelling]

Nathalis felicia Poey, 1854.

Nathalis leuteolus Reakirt, 1863.

COMMON NAMES

Dainty Sulphur (Klots, 1951:199)

Dainty Dwarf (Mitchell & Zim, 1964:35)

DIAGNOSIS

This species may be distinguished from all other pierids in the

Ohio area by its small size (wing spread less than 30mm) and the presence of dark bars of color along both the posterior margin of the fore-wing and the anterior margin of the hind-wing. 203 DISTRIBUTION

Nathalis iole occurs in tropical America, and is found across the

southern United States from Florida to southern California (Howe, 1975).

It is often found as far north as Minnesota in the central states, and

occurs in Illinois and Indiana (Klots, 1951), as well as Ohio, where

it has been found as far north as Lucas and Lorain Counties (Fig. 130).

Along the East coast it is found as far north as Virginia (Clark &

Clark, 1951).

BIOLOGY

Foodplants include fetid marigolds (Dyssodia sp.), Common

Chickweed (Stellaria media), Bur Marigold (Bidens pilosa), and

Sneezeweed ( autumnale). The species probably overwinters in

the southern portions of the United States and migrates ‘northward each year where one or more generations may occur (Klots, 1951; Howe, 1975).

Ohio records indicate a flight period from August into October (Fig. 131).

In Illinois adults have been found as early as July 4th, and as late as

November 13th (Irwin & Downey, 1973). 204 LYCAENIDAE Leach, 1815.

This is a large family of relatively small butterflies (wing span generally less than 50mm in America north of Mexico) with approximately

5000 species described worldwide. The greatest diversity of species occurs in the Tropics. Adult males of this family have incompletely developed forelegs, with reduced number of tarsal segments, which are not functional in walking. Females have fully developed, functional forelegs.

The hind-wing lacks the humeral vein, and the fore-wing lacks either one or two of the radial branches. Coloration and other characteristics of some members of this family have led to the common names usually given to the subfamilies: harvesters (), coppers (LYCAENINAE), hairstreaks (THECLINAE), blues (P0LY0MMATINAE). The common name "gossamer winged butterflies" is used by some authors for the entire family.

Key to the subfamilies of the LYCAENIDAE in North America.

1. Mid and hind tibia each with distinct, single, paired, terminal spurs...... 3.

1'. Mid and hind tibia without spurs as above, or with spurs which are very small and difficult to see...... 2.

2(1'). Fore-wing radius three-branched, no branches stalked; M^ of fore-wing separate beyond discal cell, not stalked; dorsal fore-wing of male often with patch of androconial scales; female antenna distinctly clubbed...... THECLINAE.

2'. Fore-wing radius four-branched, last two branches stalked; M^ of fore-wing stalked with R beyond the discal cell; dorsal fore-wing of male never with androconial patch; female antenna tapering to swollen club...... GERYDINAE.

3(1). Antennal club usually cylindrical; fore-wing radius three- branched (if four-branched then tarsal claws with inner tooth and hind-wing tailed and fore-wing length greater than 18mm); hind-wing usually with tornal lobe and one or more tails; male dorsal fore-wing often with androconial patch...... THECLINAE. 205 3'. Antennal club flattened or hollowed beneath; fore-wing radius four-branched (but without combination of parenthetical characters under 3(1) above); hind-wing without tomal lobe and never with more than one tail at end of vein C^; male never with androconial patch on dorsal fore-wing...... 4.

4(3’). Tarsal claw usually with inner tooth (absent only in genera and , neither in Ohio area); veins and R of fore-wing arise distinctly separately from discal cell; male dorsal fore-wing often blue...POLYOMMATINAE.

4'. Tarsal claw always without inner tooth; veins and R of fore-wing arise from discal cell very closely or together; male dorsal fore-wing usually orangeish, rarely bright blue, occaisionally grey...... LYCAENINAE.

GERYDINAE Doherty, 1886.

The taxon represented by the name SPALGINAE Toxopeus, 1929 is now

generally believed to be a tribe within the GERYDINAE. The name

MILETINAE Corbet, 1939, used by some authors for the GERYDINAE, is a junior synonym of GERYDINAE. This subfamily is mainly found in the

Oriental and Ethiopian Regions. Its single, nearctic species is discussed below. Feniseca Grote, 1869

NOMENCLATORIAL AND SYSTEMATIC STATUS

Feniseca Grote, 1869. [type-species: Hesperia tarquinius Fabricius, 1793, by original designation; valid name (Miller & Brown, 1981: 92)]

DIAGNOSIS

In addition to the characters distinguishing the subfamily in the key above, the dorsal fore-wing has wide, marginal dark brown areas surrounding a central, orange-yellow to whitish area which contains several large spots of the marginal color. Wing span 28-32mm. 207 Feniseca targuinius (Fabricius, 1793)

Harvester

NOMENCLATORIAL STATUS

Hesperia targuinius Fabricius, 1793. [type-locality: "in Indiis"]

SYSTEMATIC STATUS

Feniseca targuinius (Fabricius, 1793). [valid name (Miller & Brown, 1981:92)]

Polyommatus crataegi Boisduval & Leconte, 1833.

Polyommatus porsenna Scudder, 1864.

COMMON NAMES

Harvester (Mitchell & Zim, 1964:70; Pyle, 1981:401)

Wanderer (Comstock & Comstock, 1904:237; Macy & Shepard, 1941:161)

Alder Butterfly (Macy & Shepard, 1941:161)

DISTRIBUTION

From eastern Canada south to Florida, westward to Illinois and

Texas. Ohio records indicate distribution throughout most of the state

(Fig. 132).

BIOLOGY

Larvae of this butterfly are carnivorous, feeding on wooly aphids

of two genera, Schizoneura and . Adults are often found in

damp areas including wooded streambanks and wooded trails. The species

is multivoltine throughout most of its area. Ohio records extend from

April thorugh August indicating at least two generations (Fig. 133). 208 LYCAENINAE Leach, 1815.

This subfamily includes those butterflies commonly referred to as

the coppers because of the red-orange wings of many adults. The color

alone, however, is not sufficient to distinguish these from other

lycaenids. Other colors, including bright blue, are found in some

species. Wing venation as well as leg and antennal characters are used

in the key to subfamilies above. The LYCAENINAE are mainly holarctic

in distribution. A few species are found in the Tropics, in the

Neotropical, Ethiopian, and Indo-Australian Regions. The arrangement

of genera here follows Miller and Brown (1979), who recognize seven

genera in America north of Mexico. Three of these genera, Lycaena,

Hyllolycaena, and Epidemia are found east of the Mississippi River, and are discussed below. The characters needed to distinguish the taxa

in this subfamily are used in the keys below, with little further discussion.

Key to the genera of the LYCAENINAE in eastern North America.

1. Ventral fore-wing with distinct, thin black marginal line; fore-wing length usually greater than 16mm...... Hyllolycaena.

1'. Ventral fore-wing without line as above; fore-wing length usually less than 16mm...... 2.

2(1'). Ventral fore-wing with black spots in discal area ringed with white...... Lycaena.

2'. Ventral fore-wing with black discal spots not ringed with white...... Epidemia. Lycaena Fabricius, 1807

NOMENCLATORIAL STATUS

Lycaena Fabricius, 1807. [type-species: Papilio phlaeas Linnaeus, 1761, designated by Curtis, 1828]

Lycia Sodovskii, 1837. [preoccupied by Lycia Hllbner, 1825]

Migonitis Sodovskii, 1837. [preoccupied by Migonitis Rafinesque, 1815]

SYSTEMATIC STATUS

Lycaena Fabricius, 1807. [valid name (Miller & Brown, 1981:92)]

DISCUSSION

Two species in this holarctic genus are found in America north of

Mexico. One of these, Lycaena phlaeas, is found in the Ohio area and is discussed below. 210 Lycaena phlaeas (Linnaeus, 1761)

American Copper

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio phlaeas Linnaeus, 1761. [type-locality: Sweden]

Lycaena phlaeas (Linnaeus, 1761). [valid name (Miller & Brown, 1981: 93)]

Chrysophanus hypophlaeas; Holland, 1898.

COMMON NAMES

American Copper (Holland, 1931:251; Pyle, 1981:403)

Flame Copper (Pyle, 1981:513)

DISTRIBUTION

From Greenland, Hudson Bay and eastern Canada south to Florida, west to Alaska and California. Ohio records indicate statewide

distribution (Fig. 134).

BIOLOGY

Larvae feed on Sheep Sorrel (Rumex acetosella) and Curly Dock

(). Adults are commonly found in old fields and waste

areas. This species is multivoltine, with two generations in the North and four or more in the South. Ohio records extend from April through

October indicating two or possibly three overlapping generations (Fig.

135) . 211

Hyllolycaena Miller & Brown, 1979

NOMENCLATORIAL AND SYSTEMATIC STATUS

Hyllolycaena Miller & Brown, 1979. [type-species: Papilio hyllus Cramer, 1775, by original designation; valid name (Miller & Brown, 1981:94)]

DISCUSSION

A single species, Hyllolycaena hyllus, is included in this genus

by Miller and Brown (1979). It is found in Ohio and is discussed below.

Its large size (wing span 32-35mm) helps to distinguish it from other members of the subfamily.

Hyllolycaena hyllus (Cramer, 1775)

Bronze Copper

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio hyllus Cramer, 1775. [type-locality: "", in error (Miller & Brown, 1981:94)]

Hyllolycaena hyllus (Cramer, 1775). [valid name (Miller & Brown, 1981: 94)]

Polyommatus thoe Guerin-Meneville, 1831.

Chrysophanus thoe; Holland, 1898.

Lycaena thoe; Klots, 1951.

Lycaena hyllus; Howe, 1975.

COMMON NAMES

Bronze Copper (Comstock & Comstock, 1904:239; Pyle, 1981:411) 212

DISTRIBUTION

From eastern Canada south to New Jersey and Maryland, thence westward to Alberta and Colorado. Opler (1975:312) describes this as

an Upper Austral and Transition zone species which strays occasionally

into the Gulf states. Ohio records are limited to the northern two-

thirds of the state (Fig. 136). This species has not been found in

Hocking, Vinton, or Adams counties in the southern part of the state, all three of which have been well collected.

BIOLOGY

Larval foodplants include Curly Dock (Rumex crispus) and knotweeds

(Polygonum spp.). This species is restricted to moist grassy areas including wet meadows, stream edges and ditches, and swamps where the foodplants are found (Pyle, 1981:411). Multivoltine, with two, possibly three generations in June/July and August/October (Pyle, 1981:

411). Ohio records extend from May into October indicating at least two generations (Fig. 137). Further study may show three generations in some parts of the state. 213 Epidemia Scudder, 1876

NOMENCLATORIAL AND SYSTEMATIC STATUS

Epidemia Scudder, 1876. [type-species: Polyommatus epixanthe Boisduval & Leconte, 1833, by original designation; valid name (Miller & Brown, 1981:95)]

DISCUSSION

Of the five members of this genus known from America north of

Mexico, three are found in Ohio and are discussed below. Compared to

Hyllolycaena the five members of this genus are small (wingspan 22-32mm) and all have a dark purple sheen or gloss on the dorsal wing surface.

Key to the species of Epidemia in the Ohio area.

1. Dorsal fore-wing with postmedian row of six or more dark spots...... 2.

1'. Dorsal fore-wing without row of spots as above...... epixanthe.

2(1). Ventral hind-wing with well-developed, scalloped, orange to red submarginal band; ventral fore-wing yellowish, with at most a faint tinge of brown; ventral hind-wing pinkish- to grayish-tan, contrasting with ventral fore-wing...... helloides.

2'. Ventral hind-wing with orange band faint or lacking; ventral fore-wing brownish with yellowish to pinkish cast; ventral hind-wing similar to ventral fore-wing...... dorcas. 214

Epidemia epixanthe (Boisduval & Leconte, 1833)

Bog Copper

NOMENCLATORIAL AND SYSTEMATIC STATUS

Polyommatus epixanthe Boisduval & Leconte, 1833. [type-locality: New Harmony, Indiana, restricted to New Jersey by Rawson, 1948]

Epidemia epixanthe (Boisduval & Leconte, 1833). [valid name (Miller & Brown, 1981:95)]

Chrysophanus hypoxanthe Kirby, 1862.

Chrysophanus epixanthe; Holland, 1898:254.

Lycaena epixanthe; Klots, 1951:152.

COMMON NAMES

Bog Copper (Klots, 1951:152; Pyle, 1981:414)

Purple Disk (Comstock & Comstock, 1904)

Epixanthe Copper (Holland, 1931:250)

Marsh Copper (Macy & Shepard, 1941)

DISTRIBUTION

From eastern Canada south to New Jersey, west to Manitoba, Minnesota

and Wisconsin. This species is known only from Cuyahoga County (Opler,

1975). No specimen of this species has been found from Ohio in this

study. If present in the past, this species may now be extirpated

from Ohio.

BIOLOGY

Wild Cranberry ( macrocarpum) serves as foodplant for

this species, and restricts its distribution. It is found in acid

bogs and boggy marshes where the foodplant is found. One possible

locality of this type in Ohio is Cranberry Bog in Buckeye Lake, in

southern Licking County. This lake was formed by man in 1830 as a 215

reservoir for the Ohio and Erie Canal. A swampy area was used for the

lake, and as the lake water level rose part of the fibrous, top layer

of a large peat bed separated from the substrate and floated on the

lake surface (Dachnowski, 1912:86). A search for EL epixanthe at

Cranberry Bog failed. It may still be possible to find this species

in one or more of the remaining acid bogs in northeastern Ohio. The

species is univoltine, with adults from June to late July or August.

Epidemia dorcas (Kirby, 1837)

Dorcas Copper

NOMENCLATORIAL AND SYSTEMATIC STATUS

Lycaena dorcas Kirby, 1837. [type-locality: "Lat. 54°", restricted to The Pas, Manitoba by Ferris, 1977 (Miller & Brown, 1981:96)]

Epidemia dorcas (Kirby, 1837). [valid name (Miller & Brown, 1981:96)]

Chrysophanus dorcas; Holland, 1931:249.

COMMON NAMES

Dorcas Copper (Klots, 1951:154; Pyle, 1981:416)

DISTRIBUTION

From Newfoundland south to Maine, west to Alaska and Minnesota.

It has been reported only from Mud Lake in Williams County in northwestern

Ohio, by Price (1970) who found a single female, apparently ovipositing on Shrubby Cinquefoil (Potentilla fruticosa).

BIOLOGY

Larvae have been reported feeding on various cinquefoils (Potentilla).

Univoltine, with adults in July and August. The single date of capture known for Ohio is July 11, 1948 (Price, 1970). This species is known to inhabit moist meadows, bogs, and forest clearings. 216 Epidemia helloides (Boisduval, 1852)

Purplish Copper

NOMENCLATORIAL AND SYSTEMATIC STATUS

Polyommatus helloides Boisduval, 1852. [type-locality: San Francisco, California]

Epidemia helloides (Boisduval, 1852). [valid name (Miller & Brown, 1981:96)]

Chrysophanus helloides; Holland, 1898:254.

Lycaena helloides; Klots, 1951:153.

COMMON NAMES

Purplish Copper (Holland, 1931:249; Pyle, 1981:414)

DISTRIBUTION

From the Great Lakes region of eastern Canada south to Michigan

and Ohio, west to the Pacific coast. Ohio records are restricted to

the northwestern quarter of the state (Fig. 138).

BIOLOGY

Authors report various members of the buckwheat family ()

as larval foodplants, including docks, sorrels and knotweeds (Rumex and

Polygonum). This species may be found in disturbed areas including old

fields and urban weed fields, often when moist. Multivoltine, with adults

from May to September. Ohio records extend from April to July (Fig. 139). 217 THECLINAE Swainson, 1831.

This subfamily, commonly called the hairstreaks, consists of butterflies whose dorsal surface as adults in the Ohio area is often drab brown or gray-brown. The ventral wing surface is commonly more useful in determination of species. The THECLINAE have achieved their greatest diversity of species in the Tropics, where adults are often very brightly colored. Many species have tail-like appendages near the hind-wing tomus, at the ends of the cubitus. Many of the adults are small (wing span less than 35mm). Characters used to determine genera and species will be included in the keys below, with discussion in the text under the taxa. 218

Key to the genera of THECLINAE in the eastern United States, except Florida (after Sites & McPherson, 1980).

1. Small (wing span 19-25mm); dorsal wing surface blue with black margins; ventral wing surface bluish-green with white-rimmed, reddish spots; tailess (rare in Ohio)...... Erora.

1'. Without above combination of characters...... 2.

2(1'). Dorsal wing surface with ground color iridescent blue (May be restricted to basal half of wing); ventral wing with postmedian line, if present, edged basally with black...... 3.

2'. Dorsal wing with ground color brown, gray, or black; if ground color blue, then ventral wing with postmedian line edged basally with red band...... 4.

3(2). Ventral wing with white postmedian line, lacking red or orange basal spots (although orange line is present along base of costal margin...... Parrhasius.

3'. Ventral wing without white postmedian line; ventral fore-wing with red or orange basal spot; ventral hind-wing with two red or orange basal spots...... Atlides.

4(2'). Ventral hind-wing with clearly-defined red or orange band bordering inner edge of much narrower white and black postmedian line (evident without magnification)...... Calycopis.

4'. Ventral hind-wing with poorly-defined red or orange band bordering inner edge of wider white and black postmedian line, or band absent...... 5.

5(4'). Ventral wing with ground color green or greenish- brown; ventral fore-wing with well-developed white submarginal line; ventral hind-wing with row of white postmedian marks forming a broken, irregular line; hind-wing with tails present...... Mitoura.

5'. Ventral wing with ground color variable, but if green or greenish-brown then ventral fore-wing without well-developed white submarginal line; ventral hind-wing usually without row of white postmedian marks as above; hind-wing tails present or absent...... 6. 219

6(5'). Ventral hind-wing with row of eight red or orange submarginal spots, row of black postmedian spots ringed with white, blue anal spot lacking, ground color brownish-gray, tails absent...... Harkenclenus.

6'. Ventral hind-wing usually without row of eight red or orange submarginal spots, but if present then blue anal spot present, ground color variable, tails present or absent 7.

7(6'). Ventral hind-wing with red or orange marginal or submarginal spots; hind-wing tails present; ventral fore-wing with variable ground color...... 8.

7'. Ventral hind-wing without red or orange marginal or submarginal spots, although light bands or scattered red scales may be present; hind-wing tails present or absent; ventral fore-wing with ground color brown, dark brown, or greenish-brown...... Incisalia.

8(7). Ventral wing with dark bar at distal end of discal cell.Satyrium.

8*. Ventral wing without bar as above...... 9.

9(8’). Ventral fore-wing with postmedian line extending into cell Cu2~2A, ground color brown...... Euristrymon.

9'. Ventral fore-wing without postmedian line in cell Cu 2~2A, ground color light gray...... Strymon. 220 Atlides Hllbner, 1819

NOMENCLATORIAL AND SYSTEMATIC STATUS

Atlides Hllbner, 1819. [type-species: Papilio halesus Cramer, 1777, designated by Scudder, 1875:124; valid name (Miller & Brown, 1981:98)]

DISCUSSION

This mainly tropical genus is represented in the United States by a single species, Atlides halesus, discussed below. Atlides halesus (Cramer, 1777)

Great Purple Hairstreak

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio halesus Cramer, 1777. [type-locality: Virginia]

Atlides halesus (Cramer, 1777). [valid name (Miller & Brown, 1981:98)]

Atlides dolichos Hllbner, 1823.

Thecla juanita Scudder, 1868.

Thecla halesus; Holland, 1898.

COMMON NAMES

Great Purple Hairstreak (Holland, 1931:224; Pyle, 1981:421)

Giant Purple Hairstreak (Macy & Shepard, 1941:146)

Great Blue Hairstreak (Pyle, 1981:421)

DISTRIBUTION

This is primarily a southern species whose range extends from east to west coasts in North America. It is found as far north as New York,

Ohio, Illinois, and Oregon. The single Ohio record (Dury, 1878) is from Hamilton County in the southwestern corner of the state, and probably represents a stray.

BIOLOGY

Larvae of this species feed on various mistletoes (Phoradendron) which parasitize woody plants. Phoradendron flavescens is found in 14 counties in southernmost Ohio, from Hamilton in the Southwest to

Washington in the Southeast (Braun, 1969:139). Atlides halesus is bivoltine, with early spring and late summer generations. The.date of the single Ohio record is unknown. 222 Harkenclenus dos Passos, 1970

NOMENCLATORIAL AND SYSTEMATIC STATUS

Chrysophanus Hllbner, 1818. [name suppressed by I.C.Z.N., Opinion 541]

Harkenclenus dos Passos, 1970. [type-species: Chrysophanus Hllbner, 1818, by original designation; valid name (Miller & Brown, 1981:99)]

DISCUSSION

This monotypic genus is found from eastern Canada south to Georgia, west to British Columbia, California, and Arizona, excepting the Gulf

states and the extreme Southwest. The single species, Harkenclenus

titus, is probably distributed throughout the state of Ohio (Fig. 140).

Harkenclenus titus (Fabricius, 1793)

Coral Hairstreak

NOMENCLATORIAL AND SYSTEMATIC STATUS

Hesperia titus Fabricius, 1793. [type locality: "in Anglia", probably Newfoundland (Miller & Brown, 1981:99)]

Harkenclenus titus (Fabricius, 1793). [valid name (Miller & Brown, 1981:99)]

Thecla titus; Holland, 1898.

Strymon titus; Klots, 1951.

COMMON NAMES

Coral Hairstreak (Klots, 1951:134; Pyle, 1981:482)

DISTRIBUTION

See discussion under the genus account above. 223

BIOLOGY

Larvae of this species feed on developing fruits of plums and wild cherries. The species is univoltine, with adults from June to August

in Ohio (Fig. 141). This is a butterfly of meadows, old fields, and

roadsides. Adults are commonly found at the blossoms of Butterfly

Weed (Asclepias tuberosa). 224 Satyrium Scudder, 1876

NOMENCLATORIAL AND SYSTEMATIC STATUS

Satyrium Scudder, 1876. [type-species: Lycaena fuliginosa Edwards, 1861, by original designation; valid name (Miller & Brown, 1981:100)]

Callipsyche Scudder, 1876.

DISCUSSION

This holarctic genus contains about sixteen species in America north of Mexico. Most species have tails and well-developed anal lobes on the hind wing. Five species are known from the Ohio area, and are discussed below.

Key to the species of Satyrium in the Ohio area.

1. Ventral wing with dark postmedian lines broad, as wide as thorax, also broken and offset in some parts the full width of the lines...... liparops.

1'. Ventral wing with postmedian line not as above, much narrower than width of thorax...... 2.

2(1'). Dorsal hind-wing with orange marginal spot in cell Cu^-Cu„; ventral fore-wing with postmedian dark line consisting of disconnected rounded spots...... acadica.

2'. Dorsal hind-wing lacking orange spot, or, if present, then u-shaped with dark center; ventral fore-wing with postmedian dark line continuous, or consisting of elongate spots ...... 3.

3(2'). Ventral hind-wing with postmedian dark line broken into separate, narrow, oval, white-edged spots; orange line along inner margin extending distad beyond center of blue spot...... edwardsii.

3'. Ventral hind-wing with postmedian dark line not broken as above, although it may be irregular, consisting of parallel-sided spots; orange line as above, if present, extends only to proximal part of blue spot, not to center...... 4. 225 4(3'). Ventral hind-wing with orange spot in cell C^-Cu- just anterior to tail as thick or thicker than black spot it encloses; ventral hind-wing with anteriormost segment of postmedian line offset basad, but not so far as to be in line with dark bar in distal end of discal cell; male dorsal fore-wing with androconial pad more than half as wide as discal cell...... calanus.

4'. Ventral hind-wing with orange spot as above small, narrower than black spot it encloses; postmedian line segment as above offset so far basad as to be more or less in line with bar at distal end of discal cell; male androconial pad about h width of discal cell caryaevorus. 226 Satyrium acadica (Edwards, 1862)

Acadian Hairstreak

NOMENCLATORIAL AND SYSTEMATIC STATUS

Thecla acadica Edwards, 1862. [type-locality: London, Ontario]

Satyrium acadica (Edwards, 1862). [valid name (Miller & Brown, 1981:100)]

Strymon acadica; Klots, 1951.

COMMON NAMES

Acadian Hairstreak (Klots, 1951:136; Pyle, 1981:449)

DISTRIBUTION

This northern species ranges from Nova Scotia south to New Jersey, west to British Columbia and Washington. Ohio records are limited to the northern two-thirds of the state (Fig. 142).

BIOLOGY

Larvae feed on (Salix). Adults are often found in damp fields and streamsides. Univoltine, with adults in June and July

(Fig. 143). 227 Satyrium edwardsii (Grote & Robinson, 1867)

Edwards' Hairstreak

NOMENCLATORIAL AND SYSTEMATIC STATUS

Thecla edwardsii Grote & Robinson, 1867. [type-locality: London, Ontario]

Satyrium edwardsii (Grote & Robinson, 1867). [valid name (Miller & Brown, 1981:101)]

Thecla edwardsi; Holland, 1898.

Strymon edwardsii; Klots, 1951.

COMMON NAME

Edwards' Hairstreak (Klots, 1951:136; Pyle, 1981:452)

DISTRIBUTION

From eastern Canada south to Georgia, west to Saskatchewan and

Texas. Ohio records occur in westcentral and northwestern counties

(Fig. 144).

BIOLOGY

Larvae feed on oaks, primarily Scrub Oak (Quercus ilicifolia).

According to Braun (1969:129) the Scrub Oak is said to occur in Ohio, although she found no specimens from Ohio. It appears, then, that

Satyrium edwardsii uses another oak species as a larval foodplant. Pyle

(1981:452) describes the preferred habitat of this species as "dense scrub oak thickets among open woods and sandy barrens." Univoltine, with adults in June and July (Fig. 145). Satyrium calanus falacer (Godart, 1824)

Banded Hairstreak ■

NOMENCLATORIAL AND SYSTEMATIC STATUS

Polymmatus falacer Godart, 1824. [type-locality: near Philadelphia, Pennsylvania]

Satyrium calanus falacer (Godart, 1824). [valid name (Miller & Brown, 1981:101)]

Thecla lorata Grote & Robinson, 1867.

Thecla inornata Grote & Robinson, 1868.

Thecla calanus; Holland, 1898.

Strymon falacer; Klots, 1951.

Strymon borealis Lafontaine, 1969.

Satyrium calanus; Pyle, 1981.

COMMON NAME

Banded Hairstreak (Holland, 1931:234; Pyle, 1981:453)

DISTRIBUTION

From southeastern Canada to Georgia, west to Saskatchewan and

through western Florida and Texas to Utah. The nominate subspecies

is limited to southeastern Georgia and Central Florida. Ohio records

indicate statewide distribution (Fig. 146).

BIOLOGY

Larval foodplants include walnuts (Juglans), hickories (Carya),

and oaks (Quercus) . Univoltine in the North with adults in June and

July. Ohio records extend from mid-May to early August (Fig. 147).

Adults frequent wood edges along roads, clearings, and utility cuts. 229

Satyrium caryaevorum (McDunnough, 1942)

Hickory Hairstreak

NOMENCLATORIAL AND SYSTEMATIC STATUS

Strymon caryaevorum McDunnough, 1942. [type-locality: Merivale, Ontario]

Satyrium caryaevorum (McDunnough, 1942). [valid name (Miller & Brown, 1981:102)]

Strymon caryaevorus; Klots, 1951.

Satyrium caryaevorus; Pyle, 1981.

COMMON NAME

Hickory Hairstreak (Klots, 1951:137; Pyle, 1981:454)

DISTRIBUTION

From Quebec south to Connecticut, west to Ontario, Minnesota, and Iowa. Ohio is at the southern limit of this species, with records limited to the northern quarter of the state, except for a single record for Cedar Bog in Champaign County (Albrecht, 1974), probably representing a relict population in that relict bog (Fig. 148).

BIOLOGY

Larval foodplants include hickories (Carya) and ashes (Fraxinus).

Univoltine, adults in June and July (Fig. 149). Adults are found flying along roadsides and field edges near deciduous woods. Satyrium liparops (Leconte, 1833)

Striped Hairstreak

NOMENCLATORIAL AND SYSTEMATIC STATUS

Thecla liparops Leconte, 1833. [type-locality: Georgia]

Satyrium liparops (Leconte, 1833). [valid name (Miller & Brown, 1981: 102)]

Strymon liparops; Klots, 1951.

COMMON NAME

Striped Hairstreak (Klots, 1951:138; Pyle, 1981:456)

DISTRIBUTION

From southeastern Canada south to Florida, west to Alberta, Colorado, and Arkansas. Ohio records, although including fewer than half of the counties, indicate probable statewide distribution (Fig. 150).

BIOLOGY

Larval foodplants include a variety of woody plants such as oaks

(Quercus), plums (Prunus), willows (Salix), blueberries (Vaccinium), and hollies (Ilex). Univoltine, with adults in June and July in Ohio

(Fig. 151). Habitats include old fields, hedgerows, and woody thickets. Calycopis Scudder, 1876

NOMENCLATORIAL AND SYSTEMATIC STATUS

Calycopis Scudder, 1876. [type-species: Rusticus poeas Htlbner, 1811, by original designation; valid name (Miller & Brown, 1981:104)]

DISCUSSION

This is mainly a tropical genus with two species in America north

of Mexico. One of these, Calycopis cecrops, is found east of the

Mississippi River including the Ohio area, and is discussed below. Calycopis cecrops (Fabricius, 1793)

Red-banded Hairstreak

NOMENCLATORIAL AND SYSTEMATIC STATUS

Hesperia cecrops Fabricius, 1793. [type-locality: "in Indiis"]

Calycopis cecrops (Fabricius, 1793). [valid name (Miller & Brown, 1981:104)]

Resticus poeas Htlbner, 1811.

Thecla cecrops; Holland, 1898.

Strymon cecrops; Klots, 1951.

COMMON NAMES

Red-banded Hairstreak (Pyle, 1981:467)

Least Purple Hair-streak (Comstock & Comstock, 1904:231)

Cecrops Hair-streak (Holland, 1931:238)

DISTRIBUTION

This southern species is found from New Jersey south to Florida,

westward to Kansas and Texas. A line from New Jersey through southern

Ohio is considered by authors to represent the northern limit of this

species' range. Strays have been reported from Michigan and New York.

Ohio records are limited to the southern part of the state (Fig. 152).

BIOLOGY

Larval foodplants include Dwarf Sumac (Rhus copallina) and

(Croton). Habitats include old fields and wooded edges. Multivoltine, with three generations in mid-Atlantic states: April to May, July, and

September to October (Pyle, 1981:468). Ohio records extend from late

May into early October, indicating two generations (Fig. 153). 233 Mitoura Scudder, 1871

NOMENCLATORIAL AND SYSTEMATIC STATUS

Mitoura Scudder, 1871. [type-species: Thecla smilacis Boisduval & Leconte, 1833, by original designation; valid name (Miller & Brown, 1981:106)]

DISCUSSION

This North American genus of seven species reaches its greatest diversity in the west coast states of the United States. A single species, Mitoura gryneus, is found in the Ohio area.

Mitoura gryneus (HUbner, 1819)

Olive Hairstreak

NOMENCLATORIAL STATUS

Lycus gryneus Hllbner, 1819. [type-locality: not stated]

Papilio damon Stoll, 1782: [preoccupied by Papilio damon Dennis & SchiffermUller, 1775]

SYSTEMATIC STATUS

Mitoura gryneus (Htlbner, 1819). [valid name (Miller & Brown, 1981:107)]

Polyommatus damastus Godart, 1824.

Thecla auburniana Harris, 1862.

Thecla patersoniana Brehme, 1907.

Thecla gryneus; Holland, 1931.

Callophrys gryneus; dos Passos, 1970.

COMMON NAME

Olive Hairstreak (Klots, 1951:141; Pyle, 1981:438) 234

DISTRIBUTION

From southern New England to Florida, west to western Ontario,

Kansas, and Texas. In Ohio (Fig. 154), this species has been found in the southwestern quarter of the state.

BIOLOGY

Larval foodplants include Eastern Redcedar (Juniperus virginiana) and Southern Redcedar (Juniperus siliciola) . Of these two trees, only the Eastern Redcedar is known from Ohio and, although it is widely distributed in Ohio, it is most common on calcareous outcrops and limestone soils of the southwestern part of the state (Braun, 1969:66).

The habitats for Mitoura gryneus are dry hillsides, rocky bluffs, and old fields where the larval foodplant is established or colonizing

(Pyle, 1981:439). Adults may be found in the vicinity of redcedars.

Distribution in Ohio, then, appears to be limited by the availability of habitat, more specifically the larval foodplant, above an unknown minimal density level. M. gryneus is bivoltine, with spring and summer generations. Ohio records run from mid-April into August with a distinct break from late May through June (Fig. 155). An ecological study of this species in southern Ohio, with particular attention paid to habitat requirements both in an area of a well-established population and at the edge of its local range, should provide useful information. Such well- established populations exist on and near Fort Hill State Memorial in southern Highland County, and in Adams County. 235 Incisalia Scudder, 1871

NOMENCLATORIAL AND SYSTEMATIC STATUS

Incisalia Scudder, 1871. [type-species: Licus niphon Htlbner, 1823, by original designation; valid name (Miller & Brown, 1981:108)]

DISCUSSION

This genus has eight species in America north of Mexico, commonly

referred to as the elfins. Five of these species are known from the

Ohio area and are discussed below.

Key to the species of Incisalia in the Ohio area.

1. Hind-wing with distinct tail-like projection at end of C ^ , longer than those at ends of other veins; ventral hind-wing with extensive submarginal grayish shading...... 2.

1'. Hind-wing lacking projection as above; ventral hind-wing often lacking shading as above, but if present then ventral fore-wing with dark, submarginal line from anterior to posterior edge...... 3.

2(1). Ventral hind-wing with small but distinct, dark, submarginal spot in cell Cu^-Cu^; ventral hind-wing with projection extending from dark basal area into lighter, postmedian area, with anterior edge of this projection extending farther distad than posterior edge; male with stigma...... irus.

2'. Ventral hind-wing without dark spot as above; ventral hind-wing with basal area projection as above square-like, with posterior edge extending as far distad as anterior edge; male without stigma...... henrici.

3(1'). Ventral fore-wing with submarginal dark spots angulate and pointing basad; hind-wing with outer margin scalloped; ventral wing with irregularly contrasting light and dark colored bars and lines...... niphon.

3'. Ventral fore-wing lacking submarginal dark spots, or if present then rounded, not as above; outer margin of hind-wing smooth, not scalloped, or if scalloped at all then basal area of ventral hind-wing solidly colored without irregular, contrasting dark lines...... 4. 236 4(3'). Ventral hind-wing with postmedian area orange- brown to red-brown, no gray shading; ventral fore­ wing brownish terminally...... augustus.

4'. Ventral hind-wing with grayish postmedian area; ventral fore-wing with distinct terminal gray band...... polios. 237 Incisalia augustus (Kirby, 1837)

Brown Elfin

NOMENCLATORIAL STATUS

Thecla augustus Kirby, 1837. [type-locality: 54° N. Lat.]

SYSTEMATIC STATUS

Incisalia augustus (Kirby, 1837). [valid name (Miller & Brown, 1981:108)]

Thecla augustinus Westwood, 1852. [this name was proposed to replace Thecla augustus Kirby, 1837, wrongly believed to be preoccupied]

Incisalia augustinus; Klots, 1951.

Callophrys augustinus; dos Passos, 1964.

COMMON NAME

Brown Elfin (Klots, 1951:146; Pyle, 1981:424)

DISTRIBUTION

From Newfoundland south to Georgia, west to Alaska and Baja

California. Although considered by some to be the most common of the elfins in North America (Pyle, 1981:425) it is certainly not common in

Ohio. It has been found in only two Ohio counties (Hocking and Vinton) in the unglaciated hills of the southeastern part of the state (Fig. 156).

It may be expected to be found in other areas where the foodplants and habitat are suitable (see below).

BIOLOGY

In the East, this species feeds on blueberries (Vaccinium) and azalea (Rhododendron), and is found in open wooded areas and edges

(Pyle, 1981:425). It is univoltine, with adults in April and May (Fig.

157). 238

Incisalia polios Cook & Watson, 1907

Hoary Elfin

NOMENCLATORIAL AND SYSTEMATIC STATUS

Incisalia polios Cook & Watson, 1907. [type-locality: Lakehurst, New Jersey; valid name (Miller & Brown, 1981:109)]

COMMON NAME

Hoary Elfin (Klots, 1951:147; Pyle, 1981:427)

DISTRIBUTION

From Nova Scotia south to New York, west to Alaska, Washington,

and New Mexico. Although not known from Ohio, this species has been

found in Pennsylvania (Clench, 1971), West Virginia (Drees & Butler,

1978), Michigan (Moore, 1960), and Indiana (Shull & Badger, 1972).

It may be that the required combination of habitat and foodplant

(see below) does not exist in Ohio.

BIOLOGY

Bearberry (Arctostaphylos uva-ursi) is the reported larval food­

plant for this species. In Ohio, Bearberry is very rare, and is known

only from beaches and dunes along Lake Erie in Ashtabula and Erie

counties (Braun, 1969:296). Pyle (1981:427) describes the habitat for

this species as dry, open, often rocky areas. Authors describe it as

only locally common, favoring heathlands and barrens. Ferris & Brown

(1981:251) suggest that this butterfly may be common only sporadically, not every year. It is found above 6000 feet in the Rocky Mountains, and near sea level in the East and Northwest. If the Hoary Elfin is found 239

in Ohio, it will probably be found in a fairly restricted area, with the

foodplant, and removed somewhat from the wetter portions of beaches.

Adults may not be noticed every year even though a population exists.

Incisalia irus (Godart, 1824)

Frosted Elfin

NOMENCLATORIAL AND SYSTEMATIC STATUS

Polyommatus irus Godart, 1824. [type-locality: America]

Incisalia irus (Godart, 1824). [valid name (Miller & Brown, 1981:109)]

Thecla irus; Holland, 1898.

Callophrys irus; dos Passos, 1970.

COMMON NAMES

Frosted Elfin (Klots, 1951:148; Pyle, 1981:428)

Hoary Elfin (Comstock & Comstock, 1904:233; Holland, 1931:226)

DISTRIBUTION

From southeastern Canada south to South Carolina, west to Illinois and eastern Texas. In Ohio it has been found only in Lucas County (Fig.

158) in the northwestern corner of the state.

BIOLOGY

Larval foodplants include lupines (Lupinus) and False Indigo

(). It prefers open, second growth woods, roadsides and open brushy fields, and pine barrens (Pyle, 1981:428). Univoltine, adults from April into early June in Ohio (Fig. 159). 240 Incisalia henrici (Grote & Robinson, 1867)

Henry's Elfin

NOMENCLATORIAL AND SYSTEMATIC STATUS

Thecla henrici Grote & Robinson, 1867. [type-locality: Philadelphia, Pennsylvania]

Incisalia henrici (Grote & Robinson, 1867). [valid name (Miller & Brown, 1981:109)]

Callophrys henrici; dos Passos, 1970.

COMMON NAMES

Henry's Elfin (Klots, 1951:147; Pyle, 1981:429)

Henry's Hair-streak (Holland, 1931:226)

DISTRIBUTION

From Quebec south to Florida, west through Michigan and Illinois

to Nebraska and Texas. Ohio records are limited to the southwestern

quarter of the state (Fig. 160). Forbes (1960:132) describes this

species as "not rare but very local." Shapiro (1974:17) also says

that this species is very localized, and adds that it is poorly under­

stood. Pyle (1981:429) says that in areas where several species of

Incisalia are found together, _I. henrici is likely to be the least

numerous species. Pyle also points out that this species is poorly

known. There, this species should be looked for more carefully, in

all parts of Ohio, and more detailed biological study should be done.

BIOLOGY

Larvae of this species feed on blueberries (Vaccinium), Redbud

(Cercis canadensis), huckleberry (Gaylussacia), and wild plum (Prunus).

Adults are found in brushy areas and forest openings. Univoltine, adults

from late March through May in Ohio (Fig. 161). Incisalia niphon (Hubner, 1823)

Pine Elfin

NOMENCLATORIAL AND SYSTEMATIC STATUS

Licus niphon HUbner, 1823. [type-locality: Florida]

Incisalia niphon (H(lbner, 1823). [valid name (Miller & Brown, 1981:109)]

Thecla niphon; Holland, 1898.

Callophrys niphon; dos Passos, 1970.

COMMON NAMES

Pine Elfin (Klots, 1951:149; Mitchell & Zim, 1964:66)

Eastern Pine Elfin (Pyle, 1981:430)

Banded Elfin (Holland, 1931:227)

DISTRIBUTION

From Nova Scotia south to Florida, west to Manitoba and Texas.

Ohio records are limited to southcentral counties (Fig. 162).

BIOLOGY

Larvae of this species feed on young leaves of various pines

(Pinus). Pyle (1981:431) describes their habitat as pine woods, edges, roadsides, and old fields with young pine growth. Univoltine, adults from April into early June in Ohio (Fig. 163). 242 Euristrymon Clench, 1961

NOMENCLATORIAL AND SYSTEMATIC STATUS

Euristrymon Clench, 1961. [type-species: Papilio favonius Smith, 1797, by original designation; valid name (Miller & Brown, 1981:110)]

DISCUSSION

This holarctic genus has three species in America north of Mexico,

one of which, Euristrymon Ontario, is found in Ohio and is discussed

below.

Euristrymon Ontario (Edwards, 1868)

Northern Hairstreak

NOMENCLATORIAL AND SYSTEMATIC STATUS

Thecla Ontario Edwards, 1868. [type-locality: London, Ontario]

Euristrymon Ontario (Edwards, 1868). [valid name (Miller & Brown, 1981:110)]

Strymon Ontario; Klots, 1951.

COMMON NAMES

Northern Hairstreak (Klots, 1951:135; Pyle, 1981:460)

Ontario Hair-streak (Holland, 1931:235)

DISTRIBUTION

Southern Ontario and Massachusetts south to Georgia, west to

Colorado and Arizona. This species is relatively rare in the Northeast, becoming progressively more common toward the Southwest (Pyle, 1981:461).

Ohio records are from central and southern counties (Fig. 164). 243

BIOLOGY

Larvae feed on various species of oak (Quercus). In the Northeast

this species prefers open woodlands and coastal barrens (Pyle, 1981:461).

Univoltine, adults in June and July in the North, only June in Ohio

(Fig. 165). Parrhasius Hilbner, 1819

NOMENCLATORIAL AND SYSTEMATIC STATUS

Parrhasius HUbner, 1819. [type-species: Papilio polibetes Stoll, 1781, designated by Scudder, 1875; valid name (Miller & Brown, 1981:110)]

Eupsyche Scudder, 1876.

DISCUSSION

This tropical genus has a single species in America north of Mexico.

That species, Parrhasius m-album, ranges from Connecticut south to

Florida, west to Iowa and Texas, thence southward into South America.

Ohio records (Fig. 166) are from the southern half of the state, and

Cuyahoga County in the northeastern part of the state. 245

Parrhasius m-album (Boisduval & Leconte, 1833)

White M Hairstreak

NOMENCLATORIAL AND SYSTEMATIC STATUS

Thecla m-album Boisduval & Leconte, 1833. [type-locality: Georgia]

Parrhasius m-album (Boisduval & Leconte, 1833). [valid name (Miller & Brown, 1981:110)]

Eupsyche m-album; dos Passos, 1964.

Panthiades m-album; dos Passos, 1970.

COMMON NAMES

White M Hairstreak (Klots, 1951:154; Pyle, 1981:422)

Azure Hairstreak (Macy & Shepard, 1941:147)

DISTRIBUTION

See under the generic account above.

BIOLOGY

Larvae of this butterfly feed on various species of oaks (Quercus).

Adults may be found in open areas and trails near or in woods.

Multivoltine, Ohio records include April and May, and August suggesting

two generations (Fig. 167). 246 Strymon Hllbner, 1818

NOMENCLATORIAL AND SYSTEMATIC STATUS

Strymon Hllbner, 1818. [type-species: Strymon melinus Hllbner, 1818, designated by Riley, 1922; valid name (Miller & Brown, 1981:110)]

Callipareus Scudder, 1872.

Callicista Grote, 1873.

Uranotes Scudder, 1876.

DISCUSSION

Thirteen species of this tropical genus are found in America north of Mexico. One of these, Strymon melinus, is known from Ohio, and is discussed below. Strymon melinus Hllbner, 1818

Gray Hairstreak

NOMENCLATORIAL AND SYSTEMATIC STATUS

Strymon melinus Hllbner, 1818. [type-locality: Georgia; valid name (Miller & Brown, 1981:111)]

Thecla hyperici Boisduval & Leconte, 1833.

Thecla melinus; Holland, 1898:242.

COMMON NAMES

Gray Hairstreak (Klots, 1951:134; Pyle, 1981:472)

Common Hair-streak (Holland, 1931:235)

Cotton Square Borer (larva) (Sutherland, 1978:112)

DISTRIBUTION

From eastern Canada south to Florida, west to British Columbia and

Baja California, and south to and Venezuela. Ohio records include much of the state suggesting statewide distribution (Fig. 168).

BIOLOGY

Larval foodplants include Corn (Zea mays), oaks (Quercus), and cotton (Gossyplum). Open areas near woods, including old fields, roadsides, and parks provide habitat for this species. Multivoltine, with two generations in the North and three or more in the South (Pyle,

1981:473). Ohio records extend from mid-April to October, peaking in

May and, to a lesser extent in July (Fig. 169). 248 Erora Scudder, 1872

NOMENCLATORIAL AND SYSTEMATIC STATUS

Erora Scudder, 1872. [type-species: Thecla laeta Edwards, 1862, by original designation; valid name (Miller & Brown, 1981:112)]

DISCUSSION

Two members of this genus occur in America north of Mexico. One

of these, Erora laeta, is found east of the Mississippi River

including Ohio. Erora laeta (Edwards, 1862)

Early Hairstreak

NOMENCLATORIAL AND SYSTEMATIC STATUS

Thecla laeta Edwards, 1862. [type-locality: London, Ontario]

Erora laeta (Edwards, 1862). [valid name (Miller & Brown, 1981:112)]

Thecla clothilde Edwards, 1863.

COMMON NAME

Early Hairstreak (Klots, 1951:142; Pyle, 1981:466)

DISTRIBUTION

From eastern Canada south in the Appalachians to South Carolina, westward to Michigan and Ohio. This species has been found in Ohio in only two counties, Cuyahoga and Highland (Fig. 170).

BIOLOGY

Although the immature states of this species are undescribed, it is believed to feed on beech (Fagus) and hazlenut (Corylus). Adults are found at edges and along trails and roads in beech-maple woods. It is described variously by authors as univoltine or bivoltine, although it is generally agreed that there is at least a partial second generation in summer, in addition to the spring generation. Ohio records occur in

April and July (Fig. 171). 250

POLYOMMATINAE Swainson, 1827.

This taxon, commonly known as the blues, has been referred to by

authors by the junior synonym PLEBEJINAE Tutt, 1909, sometimes

misspelled PLEBEIINAE. The common name comes from the wing color of most males. in wing color is common in this group, with many females having some brown or gray on the wings in addition

to basal blue.

Key to the genera of the POLYOMMATINAE in the eastern United States

and Canada.

1. Hind-wing with small, delicate tail at end of vein Cu£...... Everes.

1'. Hind-wing tailess...... 2.

2(1'). Ventral hind-wing with four relatively large, black, metallic spots near wing margin (southeastern United States, not in Ohio area)...... Brephidium.

2'. Ventral hind-wing lacking four spots as above, with two or fewer metallic spots...... 3.

3(2'). Ventral hind-wing with at least one or two submarginal spots which are metallic or partly metallic...... 4.

3'. Ventral hind-wing lacking metallic scaling, although submarginal spots often present...... 6.

4(3). Ventral hind-wing with distinct metallic scales only in submarginal spots posterior to vein Cu^...... 5.

4'. Ventral hind-wing with some metallic scales in submarginal spots both anterior and posterior to Cu^.....

5(4). Ventral hind-wing with dark spots limited to submarginal area...... .

5'. Ventral hind-wing with median and postmedian as well as submarginal spots...... Hemiargus. 251 6(3'). Ventral wing surface with marginal, submarginal, and postmedian dark spots and lines, none ringed with white...... 7.

6 1. Ventral wing surface lacking marginal and submarginal dark spots and broken lines; dark spots and lines in postmedian area distinctly ringed in white...... Glaucopsyche.

7(6). Ventral hind-wing with submarginal row of orange spots, bordered both distally and proximally with black spots...... Lycaeides.

7'. Ventral hind-wing without row of orange spots as above...... Celastrina. 252

Hemiargus Hllbner, 1818

NOMENCLATORIAL AND SYSTEMATIC STATUS

Hemiargus HUbner, 1818. [type-species: Hemiargus antibubastus HUbner, 1818, designated by Hemming, 1934; valid name (Miller & Brown, 1981:114)]

Cyclargus Nabokov, 1945.

Echinargus Nabokov, 1945.

DISCUSSION

This genus has three species in America north of Mexico, one of which, Hemiargus isola, is found in the Ohio area and is discussed

below.

Hemiargus isola (Reakirt, 1866)

Reakirt's Blue

NOMENCLATORIAL AND SYSTEMATIC STATUS

Lycaena isola Reakirt, 1866. [type-locality: Vera Cruz, Mexico]

Hemiargus isola (Reakirt, 1866).' [valid name (Miller & Brown, 1981:115)]

COMMON NAMES

Reakirt's Blue (Klots, 1951:159; Pyle, 1981:490)

Solitary Blue (Pyle, 1981:490)

DISTRIBUTION

From Michigan south to Mississippi and through Mexico to Costa

Rica, west to Saskatchewan and California. In Ohio it has been found

only in Lucas and Hamilton counties (Fig. 172). 253

BIOLOGY

Larvae feed on various legumes including Alfalfa (Medicago sativa) and Sweet Clover (Melilotis officinalis). Adults have been found in

July in Ohio (Fig. 173). 254 Everes Htlbner, 1819

NOMENCLATORIAL AND SYSTEMATIC STATUS

Everes Htlbner, 1819. [type-species: Papilio amyntas Denis & Schiffermllller, 1775, designated by Scudder, 1871; valid name (Miller & Brown, 1981:115)]

Tiora Evans, 1912.

Ununcula van Eecke, 1915.

DISCUSSION

This holarctic genus, found also in the Indo-Australian Region,

has two North American species. One of these, Everes comyntas, is

common in the Ohio area.

Everes comyntas (Godart, 1824)

Eastern Tailed Blue

NOMENCLATORIAL AND SYSTEMATIC STATUS

Polyommatus comyntas Godart, 1824. [type-locality: North America]

Everes comyntas (Godart, 1824). [valid name (Miller & Brown, 1981:115)]

Lycaena sissona Wright, 1905.

Everes watermani Nakahara, 1925.

COMMON NAMES ' ' ' ( Eastern Tailed Blue (Klots, 1951:163; Pyle, 1981:491)

Tailed Blue (Comstock & Comstock, 1904; Macy & Shepard, 1941:170)

DISTRIBUTION

From southern Canada to Central America covering almost the entire

area east of the Rocky Mountains. Also, it is found west of the Rockies

to the Pacific Ocean at low elevations. Ohio distribution is state­ wide (Fig. 174). 255 BIOLOGY

Larvae feed on many legumes, especially clovers (Trifolium). Adults

are found in many open areas including roadsides, power cuts, meadows,

old fields, gardens, and some crop fields. Multivoltine, with over­

lapping generations, especially in the South. This is one of the most

common butterflies in Ohio, with adults being found continuously from

April into October (Fig. 175). 256 Celastrina Tutt, 1906

NOMENCLATORIAL AND SYSTEMATIC STATUS

Celastrina Tutt, 1906. [type-species: Papilio argiolus Linnaeus, 1758, by original designation; valid name (Miller & Brown, 1981:116)]

DISCUSSION

This Holarctic genus has two Nearctic species, both of which are

found in Ohio and are discussed below.

Key to the species of Celastrina in eastern United States.

I. Male dorsal wing blackish-brown, with occasional blue scaling at base; female dorsal wing pale gray-blue, black-bordered...... ebenina.

II. Dorsal wing deep silvery violet-blue to blanched violet-blue, female fore-wing with black border; summer generation with extensive white basally on dorsal wing, especially female...... lad on.

Celastrina ladon (Cramer, 1780)

Spring Azure

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio ladon Cramer, 1780. [type-locality: "Cape of Good Hope" (in error according to Miller & Brown, 1981:116)]

Celastrina ladon (Cramer, 1780). [valid name (Miller & Brown, 1981)116)]

Argus pseudargiolus Boisduval & Leconte, 1833.

Lycaena violacea Edwards, 1866.

Lycaena pseudargiolus; Holland, 1898.

Lycaenopsis argiolus; Klots, 1951.

Celastrina argiolus; dos Passos, 1964. 257

COMMON NAMES

Spring Azure (Klots, 1951:169; Pyle, 1981:493)

Common Blue (Mitchell & Zim, 1964:71)

DISTRIBUTION

Found throughout the United States and southern Canada, south to

Mexico and Panama in the mountains. Ohio distribution is statewide

(Fig. 176).

BIOLOGY

Larval foodplants include dogwoods (Co m u s ) , viburnum (Viburnum) , and blueberries (Vaccinium) . Adults are found in open wooded areas, edges, roadsides, and brushy areas. Multivoltine, from March to

September in Ohio, (Fig. 177), with perhaps three generations.

Celastrina ebenina Clench, 1972

Dusky Azure

NOMENCLATORIAL AND SYSTEMATIC STATUS

Celastrina ebenina Clench, 1972. [type-locality: Coalburgh, Kanawha County, West Virginia; valid name (Miller & Brown, 1981:117)]

COMMON NAMES

Dusky Azure (Pyle, 1981:495)

Sooty Azure (Pyle, 1981:495)

Dusky Blue (Sites & McPherson, 1980:109)

DISTRIBUTION

From western Pennsylvania south to North Carolina, west to Illinois and Missouri. In Ohio, this species has been found only in Vinton and 258

Highland counties (Fig. 178). The foodplant (see below) is common in unglaciated Ohio in the Southeast (Cusick & Silberhorn, 1977:72).

Further collecting in southern Ohio counties probably will find

Celastrina ebenina in more localities.

BIOLOGY

Larvae feed on 's Beard (Aruncus dioicus). Adults are found in moist, deciduous forests, especially on shaded north-facing slopes.

Univoltine, adults in April and early May (Fig. 179). Glaucopsyche Scudder, 1872

NOMENCLATORIAL AND SYSTEMATIC STATUS

Glaucopsyche Scudder, 1872. [type-species: Polyommatus lygdamus Doubleday, 1841; valid name (Miller & Brown, 1981:119)]

Phaedrotes Scudder, 1876.

DISCUSSION

This genus contains three nearctic species, one of which,

Glaucopsyche xerces, is extinct. Glaucopsyche lygdamus is widespread

in North America and is found in Ohio.

Glaucopsyche lygdamus (Doubleday, 1841)

Silvery Blue

NOMENCLATORIAL AND SYSTEMATIC STATUS

Polyommatus lygdamus Doubleday, 1841. [type-locality: Georgia]

Glaucopsyche lygdamus (Doubleday, 1841). [valid name (Miller & Brown, 1981:120)]

Glaucopsyche nittanyensis Chermock, 1944.

Glaucopsyche boydi Clark, 1948.

Lycaena lygdamas; Holland, 1898. [misspelling]

Lycaena lygdamus; Holland, 1931.

DISTRIBUTION

From Nova Scotia south to Georgia, west to eastern Alaska and

Baja California. It has been found in only four counties in southern

Ohio (Fig. 180).

BIOLOGY

Larvae feed on various legumes including lupine (Lupinus) and vetch

(Vicia). Adults are found in shaded, wooded areas including roadsides 260 and streamsides. Univoltine, adults in late April and May in Ohio

(Fig. 181). 261 Lycaeides Htlbner, 1819

NOMENCLATORIAL AND SYSTEMATIC STATUS

Lycaeides Htlbner, 1819. [type-species: Papilio argyrognomon BergstrHsser, 1779, by designation of I.C.Z.N., 1954, opinion 269; valid name (Miller & Brown, 1981:121)]

DISCUSSION

Of the two nearctic species in this genus one, Lycaeides melissa, is found in Ohio.

Lycaeides melissa (Edwards, 1873)

Orange-bordered Blue

NOMENCLATORIAL AND SYSTEMATIC STATUS

Lycaena melissa Edwards, 1873. [type-locality: Colorado]

Lycaeides melissa (Edwards, 1873). [valid name (Miller & Brown, 1981: 122)]

Plebe.jus melissa; Howe, 1975.

COMMON NAMES

Orange-bordered Blue (Macy & Shepard, 1941:172; Pyle, 1981:506)

Orange-margined Blue (Holland, 1931:264)

Melissa Blue (Klots, 1951:165; Pyle, 1981:506)

DISTRIBUTION

From New Hampshire west to Manitoba and Mexico in the Sierra Nevada, local and spotty in the East. It has been found in only two counties in northern Ohio (Fig. 182).

BIOLOGY

Larval foodplants include lupine (Lupinus) and Alfalfa (Medicago sativa). Lupine is preferred in the East according to Pyle (1981:506), 262 who describes the habitat in the East as sand barrens and dry, open woods. Multivoltine, adults in Ohio in May/June and July/August

(Fig. 183). 263 RIODINIDAE Grote, 1895.

This family, treated as a subfamily of the LYCAENIDAE by some authors, is given full family status here (following Miller & Brown,

1981 and dos Passos, 1964). This is a largely tropical family with greatest diversity of species occuring in the Neotropical Region. There are seven Nearctic genera, one of which (Calephelis) is found in eastern

North America. The common name for this family, the metalmarks, derives from the metallic markings often found on the wings.

Calephelis Grote & Robinson, 1869

NOMENCLATORIAL AND SYSTEMATIC STATUS

Calephelis Grote & Robinson, 1869. [type-species: Erycina virginiensis Gray, 1832, designated by I.C.Z.N., 1966, Opinion 775; valid name (Miller & Brown, 1981:128)]

Nymphidium Fabricius, 1807. [now restricted to some Neotropical species]

Nymphidia Boisduval & Leconte, 1833. [misspelling of Nymphidium Fabricius, 1807]

Calephalis Wright, 1908. [misspelling of Calephelis Grote & Robinson, 1869]

Lephelisca Barnes & Lindsey, 1922. [junior objective synonym of Calephelis Grote & Robinson, 1869]

Callephelis Stichel, 1930. [misspelling of Calephelis Grote & Robinson, 1869]

DISCUSSION

These are predominantly brown butterflies, darker than some of the

Neotropical genera, and with fewer of the metallic colored markings which are often found in the family. Ten species of this New World genus are known from America north of Mexico. Three of these are found in eastern North America and are discussed below. Diagnostic characters 264 for the species are given in the key below, rather than in discussions with each of the species accounts.

Key to the species of Calephelis in the Ohio area.

1. Small, wing span less than 20mm (16-19mm); dorsal wing bright rust to orange...... virginiensis.

1'. Wing span greater than 20mm...... 2.

2(1'). Dorsal wing with basal half somewhat darker in ground color than and contrasting with distal half; dorsal wing dingy orange-brown, without reddish cast; wing span 25-32mm...... borealis.

2'. Dorsal wing with mostly uniform ground color, not contrasting as above; dorsal wing mahogany brown with reddish cast (best seen in fresh specimens); wing span 22-28mm...... muticum.

Calephelis virginiensis (Guerin-Meneville, 1831)

Little Metalmark

NOMENCLATORIAL AND SYSTEMATIC STATUS

Erycina virginiensis Guerin-Meneville, 1831. [type-locality: Georgia]

Calephelis virginiensis (Guerin-Meneville, 1831). [valid name (Miller & Brown, 1981:128)]

Nymphidia pumila Boisduval & Leconte, 1833.

Calephelis louisiana Holland, 1929.

COMMON NAMES

Little Metalmark (Klots, 1951:123; Pyle, 1981:519)

Virginia Metalmark (Pyle, 1981:519)

DISTRIBUTION

From Virginia to Florida, thence west to Arkansas and Texas. Klots

(1951:123) includes Ohio in the distribution of this species, as do 265 Macy & Shepard (1941:140). A careful examination of the specimens in the American Museum of Natural History (where Klots did much of his work) failed to find a specimen of this species from Ohio. No other record has been found of this species in Ohio, neither specimen nor literature reference. It is assumed that Klots got his reference from Macy &

Shepard (op. cit.), and the basis for that reference is unknown. At the present time this species should be regarded as unknown from Ohio, although since it has been reported from West Virginia (Drees & Butler,

1978:202) it may be found in Ohio.

BIOLOGY

Larval foodplants in the East are unknown. In Texas larvae have been reported to feed on Yellow Thistle () (Pyle,

1981:419). Authors report three generations in Virginia, from April to

September. Pyle (1981:519) reports the preferred habitats as pine flats, woodland edges, and damp areas.

Calephelis borealis (Grote & Robinson, 1866)

Northern Metalmark

NOMENCLATORIAL AND SYSTEMATIC STATUS

Nymphidia borealis Grote & Robinson, 1866. [type-locality: near Upper Coldenham, Orange Co., New York]

Calephelis borealis (Grote & Robinson, 1866). [valid name (Miller & Brown, 1981:128)]

Lephelisca borealis; Klots, 1951.

COMMON NAMES

Northern Metalmark (Klots, 1951:123; Pyle, 1981:519)

Large'Metal-mark (Comstock & Comstock, 1904) 266 DISTRIBUTION

From southern New York to Virginia, west through central

Pennsylvania, West Virginia, Ohio, northern Kentucky to Indiana. Pyle

(1981:521) also includes Missouri in the known range. Ohio records come from central and southwestern counties, as well as a few northwestern and southeastern counties (Fig. 184).

BIOLOGY

Larvae feed on Ragwort (Senecio obovatus)■ Univoltine, Ohio records in June and July (Fig. 185). Habitats include open wooded areas, roadsides, and hilly meadows, often near streams.

Calephelis muticum McAlpine, 1937

Swamp Metalmark

NOMENCLATORIAL AND SYSTEMATIC STATUS

Calephelis muticum McAlpine, 1937. [type-locality: Willis, Washtenaw Co., Michigan;, valid name (Miller & Brown, 1981:129)]

Lephelisca muticum; Klots, 1951:123.

COMMON NAME

Swamp Metalmark (Klots, 1951:123; Pyle, 1981:522)

DISTRIBUTION

From western Pennsylvania through Ohio and southern Michigan, northern Indiana and Illinois, to southern Wisconsin and southeastern

Minnesota, and southward to Missouri and northern Arkansas. Ohio records are limited to four counties in the western part of the state (Fig. 186).

BIOLOGY

The Swamp Thistle (Cirsium muticum) is the larval foodplant of this butterfly. Calephelis muticum is found in wet areas including swamps 267 and bogs. A colony of this butterfly exists in Cedar Bog State Memorial, a relict, boreal, alkaline bog in southern Champaign County, where the foodplant is common. This butterfly is univoltine, with Ohio adults from

June to early August (Fig. 187). 268 LIBYTHEIDAE Boisduval, 1836.

This is one of the smallest of butterfly families with only two genera, and less than a dozen species. Nevertheless, it is worldwide in distribution, with representatives in all six zoogeographical regions.

The New World species are now generally placed in the genus Libytheana, while the Old World species remain in the restricted genus Libythea.

The common name for this group, the snout butterflies, comes from the greatly extended, snout-like, labial palps.

Libytheana Michener, 1943

NOMENCLATORIAL AND SYSTEMATIC STATUS

Libytheana Michener, 1943. [type-species: Libythea bachmanii Kirtland, 1851, by original designation; valid name (Miller & Brown, 1981: 132)]

Libythea Fabricius, 1807 (in part). [now restricted to Old World species]

Libythea (Libytheana); Forbes, 1960.

DISCUSSION

In addition to the extended labial palps, the produced, squared off anterior third of the fore-wing outer margin helps to distinguish this genus from all other Nearctic butterflies. Of the two Nearctic species, one, Libytheana bachmanii, is found in the Ohio area. 269

Libytheana bachmanii (Kirtland, 1851)

Snout Butterfly

NOMENCLATORIAL AND SYSTEMATIC STATUS

Libythea bachmanii Kirtland, 1851. [type-locality: Ohio]

Libytheana bachmanii (Kirtland, 1851). [valid name (Miller & Brown, 1981:132)] Libythea bachmanni; Holland, 1898. [misspelling]

Libytheana bachmannii; Klots, 1951. [misspelling]

Libythea (Libytheana) bachmannii; Forbes, 1960. [misspelling]

COMMON NAMES

Snout Butterfly (Klots, 1951:121; Pyle, 1981:534)

Common Snout Butterfly (Mitchell & Zim, 1964:62)

DISTRIBUTION

From Ontario and New England south to Florida, west to the Rocky

Mountains, thence southward into Mexico. Ohio records include much of the northern and western parts of the state (Fig. 188). It has not been taken in southeastern Ohio. Except for Tuscarawas County in eastern Ohio, the range of this species falls entirely within the glaciated part of the state.

BIOLOGY

Larvae of this butterfly feed on hackberries (Celtis). Adults are found in wooded areas, and open, disturbed areas where the foodplant is present. Multivoltine, Ohio records extend from June to October with a peak in July (Fig. 189). The length of the flight period suggests at least a partial second generation in Ohio. 270 HELICONIIDAE Swainson, 1827.

This mainly Neotropical family is considered by some authors to be

a subfamily of the NYMPHALIDAE. It is treated here as a family after

Miller & Brown, 1981:133. The HELICONIIDAE are sometimes referred to as

the longwings because of the elongate fore-wing of most species. In the

Tropics, members of this family are often involved in mimicry complexes

(both Batesian and MUllerian) with other butterflies and sometimes moths.

Four genera occur in America north of Mexico, and two of these,

Heliconius Kluk, 1802 and Agrauli s Boisduval & Leconte, 1833, are found

in the eastern United States, primarily in Florida. Agraulis vanillae

(Linnaeus, 1758), commonly known as the Gulf Fritillary (Ferris & Brown,

1981:292) has been reported as far north as Long Island (Shapiro, 1974)

as a stray, and Clench (1971) includes it in his list for Pennsylvania without specifying the part of the state involved. Also, it has been

reported from Illinois, where reproducing populations may occur (Irwin

& Downey, 1973). There are two records for Indiana from southwestern

counties (Shull & Badger, 1972). This species is easily recognized in

the Ohio area by its size (wingspan about 60-75mm), and the dorsal fore­ wing with orange-brown to reddish-brown ground color and two black- bordered white spots in the discal cell. Agraulis vanillae may be found

in Ohio as a stray, and should be looked for in southwestern Ohio in

sunny, open areas "with abundant flowers" (Pyle, 1981:539). 271 NYMPHALIDAE Swainson, 1827.

This is the largest of the butterfly families in terms of worldwide

species diversity, and is represented in all six of the zoogeographical

regions. In Ohio it is third in number of species, behind the

HESPERIIDAE and LYCAENIDAE. Of the 38 nymphalid genera presently

recognized in America north of Mexico (Miller & Brown, 1981), 12 are

found in Ohio. Of the 130 species, 24 are found in Ohio. In addition

to the characters used in the key to the families of the PAPILI0N0IDEA

above, these butterflies have vestigial prothoracic legs in both males

and females, a character they share with the SATYRIDAE and DANAIDAE.

Many of the NYMPHALIDAE are some shade of orange, and many adults are

of a medium size for butterflies with wingspans of about 35-75mm.

Key to the Nearctic subfamilies of the NYMPHALIDAE.

1. Eyes usually hairy; if not then dorsal hind-wing with two submarginal eyespots, anterior one very large, about twice the head size...... NYMPHALINAE.

1'. Eyes not hairy; if spots are present on hind-wing they are without contrasting centers (as in eyes) and are smaller than the head...... 2.

2(1'). Hind-wing with tail-like extensions at end of vein M ^ ...... 3.

2'. Hind-wing without tails as above...... 4.

3(2). Hind-wing tail as long or longer than the combined length of thorax and abdomen...... MARPESIINAE.

3'. Hind-wing tail shorter than the combined length of thorax and abdomen...... CHARAXIINAE.

4(2'). Mesotibia well-spined dorsally...... 5.

4'. Mesotibia dorsally unspined, or with only a few, weak spines...... 6. 272 5(4). Hind-wing with humeral vein arising opposite Rs...... LIMENITIDINAE.

5'. Hind-wing with humeral vein not arising opposite Rs, usually arising distal to Rs ...... ARGYNNINAE.

6(4’). Antennal club ventrally with one well-developed ridge that extends onto antennal shaft, or no ridge....MELITAEINAE.

6'. Antennal club with three ridges as above...... APATURINAE. 273 ARGYNNINAE Blanchard, 1827.

This group, commonly known as the fritillaries, is represented by

three genera in Ohio, including eight species. Some are known

individually as silverspots because of the ventral wing markings.

Key to the genera of the ARGYNNINAE in eastern North America.

1. Antennal club ventrally with three well-developed ridges that extend onto the shaft...... 2.

1'. Antennal club with one ridge as above, or lacking such ridges...... Clossiana.

2(1). Fore-wing with vein R_ arising from discal cell; ventral hind-wing usually with postmedian silver patches (except in Speyeria diana)...... Speyeria.

2'. Fore-wing with vein R„ stalked with beyond discal cell; ventral nind-wing without silver patches....Euptoieta. 274 Euptoieta Doubleday, 1848

NOMENCLATORIAL AND SYSTEMATIC STATUS

Euptoieta Doubleday, 1848. [type-species: Papilio claudia Cramer, 1775, designated by Scudder, 1871; valid name (Miller & Brown, 1981:135)]

Eupineta Edwards, 1873. [misspelling]

DISCUSSION

This mainly Neotropical genus has two species in America, north of

Mexico. One of these, Euptoieta claudia, is found in Ohio and is discussed here.

Euptoieta claudia (Cramer, 1775)

Variegated Fritillary

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio claudia Cramer, 1775. [type-locality: Jamaica]

Euptoieta claudia (Cramer, 1775). [valid name (Miller & Brown, 1981:135)]

Papilio daunius Herbst, 1798.

Euptoieta dodgei Gunder, 1927.

COMMON NAME

Variegated Fritillary (Klots, 1951:84; Pyle, 1981:542)

DISTRIBUTION

From Quebec south to Florida, west to British Columbia and southern

California. However, resident populations exist only in the southern part of the range. Each year adults migrate northward in the spring.

Ohic records occur mainly in the northern and western parts of the state

(Fig. 190). 275 BIOLOGY

Larvae feed on pansies (Viola) , flax (Lina), and plantain

(Plantago), and other plants. Adults are found flying in open areas including meadows, old fields, and forest openings. Multivoltine, with continuous generations from spring into fall in the southern part of the range. Since this species overwinters in the adult stage (Howe, 1975:

210), adult activity in Ohio could be expected in late March or April if the species were resident throughout the year. However, only a few adults have been found in Ohio before July (see Fig. 191). This species is probably not resident in Ohio. Its presence here is probably dependent upon annual migration into the state from more southern areas. 276 Speyeria Scudder, 1871

NOMENCLATORIAL AND SYSTEMATIC STATUS

Speyeria Scudder, 1871. [type-species: Papilio idalia Drury, 1773, by original designation; valid name (Miller & Brown, 1981:136)]

Argynnis Fabricius, 1807. [now limited to Palearctic species]

Semnopsyche Scudder, 1875.

DISCUSSION

This is a wholly Nearctic genus. Central Mexico is the southern limit of its range (Howe, 1975:210). Thirteen species are recognized by Miller & Brown (1981) although some authors split these into seventeen or more. Five species are found in Ohio and are discussed below. Larvae of all of these species in the Ohio area feed on various violets (Viola), both cultivated and wild. Klots (1951:85) says that members of this genus are univoltine, although the somewhat lengthened emergence period, extended over a period of several months, may give the appearance in some cases of a second generation. A characteristic of the distribution of this genus in Ohio is the apparent lack or scarcity of at least some of its species in the westcentral part of the state. No particular factor is known which could lead to this sort of pattern. Of course, this may be an artifact of collecting . Further study of this genus in Ohio, particularly in the westcentral portion of the state, should be carried out. 277 Key to the species of Speyeria in the eastern United States.

1. Hind-wing ventral surface with basal two-thirds nearly uniform in color (brown in females and dark orange in males)...... djana.

1'. Hind-wing ventral surface not marked as above,

with whitish or silvery spots on basal area...... 2 .

2(1'). Hind-wing dorsal surface blueish-black in distal half with submarginal and median rows of light-colored spots idalia.

2'. Hind-wing dorsal surface not marked as above; rows of light-colored spots absent, at least part of distal half of wing surface fulvous, although outer margin may be nearly solid black...... 3.

3(2'). Dorsal wing with outer margin almost solid black, with some minor interruptions, but with yellow on hind-wing less than 50% of border area...... atlantis.

3'. Dorsal wing with outer margin marked with varying amounts of black, but with yellow on more than 50% of border area of hind-wing...... 4.

4(3'). Ventral hind-wing with postmedian row of silver spots, with black spots at outer ends of silver spots; . ventral hind-wing with submarginal yellow band between two outer rows of silver spots narrower than width of postmedian row of silver spots...... aphrodite.

4'. Ventral hind-wing with postmedian row of silver spots as above lacking black spots, at most with light brownish scaling at outer ends of silver spots; ventral hind-wing with submarginal yellow band as above as wide or wider than width of postmedian row of silver spots...... cybele.

Speyeria diana (Cramer, 1775)

Diana

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio diana Cramer, 1775. [type-locality: Jamestown, Virginia]

Speyeria diana (Cramer, 1775). [valid name (Miller & Brown, 1981:136)]

Argynnis diana; Holland, 1931. 2.78

COMMON NAMES

Diana (Klots, 1951:86; Pyle, 1981:136)

Diana Fritillary (Holland, 1931:84)

DISTRIBUTION

From Maryland south to northern Georgia, west to eastern

Oklahoma and Louisiana. In Ohio this species has been found in only

three counties: Hamilton in the Southwest; Franklin in the central part

of the state; Medina County in the Northeast (a single record for

each county, not including the observation for Hamilton County cited

below). The most recent record for Ohio found during this study was

from July, 1949 (Hamilton County). Pyle (1981:545) points out that the

range of this species has decreased because of forest cutting. This

species of butterfly may now be extirpated from Ohio.

BIOLOGY

Adults are found in deciduous and pine woodlands near streams.

Univoltine, from June to September. The only date found for Ohio, other

than the July date mentioned above, is 17 July 1932. This date was for

the Franklin County record, and was with a worn specimen. The Medina

County record, reported by Hine (1898a), was for a badly worn specimen.

Hine (op. cit.) also reported that this species was often observed across

the Ohio River from Cincinnati by Charles Dury. More recently,

approximately forty years ago, this species was occasionally seen in

Hamilton County, Ohio, near Cincinnati (Charles Oehler, personal

communication). Therefore, it is likely that this species was resident 279 in Hamilton County, although it is probably no longer there. The two

Ohio specimens from the more northern counties probably are strays.

The blue female of this dimorphic species mimics the Pipevine

Swallowtail (Battus philenor). Speyeria cybele (Fabricius, 1775)

Great Spangled Fritillary

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio cybele Fabricius, 1775. [type-locality: New York]

Speyeria cybele (Fabricius, 1775). [valid name (Miller & Brown, 1981:136)]

Papilio daphnis Cramer, 1775.

Argynnis cybele; Holland, 1898.

COMMON NAME

Great Spangled Fritillary (Holland, 1931:86, Pyle, 1981:545)

DISTRIBUTION

From Quebec south to northern Georgia, thence westward to British

Columbia and New Mexico. Ohio records indicate statewide distribution

(Fig. 192).

BIOLOGY

Adults of this species are found flying in open wooded areas, wet meadows, old fields, and roadsides. Although Ohio records (Fig. 193), which extend from May to October, indicate a partial second generation, this species is considered to be univoltine (see under generic discussion above).

Speyeria aphrodite (Fabricius, 1787)

Aphrodite

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio aphrodite Fabricius, 1787. [type-locality: New York]

Speyeria aphrodite (Fabricius, 1787). [valid name (Miller & Brown, 1981:137)] 281 Argynnis aphrodite; Holland, 1898.

COMMON NAMES

Aphrodite (Klots, 1951:87; Pyle, 1981:546)

Silver-spot Fritillary (Comstock & Comstock, 1904:117)

Aphrodite Fritillary (Holland, 1931:88)

DISTRIBUTION

From Nova Scotia south to Georgia, west to British Columbia and

Arizona. Ohio records occur mainly in the eastern half of the state, with scattered records in the western half of the state (Fig. 194).

BIOLOGY

This species is found in wet meadows, and in open, wooded areas.

It is univoltine, with adults between June and early October in

Ohio (Fig. 195).

Speyeria idalia (Drury, 1773)

Regal Fritillary

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio idalia Drury, 1773. [type-locality: New York]

Speyeria idalia (Drury, 1773). [valid name (Miller & Brown, 1981:137)]

Argynnis astarte Fisher, 1858. [preoccupied by Argynnis astarte Doubleday, 1847]

Argynnis ashtaroth Fisher, 1859.

Argynnis idalia; Holland, 1898.

COMMON NAME

Regal Fritillary (Pyle, 1981:547) 282 DISTRIBUTION

From Ontario south to North Carolina, west to Manitoba and eastern Colorado. Ohio records include much of the state, except for the westcentral part of the state (Fig. 196).

BIOLOGY

As with Speyeria aphrodite this is a butterfly of open grassy areas and wet meadows. Univoltine, with Ohio records between June and

September (Fig. 197).

Speyeria atlantis (Edwards, 1862)

Atlantis Fritillary

NOMENCLATORIAL AND SYSTEMATIC STATUS

Argynnis atlantis Edwards, 1862. [type-locality: Hunter, Green Co, New York]

Speyeria atlantis (Edwards, 1862). [valid name (Miller & Brown, 1981: 143)]

COMMON NAMES

Atlantis Fritillary (Klots, 1951:86;'Pyle, 1981:554)

Mountain Silver-spot (Holland, 1931:90)

DISTRIBUTION

From eastern Canada south to Virginia in the Appalachians, thence west to eastern Alaska and the southern Rocky Mountains. Klots (1951:86) describes this as a Canadian Zone species. This is not an accurate characterization of its distribution in Ohio, since in the Life Zone system Ohio is mainly Upper Austral, with a small part of the northeastern part of the state described as Transition. Ohio records are few and are limited to central, northern, and eastern counties

(Fig. 198). Ohio is probably near the southern limit of this species

BIOLOGY

This species is found in forest openings, and along streams and in wet meadows. Univoltine, Ohio adults are found from early June to August (Fig. 199). Speyeria atlantis probably is resident in Ohio rather than being a stray. Adults are found in early June in Ohio which is earlier than reported by some authors for other parts of the

Ohio area (July 1st in Michigan by Moore, 1960:7, and July for the overall range by Pyle, 1981:555). June is the earliest month reported for Pennsylvania by Tietz (1952:7) and for New York (Shapiro

1974:10). Since individuals of this species hibernate as a newly hatched larva (Klots, 1951:86) it seems likely that individuals in

Ohio are developing through emergence in place, rather than flying into this area following emergence. 284 Clossiana Reuss, 1920

NOMENCLATORIAL AND SYSTEMATIC STATUS

Clossiana Reuss, 1920. [type-species: Papilio selene Denis & SchiffermUller, 1775, by original designation; valid name (Miller & Brown, 1981:148)]

Argynnis Fabricius, 1807.

Brenthis HUbner, 1819.

Boloria Moore, 1900.

DISCUSSION

Approximately twelve species of this genus are reported to be in

America north of Mexico. Two of these are known from the Ohio area and

are discussed below. As with the genus Speyeria, the two members of this

genus found in Ohio feed on violets (Viola) .

Key to species of the genus Clossiana in the Ohio area.

1. Ventral hind-wing with prominent silver patches; dorsal fore-wing with row of orange marginal spots...... selene myrina.

1'. Ventral hind-wing lacking silver patches; dorsal fore-wing without marginal row of orange spots...... bellona.

Clossiana selene myrina (Cramer, 1777)

Silver-bordered Fritillary

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio myrina Cramer, 1777. [type-locality: vicinity New York, N.Y.]

Clossiana selene myrina (Cramer, 1777). [valid name (Miller & Brown, 1981:148)]

Argynnis myrissa Godart, 1824.

Brenthis myrinna; Holland, 1898.

Brenthis marilandica Clark, 1941. 285 Boloria selene myrina; Klots, 1951.

Brenthis selene race myrina; Forbes, 1960.

COMMON NAME

Silver-bordered Fritillary (Klots, 1951:89; Pyle, 1981:559)

DISTRIBUTION

From Newfoundland south to North Carolina (in the mountains),

thence westward through Ohio and Illinois to Oregon, and southward in

the West in spring-fed bogs. Ohio is at the southern limit of the range

of this species and records are limited to northern and western counties

(Fig. 200). This is a Holarctic species. The nominate subspecies is

Palearctic.

BIOLOGY

This species is found in wet meadows and bogs, often near wooded

areas, sometimes straying onto roadways and dry edges. Univoltine in

the northern and multivoltine in the southern parts of its range, with

Ohio adults from May to September (Fig. 201). The September records in

Ohio, which follow a break in August, may represent a second generation, although more data are needed. A study of voltinism in this species in

the Ohio area could well be made at Cedar Bog State Memorial in

Champaign County in westcentral Ohio. This area contains a variety of habitats including a relict, alkaline, boreal bog with a population of Clossiana selene myrina. The entire Ohio seasonal range of this species is represented at Cedar Bog. Clossiana bellona (Fabricius, 1775)

Meadow Fritillary

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio bellona Fabricius, 1775). [type-locality: America]

Clossiana bellona (Fabricius, 1775). [valid name (Miller & Brown, 1981:149)]

Brenthis bellona; Holland, 1898.

Argynnis ammiralis Hemming, 1933.

Boloria toddi ammiralis; Klots, 1951.

Boloria bellona; Howe, 1975.

COMMON NAMES

Meadow Fritillary (Pyle, 1981:560)

Eastern Meadow Fritillary (Mitchell & Zim, 1964)

DISTRIBUTION

From Quebec south to North Carolina, west to British Columbia and

Colorado. Statewide in Ohio (Fig. 202).

BIOLOGY

Adults are found flying in meadows and along roadsides near streams

or swampy areas. Multivoltine, with up to three generations between

May & September according to Pyle (1981:561). Ohio records extend

from April to October (Fig. 203), with peaks in early May, June, and

September. The length of the flight period, along with the continued

relatively high level of activity and the apparent three peaks,

suggest three generations in Ohio. MELITAEINAE Grote, 1897.

This worldwide subfamily has twelve genera in America north of

Mexico (Miller & Brown, 1981), three of which are found in the East.

As a group, these are commonly called crescents, crescentspots,

checkerspots, and patches* by various authors. These names come from

the more or less mottled appearance of the wings in most species,

although there are no common patterns of wing markings in the group.

Key to the genera of the MELITAEINAE in eastern North America.

1. Dorsal wing with ground color black, with two marginal rows of orange spots...... Euphydryas.

1'. Dorsal wing with ground color orange or orange and dark brown; marginal row of orange spots

present or absent...... 2 .

2(1'). Ventral fore-wing with median black patch in

cell Cu 2 ~ 2 A; ventral hind-wing mainly yellowish with fine brownish lines...... Phyciodes.

2'. Ventral fore-wing without black patch as above, or if present, then ventral hind-wing with median band of white; ventral hind-wing strongly mottled with white, brown, and sometimes red and yellow...... Charidryas. 288 Charidryas Scudder, 1871

NOMENCLATORIAL AND SYSTEMATIC STATUS

Charidryas Scudder, 1871. [type-species: nycteis Doubleday & Hewitson, 1847, by original designation; valid name (Miller & Brown, 1981:155)]

Melitaea Fabricius, 1807.

Phyciodes Hdbner, 1819.

Chlosyne Butler, 1870.

DISCUSSION

This is a genus of nine species in America north of Mexico. Three of these are found east of the Mississippi River including the Ohio area, and are discussed below.

Key to the species of Charidryas in the Ohio area (after Sites Si

McPherson, 1980).

1. Ventral hind-wing with submarginal row of cream-colored spots, and marginal orange band...... harrisii.

1'. Ventral hind-wing without spots and band as above...... 2.

2(1'). Ventral hind-wing with white postmedian band, containing a dark zig-zag line, with acute distal angles of zig-zag line on veins...... gorgone.

2'. Ventral hind-wing usually without dark zig-zag line as above, but if line present then distal angles rounded and between veins...... nycteis. 289

Charidryas gorgone (Hllbner, 1810)

Gorgone Checkerspot

NOMENCLATORIAL AND SYSTEMATIC STATUS

Dryas gorgone HUbner, 1810. [type-locality: probably coastal Georgia (Miller & Brown, 1981:155)]

Charidryas gorgone (Httbner, 1810). [valid name (Miller & Brown, 1981: 155)]

Melitaea gorgone; Klots, 1951.

COMMON NAME

Gorgone Checkerspot (Klots, 1951:95; Pyle, 1981:580)

DISTRIBUTION

From New York (only a single locality according to Shapiro, 1974:

11) to Georgia, west to Alberta and Utah. This species is not known from Ohio. In the area around Ohio it has been found in West Virginia

(Drees & Butler, 1978), Kentucky (Covell, 1974), Indiana (Shull & Badger,

1972), and Michigan (Moore, 1960). In the first three states above it is known from only one or two records each. In Michigan it has been found in five different counties in both the upper and lower peninsulas. Its early adult flight dates in Michigan (May) correspond with the earliest flight dates for the species as a whole in the northern part of its range (Pyle, 1981:581), and suggest a permanent, breeding population in Michigan. The few records further south in the

Ohio area, for which the dates are not known, may represent strays. 290

BIOLOGY

Larval foodplants include sunflowers (Helianthus), ragweed

( trifida), and other composites. Habitats include roadsides near grassy areas, old fields, and waterways (streams and canals).

This butterfly may move into new areas forming large colonies, and

its members may vary dramatically from year to year (Pyle, 1981:581).

Irwing & Downey (1973:28) report that this species seems to have

"undergone a dramatic population explosion" in Illinois since the early 1960's. Considering the dates of adult capture, this species probably is a reproducing resident in Illinois. Adults may be found taking at goldenrod (Solidago) flowers. Univoltine northward

(May to July), bivoltine in the southern part of its range (May to

September). Although it seems to be rare in the eastern part of its range, C. gorgone should be looked for in Ohio. 291 Charidryas nycteis (Doubleday & Hewitson, 1847)

Silvery Checkerspot

NOMENCLATORIAL AND SYSTEMATIC STATUS

Melitaea nycteis Doubleday & Hewitson, 1847. [type-locality: middle states [USA]]

Charidryas nycteis (Doubleday & Hewitson, 1847). [valid name (Miller & Brown, 1981:155)]

Melitaea oenone Scudder, 1864.

Phyciodes nycteis; Holland, 1898.

Chlosyne nycteis; dos Passos, 1969.

COMMON NAMES

Silvery Checkerspot (Klots, 1951:98)

Silver Crescent (Comstock & Comstock, 1904:127)

Nycteis Crescent-spot (Holland, 1931:136)

Silvery Crescentspot (Pyle, 1981:581)

DISTRIBUTION

From Quebec south to Georgia, west to Saskatchewan and Arizona.

Statewide in Ohio (Fig. 204).

BIOLOGY

Larval foodplants include Coneflower (Rudbeckia laciniata), asters

(Aster), and sunflowers (Helianthus). Habitats include wet meadows, streams and roadways near wooded areas, and open deciduous woods.

Bivoltine in Ohio with adults from May to September (Fig. 205). Charidryas harrisii (Scudder, 1864) 2 9

Harris' Checkerspot

NOMENCLATORIAL AND SYSTEMATIC STATUS

Phyciodes harrisii Scudder, 1864. [type-locality: Norway, Maine]

Charidryas harrisii (Scudder, 1864). [valid name (Miller & Brown, 1981:156)]

Melitaea harrisi; Holland, 1898.

Chlosyne harrisii; dos Passos, 1964.

COMMON NAMES

Harris' Checkerspot (Klots, 1951:94; Pyle, 1981:583)

Harris' Butterfly (Comstock & Comstock, 1904)

DISTRIBUTION

From Nova Scotia south to New Jersey, west to Saskatchewan, North

Dakota, and Illinois. Except for Hamilton County in the southwestern part of the state, Ohio records are limited to northern counties

(Fig. 206). Ohio is near the southern limit of this species.

BIOLOGY

As with some other members of this genus, Charidryas harrisii feeds as larva on asters (Aster) and probably some other composites.

Pyle (1981:583) describes the habitat for this species as "moist meadows, edges of bogs and marshes, openings in forests, and old fields." Shapiro (1974:11) says that it is more common in acid-soil areas [than in other soil types]. If the soil acidity is a significant ecological factor in the distribution of this species in Ohio, then it should be found more frequently and in more localities in the eastern half of the state (see discussion under Hesperia sassacus above). Univoltine, with adults in June and July. Adults in Ohio have been found only in June (Fig. 207). 293 Phyciodes HUbner, 1819

NOMENCLATORIAL AND SYSTEMATIC STATUS

Phyciodes Htlbner, 1819. [type-species: Papilio cocyta Cramer, 1871, designated by Scudder, 1871; valid name (Miller & Brown, 1981:161)]

Tritanassa Forbes, 1945 (in part).

DISCUSSION

This New World genus has nine Nearctic species, two of which are

found in the Ohio area. Both of these use asters (Aster) as larval

foodplants.

Key to the species of Phyciodes in the Ohio area.

1. Ventral fore-wing with subapical black patch on costal margin equal to or larger than median black patch on posterior margin; ventral hind-wing with marginal patch dark brown or black...... tharos.

1*. Ventral fore-wing with subapical patch as above smaller than posterior patch; ventral hind-wing with marginal patch light brown or lacking as an area contrasting with ground color...... batesii.

Phyciodes tharos (Drury, 1773)

Pearl Crescent

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio tharos Drury, 1773. [type-locality: New York, New York (Miller & Brown, 1981:160)]

Phyciodes tharos (Drury, 1773). [valid name (Miller & Brown, 1981:160)]

Papilio cocyta Cramer, 1777.

Papilio euclea BergstrHsser, 1780.

Melitaea selenis Kirby, 1837. 294 COMMON NAMES

Pearl Crescent (Klots, 1951:100)

Pearl Crescentspot (Pyle, 1981:573)

DISTRIBUTION

From eastern Canada south to Florida, west to the Yukon and

southeastern California. Statewide in Ohio (Fig. 208).

BIOLOGY

This is a very common butterfly of old fields, roadsides,

streamsides, and most other open areas. Multivoltine, in Ohio adults

occur from April to November representing two or more overlapping

generations (Fig. 209).

Phyciodes batesii (Reakirt, 1865)

Tawny Crescent

NOMENCLATORIAL AND SYSTEMATIC STATUS

Eresia batesii Reakirt, 1865. [type-locality: restricted to Gloucester, New Jersey (Klots, 1951:100)]

Phyciodes batesii (Reakirt, 1865). [valid name (Miller & Brown, 1981:160)]

COMMON NAMES

Tawny Crescent (Klots, 1951:100)

Tawny Crescentspot (Pyle, 1981:574)

Bates' Crescentspot (Pyle, 1981:574)

DISTRIBUTION

From Nova Scotia south in the Appalachians to Georgia, west to the

Northwest Territories and Nebraska. This species has not been found in

Ohio. It has been reported from Pennsylvania (Clench, 1971), West 295

Virginia (Drees & Butler, 1978), Indiana (Shull & Badger, 1972), and

Michigan (Moore, 1960). It may be expected to be found in Ohio. Its similarity to Phyciodes tharos may have led to its being overlooked in the past.

BIOLOGY

Habitats include both moist open areas, and dry, open slopes.

Univoltine, adults in June and July. Euphydryas Scudder, 1871

NOMENCLATORIAL AND SYSTEMATIC STATUS

Euphydryas. Scudder, 1871. [type-species: Papilio phaeton Drury, 1773, by original designation; valid name (Miller & Brown, 1981:169)]

Melitaea Fabricius, 1807 (in part).

DISCUSSION

This Holarctic genus, as presently understood, has a single species in America north of Mexico (Miller & Brown, 1981). That single species,

Euphydryas phaeton, is discussed below. The other, western species, formerly included in this genus, are now placed in Hypodryas and

Occidryas.

Euphydryas phaeton (Drury, 1773)

Baltimore

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio phaeton Drury, 1773. [type-locality: New York]

Euphydryas phaeton (Drury, 1773). [valid name (Miller & Brown, 1981:169)]

Melitaea phaeton; Holland, 1898.

Melitaea phaetona Holland, 1931. [misspelling]

Melitaea schausi Clark, 1927.

Euphydryas borealis Chermock & Chermock, 1940.

COMMON NAME

Baltimore (Klots, 1951:93; Pyle, 1981:602)

Baltimore Checkerspot (Howe, 1975:175) 297

DISTRIBUTION

From Nova Scotia south to Georgia, west to Manitoba and Arkansas.

This species has been found in northwestern, northeastern, and

Southcentral Ohio (Fig. 210).

BIOLOGY

Turtlehead (Chelone glabra), the larval food plant usually found

to be used by this species in Ohio, grows in wet meadows and swampy

areas, also the primary habitat for Euphydryas phaeton in Ohio.

Mature larvae also have been found in large numbers on lousewort

(Pedicularis sp.), in a "bog" in Williams County (Price, 1970:11).

Other larval foodplants, reported for this butterfly by authors, are

considered rare or questionable by some entomologists (Klots, 1951:93;

David J. Horn, 1982, personal communication). NYMPHALINAE Swainson, 1827.

As presently conceived, this subfamily combines the VANESSINAE

of some other authors with the NYMPHALINAE. This is a worldwide

subfamily, mainly occuring in temperate parts of the Holarctic Region.

Nine genera are presently recognized in America north of Mexico. Five

of these are found in the East and are discussed here.

Key to the genera of the NYMPHALINAE in eastern North America (except

Florida).

1. Dorsal hind-wing with a large, submarginal eyespot, larger than head...... Junonia.

1'. Dorsal hind-wing without eyespot as above, if spots are present they are solid in color and are smaller

than the head...... 2 .

2(1'). Hind-wing with tail-like extension at end of vein M^...... 3.

2 ’. Hind-wing with outer margin mainly smooth, without extension as above...... Vanessa.

3(2). Dorsal fore-wing with at least five black spots in discal area in addition to larger, dark brown or black patches along costal edge; dorsal wing without white markings; fore-wing with inner margin concave...... Polygonia.

3'. Dorsal fore-wing as above without dark spots, or if present then dorsal wing with white spot; fore-wing with inner margin straight...... 4.

4(3'). Dorsal fore-wing with wide, submarginal orange band Aglais.

4'. Dorsal fore-wing without wide, orange band...... Nymphalis. 299

Polygonia Httbner, 1819

NOMENCLATORIAL AND SYSTEMATIC STATUS

Polygonia Hllbner, 1819. [type-species: Papilio c-aureum Linnaeus, 1758, designated by Scudder, 1871; valid name (Miller & Brown, 1981:170)]

Eugonia Hllbner, 1819.

Comma Rennie, 1832.

Grapta Kirby, 1837.

DISCUSSION

Members of this Holarctic genus are commonly referred to as the angle wings. Of the ten species in America north of Mexico, four are found in the Ohio area and are discussed below. A fifth species,

Polygonia satyrus (Edwards, 1869), is northern and western in distribution and does not enter the area covered by this study, although said to do so by some collectors. Members of this genus overwinter as adults which are sometimes seen flying during warm, sunny winter days (Klots, 1951:102). Seasonal dimorphism is common among adults of this genus, with adults of the overwintering generation often lighter colored than those of the summer generation(s) (Klots, op. cit.). 300

Key to the species of Polygonia in the Ohio area.

1. Hind-wing with tail about three times as long as narrowest width; central white line of ventral hind-wing broken with dot posterior to line, giving appearance of question mark on right wing; dorsal fore-wing with dark postmedian spots in cells

M2-M3’ ^3-^u1’ Cu^-C^, Cu2 ~ 2 A ...... interrogationis.

1'. Hind-wing with tail only about twice as long as width; central white line of ventral hind-wing complete, giving appearance of comma on right wing; dorsal fore-wing with postmedian spots in cells.

M^-Cu^, Cu^-C^, Cu 2 ~2 A, absent in 2.

2(1'). Ventral hind-wing with central white mark enlarged or hooked at both ends; dorsal fore-wing usually with two dark postmedian spots in cell C u ^ - C ^ ...... 3.

2'. Ventral hind-wing with central white mark with both ends tapered to near point, neither enlarged nor hooked; dorsal fore-wing with only one dark postmedian spot in cell Cuj...... progne.

3(2). Ventral hind-wing usually with two submarginal rows of greenish spots; dorsal wing surface sometimes with greenish sheen; outer margin of wing very irregular giving sharply scalloped appearance at ends of most veins; dark discal spots of dorsal fore­ wing large, each almost as large as spot in

cell Cu 2 ~ 2 A ...... faunus.

3'. Wings without greenish spots or sheen as above; outer margin of wing less sharply irregular, with scalloped appearance soft, not sharp; dark discal spots of dorsal fore-wing noticeably smaller

than spot in cell Cu 2 ~ 2 A ...... comma. 301

Polygonia interrogationis (Fabricius, 1798)

Question Mark

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio interrogationis Fabricius, 1798. [type-locality: America]

Polygonia interrogationis (Fabricius, 1798). [valid name: (dos Passos, 1964:79)]

Grapta fabricii Edwards, 1870.

Grapta interrogationis; Holland, 1898.

Polygonia interrogantionis Miller & Brown, 1981. [misspelling]

COMMON NAMES

Question Mark (Klots, 1951:102; Pyle, 1981:609)

Violet-tip (Comstock & Comstock, 1904:134)

Question-sign (Holland, 1931:149)

DISTRIBUTION

From eastern Canada south to Florida, west to Saskatchewan and

Texas, and south into Mexico. Ohio records indicate that probably it has statewide distribution (Fig. 212).

BIOLOGY

Larvae of this butterfly feed on various plants including nettles

(Urticaceae), hops (Humulus), hackberries (Celtis)s basswood (Tilia), and elms (Ulmus). Adults are found flying along roadways and waterways, in open grassy areas, and old fields. Multivoltine, adults in Ohio from April to October with at least two overlapping generations

Fig. 213). 302

Polygonia comma (Harris, 1842)

Comma

NOMENCLATORIAL AND SYSTEMATIC STATUS

Vanessa comma Harris, 1842. [type-locality: New England]

Polygonia comma (Harris, 1842). [valid name (Miller & Brown, 1981:170)]

Grapta harrisii Edwards, 1873.

Grapta comma; Holland, 1898.

COMMON NAMES

Comma (Pyle, 1981:610)

Hop-merchant (Comstock & Comstock, 1904:140)

Comma Butterfly (Holland, 1931:149)

DISTRIBUTION

From eastern Canada south to Georgia, west to Saskatchewan, eastern

Colorado, and Mississippi. Ohio records indicate statewide distribution

(Fig. 214).

BIOLOGY

Larval foodplants include hops (Humulus), nettles (Urtica), and elms (Ulmus) . Habitats used by this species are similar to the preceding, Polygonia interrogationis. Authors report two generations in the North, three in the South. Ohio records extend from March through

October with two or three overlapping generations (Fig. 215). 303

Polygonia faunus (Edwards, 1862)

Green Comma

NOMENCLATORIAL AND SYSTEMATIC STATUS

Grapta faunus Edwards, 1862. [type-locality: Hunter, Green Co., New York]

Polygonia faunus (Edwards, 1862). [valid name (Miller & Brown, 1981: 171)]

COMMON NAMES

Green Comma (Klots, 1951:104; Pyle, 1981:612)

Faunus Anglewing (Pyle, 1981:612)

DISTRIBUTION

From eastern Canada south in the Appalachians to northern Georgia, west to British Columbia and California. This species was reported from

Cincinnati in Hamilton County in the southwestern part of Ohio by Hine

(1898b). This record remains unsubstantiated by either later authors or specimens, and could represent an error in identification. However,

Polygonia faunus has been reported from Pennsylvania (Clench, 1971),

West Virginia (Drees & Butler, 1978), Kentucky (Covell, 1974), and

Michigan (Moore, 1960), and it is possible that it could occur in Ohio.

However, since this species is essentially limited to the Canadian

Zone (Klots, 1951:104) or the Canadian and Transition Zones (David J.

Horn, 1982, personal communication), if found in Ohio it will probably be in the northeastern corner of the state. That part of Ohio is considered by some authors (eg. Klots, 1951) to be in the Transition

Zone. 304

BIOLOGY

Larval foodplants include (Betula), (Alnus), and willow

(Salix). Adults are found in sunny glades, roadsides, and especially in coniferous woods (Pyle, 1981:613). Shapiro (1974:12) found it absent from areas of low relief and deciduous woods, and found it favoring elevated areas and coniferous forest. Horn (David J. Horn,

1982, personal communication) doubts that it will be found in Ohio because of its association with Canadian and Transition Zone areas, and its likelihood not to stray. 305

Polygonia progne (Cramer, 1776)

Gray Comma

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio progne Cramer, 1776. [type-locality: "Jamaica and New York" (Miller & Brown, 1981:171)]

Polygonia progne (Cramer, 1776). [valid name (Miller & Brown, 1981:171)]

Grapta g-argenteum Kirby, 1837.

Grapta progne; Holland, 1898.

COMMON NAME

Gray Comma (Klots, 1951:104; Pyle, 1981:618)

DISTRIBUTION

From Nova Scotia south to North Carolina in the mountains, west to British Columbia, Wyoming and Missouri. Ohio records include northern, central, and southern counties (Fig. 216).

BIOLOGY

Larval foodplants include currants and gooseberries (Ribes) and occaisonally elms (Ulmus). Habitats include wooded areas, especially along trails and in clearings. Bivoltine, adults from April to October.

Adults have been found in April, June, and July in Ohio (Fig. 217), but are too few to suggest conclusions. Further study is needed on this species in Ohio. Such a study could be carried out in the wooded area immediately west of Lake Hope in Vinton County, where a population exists. 306

Nymphalis Kluk, 1802.

NOMENCLATORIAL AND SYSTEMATIC STATUS'

Nymphalis Kluk, 1802. [type-species: Papilio polychloros Linnaeus, 1758, designated by Hemming, 1933; valid name (Miller & Brown, 1981:172)]

Vanessa Fabricius, 1807.

Aglais Dalman, 1816.

Scudderia Grote, 1873. [preoccupied by Scudderia Stal, 1873]

Euvanessa Scudder, 1889.

DISCUSSION

This genus is mainly northern and Holarctic. Three species in the area of America north of Mexico are now included in this genus (Miller &

Brown, 1981). Two of these are found in Ohio and are discussed below.

As with Nymphalis, members of this genus and Aglais following over­ winter as adults, which may be found flying on warm, winter days.

Key to the species of Nymphalis in the Ohio area.

1. Dorsal hind-wing with black spot near costal edge bordered distally by white patch...... vau-album.

1'. Dorsal hind-wing with neither black spot nor white patch as above...... antiopa. 307

Nymphalis vau-album (Denis & Schiff ermtlller, 1775)

Compton Tortoiseshell

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio vau-album Denis & Schif fermllller, 1775. [type-locality: presumably near Vienna [Austria] (Miller & Brown, 1981:172)]

Nymphalis vau-album (Denis & Schiffermliller, 1775). [valid name (Miller & Brown, 1981:172)]

Vanessa j-album; Holland, 1898.

Aglais „j-album; Holland, 1931.

COMMON NAMES

Compton Tortoiseshell (Klots, 1951:105; Pyle, 1981:619)

Compton Tortoise (Holland, 1931:152)

DISTRIBUTION

From eastern Canada south to North Carolina (in the mountains), west to Alaska, Oregon, Colorado, and Missouri. Ohio records occur in northern, central, and southwestern counties (Fig. 218).

BIOLOGY

Larval foodplants include (Betula) and willows (Salix).

Adults are found in dense, deciduous wooded areas, particularly in openings and along pathways. Univoltine, adults from March to

September in Ohio (Fig. 219). Since this species overwinters as an

adult which becomes active in the spring, it is doubtful that this

is a breeding resident of northern Ohio. The single March record

(28 March 1981, Hocking County, David J. Horn, collector) is a very worn specimen which appears to have overwintered as an adult. 308

Nymphalis antiopa (Linnaeus, 1758)

Mourning Cloak

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio antiopa Linnaeus, 1758. [type-locality: Sweden]

Nymphalis antiopa (Linnaeus, 1758). [valid name (Miller & Brown, 1981: 172)]

Papilio pompadour Pollich, 1779.

Vanessa antiopa; Holland, 1898.

Aglais antiopa; Holland, 1931.

COMMON NAMES

Mourning Cloak (Klots, 1951:106; Pyle, 1981:621)

Camberwell Beauty (Holland, 1931) [mainly in England]

DISTRIBUTION

This species is found throughout much of North America southward northern South America, and in the Palearctic Region. It probably is statewide in Ohio (Fig. 220).

BIOLOGY

Larval foodplants include willow (Salix), elm (Ulmus), cottonwood

(Populus), and hackberry (Celtis). Habitats include open areas and edges

near woodlands, old fields, and watercourses. Multivoltine, adults

active in Ohio from April to October (Fig. 221). Since overwintering

adults may fly on warm, winter days, adults may be seen in all months. 309

Aglais Dalman, 1816

NOMENCLATORIAL AND SYSTEMATIC STATUS

Aglais Dalman, 1816. [type-species: Papilio urticae Linnaeus, 1758; valid name (Miller & Brown, 1981:173)]

Nymphalis Kluk, 1802. [in Nearctic Region]

Vanessa Fabricius, 1807. [in Nearctic Region]

DISCUSSION

This Holarctic genus has a single species in North America, Aglais milberti, which ranges from eastern Canada south to West Virginia, and westward to the Northwest Territories and California. Ohio is at the southern limit of its distribution. Within Ohio records are limited to northern and western counties (Fig. 222).

Aglais milberti (Godart, 1819)

Milbert's Tortoiseshell

NOMENCLATORIAL AND SYSTEMATIC STATUS

Vanessa milberti Godart, 1819. [type-locality: United States]

Aglais milberti (Godart, 1819). [valid name (Miller & Brown, 1981:173)]

Nymphalis milberti; Klots, 1951.

COMMON NAMES

Milbert's Tortoiseshell (Klots, 1951:106; Pyle, 1981:622)

American Tortoise-shell (Comstock & Comstock, 1904:151)

DISTRIBUTION

See under the discussion of the genus Aglais above. 310

BIOLOGY

Larvae of this species feed on nettles (Urtica). Adults are found flying along roadsides, trails, waterways, and woodland openings.

Multivoltine, Ohio records extend from April through October (Fig. 223), with probably two generations. 311 Vanessa Fabricius, 1807

NOMENCLATORIAL. AND SYSTEMATIC STATUS

Vanessa Fabricius, 1807. [type-species: Papilio atalanta Linnaeus, 1758; valid name (Miller & Brown, 1981)]

Nymphalis Latreille, 1804. [preoccupied by Nymphalis Kluk, 1802]

Cynthia Fabricius, 1807.

Pyrameis Htlbner, 1819.

Ammiralis Rennie, 1832.

Phanessa Sodovskii, 1837.

Neopyrameis Scudder, 1889.

DISCUSSION

This genus is almost worldwide in distribution with four Nearctic

species. Three of these are found in Ohio and are discussed below.

Key to the species of Vanessa in the Ohio area.

1. Dorsal fore-wing with reddish-orange bar crossing wing in discal area, bar clearly defined and with nearly parallel sides...... atalanta.

1'. Dorsal fore-wing without bar as above...... 2.

2(1'). Ventral hind-wing with four or five submarginal eyespots...... cardui.

2'. Ventral hind-wing with two submarginal eyespots...... virginiensis. Vanessa virginiensis (Drury, 1773)

American Painted Lady

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio virginiensis Drury, 1773. [type-locality: Virginia]

Vanessa virginiensis (Drury, 1773). [valid name (Miller & Brown, 1981: 173)]

Papilio huntera Fabricius, 1775.

Papilio .jole Cramer, 1776.

Pyrameis huntera; Holland, 1898.

COMMON NAMES

American Painted Lady (Klots, 1951:108; Pyle, 1981:623)

Painted Beauty (Comstock & Comstock, 1904:156)

Hunter's Butterfly (Holland, 1931:154)

Virginia Lady (Pyle, 1981:623)

DISTRIBUTION

From eastern Canada south to Florida, west to British Columbia and Mexico. Statewide in Ohio (Fig. 224).

BIOLOGY

Larvae of this species feed on various composites. Habitats include sunny open areas such as old fields, waterways, and roadsides.

Bivoltine in Ohio, with adults between April and October (Fig. 225). Vanessa cardui (Linnaeus, 1758)

Painted Lady

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio cardui Linnaeus, 1758. [type-locality: Sweden]

Vanessa cardui (Linnaeus, 1758). [valid name (Miller & Brown, 1981:174)]

Pyrameis cardui; Holland, 1898.

COMMON NAMES

Painted Lady (Holland, 1931:154; Pyle, 1981:625)

Cosmopolite (Comstock & Comstock, 1904:158)

Thistle Butterfly (Holland, 1931:154)

DISTRIBUTION

From eastern Canada south to Florida, west to the Northwest

Territories and California, south to Panama. Virtually all of North

America except for the Arctic is included in the range of this species.

Vanessa cardui has been found throughout much of Ohio and is probably statewide in distribution (Fig. 226).

BIOLOGY

Larvae of this butterfly feed on many composites, but seem to prefer thistles (Cirsium). This species is found in almost any open, sunny area, especially where flowers are present. This species is a strong migrator with adults moving northward in the early spring of each year.

This species probably overwinters in Ohio, with adults being found from April through October (Fig. 227), especially from June onward.

There appear to be two generations in Ohio. Adults do not migrate southward in the fall (Pyle, 1981:626). Vanessa atalanta (Linnaeus, 1758)

Red Admiral

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio atalanta Linnaeus, 1758. [type-locality: Sweden]

Vanessa atalanta (Linnaeus, 1758). [valid name (Miller & Brown, 1981: 174)]

Pyrameis atalanta; Holland, 1898.

COMMON NAMES

Red Admiral (Pyle, 1981:627)

Alderman (Pyle, 1981:627)

DISTRIBUTION

This Holarctic species is found throughout most of North America from subarctic Canada to Central America. It is statewide in Ohio

(Fig. 228).

BIOLOGY

Larvae feed on various members of the Urticaceae, especially the nettles (Urtica). As with other members of its genus, adults of this species may be found in almost any open, sunny area. Pyle (1981:628) points out that this species overwinters only in areas with mild winters, with adults moving northward in the spring, and some flying southward in the fall. This species probably overwinters in Ohio since adults appear as early as March (Fig. 229), and since the March and April records are from both northern and southern counties. Bivoltine, with

Ohio adults from March to October. 315 Junonia HUbner, 1819

NOMENCLATORIAL AND SYSTEMATIC STATUS

Junonia Htlbner, 1819. [type-species: Papilio lavinia Cramer, 1775; valid name (Miller & Brown, 1981:175)]

Precis HUbner, 1819.

Alcyoneis Htlbner, 1819.

Aresta Billberg, 1820.

Dunonia Mabille, 1876. [Misspelling]

DISCUSSION

This mainly tropical genus has three species in America north of

Mexico, one which, Junonia coenia, is found in Ohio. Junonia coenia is

found from Ontario and New England south to Florida, west to Oregon and

Arizona. In Ohio it has been found in most parts of the state (Fig.

230), and probably will be found throughout the state.

Junonia coenia Hllbner, 1822

Buckeye

NOMENCLATORIAL AND SYSTEMATIC STATUS

Junonia coenia HUbner, 1822. [type-locality: not stated, but usually considered to be the United States; valid name (Miller & Brown, 1981:175)]

Precis lavinia coenia; Klots, 1951.

COMMON NAME

Buckeye (Klots, 1951:108; Pyle, 1981:630)

BIOLOGY

Larvae of this species feed on a wide variety of plants including plantain (Plantago), snapdragon (Antirrhinum), and false loosestrife

(Ludvigia). Adults are found in many kinds of open areas including 316 roadsides, old fields, and waterways. This species overwinters as an

adult, and is not able to overwinter in the northern part of its range.

Adults migrate northward each year. In Ohio there is a single record for

April, and then additional records from July into October (Fig. 231).

Since adults would be expected in the spring from a species that

overwinters as an adult, this species probably does not overwinter in

Ohio. 317 LIMENITIDINAE Butler, 1869.

This worldwide taxon has 11 genera in America north of Mexico.

One of these, Basilarchia, is found east of the Mississippi River

and north of Florida.

Basilarchia Scudder, 1871

NOMENCLATORIAL AND SYSTEMATIC STATUS

Basilarchia Scudder, 1871. [type-species: Papilio astyanax Fabricius, 1775, by original designation; valid name (Miller & Brown, 1981: 177)]

Limenitis Fabricius, 1807.

Callianira Hllbner, 1819. [preoccupied by Callianira Peron & Leseur, 1810]

Nymphalis Felder, 1861. [preoccupied by Nymphalis Kluk, 1802]

DISCUSSION

Of the five Nearctic species in this genus three are found in the

Ohio area. Two of these mimic other butterflies which are distasteful to avian predators, thereby gaining protection from the same predators.

Basilarchia archippus mimics the Monarch (Danaus plexippus), and

Basilarchia astyanax mimics the Pipevine Swallowtail (Battus philenor).

Two of the taxa in this genus, 13. arthemis and 13. astyanax, have been considered by some authors to be subspecies of the same species, JB. arthemis (Miller & Brown, 1981 consider them a single species, while both

Pyle, 1981 and Ferris & Brown, 1981 treat them as separate species).

They are treated as subspecies of the same species here. Key to the species of Basilarchia in the eastern United States

(except Florida).

1. Dorsal fore-wing mainly red-orange, with some darker markings...... archippus.

11. Dorsal fore-wing dark blue to black 2.

2(1’). Dorsal fore-wing solid color in discal area without contrasting band (as below)...... arthemis astyanax.

2'. Dorsal fore-wing with white, strongly contrasting band crossing discal area...... arthemis arthemis.

Basilarchia arthemis arthemis (Drury, 1773,)

White Admiral

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio arthemis Drury, 1773. [type-locality: New York]

Basilarchia arthemis arthemis (Drury, 1773). [valid name (Miller & Brown, 1981:177)]

Papilio lamina Fabricius, 1793.

Limenitis arthemis; Klots, 1951.

COMMON NAMES

White Admiral (Klots, 1951:115; Pyle, 1981:635)

Banded Purple (Klots, 1951:115; Pyle, 1981:635)

DISTRIBUTION

From Maine south to New York, west to Manitoba and Minnesota, also in Alaska and British Columbia. This northern subspecies has not been confirmed with specimens for Ohio, although several literature records exist. Kirtland (1854) reported it from Poland in Mahoning

County; Porter (1965) reported it from Seneca County; and Hine (1898a) said that it had been "taken in Ohio by Lynds Jones of Oberlin" 319

(Lorain County). All of these localities are in the northeastern quarter of the state. Presumably some collectors recently have seen specimens of this subspecies which were collected in northeastern Ohio

(Eric H. Metzler, 1982, personal communication), although none has been available for examination during this study. Basilarchia arthemis arthemis has been reported from Pennsylvania (Clench, 1971), West

Virginia (Drees & Butler, 1978), Kentucky (Covell, 1974), and

Michigan (Moore, 1960). Both West Virginia and Kentucky have only one or two records each. This subspecies, then, may be found in Ohio.

However, specimens are needed to confirm its presence in Ohio. This and the following subspecies are mainly allopatric, but they regular­ ly interbreed in nature where their ranges overlap, producing partially banded individuals of both sexes which are completely viable (Platt, 1975). This contact zone between the two subspecies runs from New Jersey, through New York and northern Pennsylvania, and may include northeastern Ohio.

BIOLOGY

Larval foodplants include birches (Betula), willows (Salix ) , and poplars (Populus). Adults are found in deciduous forests, in open areas and along edges. Bivoltine in the southern part of its range, which includes Ohio. 320

Basilarchia arthemis astyanax (Fabricius, 1775)

Red Spotted Purple

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio astyanax Fabricius, 1775. [type-locality: America]

Basilarchia arthemis astyanax (Fabricius, 1775). [valid name (Miller & Brown, 1981:178)]

Limenitis arthemis astyanax; Klots, 1951.

Limenitis astyanax; Howe, 1975.

COMMON NAME

Red Spotted Purple (Klots, 1951:116)

DISTRIBUTION

From New England south to Florida, west to the Dakotas, Arizona,

and Mexico. Ohio records for this subspecies occur throughout the

state (Fig. 232).

BIOLOGY

Larval foodplants include willows (Salix), poplars (Populus), and cherries (Prunus). Habitats include open wooded areas and edges, old fields, and roadways. Multivoltine with up to three generations in the

South. Probably two generations in Ohio with adults from May to

October (Fig. 233). 321

Basilarchia archippus (Cramer, 1776)

Viceroy

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio archippus Cramer, 1776. [type-locality: Jamaica]

Basilarchia archippus (Cramer, 1776). [valid name (Miller & Brown, 1981:178)]

Nymphalis disippe Godart, 1824.

Basilarchia disippus; Holland, 1898.

Limenitis archippus; Klots, 1951.

COMMON NAMES

Viceroy (Klots, 1951:114; Pyle, 1981:637)

Mimic (Klots, 1951:114)

DISTRIBUTION

This species is found throughout most of North America south of

Hudson Bay and east of the Great Basin, and in the eastern parts of the

Pacific Coast states. Ohio records indicate statewide distribution

(Fig. 234).

BIOLOGY

Larval foodplants include poplars and aspens (Populus), apples

(Malus), and cherries and plums (Prunus). Adults are found flying in many areas, including paths and roadways, streamsides, old fields, and wood edges. Multivoltine, with Ohio records from May to October, indicating two generations (Fig. 235). MARPESIINAE Aurivillius, 1898. t This subfamily is mainly tropical, with a single genus in

America north of Mexico. This genus, Marpesia Hilbner, 1818, is limited

in the United States to the southernmost states, Texas and Florida.

CHARAXINAE Guenee, 1865.

The CHARAXINAE are mainly tropical and are commonly known as the

leaf wing butterflies. This name comes from the leaf-like appearance

of the ventral wing surface, especially when the wings are held together

over the back of the butterfly. There are two Nearctic genera, one of which (Anaea) is found in eastern United States, and is discussed below. 323 Anaea Htlbner, 1819

NOMENCLATORIAL AND SYSTEMATIC STATUS

Anaea Htlbner, 1819. [type-species: Papilio troglodyta Fabricius, 1775; valid name (Miller & Brown, 1981:186)]

Pyrrhanaea RHber, 1888.

DISCUSSION

Of the two Nearctic species in this genus, one, Anaea andria, is

found in Ohio. Its overall range extends from Michigan south to Georgia,

west to Wyoming and Mexico. In Ohio it has been found in central and

southern counties (Fig. 236).

Anaea andria Scudder, 1875

Goatweed Butterfly

NOMENCLATORIAL AND SYSTEMATIC STATUS

Anaea andria Scudder, 1875. [type-locality: not stated]

Pyrrhanaea andria; Holland, 1898.

Anaea glycerinus Stichel, 1939.

COMMON NAME

Goatweed Butterfly (Klots, 1951:117; Pyle, 1981:652)

BIOLOGY

The common name for this butterfly comes from its larval foodplants, •

the goatweed (Croton). Adults may be found along roadways, waterways, and wood edges, and in old fields and other open areas. Multivoltine with overlapping generations (Pyle, 1981:652). Ohio records are spotty, extending from March to July (Fig. 237). More data are needed for this species in Ohio. 324 APATURINAE Boisduval, 1840.

These are known as the hackberry butterflies because of the larval foodplants (Celtis) where known. Two genera occur in America north of

Mexico, one of which (Asterocampa) is found east of the Mississippi River.

Asterocampa RBber, 1915

NOMENCLATORIAL AND SYSTEMATIC STATUS

Asterocampa Rflber, 1915. [type-species: celtis Boisduval & Leconte, 1834; valid name (Miller & Brown, 1981:187)]

Apatura Fabricius, 1807.

Chlorippe Boisduval, 1870.

Celtiphaga Barnes & Lindsey, 1922.

DISCUSSION

Two species of Asterocampa are found in Ohio, both of which are found in areas where the foodplants, hackberries, are found.

Key to the species of Asterocampa in the Ohio area.

1. Dorsal fore-wing with large, submarginal, black-centered eyespot...... celtis.

1'. Dorsal fore-wing without eyespot as above...... clyton. Asterocampa celtis (Boisduval & Leconte, 1834)

Hackberry Butterfly

NOMENCLATORIAL AND SYSTEMATIC STATUS

Apatura celtis Boisduval & Leconte, 1834. [type-locality: Georgia]

Asterocampa celtis (Boisduval & Leconte, 1834). [valid name (Miller & Brown, 1981:187)]

Chlorippe celtis; Holland, 1898.

COMMON NAMES

Hackberry Butterfly (Klots, 1951:120; Pyle, 1981:653)

Gray Emperor (Comstock & Comstock, 1904)

Hackberry Emperor (Pyle, 1981:653)

DISTRIBUTION

From Massachusetts south to Florida, west to the Dakotas and

eastern Texas. Ohio records are mainly from the western half of the

state (Fig. 238). This may be an artifact of collecting patterns.

Further collecting in eastern Ohio is needed for this and the following

species (Asterocampa clyton) to determine more accurately their status

in the state of Ohio.

BIOLOGY

Bivoltine in the northern part of the range, adults from June to

August (Pyle, 1981:654). In Ohio adults are found from June to

September indicating two generations (Fig. 239). 326 Asterocampa clyton (Boisduval & Leconte, 1833)

Tawny Emperor

NOMENCLATORIAL AND SYSTEMATIC STATUS

Apatura clyton Boisduval & Leconte, 1833. [type-locality: United State]

Asterocampa clyton (Boisduval & Leconte, 1833). [valid name: Miller & Brown, 1981:188)]

Apatura ocellata Edwards, 1876.

Chlorippe clyton; Holland, 1898.

COMMON NAME

Tawny Emperor (Klots, 1951:119; Pyle, 1981:657)

DISTRIBUTION

From New England south to Georgia, west to Wisconsin and eastern

Texas. As with the preceding species, A. clyton is found mainly in the western half of Ohio (Fig. 240).

BIOLOGY

Univoltine in Ohio, adults from June through August (Fig. 241). 327 SATYRIDAE Boisduval, 1833.

Members of this family are nearly cosmopolitan, relative to worldwide butterfly distribution, being absent from some oceanic islands

and the polar regions. Diversity and proliferation are greatest in the

Tropics (Miller, 1968). Of the seven subfamilies recognized by Miller

(op. cit.) two are found in Ohio: the ELYMNIINAE and SATYRINAE. Ten

species are known from Ohio. Characters used to distinguish the taxa within this family below are given in the keys, rather than in the

individual discussions.

Key to the subfamilies of the SATYRIDAE in the Ohio area.

1. Dorsal hind-wing with more than four ocelli; ventral fore-wing with four distinct ocelli...... ELYMNIINAE.

1'. Dorsal hind-wing with fewer than four ocelli; ventral fore-wing without four distinct ocelli...... SATYRINAE.

ELYMNIINAE Herrich-SchHffer, 1864.

This subfamily is widely represented in the Tropics other than the

the Neotropical Region. Two genera are found in North America, both of which are represented in Ohio.

Key to the genera of the ELYMNIINAE in America north of Mexico.

1. Hind-wing outer margin strongly scalloped; wings with ventral black spots at least four times diameter of their white pupils; antenna with more than 40 segments Enodia.

1'. Hind-wing outer margin only slightly scalloped; wings with ventral black spots at most three times diameter of their white pupils; antenna with about 35 segments....Satyrodes. Enodia HUbner, 1819

NOMENCLATORIAL AND SYSTEMATIC STATUS

Enodia HUbner, 1819. [type-species: Oreas andromacha Hllbner, 1809, designated by Scudder, 1871; valid name (Miller & Brown, 1981:189)]

Lethe HUbner, 1819 (in part).

Lethe.(Enodia); Klots, 1951.

DISCUSSION

This genus includes three Nearctic species, all of which are known

from the Ohio area, and are discussed below. The larvae are known to feed on several grasses, including cane (Arundinaria).

Key to the species of Enodia in the Ohio area.

1. Fore-wing (male) dorsally with dark, androconial patches of scales between veins distad to discal cell, wing produced apically; ventral fore-wing with five well-developed submarginal ocelli...... creola.

1'. Fore-wing (male) as above lacking androconial patches, rounded apically; ventral fore-wing with four well-

developed submarginal ocelli, one in cell Cu2 ~ 2 A tiny or lacking...... 2.

2(1’). Ventral hind-wing with circular white pupils in ocelli; antenna with ringed knob...... anthedon.

2'. Ventral hind-wing with ocellar pupils elongate (although the anterior one may be more or less rounded); antenna with knob unringed...... Portlandia. Enodia portlandia (Fabricius, 1781)

Pearly Eye

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio portlandia Fabricius, 1781. [type-locality: "America meridionali", probably Georgia (Miller & Brown, 1981:189)]

Enodia portlandia (Fabricius, 1781). [valid name (Miller & Brown, 1981: 189)]

Oreas andromacha HUbner, 1809.

Enodia androcardia HUbner, 1821.

Debia portlandia; Holland, 1898 (in part).

COMMON NAME

Pearly Eye (Pyle, 1981: 663)

DISTRIBUTION

From southern Virginia south to Florida, west to southern Illinois and Texas. Although not known from Ohio, it may be found in the southern part of the state where the foodplant (Arundinaria) is found.

References in the literature to this species in Ohio probably refer to

Enodia anthedon, although this cannot be known for certain. IS. anthedon was formerly considered to be a subspecies of portlandia but is now considered to be a separate species. Specimens from Ohio examined during this study are anthedon.

BIOLOGY

Larvae of Enodia portlandia feed on cane (Arundinaria). Pyle (1981:

663) describes the preferred habitat as moist, shady woodland spots near stands of cane. Braun (1969:69) has reported (Large

Cane) from seven counties across southern Ohio, where it grows on ravine slopes and flats, and in moist or swampy areas. Multivoltine, with up to three generations between May and September. Enodia anthedon Clark, 1936

Northern Pearly Eye

NOMENCLATORIAL AND SYSTEMATIC STATUS

Enodia anthedon Clark, 1936. [type-locality: Lava, Sullivan Co., New York; valid name (Miller & Brown, 1981:189)]

Enodia borealis Clark, 1936.

Lethe portlandia anthedon; Howe, 1975.

COMMON NAME

Northern Pearly Eye (Pyle, 1981:664)

DISTRIBUTION

From Maine south to Virginia, west to Manitoba and northern Arkansas.

Ohio appears to be near the southern limit of this species. Ohio records

indicate statewide distribution (Fig. 242).

BIOLOGY

Larvae of this butterfly feed on grasses, including

(dropseed) and erectum (Long-awned Wood Grass). Adults are

found flying in open areas within forests and at wood edges. Pyle (1981:

664) describes this species as univoltine, while Emmel, T.C. (1975:110)

says that there may be a second generation in the southern part of its

range. Ohio records extend from May to early September, with a decrease

in record numbers in July, thereby indicating two generations (Fig.243).

However, separation of the records into southern Ohio and northern Ohio

(based on the criteria given in the discussion on geographical patterns below) suggests a modified interpretation of the data (see Fig. 243).

Although more data are needed, it appears that there is a single generation in the northern part of the state, and two in the southern. Enodia creola (Skinner, 1897)

Creole Pearly Eye

NOMENCLATORIAL AND SYSTEMATIC STATUS

Debis creola Skinner, 1897. [type-locality: Opelousas, Louisiana]

Enodia creola (Skinner, 1897). [valid name (Miller & Brown, 1981:189)]

Lethe creola; Klots, 1951.

COMMON NAME

Creole Pearly Eye (Pyle, 1981:665)

DISTRIBUTION

From southeastern Virginia south to Georgia, west to southern

Indiana and Illinois, and Texas. Although not reported from Ohio, it may be at some time in the future.

BIOLOGY

Larvae of this butterfly feed on Arundinaria (cane), one species of which is found in southern Ohio (see under Enodia portlandia above).

Bivoltine, adults between late May and October. Pyle (1981:665) describes the habitat for this species as shaded hardwood, especially in wet areas near the foodplant. 332 Satyrodes Scudder, 1875

NOMENCLATORIAL AND SYSTEMATIC STATUS

Satyrodes Scudder, 1875. [type-species: Papilio eurydice Johansson, 1763, by original designation; valid name (Miller & Brown, 1981:189)]

Argus Scopoli, 1777. [junior homonym of Argus Bohadsch, 1761]

Hipparchia Fabricius, 1807.

DISCUSSION

Both species assigned to this North American genus are found in

Ohio, and are discussed below. Some authors still place these species, along with those included in Enodia above, in the genus Lethe (used in the broad sense). The arrangement of Miller & Brown (1981) is followed here.

The two species of this genus, until fairly recently, were considered to be subspecies of a single species, Satyrodes eurydice. Therefore, most literature records, where the two are not treated as separate taxa, are unreliable in separating the two. Also, relatively few specimens have been available for this study (about 30). Therefore, a single distribution map for Ohio and a single seasonal graph are used to include both species (Figs. 244 & 245), rather than attempting to separate them on the basis of insufficient data. Further study to determine the distribution of these two species in Ohio is needed. Taken together, the two species are found primarily in the northern part of the state.

Key to the Ohio area species of Satyrodes.

1. Hind-wing with ventral, postmedian line forming distinct "V's" pointing basad in cells Cu^-Cu„ and Cu2~2A; ventral hind-wing with outer half brown, yellowish-brown, or yellowish-white...... eurydice. 333 1'. Hind-wing with ventral, postmedian line as above bent somewhat basad, but not forming distinct "V's"; ventral hind-wing surface as above violet-brown, sometimes gray-brown...... appalachia.

Satyrodes eurydice (Johansson, 1763)

Eyed Brown

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio eurydice Johansson, 1763. [type-locality: Morris Arboretum, Philadelphia Co., Pennsylvania, designated by Carde, Shapiro & Clench, 1970]

Satyrodes eurydice (Johansson, 1763). [valid name (Miller & Brown, 1981:190)]

Papilio canthus Linnaeus, 1767.

Hipparchia transmontana Gosse, 1840.

Hipparchia boisduvallii Harris, 1862.

Satyrodes canthus; Holland, 1898.

Lethe eurydice; Klots, 1951.

COMMON NAME

Eyed Brown (Klots, 1951:68; Pyle, 1981:666)

DISTRIBUTION

From eastern Canada south to Delaware, thence westward to the

Northwest Territories and Colorado, according to authors who distinguish between this species and Satyrodes appalachia. Therefore, Ohio may be near the southern limit of its range. See under the discussion for the genus Satyrodes above for remarks relative to this species within Ohio.

BIOLOGY

Larvae feed on Carex (sedges). Both this species and Satyrodes appalachia are reported to inhabit wet areas. However, the former 334 prefers open meadows, sedge marshes, and wet prairies, while the latter

prefers wooded areas including edges (Pyle, 1981:667). _S. appalachia

is univoltine with adults between June and September. Combined Ohio

records for this and jS. appalachia extend from June into September.

Satyrodes appalachia (Chermock, 1947)

Appalachian Brown

NOMENCLATORIAL AND SYSTEMATIC STATUS

Lethe eurydice appalachia Chermock, 1947. [type-locality: Conestee Falls, North Carolina]

Satyrodes appalachia (Chermock, 1947). [valid name (Miller & Brown, 1981:190)]

Lethe appalachia; Howe, 1975:111.

COMMON NAME

Appalachian Brown (Pyle, 1981:667)

DISTRIBUTION

From Maine and Quebec south to northern Florida, west to eastern

South Dakota and Mississippi. See under the generic discussion above

' for remarks relative to the distribution of this species in Ohio.

BIOLOGY

Larvae feed on sedges (Carex). This species is univoltine

northward and multivoltine in the southern part of the range. Adults

are found from June to August in Maine, and October in northern Florida

(Pyle, 1981:667). This is roughly the same seasonal distribution as for

Satyrodes eurydice (see above) although it is not known whether the two

species overlap in Ohio. 335 SATYRINAE Boisduval, 1833.

Because of recent taxonomic work (Miller, 1968), all of the Ohio species of this subfamily are now placed in separate genera. This arrangement has been accepted by most workers since then, including

Miller & Brown (1981), and is followed here. Therefore, the key which follows is to species, as well as their respective genera. In addition to the five genera discussed below, six others are recognized in this subfamily in America north of Mexico.

Key to the genera and species of Satyrinae in the Ohio area.

1. Eyes densely hairy; wing span greater than or equal to 50mm...... Cercyonis pegala.

1'. Eyes sparsely hairy or naked; wing span less than 50mm...... 2.

2(1'). Ventral hind-wing with submarginal ocelli, each ocellus single and more or less circular; ventral fore-wing with one or more submarginal ocelli...... 3.

2'. Ventral hind-wing with two, double, elongate ocelli, lying very close to the outer margin; ventral fore-wing lacking submarginal ocelli...... Cyllopsis gemma.

3(2). Dorsal fore-wing with two ocelli Megisto cymela.

3'. Dorsal fore-wing lacking ocelli...... 4.

4(3'). Ventral fore-wing with four ocelli, at least two well-developed and of approximately equal size; ventral hind-wing with six ocelli ..

4'. Ventral fore-wing with one well-developed ocellus, others, if present, very small; ventral hind-wing with two ocelli, others if present tiny...... . 336 Cyllopsis Felder, 1869

NOMENCLATORIAL AND SYSTEMATIC STATUS

Cyllopsis Felder, 1869. [type-species: Cyllopsis hedemanni Felder, 1869, by monotypy; valid name (Miller & Brown, 1981:190)]

DISCUSSION

Four species of this genus occur in America north of Mexico. Only

one of these, Cyllopsis gemma, is found east of the Mississippi River

including Ohio. It is discussed below.

Cyllopsis gemma (HUbner, 1808)

Gemmed Satyr

NOMENCLATORIAL AND SYSTEMATIC STATUS

Oreas gemma HUbner, 1808. [type-locality: Georgia]

Cyllopsis gemma (HUbner, 1808). [valid name (Miller & Brown, 1981)]

Neonympha gemma; Holland, 1898.

Euptychia gemma; Holland, 1931.

COMMON NAME

Gemmed Satyr (Klots, 1951:70; Pyle, 1981:670)

DISTRIBUTION

From southern Virginia to central Florida, west to southern Illinois,

Texas, and Mexico. Ohio is near the northern limit of its range. Records

in Ohio are limited to four counties in the southern part of the state

(Fig. 246).

BIOLOGY

Larvae of this species feed on grasses, specifically Cynodon dactylon

(Bermuda Grass) in Texas (Pyle, 1981:670). Adults are found in wet, 337 shaded areas with long grasses. This species is bivoltine, with adults flying from April into August in Ohio (Fig. 247).

Hermeuptychia Forster, 1964

NOMENCLATORIAL AND SYSTEMATIC STATUS

Hermeuptychia Forster, 1964. [type-species: Papilio Fabricius, 1775, by original designation; valid name (Miller & Brown, 1981:191)]

DISCUSSION

Two Nearctic species, hermes and sosybius, both formerly considered subspecies of a single species (Euptychia hermes), are presently included in this genus. J3. sosybius is found in Ohio and is discussed below.

Hermeuptychia sosybius (Fabricius, 1793)

Carolina Satyr

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio sosybius Fabricius, 1793. [type-locality: not stated, possibly Georgia (Miller & Brown, 1981:191)]

Hermeuptychia sosybius (Fabricius, 1793). [valid name (Miller & Brown, 1981:191)]

Neonympha sosybius; Holland, 1898.

Euptychia sosybius; Holland, 1931.

Euptychia hermes sosybia; Klots, 1951. [misspelling]

Euptychia hermes sosybius; dos Passos, 1964.

COMMON NAMES

Carolina Satyr (Klots, 1951:69; Pyle, 1981:672)

Carolinian Satyr (Holland, 1931:180) 338 DISTRIBUTION

From New Jersey south to Florida, west to Ohio, Kentucky and Texas.

Ohio is near the northern limit of its range and has records from two counties, Adams in the South and Seneca in the North. The Adams County records are valid; the Seneca County record is from the literature

(Henninger, 1910) and the present location of the specimen(s) is unknown and the record cannot be verified.

BIOLOGY

Larvae feed on grasses. Pyle (1981:672) says that this species is bivoltine in the northern part of its range, with adults in spring and late summer. The two Ohio records for which dates are known are from the first half of August. A spring generation in Ohio, if it exists, has yet to be found. 339 Neonympha HUbner, 1818

NOMENCLATORIAL AND SYSTEMATIC STATUS

Neonympha HUbner, 1818. [type-species: Papilio areolatus Smith, 1797, designated by Hemming, 1937; valid name (Miller & Brown, 1981:191)]

DISCUSSION

Of the two Nearctic species in this genus one, Neonympha mitchellii, is known from the Ohio area, and is discussed below.

Neonympha mitchellii French, 1889

Mitchell's Satyr

NOMENCLATORIAL AND SYSTEMATIC STATUS

Neonympha mitchellii French, 1889. [type-locality: Cass Co., Michigan; valid name (Miller & Brown, 1981:192)]

Euptychia mitchelli; Holland, 1931. [misspelling]

Euptychia mitchellii; Klots, 1951.

COMMON NAMES

Mitchell's Satyr (Klots, 1951:69)

Mitchell's Marsh Satyr (Pyle, 1981:674)

DISTRIBUTION

This species is known only from southern Michigan and the northern parts of Indiana, Ohio and New Jersey. In Ohio it has been found only in Portage and Lake counties, both in the northeastern quarter of the state.

BIOLOGY

Neonympha mitchellii is found in acid, tamarack bogs, and is believed to feed on sedges. It is univoltine, with adults usually in the first two weeks of July. The only complete date found during this study for an Ohio specimen is for 4 July. It has not been reported from Ohio since 1968, in spite of attempts by several northern Ohio collectors to locate it, and may be extirpated from the state. Authors suggest that habitat destruction is probably the greatest threat to this species. 341 Megisto Hllbner, 1819

NOMENCLATORIAL AND SYSTEMATIC STATUS

Megisto Hilbner, 1819. [type-species: Papilio eurytus Fabricius, 1775, designated by Butler, 1868; valid name (Miller & Brown, 1981:192)]

DISCUSSION

Of the two Nearctic species of this genus one, Megisto cymela, is

found in Ohio. It is discussed below.

Megisto cymela (Cramer, 1777)

Little Wood Satyr

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio cymela Cramer, 1777. [type-locality: "Cape of Good Hope" (in error according to Miller & Brown, 1981:192)]

Megisto cymela (Cramer, 1777). [valid name (Miller & Br.own, 1981:192)]

Papilio eurytus Fabricius, 1775. [preoccupied by Papilio eurytus Linnaeus, 1758]

Papilio eurytris Fabricius, 1793. [misspelling]

Neonympha eurytus; Holland, 1898.

Euptychia eutyta Holland, 1931. [misspelling]

Euptychia cymela; Klots, 1951.

COMMON NAME

Little Wood Satyr (Klots, 1951:69; Pyle, 1981:675)

DISTRIBUTION

From southeastern Canada south and west throughout southern Canada and eastern United States to Saskatchewan, Colorado, and Texas. Ohio records indicate statewide distribution (Fig. 248). 342 BIOLOGY

Larvae feed on various'grasses and possibly sedges (Pyle, 1981:

675). Adults are found in wooded areas, including edges, openings, and wooded edges of old fields. Univoltine in the northern part of its range, including Ohio, where adults are found from May into August

(Fig. 249). 343 Cercyonis Scudder, 1875

NOMENCLATORIAL AND SYSTEMATIC STATUS

Cercyonis Scudder, 1875. [type-species: Papilio alope Fabricius, 1793, by original designation; valid name (Miller & Brown, 1981:195)]

Minois HUbner, 1819. [misidentification (Miller, 1968:99)]

DISCUSSION

Four Nearctic species are included in this genus by Miller & Brown

(1981). One of these, Cercyonis pegala, is found in Ohio and is

discussed below.

Cercyonis pegala (Fabricius, 1775)

Wood

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio pegala Fabricius, 1775. [type-locality: America, restricted vicinity of Charleston, South Carolina by Brown, 1966 (Miller & Brown, 1981:195)]

Cercyonis pegala (Fabricius, 1775). [valid name (Miller & Brown, 1981: 195)]

Satyrus maritima Edwards, 1880.

Cercyonis alope; Holland, 1898.

COMMON NAMES

Wood Nymph (Klots, 1951:72)

Grayling (Comstock & Comstock, 1904:188)

Common Wood Nymph (Macy & Shepard, 1941:85)

Blue-eyed Grayling (Pyle, 1981:683)

Large Wood Nymph (Pyle, 1981:683) 344 DISTRIBUTION

From southeastern Canada south to central Florida, west to central

Canada, California, and Texas. Ohio records indicate statewide distribution (Fig. 250).

BIOLOGY

Larvae feed on grasses. Adults fly in and near open, wooded areas, roadsides, wooded edges, and old fields. Univoltine, adults from June to early September in Ohio (Fig. 251). 345 DANAIDAE Duponchel, 1844.

This is a large family of mainly tropical butterflies with greatest diversity in the Indo-Australian Region. Members of the family are known for their migrations, and Williams (1930) lists thirty species which have exhibited migratory behavior. Adult danaids are often distasteful to avian predators, deriving protection from glycosides in their body fluids which are related to larval foodplants (including milkweeds in the Asclepiadaceae) (Ferris & Brown, 1981). Some of these protected adults are the models for mimicry complexes involving other, unprotected butterflies. There is a single Nearctic genus, Danaus, in this family.

This group has been placed as a subfamily in the NYMPHALIDAE by some authors.

Danaus Kluk, 1802

NOMENCLATORIAL AND SYSTEMATIC STATUS

Danaus Kluk, 1802. [type-species: Papilio plexippus Linnaeus, 1758, designated by Hemming, 1933; valid name (Miller & Brown, 1981:203)]

Anosia HUbner, 1816.

DISCUSSION

Of the three Nearctic species in this genus, two are known from

Ohio and are discussed below. 346 Danaus plexippus (Linnaeus, 1758)

Monarch

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio plexippus Linnaeus, 1758. [type-locality: Pennsylvania, designated by I.C.Z.N., 1954, Opinion 282]

Danaus plexippus (Linnaeus, 1758). [valid name (Miller & Brown, 1981:204)]

Anosia plexippus; Holland, 1898.

COMMON NAMES

Monarch (Klots, 1951:77; Pyle, 1981:711)

Milkweed Butterfly (Klots, 1951:77)

DISTRIBUTION

From eastern Canada south and west across most of North America

north of Mexico except for Alaska and the Pacific Northwest Coast.

This species does not overwinter in temperate areas, and is not resident

in Ohio. Adults from eastern and central North America fly southwest

into central Mexico in the fall and overwinter there. This includes

adults from the Ohio area. In the spring, these adults begin a return

trip northward and reproduce in southern United States. The adults

returning to the Ohio area are the offspring of those which migrated

southward the preceding year. This species is found throughout most

of Ohio, and probably will be recorded from every county with further

collecting (Fig. 252).

BIOLOGY

Larvae of this species feed on various species of Asclepias

(milkweeds) and Apocynum (dogbane). Adults are found in open fields, meadows, and along watercourses. This species is multivoltine with 347 overlapping generations in a single summer. Pyle (1981:712) describes three generations in the northern part of the range as follows: April-

June, adults migrating northward; July-August, resident in the North;

September-October, adults migrating southward. Ohio records (Fig. 253) extend from May into October with relatively few records in May and June,

and peaks of activity in August and in late September/early October,

indicating three generations in Ohio. Immature stages are found in

Ohio between the second and third generations, indicating a resident

third generation as well as transient migrants.

Danaus gilippus (Cramer, 1775)

Queen

NOMENCLATORIAL AND SYSTEMATIC STATUS

Papilio gilippus Cramer, 1775. [type-locality: Rio de. Janeiro]

Danaus gilippus (Cramer, 1775). [valid name (Miller & Brown, 1981:205)]

Anosia berenice; Holland, 1898.

Danais berenice Holland, 1931. [misspelling]

COMMON NAME

Queen (Klots, 1951:78; Pyle, 1981:713)

DISTRIBUTION

From southern Georgia and Florida around Gulf states to Kansas,

Texas and southern California, thence southward to South America. A single specimen has been found in Ohio (Gilbert, 1960) and that was taken in Franklin County, in the central part of the state. It was determined to be the southwestern form of this species, found in western and southwestern United States as well as Mexico. 348 BIOLOGY

Pyle (1981:714) reports■larval foodplants for this species as

Asclepias amplexicaulis (Blunt-leaved Milkweed) and Sarcostennna hirtellum (Rambling Milkweed). This species inhabits open areas where the foodplants occur. In the South this is multivoltine with successive generations between April and November. The single Ohio record (Gilbert,

1960) is from July 1, 1959. DISCUSSION

The list of Rhopalocera species known from the state of Ohio has been expanded by this study from the 93 species listed by Hine (1898a)

to 134 species at the present time (1982). These represent 77 genera and nine families of butterflies and skippers out of approximately 250 genera and 10 families known from America north of Mexico. Table 2 gives a summary of the taxa presently known from Ohio.

Table 2. Taxonomic composition of the rhopalocerous fauna of Ohio.

Family No. of Genera No. of Species

1. HESPERIIDAE 23 47 2. PAPILIONIDAE 3 6 3. PIERIDAE 8 12 4. LYCAENIDAE 19 29 5. RIODINIDAE 1 2 6. LIBYTHEIDAE 1 1 7. NYMPHALIDAE 14 27 8. SATYRIDAE 7 8 9. DANAIDAE _1______2 TOTAL 77 134

Additionally, 16 species have been discussed above which have been reported from part or all of the area surrounding Ohio, but not from within Ohio. These species may be expected to be found in Ohio after further study and collecting, or it may be determined that one or more have occurred in Ohio in the past and are now extirpated. Further field

349 350 collecting in selected areas (see below for discussion), as well as further examination of collections are needed to determine with greater accuracy the status of these species. Table 3 summarizes these species by state or province in which they are found. Table 8 in the appendix lists these species, and shows the neighboring state in which each is found.

Table 3. Taxonomic composition of the rhopalocerous fauna reported from the area surrounding Ohio, but not from Ohio itself, with the number of species reported from each state/province (P=Pennsylvania; W=West Virginia; K=Kentucky; 1=Indiana; M=Michigan; 0=0ntario).

Family Total Species No. of Species by State/Province

P W K M 0

1. HESPERIIDAE 7 5 4 3 2 2. PIERIDAE 1 1 1 1 1 1 1 3. LYCAENIDAE 1 1 1 1 1 1 4. RIODINIDAE ' 1 1 5. NYMPHALIDAE 4 3 4 3 2 4 3 6. SATYRIDAE 2 2 1

The species of Rhopalocera known from Ohio have different status in terms of the permanence of residence relative to annual cycles of activity. Some species are permanent residents which overwinter here and reproduce annually. Others are regular migrants, moving into

Ohio each year and reproducing at least once annually. Still others are strays, and are represented in the data which supported this study by only a very few records each. Finally, a few species are believed 351

to have been extirpated from Ohio. This group includes species for

which there is reasonably good evidence that they were residents in

Ohio in the past, and for which there are no recent specimens or

literature records to support including them in the current resident

list. Table 4 summarizes the Rhopalocera of Ohio by residnece status.

Table 9 in the appendix lists all of the butterfly and skipper species

known from Ohio with an indication of their individual present status.

Also included in this table is a notation for each species not listed

for Ohio by Hine (1898a). These are species which have been added to

the state list since 1898. At least one of these, Thymelicus lineola,

is known to have immigrated into the state since 1898.

Table 4. Summary of the Rhopalocera of Ohio by status (R=Resident; M=Regular Migrant; S=Stray; E=Extirpated).

Family Status

R M _S E

1. HESPERIIDAE 44 1 2 2. PAPILIONIDAE 6 3. PIERIDAE 6 3 2 1 4. LYCAENIDAE 27 1 1 5. RIODINIDAE 2 6. LIBYTHEIDAE 1 7. NYMPHALIDAE 24 2 1 8. SATYRIDAE 7 1 9. DANAIDAE 1 1

The position of Ohio varies within the overall ranges of the

species. There are three categories of geographical position for Ohio within a larger range: 1) Ohio lies within the range of the species

such that the species may be expected to be found in almost any direction 352 moving away from the state; 2) Ohio lies at or very near the edge of the range of a species; 3) Ohio is part of the range of a species where the range is generally made up of disjunct localities, each representing isolated populations. A fourth possible category would be for endemic species where the entire range of the species is found within Ohio, if such existed. No such endemism is known for any butterfly or skipper in the state of Ohio. The second major category above, which includes those species with a distribution edge running through or very near Ohio, may be further divided into four minor, subcategories. This is based upon which geographical limit in terms of direction occurs in or very near

Ohio: a) eastern limit; b) southern limit; c) western limit; d) northern limit. Table 5 lists all of the Rhopalocera known from

Ohio, and indicates the position of Ohio within the overall range of the species. 353

Table 5. The butterflies and skippers of Ohio with notation relative to the position of Ohio within the overall range of the species (R=Within Range; D=Disjunct Range; E=East; S=South; W=West; N=North).

Species R Range Edge D

E S W N

HESPERIIDAE: PYRGINAE 1. Epargyreus clarus X 2. Autochton cellus 3. Achalarus lyciades X 4. Thorybes bathyllus X 5. Thorybes pylades X 6. Staphylus hayhurstii X 7. Erynnis icelus X 8. Erynnis brizo X 9. Erynnis .iuvenalis X 10. Erynnis horatius X 11. Erynnis martialis X 12. Erynnis lucilius X 13. Erynnis baptisiae X 14. Erynnis persius X

15. Pyrgus centaureae X * 16. Pyrgus communis X 17. Pholisora catullus X HESPERIIDAE: HESPERIINAE 18. Ancyloxypha numitor X 19. Thymelicus lineola X 20. Nastra lherminier X 21. Hylephila phyleus X 22. Hesperia leonardus X 23. Hesperia metea X 24. Hesperia attalus X 25. Hesperia sassacus X 26. Polites coras X 27. Polites themistocles X 28. Polites origenes X 29. Polites mystic X 30. Polites vibex X 31. Wallengrenia egeremet X 32. Pompeius verna X 33. Atalopedes campestris X 34. Atrytone logan X 35. Poanes massasoit X 36. Poanes hobomok X 37. Poanes zabulon X 38. Poanes viator X Table 5(continued).

Species R

HESPERIIDAE: HESPERIINAE (continued) 39. Euphyes dion X 40.. Euphyes dukesi 41. Euphyes conspicua 42. Euphyes bimacula 43. Euphyes ruricola 44. Atrytonopsis hianna 45. Amblyscirtes hegon 46. Amblyscirtes vialis 47. Panoquina ocola PAPILIONIDAE: PAPILIONINAE 48. Battus philenor 49. Eurytides marcellus 50. Papilio polyxenes asterius 51. Papilio cresphontes 52. Papilio glaucus 53. Papilio troilus PIERIDAE: PIERINAE 54. Pontia protodice 55. Artogeia virginiensis 56. Artogeia rapae PIERIDAE: ANTHOCHARINAE 57. Falcapica midea PIERIDAE: COLIADINAE 58. Colias philodice 59. Colias eurytheme 60. Zerene cesonia 61. Phoebis sennae eubule 62. Phoebis philea 63. Eurema lisa 64. Eurema nicippe 65. Nathalis iole LYCAENIDAE: GERYDINAE 66. Feniseca tarquinius LYCAENIDAE: LYCAENINAE 67. Lycaena phlaeas

68. Hyllolycaena hyllus xx x x xxxxxx x x xxxxxx xxxx 69. Epidemia epixanthe 70. Epidemia dorcas 71. Epidemia helloides 355 Table 5(continued).

Species R Range Edge

jS W N

LYCAENIDAE: THECLINAE 72. Atlides halesus X 73. Harkenclenus titus X 74. Satyrium acadica X 75. Satyrium edwardsii X 76. Satyrium calanus falacer X 77. Satyrium caryaevorum X 78. Satyrium liparops X 79. Calycopis cecrops 80. Mitoura gryneus X 81. Incisalia augustus X 82. Incisalia irus X 83. Incisalia henrici X 84. Incisalia niphon X 85. Euristrymon Ontario X 86. Parrhasius m-album 87. Strymon melinus X 88. Erora laeta X LYCAENIDAE: POLYOMMATINAE 89. Hemiargus isola X 90. Everes comyntas X 91. Celastrina ladon X 92. Celastrina ebenina 93. Glaucopsyche lygdamus X 94. Lycaeides melissa X RIODINIDAE 95. Calephelis borealis X 96. Calephelis muticum X LIBYTHEIDAE 97. Libytheana bachmanii X NYMPHALIDAE: ARGYNNINAE 98. Euptoieta claudia X 99. Speyeria diana X 100. Speyeria cybele X 101. Speyeria aphrodite X 102. Speyeria idalia X 103. Speyeria atlantis X 104. Clossiana selene myrina X 105. Clossiana bellona X 356 Table 5(continued).

Species R Range Edge

E S W N

NYMPHALIDAE: MELITAEINAE 106. Charidryas nycteis X 107. Charidryas harrisii X 108. Phyciodes tharos X 109. Euphydryas phaeton X NYMPHALIDAE: NYMPHALINAE 110. Polygonia interrogationis X 111. Polygonia comma X 112. Polygonia progne X 113. Nymphalis vau-album X 114. Nymphalis antiopa X 115. Aglais milberti X 116. Vanessa virginiensis X 117. Vanessa cardui X 118. Vanessa atalanta X 119. Junonia coenia X NYMPHALIDAE: LIMENITIDINAE 120. Basilarchia arthemis astyanax X 121. Basilarchia archippus NYMPHALIDAE: CHARAXINAE 122. Anaea andria X X NYMPHALIDAE: APATURINAE 123. Asterocampa celtis X 124. Asterocampa clyton X SATYRIDAE: ELYMNIINAE 125. Enodia anthedon X 126. Satyrodes eurydice X 127. Satyrodes appalachia X SATYRIDAE: SATYRINAE 128. Cyllopsis gemma X 129. Hermeuptychia sosybius X 130. Neonympha mitchellii X 131. Megisto cymela X 132. Cercyonis pegala X DANAIDAE 133. Danaus plexippus X 134. Danaus gilippus 357

The relative abundance of the taxa included in this study was rejected for inclusion in this report because of the bias which exists

in collections and publications. This bias results from student or

collector preferences, sometimes prejudices, and leads to unrealistic relations between numbers of specimens preserved in different taxa.

For example, the swallowtail butterflies are fairly common in Ohio, except for Papilio cresphontes. Nevertheless, these species are rather poorly represented in most private and many public collections in terms of quantity of specimens. This is largely because of the cost and space required to house them in a collection. Also, the familiarity of these species leads to a sort of disdain toward any collection of them in the field. On the other hand, if a collector becomes interested in the

LYCAENIDAE or HESPERIIDAE he is likely to collect almost every specimen he finds, creating long series for his and ultimately, perhaps, a public collection. A student of a particular group will concentrate his efforts on that group, and a collection will reflect this interest.

Therefore, a specialized study of abundance of these species within

Ohio is needed if any sort of accurate inferences are to be drawn.

Such a study would need to include proper sampling of all parts of the state with relative abundance of taxa as a predetermined goal of the work. This was beyond the scope of this study, and was not done.

A cursory examination of the literature shows that all four life stages of butterflies and skippers are involved in overwintering by

Ohio's species. The larval foodplants utilized by these species are reasonably well documented in the literature, and are included in the 358

discussions above. However, more data are needed relative to the specific

larval foodplants utilized in Ohio. Little has been recorded and reported

relative to adult nectaring sources in the Ohio area. Price (1970) has

reported on a few of the plants he found being utilized as nectar

sources during his study of northwestern Ohio. Again, further study is

needed. Such a work is presently underway (1982, Susan Ahearn, personal

communication) in connection with a more general survey in Franklin

County in central Ohio. Data concerning overwintering stages, larval

foodplants, and adult nectar sources could be correlated with seasonal

patterns of adult activity, as reported in the species discussions above,

such that the ecological requirements of these species in the Ohio area

could be better understood.

Within Ohio a number of counties are well-represented in museum

collections and in the literature, and have been frequently collected by

various collectors. These counties, of course, have contributed the most

data which support this study. A simple measure of the effectiveness of

these records, in terms of this study, is a percentage of the total

species known for the state as represented by the total species known

for a given county. The following counties have had 50% or more of the

Ohio Rhopalocera species recorded for each: Williams, Lucas, Defiance,

Paulding, and Seneca in the Northwest; Cuyahoga and Wayne in the

Northeast; Marion, Franklin, and Licking in the central part of the

state; Montgomery, Hamilton, and Adams in the Southwest; and Hocking and

Vinton in the Southeast (see Fig. 254). 359

Additionally, the following counties have had between 25% and 50%

of the species known from Ohio reported from them: Van Wert in the

Northwest; Erie, Lake, Geauga, Richland, Portage, Stark, and Carroll in

the Northeast; Champaign, Delaware, Pickaway, and Fairfield in the

central part of the state; Greene, Butler, Ross, Highland, and Scioto

in the Southwest; and Morgan, and Jackson in the Southeast.

It is unfortunate for the sake of completeness of this study that

not all Ohio counties are represented to the same extent. Museum records,

literature records, and collections made by private individuals usually

reflect the specific interests of the persons or institutions involved.

These interests are often limited by geographical circumstances. For

example, the Lepidoptera collections of the Cleveland Mus,eum of Natural

History primarily have material from Cuyahoga County and its neighbors, while the collections in the Cincinnati Museum of Natural History

Primarily show the fauna of southwestern Ohio. Similarly, published

reports of Ohio butterflies often are limited to the area close to the

residence of the author (eg. Henniger, 1910 and Price, 1970). During

this study I have been able to collect in approximately two-thirds of

Ohio's counties, but not to the same extent in each county. The result

is bias in terms of knowledge of the rhopalocerous faunal diversity within Ohio. Some counties are better known than others.

The maximum number of butterfly species known for a single county

is 94, in both Cuyahoga County (home of the Cleveland Museum of Natural

History) and Franklin County (home of The Ohio State University and The

Ohio Historical Society's collections). At the low end of the scale 360

Hardin County in northwestern Ohio was inadvertently overlooked during

this study, and has no known butterflies or skippers.

Therefore, additional collecting of butterflies and skippers is needed in several parts of Ohio. Eastcentral Ohio probably is the least well known part of the state. This is the area east of Licking and

Hocking counties, and south of Wayne County to the Ohio River. In all of these counties fewer than forty species (less than 30% of the state's

total) are known for each county. The far northeastern corner of the

state needs further collecting. This is the part of Ohio which includes part of the Transition Life Zone in portions of four to six counties, according to some authors (see under discussion of distributional patterns below). In the southeastern part of the state the counties along the

Ohio River are rather poorly known, and need further data. Far western

Ohio, and portions of the northwestern and southwestern quarters of the

state, need more collecting as well.

As has been noted above, Ohio's, rhopalocerous fauna is much better known as a result of this study than it has been in the past. If all of the possible species listed above eventually are to be found in Ohio,

then the known fauna today is about 90% complete. With further gathering of data, many of the apparent gaps in the ranges of various species undoubtedly will be filled. In the case of a few species, as noted in the individual discussions above, more data are certainly needed and the understanding of the ranges of those species within Ohio may very well change. During the course of this study attention was paid to possible

correlations between species distribution and physiographic or other

subdivisions of the state. No patterns of distribution relative to

these subdivisions were found. Therefore, except for a few specific

references to one or more of these areas in the species accounts above,

no use was made of the various existing systems.

In the Life Zone system of C. Hart Merriam, Ohio includes portions

of two of these Zones: the Transition Zone occurs in a small portion of

the northeastern corner of the state; the remainder of the state lies

within the Upper Austral Zone (Klots, 1951:22-26; Clench, 1975:34-38).

Of the species listed by Klots and Clench (op. cit.) as being

characteristic of the Transition Zone, several are known from Ohio as

follows: Hesperia sassacus, Polites mystic, Poanes hobomok, Artogeia

virginiensis, Satyrium acadica, Styrium edwardsii, Erora laeta,

Clossiana selene myrina, Clossiana bellona, Charidryas harrisii,

Euphydryas phaeton, and Neonympha mitchellii. Of these species, only

Neonympha mitchellii is limited to that portion of the state which is

included in the Transition Zone. All of the others have been reported

from well outside that portion of the state. Therefore, the Life Zone

system seems to be of no utility in describing distribution patterns of butterflies and skippers in Ohio. Indeed, it could well lead to wrong

assumptions and inaccuracies.

Although occasional species have individual distributional limits within Ohio that appear to correspond with other systems of regionalization, there are no apparent patterns of distribution involving a group of species. These other systems include the physiographic 362 sections, glacial deposits, and geological (bedrock) system. Therefore,

no existing system of regions, sections, or other divisions of the

state of Ohio is of any use in generalizing about butterfly and skipper

distribution within the state.

An examination of the maps showing distribution of butterflies and

skippers in Ohio will show that the distributions of the various species

are not consistent. Many species are found, or probably will be found,

throughout the state. Others show distributional limitations to one

part of the state or another. Additional collecting in various parts of

the state may extend the ranges of some of these species. However, it is

believed that enough data have been examined (over 8000 records of the

butterflies and skippers, averaging to 91 per county or 59 per species)

to give a usable indication of specific range boundaries.

The distribution of each species known from Ohio has been discussed

briefly above. Those species whose distribution appears to include less

than the entire state, when examined individually, provide information

relative to their own distribution, but little more. However, by

examining these limited distributions together, it is possible to show

patterns of distribution involving multiple species. These patterns

are distinct enough that the state may be divided into sections, each with some distinct characteristics relative to its butterfly fauna.

After completing the Ohio distributional maps for each of the

species included in this study, those whose distribution appeared to

be limited to less than the whole state were removed from those with

apparent statewide distribution. A transparent, acetate copy was made of each map. These acetate copies were stacked in groups of about 25

and were looked at with back lighting. In each group it was found that

a minority of maps showed dots outside of the area of coverage for the

majority. These minor maps were removed, shifted to another group, and

the process was repeated. After a series of selections, several groups

of maps were determined. Each group shows butterfly distribution of

two or more species within a fairly well defined area, and absent from most of the remainder of the state. Two of these areas are discussed below.

The state of Ohio can be divided into two major areas based upon

apparent patterns in butterfly and skipper distribution, as determined with the process described above. A northern area, called here the

Northern District, consists of the northeastern and northwestern parts of the state as well as portions of the central and western parts. The

Southern District takes up the remainder of the state, and consists of the southeastern part of Ohio along with portions of the southcentral and southwestern parts. A line separating these two Districts extends from the southwestern corner of the state (between Preble and Butler

Counties) more or less northeastward through Columbus to the juncture of the states of Ohio, West Virginia, and Pennsylvania (see Fig. 255).

This line corresponds roughly with the line dividing Ohio into northern and southern parts, as suggested by Webster (1893). Webster extended his line from New York City (latitude 40°43' north) through

Ohio to St. Louis, Missouri (latitude 38°38' north), and from there on to Colorado. It was his belief that this line approximated a natural 364 divide between northern and southern insect faunas. He based this on

insect species which had immigrated into Ohio during the 19th century.

Webster's line passes approximately through Columbus, Ohio (latitude

40°0' north), as does the line for butterflies and skippers described

above. At its eastern end in Ohio, however, Webster's line lies further

south than does the line described for Rhopalocera, and at its western

end it lies further north.

In the past, several workers have divided a geographical unit into

parts based on faunal distributional patterns. Hagmeier and Stults

(1964) used numerical methods in analysing mammalian patterns in North

America. The state of Washington has been divided into butterfly

provinces on the basis of patterns of distribution, first intuitively,

then analytically using numerical techniques (Pyle, 1976). Pena (1966)

has divided Chile into regions, using especially members of the family

TENEBRIONIDAE (Coleoptera). Lee (1980) has established faunal regions in

the Yucatan Peninsula based upon the distributions of reptiles and

amphibians.

Analysis of possible faunal areas within Ohio, including the

Northern and Southern Districts suggested above, should be conducted

using statistical or other numerical techniques. Baroni-Urbani and

Buser (1976) present a technique for establishing reliable similarity

coefficients which can be used to compare two units, whether they be

taxonomic, geographical, or other. The technique is based upon binary

data derived from a set of attributes. They also provide a theoretical model which can be used to test the significance of the comparisons.

Lee (1980) used this procedure in part of his faunal analysis of the 365 Yucatan Peninsula. Such a technique can be used in analysing faunal

areas in Ohio. However, more data need to be collected before

undertaking such an analysis. There are still too many gaps in the

distributional maps of Ohio species of butterflies and skippers. As

noted above, eastcentral Ohio is rather weak in terms of data, and

the line proposed for separating the Northern and Southern Districts

runs through this part of the state. Also, there are indications that

both the Northern and Southern Districts can be subdivided further.

This would require even more careful analysis, since fewer species

appear to be involved.

As tentatively understood, the Northern and Southern Districts of

Ohio contain species of Rhopalocera that are more or less restricted to

one or the other area within the state. Table 6 lists these species

for the Southern District, and Table 7 lists them for the Northern

District. All of these are considered residents of the state, as

described above. Care must be taken, however, in any use of these

Districts until they are better understood. Table 6. Resident Rhopalocera in Ohio which are more or less restricted to the Southern District of the state.

HESPERIIDAE: PYRGINAE 1. Autochton cellus HESPERIIDAE: HESPERIINAE 2. Amblyscirtes hegon PIERIDAE: ANTHOCHARINAE 3. Falcapica midea LYCAENIDAE: THECLINAE 4. Calycopis cecrops 5. Mitoura gryneus 6. Incisalia augustus 7. Incisalia henrici 8. Incisalia niphon 9. Euristrymon Ontario 10. Parrhasius m-album LYCAENIDAE: POLYOMMATINAE 11. Glaucopsyche lygdamus NYMPHALIDAE: CHARAXINAE 12. Anaea andria SATYRIDAE: SATYRINAE 13. Cyllopsis gemma

Table 7. Resident Rhopalocera in Ohio which are more or less restricted to the Northern District of the state.

HESPERIIDAE: PYRGINAE 1. Erynnis baptisiae 2. Erynnis persius HESPERIIDAE: HESPERIINAE 3. Thymelicus lineola 4. Hylephila phyleus 5. Polites mystic 6. Poanes massasoit 7. Poanes viator 8. Euphyes dion 9. Euphyes dukesi 10. Euphyes conspicua 11. Euphyes bimacula 12. Atrytonopsis hianna PIERIDAE: PIERINAE 13. Artogeia virginiensis LYCAENIDAE: LYCAENINAE 14. Epidemia dorcas 15. Epidemia helloides Table 7(continued)•

LYCAENIDAE: THECLINAE 16. Satyrium acadica 17. Satyrium edwardsii 18. Satyrium caryaevorum 19. Incisalia irus LIBYTHEIDAE 20. Libytheana bachmanii NYMPHALIDAE: ARGYNNINAE 21. Speyeria atlantis 22. Clossiana selene myrina NYMPHALIDAE: MELITAEINAE 23. Charidryas harrisii NYMPHALIDAE: NYMPHALINAE 24. Nymphalis vau-album 25. Aglais milberti SATYRIDAE: ELMNIINAE 26. Satyrodes eurydice SUMMARY

The primary objectives of this study were threefold: 1) to determine and describe the taxonomic composition of the Rhopalocera in

Ohio; 2) to describe the geographical distribution of each of the

species within the state, and to establish its relative position in the overall context of the species' distribution; and 3) to describe the

seasonal distribution of the adult stage of each species within the

state.

In addition, diagnostic characters were given for each of the taxa included in this study. These appeared either in the discussion of the respective taxa, or in the taxonomic keys prepared to aid in their determination. Also, information relative to foodplants, habitat, and other selected biological factors were included.

The Rhopalocera known to occur in Ohio include 134 species which represent 77 genera and nine families. An additional 16 species are known from the area surrounding Ohio, and may either be found to be present in the state, or to have been extirpated. Of the 134 species recorded from the state, 117 are residents, seven are regular migrants into the state, six of these species are strays, and four are believed to have been extirpated. In terms of Ohio's position in the overall range of the individual species, Ohio lies well within the

368 369

range of 85 species, is at or very near the range limit of 46 species, and is involved in the disjunct range of three species.

In terms of knowledge of species within Ohio, the Rhopalocera probably are about 90% catalogued. Suggestions are made concerning where additional collection of data is needed. Fifteen of Ohio's

88 counties have been well collected and are well represented on the maps which show geographical distribution of the species. These 15 counties are distributed throughout the state. Eastcentral Ohio, as well as other portions of the state, should be studied further. Also, studies relative to the relative abundance of these species are needed, as well as studies of ecological factors.

Ohio is tentatively divided into two Districts, the Northern and the Southern. Species of Rhopalocera are listed for each, 13 for the

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Drees, Bastiaan M. & Linda Butler. 1978. Rhopalocera of West Virginia. J. Lep. Soc. 32(3):198-206.

Dury, Charles. 1878. Catalogue of the Lepidoptera, observed in the vicinity of Cincinnati, Ohio.... J. Cin. Soc. Nat. Hist. 1(1):12-23. 372

Dyar, Harrison G. 1902'. A list of North American Lepidoptera. Smithsonian Inst. Bull. No. 52. xx + 723 p.

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Fig. 1. Parts of Ohio included in regional or county reports of Rhopalocera (included counties are marked).

378 379

Ro

Cu

Cu

2A

c+R Rs

3A Cu Cu 2A

Fig. 2. Wing vein terminology used in this work (A=anal; Cu=cubital; h=humeral; M=medial; R=radial; Rs=radial sector; Sc=sub- costal veins) 380

A8HTABULA

PULTON LUGA9 < j n WILLIAMS JSCAUQA | I OTTAWA WOOD ’trumqull iRAIN i DEFIANCE ERIC

HURON 8EMB0A

PUTNAM HANGOOK

| A U OLAIZE KNOX t — I LOOAN UNION I SHELBY G 08K 0G T 0N HARRISON

IA 0I80N

CLARK PERRY

PAYETTE WASHINGTON BUTLER CLINTON |

sw J—/ VINTON

HAMILTON /(mmlamd (

Fig. 3. Ohio's counties and their names (collecting divisions: _C=Central; NE=Northeastern; SE=Southeastern; NW=North- western; SW=Southwestern). 0 0 LO

Fig. 4. Geographical distribution of Epargyreus clarus in Ohio. (4) (7) (9) (2) (8) (2) (2) (7) (15) (8) (8) (2) (4) (3) No. of Adult Records t t i i » » i ********* ** ******* ** **** ******** **** ** ******* *************** ******** ******** ** ft** 1-10 11-20 21-31 11-20 11-20 21-30 11-20 21-31 21-30 Sep SepSep 1-10 11-20 Oct OcCOct 1-10 11-20 Aug 21-31 Aug Jul Jul 21-31 JunJun 1-10 11-20 Jun Aug 1-10 Jul 1-10 Apr 21-30MayMay 1-10 Apr 11-20 May Apr Mar 21-31 Mar Mar 1-10 Q. TJ (0 *1 H- O 03 o at Cfl ra Co

Fig. 5. Seasonal distribution of adult Epargyreus clarus in Ohio. 382

Fig. 6. Geographical distribution of Autochton cellus in Ohio.

35 -

30 -

'E 25 - ao & u 20 • rH TJ3 < 15 -

10 - zo

HNfOHNnHojPiHNnHNM-^Nni-iNnHNnOO'-HOOOOO'-HOOOOOfHOO^OOOOOrH i I I I I i I I i I I I i i i i I I I i i I I I H H H H H pH H H H H H H H H *“H H *H H pH pH H pH H H f—i f\J CNJ *-H tM H N ■-< CM •—I fVl p-H CN p-« CNJ SX»<:n'-)'-)'-)<5Q)a)UUU Seasonal Time Period

Fig. 7. Seasonal distribution of adult Autochton cellus in Ohio. 00 LO LO

Fig. 8. Geographical distribution of Achalarus lyciades in Ohio. (2) (4) (14) (1) (1) (1) (6) (3) (i) ►— ►— •— ro to u> No. of Adult Records t t t t i v t I in in O in o in © in * ************** **** * * * ** *** ****** 1-10 11-20 11-20 11-20 11-20 21-30 21-31 21-31 21-31 21-31 21-31 Sep Sep Sep 1-10 Oct Oct Oct Jul 11-20Jul Jun 21-30 JunJun 1-10 11-20 Aug Aug Apr 11-20 Jul 1-10 Aug 1-10 May Apr 1-10 Apr 21-30May 1-10 Mar Mar May 11-20 Mar 1-10 H- a ro o T) ►1 m o g IB CO

Fig. 9. Seasonal distribution of adult Achalarus lyciades in Ohio. 384

Fig. 10. Geographical distribution of Thorybes bathyllus in Ohio.

ro in r** i

35 -

30 -

25 -

20 -

3 15 -

10 * zo «* * 5 - * * * * * * * * * •K * * HNMHNnHN«OHNnHNouo Seasonal Time Period

Fig. 11. Seasonal distribution of adult Thorybes bathyllus in Ohio. 385

Fig. 12. Geographical distribution of Thorybes pylades in Ohio.

35 -

30 - a "S 25 - uo & * 2°-

3 15 - mo . 1 0 - •K o * * z * * * * * * * * * * * * * * * * * * * « * * * * * * *« ■** * fHCMn^NnHNnMC'j(n^Nfn* >>>* a a PHH'-t boooboap

Seasonal Time Period

Fig. 13. Seasonal distribution of adult Thorybes pylades in Ohio. 386

Fig. 14. Geographical distribution of Staphylus hayhurstii in Ohio.

35 -

30 -

25 -

20 - •q9 < 15 '

10 .

5 •

o©»-tooooo«—*oooo©»"“<©c>rHooo©o«—i I I I I I I I I I I I I I I I I I I I t I I I I *-< fN| OJ «—l CM HN

Fig. 15. Seasonal distribution of adult Staphylus hayhurstii in Ohio. 0 0 u> HE

Fig. 16. Geographical distribution of Erynnis icelus in Ohio. (3) (8) (5) (2) (2) (4) (2) (3) »-• »-• t— K9 N) U» U3 No. of Adult Records t i i « t » » Cn Cn O O Ui © tn ** ******** **** ***** *** ** ** *** 1-10 1-10 21-30 11-20 21-31 11-20 21-31 21-31 Sep 11-20 Sep Oct 11-20 Oct 21-31 Sep 1-10 Oct Jul Jul Aug 11-20Aug JunJun 1-10 11-20Jun 21-30 Aug 1-10 Apr 21-30 Jul AprApr 1-10 11-20 May 11-20May May 1-10 MarMar 1-10 Mar 21-31

Fig. 17. Seasonal distribution of adult Erynnis icelus in Ohio 00 00 u>

Fig. 18. Geographical distribution of Erynnis brizo in Ohio. (12) (7) (10) (3) (2) (1) No. of Adult Records « « I 1 1 I 1 1 w* w* o u» o Ui o *** ******* ********** ************ ** * 1-10 1-10 1-10 1-10 1-10 1-10 1-10 11-20 11-20 11-20 11-20 21-31 11-20 21-30 11-20 21-31 21-30 21-31 21-31 Mar 11-20 Mar Apr 11-20 Mar 21-31Apr Jun May Sep Sep Jul Apr 21-30May 1-10 May Jul Jul Jun Jun Sep Oct Aug Aug Oct Oct Aug

Fig. 19. Seasonal distribution of adult Erynnis brizo in Ohio 389

Fig. 20. Geographical distribution of Erynnis juvenalis in Ohio.

* © P** VO /“s h vo n n 35 -

30 •

25 -

20 -

* « * 15 * * * * « * * * * * * 10 * * * * * ■K * •K* * * it *•K n it *« n H « * it it * * it * * ** it * * * - K * * H it * * HNcOHisnHNnMNnHNmHNnHNnHNnOOi-tOOOOO’-'OOOOOi-HOO’-'OOOOO’-H I I I I I I l I I I l I I I I I t I I l I l I I •-» CN •—< fsj 1 ^ >v fl g fl H rj i GO______CO 00__ C u _ P.______CX *J SanS

Fig. 21. Seasonal distribution of adult Erynnis .juvenalis in Ohio. LO O VO

Fig. 22. Geographical distribution of Erynnis horatius in Ohio. (1) (A) (7) (2) (1) (2) (2) (2) (5) (5) (1) (1) (1) - No. of Adult Records , * ft*** ******* ** * ** ft* ft* ***** ***** ft ft ft Sep 11-20 Sep 21-30 Sep 1-10 Oct 11-20 Oct 21-31 Jun 1-10Jun 11-20 Jul 11-20 Jul 21-31 Oct 1-10 Aug 21-31 Apr 1-10 Apr 11-20 Apr 21-30 Jun 21-30 Jul 1-10 Aug 11-20 May 1-10May 11-20 May 21-31 Aug 1-10 Mar 1-10Mar 11-20 Mar 21-31 H (T> to CD o s 3 00

Fig. 23. Seasonal distribution of adult Erynnis horatius in Ohio. CO VO

Fig. 24. Geographical distribution of Erynnis martialis in Ohio. (i) (i) (2) (1) (1) (1) (i) (3) (1) ■ ■ (1) i—■ to NJ UJ U> - n Mo. of Adult Records t I I I I t I isi isi O Ln O ui O Ui * * AAA A A A * ** A A 1-10 1-10 1-10 1-10 1-10 1-10 11-20 11-20 11-20 11-20 11-20 11-20 11-20 11-20 21-31 21-31 21-30 21-31 21-31 21-31 21-30 Sep Sep Sep Oct Oct Oct Jul Jun 21-30 Jul Aug Aug Jun Jun Jul Mar Apr Apr May May Aug Apr May 1-10 MarMar 1-10

Fig. 25. Seasonal distribution of adult Erynnis martialis in Ohio N5 OJ

Fig. 26. Geographical distribution of Erynnis lucilius in Ohio. ) 1 (l) ( No. of Adult Records 1-10 1-10 1-10 11-20 11-20 11-20 11-20 11-20 21-31 21-31 21-30 21-30 Sep 1-10 Sep Sep Jun Jul Jul AprApr 1-10 11-20 Jun OctOct 1-10 11-20Oct 21-31 Aug 11-20 Jun Jul Mar 21-31 Apr 21-30May 1-10 May 21-31 MarMar 1-10 Aug Aug May w (D © CD O 9 0 H- a

Fig. 27. Seasonal distribution of adult Erynnis lucilius in Ohio LO OJ vo

Fig. 28. Geographical distribution of Erynnis baptisiae in Ohio. CD (7) (3) (4) (3) (i) (2) (1) (1) nj *— *— ►- ro No. of Adult Records * * ** ■kick **** *** * ******* * Sep 21-30 Sep 1-10 Sep 11-20 Jun 21-30 Jun 1-10 Oct 11-20 Oct 21-31 Jul 21-31 Aug 1-10Aug 11-20 Aug 21-31 May 11-20 May 21-31 Jun 11-20 Jul 11-20 Mar 11-20 Mar 21-31 Apr 1-10Apr 11-20 Apr 21-30 May 1-10 Oct 1-10 Jul 1-10 Mar 1-10 re H H- 3 w re

Fig. 29. Seasonal distribution of adult Erynnis baptisiae in Ohio. 394

Fig. 30. Geographical distribution of Erynnis persius in Ohio.

35 -

30 -

m 25 o

* 2 0 r-f -3 < 15 iO M o. 10 ■ 85 5 -

00*-<00000r-t00000^00»-«00000»-< I I I I I I I I I I I I I I I I I I I I I I I I

asg

Fig. 31. Seasonal distribution of adult Erynnis persius in Ohio. 395

Fig. 32. Geographical distribution of Pyrgus centaureae in Ohio.

35 -

30 ■

w 25 - uo 5 20 -

15 -

10 -

5 -

OOf-HOOOOOr^OOOOO^OO^OOOOO-H HNnHNtnHNnHNnHNflHNnHNnHNfOI I I I I I I I I I I I I I I I I I I I I I I I H H H H H H H H H H H H H H i-H H H H H H H H H HN «-H

H C a fl H H H DIEiOU)&p.aiJllU d d d P i C u Q- i q -) d-- d ggg^^gaB3Q)a)Q)ooo_

Seasonal Time Period

Fig. 33. Seasonal distribution of adult Pyrgus centaureae in Ohio. O' u> v£>

Fig. 34. Geographical distribution of Pyrgus communis in Ohio. (1) (2) (1) Cl) (2) (3) (2) (3) (5) (5) (2) (1) (6) (2) (1) ►— ►— »— NJ NJ u> UJ No. of Adult Records r t f » t i » Ut O Ln O Ui O Ui ** *** * ** * *** * ***** ***** ****** ** * ** ** * Sep 1-10 Sep 21-30 Sep 11-20 Oct 11-20 Oct 21-31 Mar 11-20 Mar 21-31 Apr i—Apr 10 11-20 Apr 21-30 Jun 1-10 Jun 11-20 Jun 21-30 Jul 1-10Jul 11-20 Jul 21-31 Aug 1-10 May 21-31 Aug 21-31 Oct 1-10 Mar 1-10 May 1-10May 11-20 Aug 11-20

Fig* 35. Seasonal distribution of adult Pyrgus communis in Ohio 397

Fig. 36. Geographical distribution of Pholisora catullus in Ohio.

m o\ m cm r- «—<

35 -

30 -

25 -

20 - *33 < 15

10 * 5 ■ * * * * * * * * * oo»~«ooooo—*ooooo»-

U M M o.o.p.&&&§3S'dd 3 3330)a)aiboo s s s 3*>!

Fig. 37. Seasonal distribution of adult Pholisora catullus in Ohio. 398

Fig. 38. Geographical distribution of Ancyloxypha numitor in Ohio.

O /—n /*. /v /-N

35 -

30 -

25 -

20 -

15

. 10- * * *« * * * *. *. 5 - * * * * * * * * * * * * * ****** * * * * * * * * * * * * * ********** ******* 0©»“'0©000«-«0000©*-*00»""»0000©»-<^N(O^NnMNtn^NnHNfnf^Nn<-tNni-'Nn ijliiiilllllllllilllllil —« fvj >“• (SI —I f\| H N fs| r - t rsj H N

U M M U M U ^ a a HHH OObOCC(XO'Cb4->4J4J Ssz

Fig. 39. Seasonal distribution of adult Ancyloxypha numitor in Ohio. 399

Fig. 40. Geographical distribution of Thymelicus lineola in Ohio.

35 -

30 -

25 -

20 -

15 -

10 - * 5 - * * * * * ******* * * * *

*-,>» 0 a dHHH 00 to 00 & p. a 4J u u

Seasonal Time Period

Fig. 41. Seasonal distribution of adult Thymelicus lineola in Ohio. 400

Fig. 42. Geographical distribution of Nastra lherminier in Ohio.

35 -

30 - to m 25 ■ o

10 -

•K 5 - * * *s s* * * * * © O —»00000*-<00000^00*-<00000'-^ -jNci^Nro^NnHNnHNnHNnHNnHNnI i I I I I I I i I I I I I I I i I I I i I I I I T-* CNJ

U to U ^ u „ _ , H H u do oo a p. a u p u aaan)(drt333d33333a)o)a)uooI e r l «

Seasonal Time Period

Fig. 43. Seasonal distribution of adult Nastra lherminier in Ohio. 401

Fig. 44. Geographical distribution of Hylephila phyleus in Ohio.

35

30 -

25 -

20 -

c 15 -

10 -

* « * * * oo^ooooo^ooooo^oo^ooooo*-* I t t I I l I I f .1 I I I ( t I I 1 I I 1 I I t H N f* CM • CM • < 6 1 bO bO bO a CL a 4 J StftfSSS'j'S^flsaiidittooo s hJh*nhh,i

Fig. 45. Seasonal distribution of adult Hylephila phyleus in Ohio. 402

Fig. 46. Geographical distribution of Hesperia leonardus in Ohio.

35 -

30 -

Ot. 25

20 - 3 73 c 15 -

10 - « *« * «* * - 5 - * * * * * * * * * * * OO^OOOOOw w — wwww0^00000*-<00^00000^ p^NfOHNn^b' I I I I I I I II I I I <—4I rHI HI HI HI rHI I I *— 4 ■ I 1 - 4 < CN| H N • C l uM mM u u u u >s a a c h h h m m m a o- aw u u rtcdrt a a a rt « « 3 3 3 3 3 3 3 3 3 0 « ai o o o

Seasonal Time Period

Fig. 47. Seasonal distribution of adult Hesperia leonardus in Ohio. 403

Fig. 48. Geographical distribution of Hesperia metea in Ohio.

35 -

30 -

25 -

20 ■ 3 3 15 -

10

5

O O o o © o pH O o pH O o o o o ■h CM ro CM m •H CM m CM m H CM m pH n *7* | CM * f I I I I I I I I I i 1 1 H h .1* •—4 H H pH >H«HH H pH H CM — *-> •“3

Fig. 49. Seasonal distribution of adult Hesperia metea in Ohio. 4^ O 4>* TL

Fig. 50. Geographical distribution of Hesperia sassacus in Ohio. ) ) ) 1 1 2 ( ( (3) (9) ( N 3 No. of Adult Records * *** ********* ** * Apr 1-10 Apr 21-30 Sep 1-10 Sep 11-20 Mar 1-10Mar 11-20 Mar 21-31 Apr 11-20 May 1-10 Jun 1-10Jun 11-20 Jul 11-20 Jul 21-31 Sep 21-30 May 11-20 May 21-31 Jun 21-30 Jul 1-10 Aug 1-10Aug 11-20 Aug 21-31 Oct 1-10Oct 11-20 Oct 21-31

Fig. 51. Seasonal distribution of adult Hesperia sassacus in Ohio. -C- o U1 H' ro Ln O to o l-i cr OP H* O ID I-1 O. H- rt CO H H- O' C OP rt H- O Hi hd O I—* rt H- CD CO O O H CD O CO S' O H-

) ) ) ) ) ) 8 2 6 1 2 1 (1) (ii) (ii) (ii) ( (4) (5) ( ( (5) (5) (9) (9) ( ( ( No. of Adult Records ******* *********** *********** * **** ** ***** ****** ***** ********* ** * *

1-10 11-20 Mar 11-20 Mar 1-10 Apr .1-10 M a r 21-31 Apr 11-20 Apr 21-30 May 1-10 May 11-20 May 21-31 Jun 1-10Jun 11-20 Jun 21-30 Jul 1-10Jul 11-20 Jul 21-31 Aug 11-20 Aug 21-31 Sep Sep. Aug I-10 Sep 21-30 Oct 11-20 Oct 1-10 Oct 21-31

Fig. 53. Seasonal distribution of adult Folites coras in Ohio. 406

Fig. 54. Geographical distribution of Polites themistocles in Ohio.

35 -

30 -

m 2 5 -

20 - T33 < 15

10 - * *« * * 5 - * « * * * *« * * * * * * * * * * *

OO'-HOOOOO^OOOOOr-iOO'-iOOOOO*-*HNinHNm Itllllll* I I I I I t I I I I I I I | »

b m u u m u > * > > a a C H r 4 H no oo o o ap . u w SSS

Fig. 55. Seasonal distribution of adult Polites themistocles in Ohio. 407

Fig. 56. Geographical distribution of Polites origenes in Ohio.

30 -

25 -

20 -

15 -

. lo -

5 - * « * * «* *« * ******** * * « * O0*-H00000*-'00000**HOO*-‘0 0 0 0 0 ' “1 I I t I I I I I I I f I I I I t I I I I I I I I

m ^ u aaartnJdDddddddddaJaJiDaoo M u m d cj d ^ ^ M£pu>a.ci.a*4J4j

Seasonal Time Period

Fig. 57. Seasonal distribution of adult Polites origenes in Ohio. 00 o

Fig. 58. Geographical distribution of Polites mystic in Ohio. (i) (11) (3) (12) (6) (1) No. of Adult Records t t » t t i » AAAAAAAAAAA * AAA AAAAAA A AAAAAAAAAAAA 1-10 1-10 1-10 1-10 1-10 1-10 1-10 1-10 11-20 11-20 11-20 11-20 11-20 21-31 21-30 11-20 21-31 11-20 21-31 11-20 21-30 21-31 21-30 21-31 Apr Mar Mar Apr Jun May Apr Mar Sep Sep Sep Jun Jun May May Jul Jul Jul Aug Aug Oct Oct Aug Oct ■ « 2 3 r—t 'Ji H* O 3 2 3 0 o . > 1 ill

Fig. 59. Seasonal distribution of adult Polites mystic in Ohio. 409

Fig. 60. Geographical distribution of Polites vibex in Ohio.

35 -

30 -

25 -

20 ■

10

5 ■

OO—«0 0 0 0 0 -h© 0 0 0 0 t-h0 0 -« © 0 0 0 0 -h I I I I t I I I I I I I I I I I I I I I I I I I H H H H H H H H H H H H H H pH >-< H H H H H H H pH H N H N -H CN H N <-H CM pH > >> a d d h h h wu)boaaa4J4J4J S SS<>n>Dfl)ooy Seasonal Time Period

Fig. 61. Seasonal distribution of adult Polites vibex in Ohio. 410

Fig. 62. Geographical distribution of Wallengrenia egeremet in Ohio.

35

30 ■ m U 25

20 -

15 -

10 - * •K« * * «* * * * * * * * *

00*-»00000'-<00000r-<00**-<00000*H —«cvim— I I I I I I I I I I I I I I ( I I I I • I I I t

I (M i cm i cm CM r-H CM i CM l CM u u m u u u a d 0HH»h »M tioaaaiJiJu ssac(DUOU Seasonal Time Period

Fig. 63. Seasonal distribution of adult Wallengrenia egeremet in Ohio. 411

Fig. 64. Geographical distribution of Pompeius v e m a in Ohio.

35 -

30 -

« * u 20 - * r-4 * D *■K < 15 - * •K* ■K * 10 * * * * * * * * * 5 * * * * * * * * * * * * * * 0 0 —< 0 0 0 0 0 —<00000'-<00'-«00000—'

I I I I I I I I I I 1 I 1 I I I I I I I I I I t

60t*>boaa

Seasonal Time Period

Fig. 65. Seasonal distribution of adult Pompeius verna in Ohio. 412

Fig. 66. Geographical distribution of Atalopedes campestris in Ohio.

*-H PI N H H N N

35 -

30 -

25

20

15 - o . 10 . zo 5 - « * * * * 00-^00000-^00000-<00^00000-H I I I I I I ! I I I I I I I t I I I I I I I I I

U U H t0M)00O.p.O.4J £ £

Fig. 67. Seasonal distribution of adult Atalopedes campestris in Ohio. H-4 LO ■O

Fig. 68. Geographical distribution of Atrytone logan in Ohio. (1) (4) (8) (2) (2) (4) ro - h No. of Adult Records **** ** * ******** ** **** 1-10 1-10 1-10 1-10 1-10 1-10 11-20 11-20 11-20 11-20 11-20 11-20 11-20 21-31 21-30 21-31 21-31 21-31 21-31 21-30 Sep Sep Sep Oct OcC Oct Jul Jul Aug Aug Aug Mar Jun Jun 11-20 AprApr 1-10 Apr 21-30 Jun Mar Jul Mar 1-10 May May May ft

Fig. 69. Seasonal distribution of adult Atrytone logan in Ohio. 414-

Fig. 70. Geographical distribution of Poanes massasoit in Ohio.

35 -

30 -

25 -

u 20 - rH3 < 15

10 ■

5 ■ * •K •K * *

oo*-»ooooo--^ooooo»-in-^Nr)HNro*-«-iNr,»H,^ j >,j Cd35dD3333Q)

Seasonal Time Period

Fig. 71. Seasonal distribution of adult Poanes massasoit in Ohio. 415

Fig. 72. Geographical distribution of Poanes hobomok in Ohio.

35 -

30 - 0} * * "S 25 ■ * * o « * u •K * 040) *« *« .. 20 « * * * * «* *« •K * «* * * * 10 - * * * * * * 5 - « « * * * * ****«««******* * * *

O © -<00000*-<00000»-*00»-<00000'H •-» ror)l-tNn>-*Nn,»M N n->>,>, 0 d CHHH Mtt060CLp.a4J.WiJ S S S H7»n»-)'-)'^*->

Fig. 73. Seasonal distribution of adult Poanes hobomok in Ohio. 416

Fig. 74. Geographical distribution of Poanes zabulon in Ohio.

35 -

30

25 -

20 -

15 - « 10 - * * •k* * * 5 - * * * •K* ** * * * * * * * * * .* * * * * * *

o o H O o o o o O o o o o f-H O o o o o o O -H eg co CM cn CM co •—< CM CO CM CO CM CO CM CO * CM co 1 * 1 1 1 1 1 1 i 1 1 1 1 1 1 1 1 1 * H H H H H H ^™4 H H i 1 CM *—4CM CM CM **4 CM CM CM CM u u U 60 60 a a JU 41 a S' OP <0 a O Sep Jun Jul Occ Jun Jul Aug Mar Mar Mar May Jun < < < S3 1 Jul •3 ■3 to to o O Seasonal Time Period

Fig. 75. Seasonal distribution of adult Poanes zabulon in Ohio. 417

Fig. 76. Geographical distribution of Poanes viator in Ohio.

35

30 -

25

Hu 20 - T)3 < 15

10 .

5 ■ « * * * * * * * oo»-«ooooo-'-«rsim>-«cMr-»—

I f-t rH bOM&oaaa*J4Ju 9 8*3'I 3 3 3 3 q 0) JU

Seasonal Time Period

Fig. 77. Seasonal distribution of adult Poanes viator in Ohio. 418

Fig. 78. Geographical distribution of Euphyes dion in Ohio.

r n o m h 35 -

30 -

Ot 25 - Oa> os » 20 - 3 <3 15 -

10 - * «* «* * * * * 4C « *

o O O o o o o H I •-» CM C O •-« C M I CMco H H I I I I I CMCO CMco H »i »—ii-H *-* C M H N — » C M H CM •“* CM H CM

Of) 60 M o. P* a u «J 3 3 3 01 a) 01 o C> O <<

Fig. 79. Seasonal distribution of adult Euphyes dion in Ohio. M -P- KD

Fig. 80. Geographical distribution of Euphyes dukesi in Ohio. (3) (3) (16) (9) (7) (4) No. of Adult Records *** ************************* ******* **** 1-10 1-10 1-10 1-10 1-10 1-10 11-20 11-20 11-20 11-20 11-20 11-20 11-20 21-30 21-31 21-31 21-30 21-31 21-31 21-31 Sep Sep Oct Oct Sep Oct 1-10 Jun Jun 21-30 Jul Jul Jun AprApr 1-10 11-20Apr Jul Aug Aug Aug Mar May May May Mar Mar H- a n> *0 to H

Fig. 81. Seasonal distribution of adult Euphyes dukesi in Ohio. 420

Fig. 82. Geographical distribution of Euphyes conspicua in Ohio. (1) (5) (1) (3) (1) 35 -

30 - N3 U l o f f I O U l No. of Adult Records t U l * ***** * * *** 1-10 1-10 1-10 1-10 1-10 1-10 1-10 11-20 11-20 11-20 11-20 11-20 11-20 11-20 21-31 21-30 21-31 21-30 21-31 21-31 21-30 21-31 Sep Sep Sep Jun Mar Apr Apr Jun Apr Mar Mar May May May Jun Jul Jul Jul Aug Aug OctOct 1-10 11-20Oct Aug a- ru a ■i

Fig. 83. Seasonal distribution of adult Euphyes conspicua in Ohio. 421

Fig. 84. Geographical distribution of Euphyes bimacula in Ohio.

35

30 -

25 -

20 -

15 -

10 -

* * * «* * * * * *

O"-4C'l *-«n H N H N u u m u m m >s d a dhhh boooooa&auuu ididdfra&nl(IIJ33S9333994lilltlOO.U

Seasonal Time Period

Fig. 85. Seasonal distribution of adult Euphyes bimacula in Ohio, K> -O

Fig. 86. Geographical distribution of Euphyes ruricola in Ohio. (3) (5) (1) (1) (1) (1) U > (4) - S3 h No. of Adult Records * * **** *** ***** * * * 1-10 1-10 1-10 1-10 1-10 1-10 1-10 1-10 11-20 11-20 11-20 11-20 11-20 11-20 11-20 21-30 21-31 21-31 21-31 21-31 21-31 21-30 Sep Sep Sep Oct Aug Oct Jul Jul Aug Oct Aug Jun Jun Jun Jul Apr Apr Apr 21-30May Mar 11-20Mar Mar May May o 3 fi) H* 3 ro ro H- O

Fig. 87. Seasonal distribution of adult Euphyes ruricola in Ohio 423

Fig. 88. Geographical distribution of Atrytonopsis hianna in Ohio.

35

30

25 -

20 ■

15 -

10 -

5 -

O O — ,> .> , a a g h h h booot>oap.a«i4J4J

Seasonal Time Period

Fig. 89. Seasonal distribution of adult Atrytonopsis hianna in Ohio. 424

Fig. 90. Geographical distribution of Amblyscirtes hegon in Ohio.

35

30

25

u 20 ' r-H •O3 < 15 ’

10

5

00rH 000© 0'-'00© 00«-»© 0'-<00000'“-«

I I t I I I I I I 1 I I I I I I I I I I I t I I

a drjriH u t t u & a a iJ u u «}(d3S3333333a)UtJU

Seasonal Time Period

Fig. 91. Seasonal distribution of adult Amblyscirtes hegon in Ohio, 425

Fig. 92. Geographical distribution of Amblyscirtes vialis in Ohio.

30 -

25 -

20 - 3 TJ < 15

10 -

5 _ -K 4C J * * * * * * * __* * * * *«K * «*** * * OO^OOOOO^OOOOO'-'OO'-'OOOOO-H ^Nn'-»Nr>'-'Nn*HNn*^Nrn<-4NfnHNr>'-

Seasonal Time Period

Fig. 93. Seasonal distribution of adult Amblyscirtes vialis in Ohio. 426

®r

Geographical distribution of Panoquina ocola

35 -

30 -

C 2 5 -

20 -

15 -

10 •

5 •

O0*-»00000»r«00000*-*00*-^00000*-* l i | * l I l I l l I I I I I I l I I I I I I I — 1 f\| -4 > n a a h h h bObDboaaauww

Seasonal Time Period

Fig. 95. Seasonal distribution of adult Panoquina ocola in Ohio. 427

Fig. 96. Geographical distribution of Battus philenor in Ohio.

i-H t*> oo co CM 00 CO VO W> » 35 -

30 -

25 -

20 -

15 -

10 - * « * * * 5 - * * * * * * * * * * * * * * * * * ***** * *

•^NM>-*(MnHNn^Nr,>HN

rHI H I HI I HI HI HI I H IH HI HI HI H I HI HI HI HI HI *“4I »“4 I IH H I IH HI •—» CM *-• CM *-• CM *—* CM H CM H N ^ CM CM m u u m u m >>$»>> a a a h h m Meot>ofraaw4J*j ifliddD.ao

Seasonal Time Period

Fig. 97. Seasonal distribution of adult Battus philenor in Ohio. 428

O I A

Fig. 98. Geographical distribution of Eurytides marcellus in Ohio.

t CO t-s /*—s /-> \ <-h ^Hrimco V W N—<* »w> O S_> 35 -

30 -

25 -

20 - * « * 15 - * * * « * ** 10 - * 4c o * « z 4C * * * 4C * 4C •K 4C 4C * 4C * 4C 4C 4C 4C 5 - 4C «* * 4C 4C 4C 4C 4C 4c 4C * 4C * 4C 4C 4C 4C 4C * * * 4C « 4C 4C 4c 4C 4C 4C * * * 4C 4c 4C 4C 4C 4C 4C 4C «** * 4C 4C 4C * 4C 4C 4C OO-^OOOOO^OOOOOfiOO'HOOOOO^ I I f I I l I I I I I I I I I I I I I I I I I I

u m p p p p p*. a a dpiHH Dcttobcaaau^u aMid(qaD,a'o,-><

Fig. 99. Seasonal distribution of adult Eurytides marcellus in Ohio. 429

Fig. 100. Geographical distribution of Papilio polyxenes asterius in Ohio.

s s cbCbincocsoio^HHcooirioironn »>“N / —\ s / " s r H O /■**» N «■—>.

35 -

30 -

25 -

20 -

15 -

10 - «« ** « ** * ** «*** * * * **« « * * *« * **«* * • 5 - * ***« *** *** *** * «* **« « «*«* **« * *«* **««** ** «*** **«*««** ******«** ** ■K« ******« * o o —»ooooo^ooooo*-*oo—*ooooo-< HNrn*HNn»-

Seasonal Time Period

Fig. 101. Seasonal distribution of adult Papilio polyxenes asterius in Ohio. 430

r1 1

Fig. 102. Geographical distribution of Papilio cresphontes in Ohio.

35

30 - m *2 25 *■ 0o 01ai .. 20 - 9 <1 15 -

10 -

5 - «* •K* * * * * * * * * a u u u l 00»--t©0000»-‘00000«-*00—‘©OOOO-t ■^Nfo^Nn^Nni-tNnHNn^Nn^Nn^NP1)I I I I I I I I I I I I I ! I I I I t I I I I I H H H »-< H H H H H H H r—4 H H M H H r—I H H f—t H H »-t C4 •—* C\| H N •-< N N ts I-I N «-l CM

ft h h iu >s >s >, d S (9HHH bO 60 00 (X o> a 4J . 99999999Q>a>uot>

Seasonal Time Period

Fig, 103. Seasonal distribution of adult Papilio cresphontes in Ohio. Fig. 104. Geographical distribution of Papilio glaucus in Ohio. (2) (7) (11) (11) (10) (10) (17) (17) (8) (8) (11) (5) (4) (7) (8) (19) (15) (11) (11) (4) (1) “ ~ N > ISJ o Wi o ui Ho. of Adult Records ** *********** ******* ***************** ********** ***************** ******** *********** ***** **** ******* ******** *************** ******************* *********** **** * 1-10 1-10 1-10 1-10 1-10 I-10 1-10 1-10 I 11-20 21-31 11-20 11-20 11-20 21-30 11-20 21-31 11-20 21-30 11-20 | 21-31 I Mar Mar Mar Apr Apr 11-20 Apr 21-30 May May Jun May 21-31 Jun Jun Jul Jul Jul Sep Aug Aug Sep Sep Aug 21-31 Oct Occ Oct CO ra » CO o g 01 H- g CD 'TO CO n 0 r>

Fig. 105. Seasonal distribution of adult Papilio glaucus in Ohio. ro x> U) K

Fig. 106. Geographical distribution of Papilio troilus in Ohio. (11) (1) (1) (2) (12) (7) (9) (11) (3) (5) (1) (12) (6) (7) (6) (2) (2) N) tsJ Ui UJ - l t » » r h t No* of Adult Records » Ul Ul O US O Ul O Ui ************ * *********** ********* *********** *** ******* ****** * ** ** 1-10 * I-10 1-10 1-10 1-10 ***** 11-20 11-20 11-20 ******* 21-31 21-30 ** Oct SepSep 1-10 11,-20 Sep 21-30 Oct 11-20 Oct Aug 11-20 ************ Jun Jun Jun 21-30 Jul Jul 21-31Aug ****** Aug 21-31 Apr 1-10 Apr May May 21-31 Apr 11-20 Jul Mar 11-20Mar 21-31 May Mar 1-10

Fig. 107. Seasonal distribution of adult Papilio troilus in Ohio. 433

Fig. 108. Geographical distribution of Pontia protodice in Ohio.

IT) N N rl rl CM in H H H N Cl H N

35 -

30 -

25 -

20 - d < 15 -

10 -

5 - * * * •a * * * * * •K * * * * * * * * * ** * * * * ' f-l O o O o o o o «“4 H N C O H N co CMcn ■CMr>• *-4 i CM■m »“4I I H I H I H tH H1 1 1 1 1 *4HI H i H1 H1 —4 CM *-4 CM r4CM •—4CM H CM MMMWHH£spsr*dfldHH to toto& a CL 4J4JV «iflifl&aartnlrt9333D 3 d 3 3 0) 0) ai o O 0 < toCOw o o o Seasonal Time Period

Fig. 109. Seasonal distribution of adult Pontia protodice in Ohio. 434

Fig. 110. Geographical distribution of Artogeia virginiensis in Ohio.

/-s fl O /"N , <9 H <-» fO I 35 -

30 -

25

20 *d3 <15*

10 - * * * * « * 5 - * * * ■K* *« *« *« m n c s i n B m B B 00-400000* lOOOOO-^OO—‘00000-* --iMnHNnl I I I I I I l I I I I I I I i r-j H t>0to0bDaaa4J*J saa-) b b b Seasonal Time Period

Fig. 111. Seasonal distribution of adult Artogeia virginiensis in Ohio. 435

Fig. 112. Geographical distribution of Artogeia rapae in Ohio.

/■“s /*■*>. / —N s />“ S («*> /* * / - s /•s rs vO P") H H <*“s CO *•“* O Is* O 0*1 ^ i^s H'd'lOMHvDHNHNnH^HHHH^fNHCNKr W 'w' W W W 'w' X_»- V^> ^ 'w ' <«/ V./ W' >k/ V W V 35 -

30 -

X 25 - X « * * * 20 - * * X * XX X XXX X X XX 15 - XXXX X *XXX X X * XXXX X*XXXX *X * XXXXX 10 - «* * XXXXXXX X* * XXXXXXXX X*X X X X XXXXX * X X •X * X X XXXXXXX * X * XX XXX XXXXXXX 5 - X X X XX X XX X XXX XXXX X X * X XX X XXXXXX XXXX X X * X ■X •X XXXXXXXX XX X XX * XXXXXXXXX X X X X XXXX * * ** * •X XXXXXXX XX X XXX X X 00*-»00000*-«00000*-i00'-<00000*-» <-tNn^Nr>*-«c'inHNm^Nn<-iNn^Nn'-*NcnI I I I I I I I I I I I I I I I I I I I I I I I

1333300)0)000M b&boua&aiJU W W WOOO

Seasonal Time Period

Fig. 113. Seasonal distribution of adult Artogeia rapae in Ohio. 436

Fig. 114. Geographical distribution of Falcapica midea in Ohio.

35 -

30 -

25 -

* « it 20 - * * •K * •K * « * * * * * * 15 - « * •K -K « * * * * * * * 10 • * * * * « * * * * « * * * * * * * * 5 - * « i t H * « * « « * * « * * * * * * * * * * * *

©0*-<©©Q©0*-<©0©©0*-<00»-«000©©»-« HI H I Ht i-4 I »—I I H I H I »—4I *«< I *-4 I HI H t H I H I H I •—I I H I H t H I H I H I H I H I H I H N H N •—t CM f-4 N *-4 N <-4 CM H N «-< CM ...... « 6H H H bOttfibO&p.&4JiJU aagih'ih*i

Fig. 115. Seasonal distribution of adult Falcapica midea in Ohio. 437

Fig. 116. Geographical distribution of Colias philodice in Ohio.

35 -

30 -

« * *« 20 - * * * * « *« * * * * * «* * « * « * « « * « * * * * 10 * * * * * * * * * * * * « * * * * * * * « * * * * * « * * * * * * * * * * * « * * * « * * * « * « * * * * * * * * * * * * * « * * * * * * 5 ■ * « * « * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * « « « * * * * « « * * * * * * * * * * * * * * * * * *« * «* « * * « « * *

OOrHOOOOO—'OOOOOtHOO^OOOOO*-! •-

Fig. 117. Seasonal distribution of adult fjlias philodice in Ohio. 438

Fig. 118. Geographical distribution of Colias eurytheme in Ohio.

/—s v.—s,—<.HoQr'-roiovoco

30 -

25 -

20 * 15 “ * * * **' * * *« * * * 10 ■ « * « * * * * * * * * * * * * * *...... * * * * «* * * * * 5 ■ * * * * * * * * * * * * * *********** * * * * * *********** * * * * *

OO—'OOOOO-HOOOOO^OO-^OOOOO-* -+-j,-5^

Fig. 119. Geographical distribution of Colias eurytheme in Ohio. 439

Fig. 120. Geographical distribution of Zerene cesonia in Ohio.

35 -

30 -

25

20 -

15 -

10

5 ■

HNn-^Nn-tNn-iNnHNPiMNnHNfOHMn)oo^ooooo^ooooo^oo^ooooo-^ f I I « I 1 I I I f I I I I I I I I I I I I I ! —I CNl » >> io^ io>> § § §o I"3 'a 3 3 3 a> Si 3 u u u

Seasonal Time Period

Fig. 121. Seasonal distribution of adult Zerene cesonia in Ohio. 440

Fig. 122. Geographical distribution of Phoebis sennae eubule in Ohio.

rH CM CM rH

35 -

30 -

25 -

20 -

15 -

10 -

5 - * «* * *

-^Nn-'Nn"(Nnf*tNnHNni-*Nf'i"

Fig. 123. Seasonal distribution of adult Phoebis sennae eubule in Ohio. 441

Fig. 124. Geographical distribution of Phoebis philea in Ohio.

35

30 -

25 -

20 •

15 -

10 - 55o 5 -

-^Nn<-

Fig. 125. Seasonal distribution of adult Phoebis philea in Ohio. 442

Fig. 126. Geographical distribution of Eurema lisa in Ohio.

35

30 -

25 -

20 -

15 -

10 - sso * * «* *« 5 - * * * * « * * * * * * * * * * * « * oo-^ooooo-^ooooo^oo^oooo©*-* i i t i i i i i i fM'Nr. i M<-«N(ni-

Fig. 127. Seasonal distribution of adult Eurema lisa in Ohio. 443

Fig. 128. Geographical distribution of Eurema nicippe in Ohio.

35 -

30 - (0 "S 25 ■ uo 020) . 20 ■

15 -

10 - « * * 5 - * * *« ******** * * <*«««««« «. 00*-»00000*-'00000^00*-'00000^ ^NfriHrjn'-'Nro-iNni-'Nco^Nn^C'jn'^MrO I I I I 1 I I I t t I I t I I I 1 I I I I I I I H H H H H H *—I H H H H H H i— t H M H H H H H H H fs| — < rs| {S —♦ CVJ t-<«M N *-• N *-4rl <9 .3 *3 o* o. p. « r t & 3 9 3 p p p 3 d 3 Q ) 0 ) a ) 0 0 0 SSS

Fig. 129. Seasonal distribution of adult Eurema nicippe in Ohio. -P*

Fig. 130. Geographical distribution of Nathalis iole in Ohio. (1) (1) (1) (2) ■— ■— N> No. of Adult Records * * * ** Sep 1-10 Sep 11-20 Sep 21-30 Mar 1-10Mar 11-20 Apr 1-10 Mar 21-31 Apr 11-20 Apr 21-30 Jun 1-10Jun 11-20 Jul 11-20 Jul 21-31 Aug 21-31 Oct 1-10Oct 11-20 Oct 21-31 May 1-10May 11-20 May 21-31 Jun 21-30 Jul 1-10 Aug 11-20 Aug 1-10 C/5 to a co o 3 H* H 3 (D US to n

Fig. 131. Seasonal distribution of adult Nathalis iole in Ohio 445

Geographical distribution of Feniseca tarquinius in Ohio

r-t r-H NMNNNNVO«-l •fHP'J 35

30 0} m 25 o

*,20' pH T39 < 15 ‘ U-lO - 10 -

*************** **** *

-(('in^cNfo^Nn^Nn-'NOHNn-NNnHNrt00^00000*^00000^00^00000*-^ i i I I I I t I i i t t I i I t i I t l i I I I

n u u u u m >,>*>, e c b h h h &otio&oa&a4Jut) (drecaao.&.(0{0(0999999999a)aia)uuo SSS<<

Fig. 133. Seasonal distribution of adult Feniseca tarquinius in Ohio. 446

Fig. 134. Geographical distribution of Lycaena phlaeas in Ohio.

35 *

30 -

*2 25

20 - T33 < 15 - U* « o ■K« . 10 - « o * * * « z XXX * -X X X X X ■X -X -X X XXX X 5 - XXX X X X X X xxxxxxx X X xxxxxxxxx X X X xxxxxxxxx X X X x xxxxxxxx X X X X

o © © © OO p 4 © o © o © r-4 h cs n n n i CMm CMm CM Y CMcn Y CM cn 1 1 t T t l I I I I I I ^4 r*4 pN CM CM Zl CM CM CM c C rH •H 000000a a a 4->iJ 4J Riffifljfli&Oitiininis 3 3 3 3 3 3 3 3 0) a) ai U u O n •—} *-> •“5 < < < COCOCO o o o Seasonal Time Period

Fig. 135. Seasonal distribution of adult Lycaena phlaeas in Ohio. 447

Geographical distribution of Hyllolycaena hyllus in Ohio

35 -

30 - 0) *20 25 - 01u 06 u 2 0 - ■K iH * 3 * < 15 - «* U-l «* * O * ■# . 10 - * * o « * Z « * * •K 5 - * *« * * * * *

OO-^OOOOO-HOOOOO^OO'^OOOOO'- t i i i t i t i i t i i i t t t t i i i i i i i

M U U Ul U U ^ > n S s C C C H H H 60 00 00 & f t Oi 41 44 4J ranjajcua.o-roaJrt=>3333D333a>a>a)uou SES<:<

Fig. 137. Seasonal distribution of adult Hyllolycaena hyllus in Ohio. 448

Fig. 138. Geographical distribution of Epidemia helloides in Ohio.

35 -

30 -

25

20 -

15 -

. 10 -

5 -

© o O O O © © -<©©0©©»-«©©-«©©©©©i-* --Nn-NfnH(NnHCNnHNn-«fNnHCNn--CMn i i i i i t i i i i i r i i i i i i i i i t i i ■—* CM •-* CM •■M CM CM

M U U U U U >> >> >s G G B W H H OOOOOOD.O.D.4J4J4J tdrtfl}&.CL&.nJcOrt333333333Q)

Seasonal Time Period

Fig. 139. Seasonal distribution of adult Epidemia helloides in Ohio. 449

Fig. 140. Geographical distribution of Harkenclenus titus in Ohio.

/■“s iri rs © /«“s /»s

35

30 -

"2 25

« 2 0 p H 3 •K« < 15 ■K* * «H « * o * * . 10 - * * * * * * «K * * 5 *» * * * * * * * * * * * * * ********

oo^ooooo«-*ooooo^oo-*ooooo»-< NNnNNn-tMn^NnrtNinNfMnHNnHNn i I I I I i i I I i i I I i i i i i i i i i t i ^ ^p H »H pH »H pH pH pH pH pH H >H pH pH Pi pH pH pH pH pH »H pH pH pH CM pH CM pH CM H N H N pH CM pH CM pH CM u u u u u u >,>>>> c e ch hh ueob&a&auuu nmna&D.o]iiij333333333aiiiiiiuuu

Seasonal Time Period^

Fig. 141. Seasonal distribution of adult Harkenclenus titus in Ohio. Fig. Geographical distribution of Satyrium acadica in Ohio

H H VO M VO

35

30

K 25

oc* « 20' "O < 15

zo * * «* « * * *

00*-^00000^00000»-<00-^00000'-* ^Nn<-»N«HNr)'-'Nn^N

l CM «-a>a)uuu n u u u u >, >> >n c a c h h h botio&oaao.4Ji>u

Seasonal Time Period

Fig. 143. Seasonal distribution of adult Satyrium acadica in Ohio. Ln 4>

Fig. 144. Geographical distribution of Satyrium edwardsii in Ohio. (2) (2) (4) I O f Ln No. of Adult Records ** ** **** 1-10 1-10 1-10 11-20 11-20 11-20 11-20 11-20 11-20 21-31 21-30 21-31 21-30 21-31 21-31 Oct 21-31 SepSep 1-10 Sep 21-30 Oct 11-20 Aug Oct Jul JunJun 1-10 Jun Jul AugAug 1-10 May Jul 1-10 MayMay 1-10 Apr 11-20Apr Apr Mar Mar Mar o n 3 9 K* ro W O

Fig. 145. Seasonal distribution of adult Satyrium edwardsii in Ohio ) ) ) ) 1 21 8 6 Cl) (18) ( (1) (4) (19) ( ( ( to m 1— 1— No. of Adult Records * * AAAA AAAAAAAAAAAAAAAAAAA aaaaaaaaaaaaaaaaaaaaa AAAAAAAAAAAAAAAAAA AAAAAAAA AAAAAA * 1-10 1-10 1-10 1-10 11-20 11-20 11-20 11-20 11-20 21-31 21-31 21-30 21-31 21-31 Sep 1-10 Jul Sep Sep 21-30 Oct Apr 11-20 Apr 21-30 Jun Jun Jun Jul Aug Aug OctOct 1-10 11-20 May 11-20 Mar Apr 1-10 May May 21-31 Jul MarMar 1-10 Aug to (Q o 3 Cb H* H S' re re <0

Fig. 147. Seasonal distribution of adult Satyrium calanus falacer in Ohio. 453 1

Fig. 148. Geographical distribution of Satyrium caryaevorum in Ohio. ) 2 (4) (3) ( No. of Adult Records **** *** ** 1- 1- 1- Sep 1 Sep 11 Sep 21 Apr Apr U Apr 21 Jun 11 Mar Mar May May 11 Jun 1 Jun 21 Jul 11 Jul 21 Aug 11 Aug 21 Oct 1 Oct 11 Oct 21 May 21 Jul 1 Aug (t » to o a p. H S’ It U) It c/> Marc/> 21- H

Fig. 149. Seasonal distribution of adult Satyrium caryaevorum in Ohio. 454

Fig. 150. Geographical distribution of Satyrium liparops in Ohio.

35

30 -

CO u 25 - o

10 -

5 -

oo^ooooo-^ooooo^oo^ooooo^ i I I I I i I i I I i I I t i I I I I I I ( I I

M U U U U U >> >s C C 3 f-4 *H pH CO&OfiOaa&UUU SSS

Fig. 151. Seasonal distribution of adult Satyrium liparops in Ohio. 455

Fig. 152. Geographical distribution of Calycopis cecrops in Ohio.

35 -

30 ■

"S 25

« 20 - 1^3 TJ < 15 -

10 -

5 -

<—♦rgfO^rsim^cNcnt-'cNpn^cNiPn^cNP^^cNrn^cMmOO^OOOOO'-HOOOOO^OO-^OOOOO'-* I I I I I l l I I I ) I I I I I I I I I ( I I I *-4 CM CN -H tN CN —* CM CN CM —4 fs|

U u u W- U M >> fN >, C G C rl rl H 60&ObOO.O.O.iJ4JlJ £ass<<

Fig. 153. Seasonal distribution of adult Calycopis cecrops in Ohio. 456

Fig. 154. Geographical distribution of Mitoura gryneus in Ohio.

M m ( M M

35 -

30 -

3

O . 10 - o z

« * « •K * * * * * * •K 01- o © o © © o o © O H O o o o CMm CMr n CN ? CMrn CMcn CMrn CMcn 10 CMcn -10 -10 I 1 -10 -10 l — ^■4 *«■# - cm CM 2 CM CM CM CM CM CM u u u M > 1 >> >% c a T—| •“ 4 00 00 00 a a. u JJ co a o . o . a CO CO CO 3 3 3 3 3 3 3

Fig. 155. Seasonal distribution of adult Mitoura gryneus in Ohio. 457

Fig. 156. Geographical distribution of Incisalia augustus in Ohio.

CS *3- CN 35 -

30 -

m 25*

20 - •O3 < 15 1

10 - o z 5 -

•HC'ifi^Nn-NNn^Nrt^Nn*^c'i<,n

u u u u m m ss >■»>■» c s C r t w w oooooofraawu U zss<

Fig. 157. Seasonal distribution of adult Incisalia augustus in Ohio. 458

Fig. 158. Geographical distribution of Incisalia irus in Ohio.

35 -

30 -

25 -

« 20 - •33 < 15 1

10 - zo 5 - «* « *« HN

—« CM pm n —« CM fvj U U U M U U >,>,>,£ C CHHH 00 00 00aCUQ.a)

Fig. 159. Seasonal distribution of adult Incisalia irus in Ohio. 459

Fig. 160. Geographical distribution of Incisalia henrici in Ohio.

35 -

30 -

U 25 o

« 20- ^4 T)3 < 15 -

10 • o z -K « « « 5 - -K •4C* ** * « * « «* * *« *« *«

o o ^4 o OO O o ^4 © o OO O o O o o o o o CN m CN cn CN CO 71CN cn CN cn *■* CN cn r* CN cn CN cn 1 I | 1 t 1 l 1 1 H1 1 i, i 1 1, FHi il i J. 1 H —♦CN CN CN —< CN ^4 CN «4 CN CN CN u M U u UU Sn 5K Jn e c e •H H rH 00 00 00 a & a. 0) a> a O O £ £ £ < < < £ £ £ •“J •n •X *n < < < CO CO CO o © o Seasonal Time Period

Fig. 161. Seasonal distribution of adult Incisalia henrici in Ohio. O O'

Fig. 162. Geographical distribution of Incisalia niphon in Ohio. (3) (3) (3) cn (2) (5) (6) (3) No. of Adult Records ****** *** *** *** * ** ***** 1-10 11-20 11-20 11-20 21-31 11-20 21-31 11-20 11-20 21-31 21-31 21-30 Sep Oct Sep Sep 21-30 OctOct 1-10 Aug Jul Jun 21-30 Jul 11-20 AugAug 1-10 JunJun 1-10 Apr May Jul 1-10 Apr 11-20 May 1-10 Mar Apr 1-10 May Mar 1-10' Mar 21-31 y ft ►tj ft ►i H

Fig. 163. Seasonal distribution of adult Incisalia niphon in Ohio 461

Fig. 164. Geographical distribution of Euristrymon Ontario in Ohio

30 -

u 25 -

20 - a < 15

10 • o z 5 -

00^00000^00000*^00^00000^ HCMnHCMnHNm-NnHcsfiHNnHNnHMni i I i I I I I i i i I i i I I i I i i l I I i

u U U U U >■» >, >, C c C*-4r4i-i UbOCiO&aailtJU SS23<<'->*-)

Fig. 165. Seasonal distribution of adult Euristrymon Ontario in Ohio. 462

Fig. 166. Geographical distribution of Parrhasius m-album in Ohio.

35 -

30 - 0) u 25 ■ uo 0&Qi " 20 • •O3 < 15

10 - o z

o o ^ o o o o o —»ooooo*^oo«-*ooooo^ I ( i i i i i I i i i I I i I i I I I I I » I I —* CM «-4

U u U H U U >S > S >s G G C H H H fiO&OOOa&aUiJiJ SCDn)«aO.O.Cdrta)333333333

Fig. 167. Seasonal distribution of adult Parrhasius m-album in Ohio. 463

Fig. 168. Geographical distribution of Strymon melinus in Ohio.

35 -

30 -

25 -

H* 20 " TJ3 < 15

10 - * o * * •K* -K« « * «•K 5 - « * « •K * •K * « « «* « *« *« «* * * * « « « « « « *•K « «* «* ©©'-0©©©©*-*©©00©«"*©©-*©©©©©«-* i i I i i i I i I I I i I I I I i I i I I i i I

u u u m m u >%>>>> a c c h h h ooooooaaauuu sss<<<:sss,n^*->nr-)>-5<<

Fig. 169. Seasonal distribution of adult Strymon melinus in Ohio. 464

Fig. 170. Geographical distribution of Erora laeta in Ohio.

35 -

30 •

a 25 -

20 • •O3 < 15' tt* o . 10 ■ o a s

00»-*00000—'OOOOO^OO-^OOOOO-^ I I I I I I I I I l I l I I l I I I I I I I I l

„ u „ „ u_ U 5** J S P> G d C H H H. . QOooooabfruiJiJ ««flJQ.a.Cl.n)tOnJ33333Qi c l a q) cd id 3 *-j

Fig. 171. Seasonal distribution of adult Erora laeta in Ohio. 465

Fig. 172. Geographical distribution of Hemiargus isola in Ohio.

35 -

30 -

25 -

20 - s < 15

10 •

5 -

o o -h o o o o o -

m N *-** >> >n C C C H H H 00 00 00 O. Q. O. -U 4J 4J

Seasonal Time Period

Fig. 173. Seasonal distribution of adult Hemiargus isola in Ohio. O' a\

Fig. 174. Geographical distribution of Everes comyntas in Ohio. (8) (8) (15) ( i ) (10) (10) (5) (8) (10) (9) (19) (23) (2) (3) (15) (15) (10) (3) (1) (1) (7) H- »-« W NJ CO CO No. of Adult Records I I f i i i i t Cn O Cn O Cn O Cn *************** ***** ******** ********** * ******** ******** ********** ********* ******************* ********** ********** * *** ** *** ******* *************** *************** *********************** * 1-10 1-10 1-10 1-10 11-20 11-20 11-20 11-20 11-20 11-20 11-20 21-31 21-31 21-31 21-31 Sep 21-30 Sep Sep JunJun 1-10 Jun 21-30 Jul Jul 21-31 Oct Oct Oct 1-10 Aug Apr 21-30May May Aug Apr 1-10 Apr 11-20 May Jul Aug MarMar 1-10 Mar

Fig. 175. Seasonal distribution of adult Everes comyntas in Ohio. 467

© I ©

Fig. 176. Geographical distribution of Celastrina ladon in Ohio.

^ GO V 0 / “ s N ^ H / - s O •"*s

35 -

30 - * * u 25 ■ * o *•K * u 20- * iH « •K 9 * * •O •* * < 15 - « « * « * * « « * * * * 10 - « « * * -K * * *« *« * * * * * *« *« * * * * 5 - * * * * ****** * * * * *********** * * * * ************ * * * * _«o O o o © o © o O O © O pH O o o o o H m CMcn CM cn CM cn CM cn CM cn CM cn »*** CM cn I 1 i l 1 1 I 1 1 I 1 l 1 I T I I | H H w * *■4 pm p4 w pM ^4 CM CM PM CM CM CM CMCM u U M >* Ps c a C rH 00 00 00 a a a. u 4J JJ a Qu cu CU CO CO CO 9 9 9 9 9 9 9 9 9 0) 0J 0) o a O X < < < X X X *-) *-> *") •“3 •“} < < < COCO CO- o o o Seasonal Time Period

Fig. 177. Seasonal distribution of adult Celastrina ladon in Ohio. 468

Fig. 178. Geographical distribution of Celastrina ebenina in Ohio.

35 -

30 •

25

15

10 -

5 -

o © 1-4 O o o o O — i o © o o o •M o o O o o o o *—4 CM cn CM cn CM cn H CM cn CM cn CM cn CM CM | cn cn V 1 l I ( 1 l l 1 1 l 1 1 1 t t l «—4 ^4 — —N <—4 •*4 ^4 CM •—4 CMCM CM z CM CM *—4 CM CM

U U u u U u >, 5s C e C H H H &OOOOOQ.aO.TTTTl<<

Fig. 179. Seasonal distribution of adult Celastrina ebenina in Ohio. 469

Fig. 180. Geographical distribution of Glaucopsyche lygdamus in Ohio.

35

30 -

"2O 25

« 2 0 ' H "O3 < 15 1

10 < o SB * * *« « * *« *

o o -h o o o o o ^ o o o o o — o o - < o o o o o ^ I i i i t I i i i i l i l t i i i i I i t I t t

U u M M u U >>>>>, C C C H H H OOfiO&OaaO.U'U Era(Q(aa.o.o.racon99999SS999a)a>a)ucju S 2 3 < -)'-3'-3'r)>r><<

Fig. 181. Seasonal distribution of adult Glaucopsyche lygdamus in Ohio. 470

Fig. 182. Geographical distribution of Lycaeides melissa in Ohio.

35 -

30 ■

25 -

20 • •d3 < 15

10 -

5 - *« * — —.— — warn— — WM— WUJ-LUII'BIUJMII.IIMHWIMWM—* * * * B t* — — — — — — m — . •NCMn-tMnHfsjcnHNnHNnrtfMnHpgnHNnO O — 00000-^00000-<00-H00000^ i I i i I i I f i I i ( I i I I i I i I i I I I

u u u u u m >>>>>> c c c h h h ooooooa&buuu s«njrta.o-o-rtfln}333333333a>a)uoo s s < c < s s s ,n>-j^,-)^^<;<

Fig. 183. Seasonal distribution of adult Lycaeides melissa in Ohio. Fig. 184. Geographical distribution of Calephelis borealis in Ohio.

) 10 (2) (2) (1) (26) (27) ( (3) No. of Adult Records ** ************************** *************************** * ********** *** 1-10 1-10 1-10 1-10 1-10 1-10 1-10 1-10 11-20 11-20 11-20 11-20 11-20 11-20 11-20 11-20 21-31 21-30 21-30 21-31 21-31 21-31 21-30 Jun Jun Jun Mar Sep Sep Sep Apr Jul Jul Apr Apr Mar Jul Mar Oct Oct Aug Aug Oct May 21-31 May May Aug J ro o o M ID *T} cn It w p 1-0 H- g

Fig. 185. Seasonal distribution of adult Calephelis borealis in Ohio. 472

Fig. 186. Geographical distribution of Calephelis muticum in Ohio.

m u i en 35 -

30 -

2 25 -

20 -

15

• 10 -

5 -

0 0 ^ 0 0 0 0 © ' - * 0 0 © 0 0 ,--<00'-'00000'--' _L_L_L_L_L_L* 1 r * 1 1 1 1 * * * i i * i i i » —• CM — CM CM pH CM U U U U U U > » >N 5 s c e ci i-i h h uifioaoa&auuii re rere o< o< d < to re re 33D333D3Da)a)0)OOU z z z < < < z z z h)TTTTl<<<{WMKIOOO Seasonal Time Period

Fig. 187. Seasonal distribution of adult Calephelis muticum in Ohio. 473

Fig. 188. Geographical distribution of Libytheana bachmanii in Ohio.

35 -

30 -

u 20 - 3 T 3 < 15

10 •K *« 5 * * *•K *« «* «* * •K * * * * •K "K * * * *

-NnHNMHNm-'Nn-itvjnHNn-tNn'-NriO O — O O O O O — O O O O O — OO — O O O O O — I I I I I l I I I I I I I I I I I I I I l I I I

e e *-t *h *h 6aoo6oaaauuu fljtonjaa&Qconis 33333333'->'-}|-J ' - 3 < < < C / J W W O O © Seasonal Time Period

Fig. 189. Seasonal distribution of adult Libytheana bachmanii in Ohio. 474

Fig. 190. Geographical distribution of Euptoieta claudia in Ohio.

i-« i- i

35 -

30 -

25 -

x, 20 * *H3 < 15 '

10 ■

5 ■ •K * * * 00-^00000—^00000' -Nn--Nn-HNn-Nrni-ifsi i-*Nn-HNn-*Nn<0©-*0©000—« I I I I I I I i I I I I I I i i i i I I i i i

u u u u m >»>»>% e e ooQOfioaaauuiJ fljC0aa.a.CdnjCU333333333a>

Fig. 191. Seasonal distribution of adult Euptoieta claudia in Ohio. -P- ^sl Ln w 7 0 H y i — v 0 1 m l $ I $ I L @T _ r I W' ® 1 t

Fig. 192. Geographical distribution of Speyeria cybele in Ohio. ) 21 (2) (24) (9) (3) (7) (8) (31) ( (5) (9) (1) (2) (2) (1) • h- h- h- hJ N> No. of Adult Records ** ******** ************************************************************************************* *** ***** ******* ** ********* ** 1-10 1-10 1-10 1-10 1-10 1-10 * 1-10 * 11-20 11-20 11-20 11-20 11-20 11-20 11-20 11-20 21-31 21-30 21-31 21-30 21-31 21-31 21-31 21-30 Sep Sep Jun Sep Mar May JunJun 1-10 Jul Jul Mar Mar May Apr Apr Apr May Jul Aug Aug Aug Oct Oct Oct o 3 P> H H* 3 ro nd (t) H c/i ro » to

Fig. 193. Seasonal distribution of adult Speyeria cybele in Ohio. 476

Fig. 194. Geographical distribution of Speyeria aphrodite in Ohio.

35 -

30

20 3 < 15

10 ■ * * * * * * * * * * * * * * * 5 * * * * * * *******. . * * * * * * * ********* * * * * *

OO-^OOOOO'-HOOOOO-^OO^OOOOO — «-iNri<-*Nn^(sn-HNDrtCNn»HNn<-iNn"

— H %>>>, C C C H H H &OC1OC1C1Q.&&UUU sss:<;<^^^’n»-3<<

Fig. 195. Seasonal distribution of adult Speyeria aphrodite in Ohio. 477

Fig. 196. Geographical distribution of Speyeria idalia in Ohio.

35 -

30 • w U 25 ■ o 0)u OS « 20 ■ fH3 < 15 '

. 10 - o z 5 - * * * * * * * * * ******* * * * -K *

OO-HOOOOO — O O O O O -^O O —'O O O O O — 1 I I I I I I I I I I I I I I I I I I I I I I I

UHraraaaQ.l5U(lIDD33333DDllllia)UOU M M U M M >> >X >S G C C rl H H QOOOQO&&&4JU1J ESZ<<'-)^'-)^^<<

Fig. 197. Seasonal distribution of adult Speyeria idalia in Ohio. Fig. 198. Geographical distribution of Speyeria atlantis in Ohio.

30 -

u 25 -

« 20 • i-H ■O3 < 15 -

. 10 -

5 -

0 0 — 0 0 'O o o — o o o o o — o o — o o o o o — I I I I I i I I I I I I I I I I I 1 I i I I I I

U M U U ^ U C e C H H H ClOQOClO&a&UiJiJ cT3rofl3D.a.CLroroco3=iD33DDZ3na)CL)(Dauu SSS<<-3*n<;<

Fig. 199. Seasonal distribution of adult Speyeria atlantis in Ohio. 479

Fig. 200. Geographical distribution of Clossiana selene myrina in Ohio.

cn cn in m 35 -

30

ou 25

« 20 ' »—I9 TJ < 15 '

. 10 -

5 - * «* * « * * * •K © o — ©©©©©^©©ooo^oo-'ooooo — -*cM(n-HtNnHNri-

i_, u u u u u >>>,>> c c c h h h doootiaaa&uiiu njcoraa.o.o.rtronJ999999999

Seasonal Time Period

Fig. 201. Seasonal distribution of adult Clossiana selene myrina in Ohio. 480

Fig. 202. Geographical distribution of Clossiana bellona in Ohio.

35 -

30 - 0) *2 25 - o a)u p- u 20 - •H3 T3 < 15 - U-lO . 10 - o z * *K* * * * 5 - * * * * * * * -K * * * ************** ■It * < « 4! < Jf ■* Jf .*■

o o ft o o o © o •“4 o o o O O o O O o o o o ft CO CM •H CM CO CM co CM co *7 CM co CM CO T CM CO *7 CM *7 co | 1 1 l JL •it J* J* •it J* 1 .i, •it JL •it 1 —*i, •it ft s< •it CM ft CM *-1CM CM CM -4 CM -< CM CM u u M J-l u so So e e s •H •H bObOb0 a a a 4J u u « « to o. a. a ro ro ca o 3 3 3 3 3 3 3 3 tv a> IV CJ u a z z Z < < < Z Z Z *-) •“5 *-> < < < CO co CO o o o Seasonal Time Period

Fig. 203. Seasonal distribution of adult Clossiana bellona in Ohio. 481

Fig. 204. Geographical distribution of Charidryas nycteis in Ohio.

35 -

30

oU 25 os - 20 *o < 15

. 10

•K * * * * *«K * * * *********** * * ** ..... ********* * ****** U. o OO © © © © © o © © © © O pH © © © © © oH CM cn CM cn CM cn 71CM cn CM cn *■* CM m CM cn CM cn 1 1 l 1 I T .i, ,1,pi, oi pi, H pi. pi,pi,pi,pi,pi pi, OH pi pi, pi pi pi pi CM —< CM oh CM CM CM »“* CM •"* CM —‘CM u U M u >■%5N c e B rH «—trH 60 60 60 a CL a

Fig. 205. Seasonal distribution of adult Charidryas nycteis in Ohio. 482

il f

Fig. 206. Geographical distribution of Charidryas harrisii in Ohio.

35 -

30 -

ou 25

20 »H« •O3 < 15

10 -

5 -

oo — o o o o o — ooooo-*oo-

u m u m u u ts >* >, c c c i—i ■—i h ooooooaaauuu rc{0cort3333S33Ssa>Q>aJOotJ SSS<<

Fig. 207. Seasonal distribution of adult Charidryas harrisii in Ohio. LO 00

Fig. 208. Geographical distribution of Phyciodes tharos in Ohio.

) ) ) )

) ) ) ) ) 12 1 12 10 2 22 8 2 6 (5) (5) ( ( ( ( (13) (30) (14) ( (18) (16) (16) (16) (14) (13) (19) ( (15) ( ( ( . No. of Adult Records *************** ** * ************* *********************************************************** ******** ************ ************** **************** **************** ****** ***** ** ************************* ******************************************** ********** 1-10 1-10 1-10 1-10 1-10 1-10 1-10 11-20 11-20 11-20 11-20 21-30 11-20 11-20 11-20 21-31 21-31 21-31 21-30 21-30 21-31 21-31 Sep Sep Sep Aug Oct Oct Oct Jul Jul Aug Aug Apr 11-20 Jul Apr May Jun Jun May May Jun Apr 1-10 Mar Mar Mar *-3 H* 3 rt> ft) ►1 H*

Fig. 209. Seasonal distribution of adult Phyciodes tharos in Ohio. 00 4>

Fig. 210. Geographical distribution of Euphydryas phaeton in Ohio. ) ) ) 12 1 1 (4) (16) (14) ( ( (3) ( No. of Adult Records **** ************ * ************** *** **************** * 1-10 1-10 1-10 1-10 11-20 11-20 11-20 11-20 11-20 11-20 11-20' 11-20 21-31 21-30 21-30 21-30 21-31 21-31 21-31 Sep Sep Sep Jul Jun Jul Jun 1-10 Jun MayMay 1-10 Jul Apr May AugAug 1-10 Aug Oct Oct Oct Mar 1-10 1 Apr Apr Mar Mar 21-31 P. H- a p 1—I •u p M H* O p o 3 cn p p CD

Fig. 211. Seasonal distribution of adult Euphydryas phaeton in Ohio. 485

Fig. 212. Geographical distribution of Polygonia interrogationis in Ohio.

i vO O u*i r>» n 35 -

30 •

ou 25

« 20 - f—I3 T3 < 15 ' U-4 o * * * * * . 10 . * * * o * * * z ******* * * * ****** * * 5 - ****** * * * « ****** * * . * * * * * *_*************.*** * *

o o o © © © © *-4 © © © © © _ O © © © © © o -M CN cn CN cn CN cn CN cn CN cn CN cn CN m CN cn | | •■p -p -II T T T ! l 1 l i 1 - 4 J* CN CN n CN ZJ CN z; CN Z! C'J CN CN u u U u u u J>s>% >N c c C rH r*4 GOGOGOa CLCLLi 4J Li n co CO CLCL Cl m ra co3 3 3 3 3 3 3 3 3 0)a>a) v o u x X X < ■ < < XXX *“3•n•n *n •n•n< < < C/3 co CO o © o Seasonal Time Period

Fig. 213. Seasonal distribution of adult Polygonia interrogationis in Ohio. 486

o I o o

Fig. 214. Geographical distribution of Polygonia comma in Ohio.

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30 -

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Fig. 215. Seasonal distribution of adult Polygonia comma in Ohio. 487

Fig. 216. Geographical distribution of Polygonia progne in Ohio.

35 -

30 -

25 -

20 —« - 3 < 15

10 -

5 -

© O — O O O O O — © O O O O - © O — © O O O O — -HNn-Nn-Nn-NrtMfMnMNn-Mn-NOI i i I i i I I i i i I I i i i i I i I i i i i

u u u u u u ►» >•» ss c e c h h h &o&o&oaaauuu SSSCCCESE'-J'-j'n^^^CCCwcowoOOajnjRjcucuacororaassssDsazjajaiaiuou Seasonal Time Period

Fig. 217. Seasonal distribution of adult Polygonia progne in Ohio. 488

Fig. 218. Geographical distribution of Nymphalis vau—album in Ohio.

35 -

30 -

25

« 20 ' •O3 < 1 5 .

10

5 -

* * * * *

OO—'OOOOO—'OOOOO^OO-^OOOOO^ i i i i I i I I i i i i i t i i i i i i i i i i - - fvl CvJ H N <—« C N — I C N — < C O 1-4 N — < CM h do&ocicaaauut) (QninO.aQ.(Q(0R333333 3D33Q}QJQ)OaO SSS<<

Fig. 219. Seasonal distribution of adult Nymphalis vau-album in Ohio. 489

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Fig. 220. Geographical distribution of Nymphalis antiopa in Ohio.

35

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w&m8m O O — O O O O O — © O O O O — © O — O O O O © — •HCMnMNn^NnrHNnHCMnHrgn-'CMn^tMnI I I ! I I I I I I I I ! I I I I 1 ...... CN cnI •m N —l CM -H CN N —* CN — CN c h h H bObODoaao-uuii snniioaaanni(ii33 s s < < < s s s •-> n "iT^,n<:<

Fig. 221. Seasonal distribution of adult Nymphalis antiopa in Ohio. 490

Fig. 222. Geographical distribution of Aglais milberti in Ohio.

35 -

30 -

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15 -

• 10 -

* * * * * * * * * * 5 - ** * ******* *** * ********* *** * ********* ****************

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Seasonal Time Period

Fig. 223. Seasonal distribution of adult Aglais milberti in Ohio. 491

Fig. 224. Geographical distribution of Vanessa virginiensis in Ohio.

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« 2 0 - •O3 < 15 -

10 - * * * * * * * -K 5 - * * * * * * «* * * * « * * * * * * * * * « * * * * * * * * * * * * * « * * * * * * * * *

o © © © © © © © © 0 © © © O ~4 © © © © © ~ -i C M c n CM CO CM COCM CO i-H CM CO -H CM CO CM CO C M c o 1 T 1 1 J, ,1 , J* pN .I4 J* J x J x JL .1 J 4 CM « - 4 CMCM —* CM — CM —• CMCMCM u u U u u ►» >% So c c C pH f H 0000 00O . a . a . u UU COCO 10 c l CL a CO COCO 3 3 3 3 a 3 3 3 3 a > a) 0 0 OO XXX < < < X 2: X •n • o ■ o n *0 n < < < CO CO CO 0 © O Seasonal Time Period

Fig. 225. Seasonal distribution of adult Vanessa virginiensis in Ohio. 492

Fig- 226. Geographical distribution of Vanessa cardui in Ohio.

35

30 -

cn u 25-

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< 15 -

10 -

5 - * * * ******* * * * * * * * *

-00 H N — n -ooooo-*ooooo^oo^ooooo-< N M n ^ N n « - c M r |,i>-icM(,n"

u u u u m u >s ^ >> c e c h h h so oo co a a a u u u sss:<<^*-3^3^-)<<(uuutJ Seasonal Time Period

Fig. 227. Seasonal distribution of adult Vanessa cardui in Ohio. 493

Fig. 228. Geographical distribution of Vanessa atalanta in Ohio.

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30 -

"2 25

- 20 1 rH *33 < 15 ■ U-lO . 10 ■ o z * * * * * * 5 ■ * * * * * * * * * * * *************- ************ * * * * *************** * * * ********** * * * * * o © © o © © o © © O © © © © © o o o o HH —*CM CMcn CMcn CMcn CMcn CM CM CM m T T cn cn -* cn ,1 Jh J. J* Jh J, Jh J, Jh J, Jh Jh J. Jh 1 1 1 I 1 1 1 Jh Jh I CM CM CM h-h CM -* CM —lCM CM CM u U U u u u >> >N Ps c s c w rH rH 00 00 00 a a a 4-1 4J 4J (0 TO TO CL Ou a TO TOTO 3 3 3 3 3 3 3 3 3 to TO TO TO U TO XXX < < < X X X *-> •n •n *“> •n < < < COCOCO O o O Seasonal Time Period

Fig. 229. Seasonal distribution of adult Vanessa atalanta in Ohio. 494

Fig. 230. Geographical distribution of Junonia coenia in Ohio.

35 -

30 -

25 oS

« 2 0 - s TJ < 15 - IMO . 10 - o z 5 - ******** ******************* © © — ©©©©©-<©©©©©«-<0©'-©0©©© — HN(n>-<(MnH(Nn-{\jnHNi,4-

u U U U U U SN >> X C C C H H H OOQOOOD.&auuu SSS<<

Fig. 231. Seasonal distribution of adult Junonia coenia in Ohio. 495

Fig. 232. Geographical distribution of Basilarchia arthemis astyanax in Ohio.

I /-s x—v O W N N

35 -

30

■2O 25 ■

u 2 0 - i-H T3a < 15 - * 10 - * * * * * * * * * * * * * * * * * * * * 5 - * * * * * * * * * * ***** * * * * * * * ***** * * * * * * * ***** * * * * * * * ***** *******

© O — O O O O © —OO O O © — OO — O O O O © — — c m cn — c m cn — c m cn — n n - c m cn — c m cn — c m cn — c m cn 1 1 1 Jj.l.lt'Ij'li.lt.L.l. 1 1 1 1 1 1 1 1 1 1 1 1 1 — CM — CM — CM — CM — C M — CM — CM — CM M U b i m m u > , > , > , e c c h h h ooooQoaaaiiuii ron3ma.acLci3CTjnj33D333333(i)a>aJuuo s : s s < < < 2 : s 2 : ' n ' - 5 ,n*-j^3,n< 3 < < : v 3 w c n o o o Seasonal Time Period Fig. 233. Seasonal distribution of adult Basilarchia arthemis astyanax in Ohio. Ov vO

Fig. 234. Geographical distribution of Basilarchia archippus in Ohio. ) ) ) ) ) ) ) 2 1 1 6 1 1 8 6 (5) (4) ( ( ( (3) ( (7) (13) (7) ( (4) ( ( ' (4) ' No. No. of Adult Records **** ****** *********** *** **** ** ******** ***** ******* ******* ****** ************* **** * * - 10 1-10 1-10 1-10 1-10 1-1 0 I 1-10 1-1 0 11- 2 0 11- 2 0 11- 2 0 11- 2 0 1 1 - 2 0 11- 2 0 11 - 2 0 21-30 21-31 21-31 1 1 - 2 0 21-30 21-31 21-31 21-30 21-31 Jul Jun Jun Jun Apr Apr Jul Jul Sep Sep Sep Mar Mar Apr Mar May May May Aug Aug Aug Oct Oct Oct o 3 to H* 3 to n> to to to H

Fig. 235. Seasonal distribution of adult Basilarchia archippus in Ohio. J— f

Fig. 236. Geographical distribution of Anaea andria in Ohio.

35 -

30 -

Ou 25

20 - «—44-1 •o3 < 15 -

10 -

5 -

-HCNm-HfNm—'CNm—'Csiro-HCMro—

Fig. 237. Seasonal distribution of adult Anaea andria in Ohio. 498

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Fig. 238. Geographical distribution of Asterocampa celtis in Ohio.

35 -

30 - W I 25

* 20 - *o3 < 15 -

10 - o z 5 - *«K * * •K .. * •K *

'OOOOO-^OOOOO'-'OO'-OO © — cm n*> I I i —■« CM —l CNj H CN —* CN «-< CM1 pi* JL •-* U U M <0 to nj ifa. a.bf clu >>to as>* s c g C C H H H OP M 6 0 ft. Q. a. aj XXX < C < X X X ^ 333333330)41 a> u C/3 © o o Seasonal Time Period

Fxg. 239. Seasonal distribution of adult Asterocampa celtis in Ohio. 499

Fig. 240. Geographical distribution of Asterocampa clyton in Ohio.

35 -

30 -

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u 20 - "os < 15

o 10 - z 5 -

-ft***-***** oo — o o o o o — o o o o o — oo — o o o o o — n ri n n h c s i p o — cm c n — n n n ^ cm co — cm en l I I I I I I I I I I I l I I I I I I I I I I I — CM — CM — CM — CM — CM — CM — CM — CM V4 u m u m u >n>s>nC c c h h ih Qoooooaaauuij ,->,,"> ^ ,n^ < < < : c n c O O T O © © Seasonal Time Period

Fig. 241. Seasonal distribution of adult Asterocampa clyton in Ohio. Ln O O

Fig. 242. Geographical distribution of Enodia anthedon in Ohio. ) ) ) ) ) 1 1 2 2 6 (A) ( (A) (15) (A) ( ( (5) (13) ( (A) ( »— »— l~* fsj OJ U> No. No. of Adult Records I I I I I I I f Ul Ul O Ul O Ui O Ul ■HI 11 1 1 1 11 11*** 11 1 1 ++++ 1 11 ■HI **** ** * +* ++*** + ■f | | | ********* | | | ■f ++**** -H-H- * -Northern* Ohio + - SouthernOhio - 1 0 - 2 0 - 1 0 -30 - 1 0 - 2 0 -31 - 2 0 - 2 0 -31 - 1 0 -31 -30 - 1 0 -31 -30 - 1 0 -31 - 1 0 - 2 0 - 1 0 1 1 1 1 - 2 0 1 1 - 2 0 1 1 - 2 0 2 2 2 2 2 2 2 2 Sep Sep Sep Jun Jun Oct Aug Jul Jul Jun Oct Aug Oct May Aug May Jul Mar Apr Apr Apr Mar Mar May

Fig. 243. Seasonal distribution of adult Enodia anthedon in Ohio 501

Fig. 244. Geographical distribution of Satyrodes eurydice + appalachia in Ohio.

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30 - •a u 25 -

« 20 - •O3 < 15

10 -

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u U U U U >» > S c c C H H H &OfiOQO&&Q.UUU n3«roOpa.CLnJa3(T333DD3D333(l)aJ(UUOU s s s

Fig. 246. Geographical distribution of Cyllopsis gemma in Ohio.

35 -

30 -

U 25 -

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10 - o z 5 - «* « •K * * He * *

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Fig. 247. Seasonal distribution of adult Cyllopsis gemma in Ohio. ) ) ) ) ) ) 1 1 1 22 11 1 (24) (3) ( (27) ( ( ( ( Ul) (13) ( (5) No. No. of Adult Records *** *************************** ********************** * *********** ************* *********** * * ***** * 1-10 1-10 1-10 1-10 1-10 1-10 1-10 1-10 1 1 - 2 0 11- 2 0 11- 2 0 11 - 2 0 11- 2 0 11- 2 0 1 1 - 2 0 1 1 - 2 0 21-31 21-30 21-30 21-30 21-31 21-31 21-31 21-31 ************************ Sep Apr Apr Sep Mar Mar Apr Jun Jun Jun Sep Oct Oct Mar May Aug Jul Jul Jul Aug Aug May Oct May n> (0 n H- o o. H IT

Fig. 249. Seasonal distribution of adult Megisto cymela in Ohio. ) ) ) ) 1 2 2 6 8 (i) (i) ( (27) ( ( (26) (14) (9) ( Ui Ui O Ln o to to ro u) u> No. No. of Adult Records O Ln I I t I I I » I 1/1 * ****** * *************************** ************************** ******** ********************** ************** ********* * 1-10 1-10 1-10 1-10 1-10 1-10 1 -10 1-1 0 1 1 - 2 0 1 1- 2 0 1 1- 2 0 1 1- 2 0 1 1- 2 0 11 - 2 0 1 1 - 2 0 1 1 - 2 0 21-31 21-31 21-30 21-31 21-31 21-31 Sep Apr Jun Jun 21-30 Jul Sep Jun Sep 21-30 Mar Mar Apr Apr Jul Jul Aug May May May Oct Oct Mar Aug Aug Oct

Fig. 251. Seasonal distribution of adult Cercyonis pegala in Ohio. 505

Fig. 252. Geographical distribution of Danaus plexippus in Ohio.

» /"S /“S 04 C*") H o /*N U*l N r** ON pH

35 -

30

25 -

20 - •O3 < 15 * •K* * 10 * * *K« *K« •K4C *

o O _4 o o o o O pH O 0 0 OO *H OO —4 O 0 0 0 0 pH 04 n 04 cn 04 cn 04 cn 04 cn pH 04 m 04 cn 04 cn T* '-4 Y i T 1 pH ,1* ,1, J* -h pL, *h p L pH m H pL pH pH pH 04 *>4 04 Ol —1 04 p H 04 04 04 ** 04

U u u M u >s c e a 1—i pH pH 00 GO GOa a a 4J u 4J to a O. a a a a to 3 3 3 3 3 3 3 3 3 04 a) 0) u 0 CJ £ ££ < < < £ £ £n n • n -) n •“3 < < < COCOCO 0 0 0 Seasonal Time Period

Fig. 253. Seasonal distribution of adult Danaus plexippus in Ohio. 506

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Fig. 254. Ohio's counties, with number of Rhopalocera species known from each of three classes of counties shown as a percentage of the total number of species known from the entire state (large dots = 50% or more known; small dots = 25% through 49% known; blank = less than 25% known) 507

NO CTHERN -D IS tR IC '

so; CT

Fig. 255. The state of Ohio showing tentative division into two districts based upon distribution of Rhopaloeera species within the state: the Northern and Southern Districts. 508

Table 8. Species of Rhopalocera, not known.from Qbio, but reported from neighboring states (P=Pennsylvania; W=West Virginia; K=Kentucky; I=Indiana; M=Michigan; OOntario).

Species State/Province

I W K I M 0

HESPERIIDAE: PYRGINAE 1. Urbanus proteus (Linnaeus, 1758) X X X 2. Thorybes confusls Bell, 1922. X X 3. Erynnis zarucco (Lucas, 1857). X X X HESPERIIDAE: HESPERIINAE 4. Lerema accius (Smith, 1797). X X X 5. Hesperia ottoe Edwards, 1866. X 6. Lerodea eufala (Edwards, 1869). X 7. Calpodes ethlius (Stoll, 1782). X PIERIDAE: ANTHOCHARIINAE 8. Euchloe olympia (Edwards, 1871). X X X X X X LYCAENIDAE: THECLINAE 9. Incisalia polios Cook & Watson, 1907. X X X X X RIODINIDAE: RIODININAE 10. Calephelis virginiensis (Guerin- Meneville, 1831). X NYMPHALIDAE: MELITAEINAE 11. Charidryas gorgone (HUbner, 1810). X X X X 12. Phyciodes batesii (Reakirt, 1865). X X X X X NYMPHALIDAE: NYMPHALINAE 13. Polygonia faunus (Edwards, 1862). XX'X XX NYMPHALIDAE: LIMENITIDINAE 14. Basilarchia arthemis (Drury, 1773). X X X ; X X SATYRIDAE: ELYMNIINAE 15. Enodia portlandia (Fabricius, 1781). X 16. Enodia creola (Skinner, 1897). X X 509

Table 9. The butterflies and skippers of Ohio with notation relative to the status of each species in the state (R=Resident; M=Migrant; S=Stray; E=Extirpated; N=Not in Hine, 1898a).

Species Status

R M _S E N

HESPERIIDAE: PYRGINAE 1. Epargyreus clarus X 2. Autochton cellus X X 3. Achalarus lyciades X X 4. Thorybes bathyllus X 5. Thorybes pylades X 6. Staphylus hayhurstii X X 7. Erynnis icelus X X 8. Erynnis brizo X 9. Erynnis juvenalis X 10. Erynnis horatius X X 11. Erynnis martialis X X 12. Erynnis lucilius X X 13. Erynnis baptisiae X X 14. Erynnis persius X 15. Pyrgus centaureae X X 16. Pyrgus communis X 17. Pholisora catullus X HESPERIIDAE: HESPERIINAE 18. Ancyloxypha numitor X 19. Thymelicus lineola X X 20. Nastra lherminier X X 21. Hylephila phyleus X 22. Hesperia leonardus X 23. Hesperia metea X 24. Hesperia attalus 25. Hesperia sassacus X 26. Polites coras X 27. Polites themistocles X 28. Polites origenes X 29. Polites mystic X 30. Polites vibex X X 31. Wallengrenia egeremet X 32. Pompeius verna X 33. Atalopedes campestris X 34. Atrytone logan X 35. Poanes massasoit X X 36. Poanes hobomok X 37. Poanes zabulon X 38. Poanes viator X X 510 Table 9(continued).

Species Status

R M E N

HESPERIIDAE: HESPERIINAE (continued) 39. Euphyes dion X 40. Euphyes dukesi X 41. Euphyes conspicua X 42. Euphyes bimacula X 43. Euphyes ruricola X 44. Atrytonopsis hianna X X 45. Amblyscirtes hegon X X 46. Amblyscirtes vialis X 47. Panoquina ocola X X PAPILIONIDAE: PAPILIONINAE 48. Battus philenor X 49. Eurytides marcellus X 50. Papilio polyxenes asterius X 51. Papilio cresphontes X 52. Papilio glaucus X 53. Papilio troilus X PIERIDAE: PIERINAE 54. Pontia protodice 55. Artogeia virginiensis X 56. Artogeia rapae X PIERIDAE: ANTHOCHARINAE 57. Falcapica midea PIERIDAE: GOLIADINAE 58. Colias philodice X 59. Colias eurytheme X 60. Zerene cesonia X 61. Phoebis sennae eubule X 62. Phoebis philea X 63. Eurema lisa X 64. Eurema nicippe X 65. Nathalis iole X X LYCAENIDAE: GERYDINAE 66. Feniseca tarquinius X LYCAENIDAE: LYCAENINAE 67. Lycaena phlaeas X 68. Hyllolycaena hyllus X 69. Epidemia epixanthe X X 70. Epidemia dorcas X X 71. Epidemia helloides X X 511 Table 9(continued).

Species Status

R M S. E N

LYCAENIDAE: THECLINAE 72. Atlides halesus X 73. Harkenclenus titus X 74. Satyrium acadica X X 75. Satyrium edwardsii X 76. Satyrium calanus falacer X 77. Satyrium caryaevorum X X 78. Satyrium liparops X 79. Calycopis cecrops X X 80. Mitoura gryneus X X 81. Incisalia augustus X X 82. Incisalia irus X X 83. Incisalia henrici X X 84. Incisalia niphon X X 85. Euristrymon Ontario X X 86. Parrhasius m-album X 87. Strymon melinus X 88. Erora laeta X X LYCAENIDAE: POLYOMMATINAE 89. Hemiargus isola X X 90. Everes comyntas X 91. Celastrina ladon X 92. Celastrina ebenina X X 93. Glaucopsyche lygdamus X 94. Lycaeides melissa X X RIODINIDAE 95. Calephelis borealis X 96. Calephelis muticum X X LIBYTHEIDAE 97. Libytheana bachmanii X NYMPHALIDAE: ARGYNNINAE 98. Euptoieta claudia X 99. Speyeria diana X 100. Speyeria cybele X 101. Speyeria aphrodite X 102. Speyeria idalia X 103. Speyeria atlantis X 104. Clossiana selene myrina X 105. Clossiana bellona X 512 Table 9(continued).

Species Status

R MN

NYMPHALIDAE: MELITAEINAE 106. Charidryas nycteis X 107. Charidryas harrisii X 108. Phyciodes tharos X 109. Euphydryas phaeton X NYMPHALIDAE: NYMPHALINAE 110. Polygonia interrogations X 111. Polygonia comma X 112 . Polygonia progne X 113. Nymphalis vau-album X 114. Nymphalis antiopa X 115. Aglais milberti X 116. Vanessa virginiensis X 117. Vanessa cardui X 118. Vanessa atalanta X 119. Junonia coenia X NYMPHALIDAE: LIMENITIDINAE 120. Basilarchia arthemis astyanax X 121. Basilarchia archippus X NYMPHALIDAE: CHARAXINAE 122. Anaea andria X NYMPHALIDAE: APATURINAE 123. Asterocampa celtis X 124. Asterocampa clyton X SATYRIDAE: ELYMNIINAE 125. Enodia anthedon X 126. Satyrodes eurydice X 127. Satyrodes appalachia X X SATYRIDAE: SATYRINAE 128. Cyllopsis gemma X X 129. Hermeuptychia sosybius X X 130. Neonympha mitchellii X X 131. Megisto cymela X 132. Cercyonis pegala X DANAIDAE 133. Danaus plexippus X 134. Danaus gilippus X X